Ecologically or Biologically Significant Areas (EBSAs)
Ridge South of the Azores
This area encompasses a part of the Ridge South of the Azores, with different features such as the axial valley, ridge crests and several hydrothermal vent fields, either active or inferred. The area is bordered at the north by the Menez Gwen hydrothermal vent field area and at the south by the Haynes fracture zone. At the east ridge crest, this area incorporates part of the Alberto Monaco Ridge and the seamount-like features associated with the ridge at its west. The area also includes three high-seas marine protected areas (part of the OSPAR Network of Marine Protected Areas) –Lucky Strike, Menez Gwen and Rainbow vent fields. This area has structures at depths ranging from the deepest 3460 m (inferred depth – south Oceanographer FZ), to the mid-range at 2320 m (measured depth – Rainbow), to the shallowest at Albert Monaco Ridge. The hydrothermal temperatures range between 10ᵒ C (Menez Hom and Saldanha) and 362ᵒ C (Rainbow). The uniqueness of each vent, due to the diversity of hydrothermal settings, the depth range and water mass distribution over oceanic ridge crests, significantly influences biomass production rates in the vicinity of these areas (LeBris et al., 2019).
The presence of a mid-ocean ridge with a truncated water column disrupts the general oceanographic circulation, potentially creating regions of high biomass that may arise from topographic influences on water circulation (St Laurent and Thurnherr 2007), upwelling nutrient-rich deep water as well as concentrating biomass over summits, creating mid-ocean regions of high productivity (Priede et al, 2013).
Knowledge of the Ridge South of the Azores area is based on the analysis of more than 500 scientific articles. Several of the structures are well known and have been the subject of a great number of geological and biological studies. The total number of hydrothermal vent species reported was estimated from scattered taxonomical literature and online species database. The species total derived, 342, is probably an underestimate.
A large number of species living in the area were discovered or described relatively recently (around 40 per cent of the species in the last 30 years), and a great many of them have their distribution restricted to the hydrothermal vents. The species studied in this area are in the larger majority dependent on the carbon produced at the hydrothermal vents, with the symbiotrophic species living closer to the fluid exits, and then a zonation with decreasing dependence, but always on the sphere of the increased production and chemical balance of the vents (Levin et al, 2016). There is no legislation or protection figure for the species, except for the sharks surrounding this area. Of the 342 species, shark species (Centrophorus squamosus, Centroscymnus coelolepis and Centrophorus granulosus) are the only three protected under the OSPAR Convention. Among the benthos, several species are indicators of Vulnerable Marine Ecosystems, namely the cold-water corals Lophelia pertusa and Madrepora oculata at the Menez Gwen, and several dispersed anthipatharian corals observed on the inactive structures at the outskirts of the vent fields and on the pillow lavas (Colaço pers. obs Tempera et al, 2013). At the Ridge crests and associated seamounts, which remain very poorly explored, the global habitat suitability models and distribution maps for the North Atlantic modelled the distribution of seven suborders of Octocorallia (Yesson et al. 2012) and five species of framework-forming scleractinian corals (Davies & Guinotte 2011). Both studies revealed the areas as containing important suitable habitats for these taxa.
The Ridge South of the Azores is located in the Atlantic Ocean, south of the Azores. This area has structures at depths ranging from 3460 m (inferred depth – south Oceanographer FZ), to the mid-range at 2320 m (measured depth – Rainbow), to the shallowest at Albert Monaco Ridge. The datum used is World Geodetic System 1984 (WGS84).
Cold-water coral reefs, gardens, sponge grounds and massive sponges support and enhance highly diverse benthic communities, comprising faunal biomass that is orders of magnitude above that of the surrounding seafloor (Henry and Roberts, 2007; Roberts et al., 2008; Lindsay et al., 2013). The composition of megafauna also significantly differs between sponge grounds and non-sponge grounds and between different sponge morphologies (Lindsay et al., 2013).
Since the discovery of the first hydrothermal vent field in 1977, an increasing number of fields have been found, all with different characteristics. However, there is still very little known about most of the 50,000 km of ocean ridges (Charlou et al., 2002; Kelley et al., 2002; Hein et al., 2013; https://vents-data.interridge.org/).
As tectonic plates separate in the ridge areas, magma migrates in the subsurface and erupts at the seafloor. Due to rock deformation, seawater penetrates to great depth before it is ejected to the seabed, enriched with dissolved material, especially hydrogen sulfide (H2S), various sulfide minerals, metals, carbon dioxide (CO2) and methane. Depending on ejection pressure and ambient temperature, crystallization of the sulfide minerals forms chimneys known as “black” or “white smokers” on the basis of the mineral colours precipitated (Ohmoto et al., 2006; Gold, 2013).
The species and communities present in this area belong not only to the deep-sea, but also to mid-water upper bathyal systems. The vents are characterized by extreme conditions with unique physical properties (temperature, pressure), chemical toxicity and absence of photosynthesis (Edmond et al., 1979; Mottl & Wheat, 1994; Kadko et al., 1995; Elderfield & Schultz, 1996; Minic et al., 2006). The venting dynamic of hydrothermal fluids back into the ocean is of major importance as it is associated with enhanced cooling of the ocean floor, formation of deep-sea mineral deposits, and unique ecosystems that exist around vent sites in extreme environmental conditions (Lister, 1980; Tufar et al., 1986; Haymon et al., 1989; Fouquet et al., 1995; Cathles et al., 1997; Boetius, 2005; Kelley et al., 2005; Marques et al., 2007).
The active vents are hosted by a range of different rock types, including basalt, peridotite and felsic rocks. The associated hydrothermal fluids exhibit substantial chemical variability, which is largely attributable to compositional differences among the underlying host rocks (Amend et al., 2011). Vent circulation accounts for approximately one third of the global geothermal heat flux to the oceans and strongly affects seawater chemical composition (Elderfield & Schultz, 1996). In this area there are many types of hydrothermal sites: high-temperature (250ᵒ–365ᵒ C) and low pH (<4) sites; metal-rich chimneys (i.e., Bubbylon, Lucky Strike, Menez Gwen and Rainbow); and diffuse and pervasive seepages, with apparently low temperatures (<30ᵒ C), and unknown pH (e.g., Menez Hom and Saldanha) (Barriga et al., 1998; Charlou et al., 2010). The Lucky Strike, Menez Gwen and Bubbylon are magamatic-hosted, while Menez Hom and Saldanha are ultramafic-hosted, and Rainbow presents both types (Charlou et al., 2000; Desbruyères et al., 2000; Fouquet et al, 2010).
In terms of biology, the vent fields also play a primordial role sustaining abundant populations of faunal species in the deep sea through autochthonous chemosynthetic primary production (e.g., Lutz & Kennish, 1993; Bemis et al., 2012). This process uses reduced compounds (typically hydrogen sulfide, methane or hydrogen) in vent fluids to fix inorganic carbon (Karl et al., 1980) that can be oxidized by microbes to release energy for the formation of organic carbon from carbon dioxide, carbon monoxide and/or methane (Van Dover et al., 2002). The chemosynthetic organisms may be present in the water column, at the seafloor as microbial mats, within sediments, fractures of crustal rocks or the sub-seabed, or/and in symbiosis with larger multi-cellular organisms (Dubilier et al., 2008).
Initial microbial colonization facilitates the development and maintenance of densely populated ecosystems in which both biomass and faunal abundances are larger than is typical at the deep seafloor (e.g., Lutz & Kennish, 1993; Smith et al., 2008).
Hydrothermal communities have been studied worldwide, leading to the description of more than 400 new species (Desbruyères et al., 2006), greatly enhancing our knowledge of marine biodiversity (Van Dover et al., 2002). However, knowledge about these animal communities and the biology and ecology of individual species in these waters remains limited.
This area covers a section of the Mid-Atlantic Ridge (MAR) south of the Azorean archipelago. Five major vent fields are described here:
1. Rainbow
The Rainbow vent field was discovered in 1997 (German et al., 1996b). It forms a high temperature (365°C) field of black smokers located on the western flank of the Rainbow massif along the Mid-Atlantic Ridge (MAR) (German et al.,1996a, 1999; Charlou et al., 1997; Fouquet et al., 1997, 1998). The hydrothermal vents are localized between 2270 - 2320 m depth in international waters where they comprise >30 groups of active small sulphide chimneys over an area of 15 km2. There are many inactive structures among a large number of rather short-lived active venting sites (German et al., 1996b; Charlou et al., 1997; Fouquet et al., 1997).
Around the site and through the nontransform discontinuity, a relative chronology of normal dip-slip extensional faulting, the conjugate transtensional faulting and Riedel shears are evident. The western border of the vent field is a 25 m high fault scarp where extensive stock work mineralization and replacement of ultramafic rocks by sulfides are observed (Marques et al., 2006; 2007).
Local hydrography and flow regimes dictate that the non-buoyant plume, which reaches neutral buoyancy at 2100 m depth, disperses following local topography to flow north-eastward, clockwise, along and around Rainbow ridge and into the adjacent rift valley (German and Parson, 1998; Thurnherr & Richards, 2001; Thurnherr et al., 2002).
At many places within the Rainbow vent field, unusual sediment lithification around the active field and near the top of the ridge, together with several places with dead mussels, may be related to diffuse low temperature of methane-rich fluid through the sediment. Similar processes were also proposed at low temperature Saldanha and Menez Hom sites, where large amounts of methane discharge through the sediment cover at the top of the ultramafic ridge (Schroeder et al., 2002; Ribeiro da Costa et al., 2008).
Together with the Lucky Strike segment and Menez Gwen vent fields, the Rainbow field forms a group of northern bathyal vents fields. The underlying basement and vent fluid compositions differ from those in basalt-hosted systems due in part to serpentinization of the host rocks at Rainbow. Key characteristics of the Rainbow fluids include high chlorinity (750 mM), low pH (2.8), high methane, and extremely high Fe concentrations (24 mM), resulting in a Fe/H2S molar ratio of 24 (Charlou et al., 1997; Douville et al., 2002). The high temperature vents occur along the shoulder of a W-facing hanging wall of the tilted ultramafic block and are associated with one of the largest hydrothermal plumes in terms of methane output (Charlou et al., 1996a), manganese (Aballea et al., 1998), sulfide (Radford-Knoery et al., 1998), helium and heat (Jean-Baptiste et al., 1998), and particles (German et al., 1998).
Since its discovery, Rainbow has been a frequent focus of scientific expeditions and is the only vent field on the Mid-Atlantic Ridge that has been visited by tourist operators. Scientific investigations have included long-term monitoring, manipulative experiments and geological sampling (McCaig et al., 2007; Baker et al., 2010; Crawford et al., 2010).
2. Lucky Strike
The Lucky Strike vent field was discovered in 1993. Since then, this field has been extensively studied, particularly during expeditions DIVA1 and FLORES, 1994; LUSTRE, 1996; MoMARETO and Graviluck, 2006; MoMAR, 2008; Bathyluck, 2009 and MoMARSAT 2010 and 2011. It is also the object of long-term monitoring (e.g., Ballu et al., 2009; Colaço et al., 2011), including a seafloor observatory (ESONET-EMSO European project) (Ruhl et al., 2011).
Lucky Strike is one of the largest hydrothermal areas known to date, with 21 active chimney sites distributed over an area of approximately 150,000 m2 at depth range of 1620-1730 m. Despite its proximity to the Azores hot spot, the Lucky Strike segment exhibits a morphological and tectonic architecture with many of the characteristics of a slow-spreading ridge. The Lucky Strike segment is characterized by a well-developed 13–20 km wide axial rift valley, whose depth increases from 1550 m at the segment center to 3700 m at the nodal basins near the segment ends.
Beyond the rift walls, the seafloor morphology is dominated by fault-controlled abyssal hills (Detrick et al., 1995). The centre of the segment is dominated by the 8 km wide, 15 km long, and 500 m high Lucky Strike volcano, one of the largest central volcanoes along the MAR axis. The crust is 7.5 km thick beneath the volcano and has thinned to less than 5.5 km at 20 km from the segment centre (Crawford et al., 2010; Seher et al., 2010).
The hydrothermal activity is located on the periphery of the lava lake. Submersible dive programmes documented the presence of high temperature black smoker chimneys, extensive areas of diffuse flow and sulfide deposits distributed around the lava lake margins (Fouquet et al., 1994; Langmuir et al., 1997; Ondreas et al., 2009). The presence of a lava lake at the summit also suggests recent magmatic activity and the potential for an active magma chamber directly beneath the edifice (Singh et al., 2006).
The physical/chemical qualities of the vent gases and waters are distinct from other MAR sites due to low sulphur/high methane contents. Vent fluid temperatures range from 330º C in black smokers, to 200-212ºC and even 20ᵒC in diffuse emissions (Von Damm et al., 1998; Charlou et al., 2000; Cooper et al., 2000). The larger active edifices exhibit small zones of high temperature discharge. Elsewhere in the chimneys, discharge is mostly diffuse, as leakage of transparent fluid, through the mussel-covered outer walls of the chimneys.
The chimneys show clear evidence of oxidation caused by seawater. In the more active chimneys oxidation is restricted to an outer layer of oxides a few millimeters thick, mainly of iron. Once fluid flow ceases, oxidation progresses inwards. Primary sulphides are replaced by secondary sulphides and subsequently by oxides. Chimneys become rapidly friable, fall and break into progressively less recognizable fragments. Nearly half of the area of the Lucky Strike field is covered with deeply oxidized chimney debris, with most of the remaining area composed of exposed “slabs” (Barriga & Santos, 2003).
3. Menez Gwen
Menez Gwen was discovered in 1991, during submersible dives on the ridge segment north of the Lucky Strike segment (Fouquet et al., 1994). This segment is characterized by the absence of a central rift and volcano. Circular in shape, it has a diameter of 17 km and height of 700 m, while at its summit there is an axial graben, 6 km long, 2 km wide and 300 m deep. At the graben’s northern end there is a new volcano of 600 m diameter and 120 m height, composed entirely of fresh pillow lavas with no sediment cover.
Menez Gwen is located near the top of this new volcano at the bottom of the graben at 840-870 m depth. Its hydrothermal fluids are characterized by temperatures ranging between 265ᵒC and 281ᵒC, and these temperatures mark its characteristic physiochemical diversity and presence of anhydrite and barite. The vent is in a basaltic environment, and methane is produced by outgassing of carbon from the mantle and is related to the carbon-enriched character of basalt (Charlou et al., 1997). In addition, the low pH and low Fe and Si concentrations are consistent with the short duration of fluid-rock interaction linked to a shallow circulation system (Douville et al., 1999).
This shallow system can be affected by explosive volcanic activity (Fouquet et al., 1999) on an area of several square kilometres, as disclosed by the distribution of volcanic ejecta on the bottom (ash, sand and lapilli). According to Fouquet et al. (1994), Menez Gwen is, geologically speaking, very young (probably a few decades old); its chimneys are very small, growing directly on fresh pillow lava. Its relatively young age provides an excellent opportunity to monitor the early stages of hydrothermal vent activity and thus yield new knowledge on the development of vents and their associated animal communities (Marcon et al., 2013; Sarrazin et al., 2014; Konn et al., 2015). The vent fluids are the least toxic of the sites along the MAR and make it possible for non-endemic deep-sea species to live here (Desbruyères et al., 1997; Tunnicliffe et al., 1998; Colaço et al., 1998; 2002).
4. Saldanha
The Saldanha hydrothermal field was discovered in 1998 during the Saldanha Cruise (Barriga et al., 1998). It is located between the FAMOUS and AMAR second-order segments and consists of a faulted peridotite massif detached from its segment flanks, almost parallel to the ridge segment. It is composed mainly of ultramafic and gabbroic rocks and a strong methane anomaly within the overlying water column (Charlou et al. 1997; Dias & Barriga, 2006).
Although no vent chimneys are present, hydrothermal activity is expressed as discharge of clear fluid from several small orifices through sediment over an area of at least 50 m2, and micro chimneys with silica and sulfides have been observed (Dias, 2001; Dias et al., 2002).
The discovery of this diffuse venting confirmed the presence of hydrothermal activity related to serpentinization processes, which had been inferred from the detection of geochemical (intense CH4) anomalies in the water column (Charlou et al., 1997; Bougault et al., 1998). During the serpentinization of the ultramafic rock, overlying rocks were pushed upward, generating the observed mélange of talc-rich rocks (steatite) and spilite (Costa, 2001; Costa et al., 2002). Diving operations (Fouquet et al., 1997, 2000; Barriga et al., 1998) revealed intensely altered and locally silicified ultramafic and basaltic rocks consistent with low magma budgets, relatively thin crust and irregular faulting patterns (Gràcia et al. 2000) at the top of the massif. Discrete low-temperature diffuse discharge (<6°C) from the sediment was observed near the top of the structure (Biscoito et al., 2006).
Studies to date have discovered that the site is hosted in a mélange of folded lithified sediment, relatively fresh to deeply altered basalt, variably deformed ultramafic rocks and some gabbroic rock, in large part covered by sedimentary ooze. The ensemble is interpreted as resulting from active serpentinite protrusion. Sulphide precipitation is taking place within the top of the rock pile, under a blanket of sediment (Dias, 2001; Barriga 2003).
5. Menez Hom
Like the Saldanha, the Menez Hom ultramafic dome is situated at an inside corner position relative to the non-transform offset at the south of the Lucky Strike segment. Diving operations have revealed the general outcrop of ultramafic rocks at the top of the dome. No active vents were seen. However, one small carbonate chimney was sampled and anomalous rapid lithification of the sediment covers was observed at the northern side of the dome, near the limit between the ultramafic rocks and the basalt coverage. This may indicate a preferential discharge of diffuse low-temperature CH4-rich fluids at the contact between the ultramafic and the basalt cover (Fouquet et al., 2010).
There are two attributes in common to the deep-sea hydrothermal systems in the area described: their insularity and their gradient regimes of fluid flow and chemistry suggested a priori that measures of community structure and similarity at vents would be especially sensitive to the degree of proximity between sites being compared, to the age of the sites and to within-site heterogeneity (Mullineaux & France, 1995, Marsh et al. 2001, Van Dover et al. 2001). These different vent characteristics “create” distinct habitats dominated by different chemosynthetic bacterial mats, and endemic and non-endemic species of tubeworms, mussels, gastropods, clams, shrimp and crabs. In turn these habitats support further associated invertebrate and vertebrate species.
The majority of organisms found in this area developed different strategies to adapt to its extreme environments, e.g., biological stabilization of metal (e.g., iron, copper) from hydrothermal vents under dissolved or colloidal organic (Wu et al., 2011; Hawkes et al., 2013). In the absence of photosynthesis, the food chain is based on primary production of energy and organic molecules by chemolithoautotrophic bacteria. Hydrothermal vent plumes sustain rich microbial communities with potential connections to zooplankton communities and important deep ocean biogeochemical fluxes (Dick et al., 2013). These microbial communities extract chemical energy starting from the oxidation of reduced mineral compounds (Minic et al., 2006; Boetius & Wenzhöfer, 2013). Studies in community hydrothermal evolution, initial colonization, growth, development and demise, show that colonization at vents is rapid (Lutz et al. 1994, Tunnicliffe et al. 1997, Shank et al. 1998).
The area has an uneven number of studies for its different structures. Nevertheless, there are many studies focused on the communities and species of these structures. To date a total of 342 species have been identified within this area (see “introduction”).
Since the discovery of this area, most studies have been qualitative and often focus on specific taxonomic groups, such as amphipods (e.g Myers and Cunha, 2004; Bellan-Santini et al., 2007), cirripeds (e.g., Young, 1998; 2001), Copepoda (e.g., Ivaneko and Defaye., 2004; Komai & Segonzac, 2005; Komai & Chan, 2010), Cumeacea (e.g., LeBris et al., 2000; Corbera et al., 2008), echinoderms (e.g., Stöhr & Segonzac, 2005), elasmobranchii (e.g., Biscoito et al., 2002; Biscoito, 2006; Linz, 2006), mussels (e.g., Colaço et al., 2006a; Duperron et al., 2006; 2013), polychaeta (e.g., Desbruyères & Hourdez, 2000; Hourdez & Desbruyères, 2003), shrimps (e.g., Shank & Martin, 2003; Nye et al., 2012) and tanaidacean (Larsen et al., 2006). Most research cruises that have visited the area were focused in the deep-sea hydrothermal vent fields south of the Azores (i.e., Menez Gwen, Lucky strike, Rainbow and Saldanha), that were part of the MoMAR concept (“Monitoring the Mid-Atlantic Ridge”). The OSPAR MPAs (Lucky Strike, Menez Gwen and Rainbow) have a higher number of scientific articles and reports, and consequently are thus far the best studied. The vent fields inside the NAFO/NEAFAC areas were also subject to ICES report of the WGDEC (Working Group on Deep-Water Ecology) (Auster et al., 2013). Studies have also focused on the distribution of species (e.g., Cuvelier et al, 2011a; Sarrazin et al, 2015 ), temporal evolution (Cuvelier et al, 2011b), foodwebs (Colaço et al., 2002; 2007, De Busserolles et al., 2009, Portail et al., 2018), physiology (e.g., Bettencourt et al., 2010; Martins et al., 2008; Husson et al., 2016), reproduction (e.g., Colaço et al., 2006a; Dixon et al., 2006), ecotoxicological aspects (e.g., Colaço et al., 2006b; Martins et al., 2009, 2011; Company et al., 2008), behaviour (Matabos et al., 2015) and microbiology (Crepeau et al., 2011 and references therein).
The dissolved constituents of the venting fluids of the hydrothermal vents play an important role in the geochemical mass balance of the oceans (Levin et al, 2016 and references therein). The unusual nature of the marine communities that occur around hydrothermal vents makes them particularly important areas in terms of the biodiversity of the deep sea as well as being a focus for deep-sea research. This type of ecosystem is sensitive because of its high percentage of endemic species and the unique nature of many of the species found there (e.g., Vrijenhoek, 2010; Ramirez-Llodra et al., 2011; VanDover et al., 2018).
There is a biological balance in the vents. Well documented examples of biological interactions are predation and competition, based, for instance, on trophic (e.g., access to hydrogen sulfide or other resources) and topographic (optimal positioning on the structure or limitation on available space) grounds (Hessler et al., 1985; Fustec et al., 1987; Comtet and Desbruyères, 1998; Colaço et al., 2002; 2007, Riou et al., 2008, 2010a,b, deBusserolles et al., 2009, Portail et al., 2018, Sarrazin et al., 2015, Cuvelier et al., 2011).
All five (Menez Gwen, Lucky Strike, Menez Hom, Saldanha and Rainbow) hydrothermal vent fields are included in the Azores Marine Park, created in 2007 and expanded in 2016. Lucky Strike, Menez Gwen and Rainbow are included in the OSPAR Network of Marine Protected Areas. Lucky Strike and Menez Gwen have been a part of the Natura 2000 network since 2009. All fields are classified under the reef habitat type of the EU Habitats Directive. Lucky Strike and Menez Gwen (MPAs) are also recognized by WWF as a Gift to the Earth (GttE).
The areas comprising the Azores Marine Park, and all the regional protected areas beyond the territorial sea, are classified under IUCN criteria. Lucky Strike (288 km2) and Menez Gwen (95 km2) have zoning plans ranging from “full protection” (Category 1) to “sustainable exploitation” (Category IV and VI), while Rainbow, a smaller vent field, is classified under IUCN Category IV. Lucky Strike has also been selected as a target field for the installation of the long-term seafloor former MoMAR observatory, and now EMSO-Azores (Santos et al., 2002; Person et al., 2008; Colaço et al., 2012).
The Contracting Parties to OSPAR Convention committed themselves to establish an ecologically coherent network of MPAs in the OSPAR Maritime Area by 2010 (the OSPAR Network of Marine Protected Areas). The regional delivery mechanism is based on Annex V to the OSPAR Convention. The first national MPA designated under the high seas is the Rainbow vent field, located in the High Seas sector of the OSPAR Maritime Area (Santos and Colaço, 2010; Ribeiro, 2010).
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The Ridge South of the Azores has:
1-Deep-sea vents, which represent one of the most physically and chemically unusual biomes on Earth (Takai & Nakamura, 2011).
2-The hydrothermal vents of the North MAR may represent a unique biogeographic region of invertebrate species (Van Dover, 2010). They have relatively high proportions of endemic species (Tunnicliffe & Fowler, 1996; VanDover et al, 2018) that cannot live anywhere else, dominated by the blind shrimp Rimicaris exoculata and the mussel Bathymodiolus azoricus (Desbruyères et al., 2001).
3-The uniqueness of each vent, due to the diversity of hydrothermal settings, the depth range and water mass distribution over oceanic ridge crests, significantly influences biodiversity and species composition (VanDover et al., 2018). The hydrothermal biota are characterized also by a high level of endemism, with common specific lineages at the family, genus and even species level, as well as the prevalence of symbioses between invertebrates and bacteria (Dubilier et al., 2008; Kiel, 2010).
4-In addition to the endemic vent fauna, there are also several topographical elevations associated with the flanks of the MAR, with reported endemic cold-water corals in the region of the Azores (Braga-Henriques et al., 2013; de Matos et al 2014; Sampaio et al., 2019), including a species of black coral (Heteropathes opreski) that is known exclusively from the North MAR south of Azores at depths of 1,955–2,738 m at the Oceanographer fracture zone, (de Matos et al., 2014, Molodtsova, 2016).
5- The vent communities are unique, and the species living in these areas have specialized adaptations. Such features allow the organisms to exploit vent habitats, endowed with major reorganization of internal tissues and physiologies to house microbial symbionts, biochemical adaptations to cope with sulphide poisoning, behavioral and molecular responses to high temperature, presence of metal-binding proteins and development of specialized sensory organs to locate hot chimneys (Tunnicliffe et al., 1998).
1-Most of the organisms colonizing these habitats are invertebrates and have larval stages that are subject to dispersal in an open system, although mechanisms of larval retention are developed to account for the large settlement events observed (Mullineaux & France 1995, Marsh et al. 2001, Van Dover et al. 2001).
2-The dominant symbiotrophic species span late winter so their planktotrophic larvae can eventually profit from the increased productivity in the marine environment each spring (Colaço et al, 2006a; Dixon et al. 2006).
3-Connectivity among vent fields is poorly known, with just two or three studies showing that there are genetic exchanges, however without knowledge of the time it takes for the exchanges to take place (Teixeira et al., 2012, Breusing et al., 2016)
4-Blue shark nursery at the Central North Atlantic, roughly delimited by the Azores archipelago in the North, the Atlantis–Great Meteor seamount complex in the South and the Mid-Atlantic Ridge in the South-West (Vandeperre et al., 2014)
5- Several species of seabirds use these areas as foraging grounds during their breeding (e.g., Calonectris borealis, Puffinus lherminieri baroli, Pterodroma deserta, Pterodroma madeira, Bulweria bulwerii) or non-breeding season (Calonectris diomedea, Puffinus puffinus, Rissa tridactyla, Catharacta maccormicki, Catharacta skua and Stercorarius longicaudus) (BirdLife International 2019).
6-Corals and sponges of topographic highs (e.g., Albert de Monaco Ridge) also serve as important spawning, nursery, breeding and feeding areas for a multitude of fishes and invertebrates (Pham et al. 2015, Pereira et al. 2017, Porteiro et al., 2013 Ashford et al., 2019).
1- The area contains one threatened and/or declining habitat, contained in the OSPAR List (OSPAR publication 2008/358): Oceanic ridges with hydrothermal vents/fields (OSPAR, 2014).
2-Cold-water coral species of the order Antipatharia (e.g., black corals Leiopathes sp, Bathypathes sp), Scleractinia (e.g., reef-building corals Lophelia pertusa, Madrepora oculata) and family Stylasteridae (e.g., Errina spp, Stylaster spp), are listed under CITES Appendix II (https://www.cites.org/eng/app/appendices.php). Many of these habitats, including the cold-water coral gardens and sponge aggregations, sea-pen and burrowing megafauna communities, as well as oceanic ridges with hydrothermal vents and seamounts are all listed on the OSPAR List of Threatened and/or Declining Species and Habitats (OSPAR 2009; 2010a.,b,c,d).
3- The occurrence of three species under OSPAR legal protection was recorded in the area: Centrophorus granulosus, Centrophorus squamosus and Centroscymnus coelolepis. These three shark species are included in the OSPAR list of Threatened and/or Declining Species and Habitats (BDC/MASH, 2007).
4-Four globally threatened seabird species occur in the area - Rissa tridactyla (VU), Pterodroma deserta (VU), Pterodroma madeira (EN) and the OSPAR listed Puffinus lherminieri baroli (BirdLife International 2019).
1-The Mid-Atlantic Ridge is a slow-spreading ridge, and hydrothermal vents are estimated to be up to thousands of years of age, although possibly not active continually. However, some of the individual vents are only short-lived naturally. In this case, non-consolidated structures that cannot support eukaryote life are formed easily. Therefore, the vent fields in the area described are relatively stationary in position, but dynamic regarding the individual smokers and long-term activity (Hannington et al., 1995).
2- Time series studies over 14 years show that these communities are stable over time, and that big changes might disrupt the stability (Copley et al., 2007; Cuvelier et al., 2011). The occurrence of three species under OSPAR legal protection was registered in the area: Centrophorus granulosus, Centrophorus squamosus and Centroscymnus coelolepis. These species have particular features attending to biological factors such as longevity, low fecundity, and slow growth rates characteristic to these shark species (e.g., Clark, 2001; Morato et al., 2008).
3-Cold-water corals have life history traits such as slow growth, high longevity, low reproductive potential (Clark et al 2016; 2019). Octocorals and black corals, which dominate benthic assemblages in the MAR region, have growth rates of less than 1 cm a year and age spans of hundreds of years (e.g., bamboo coral Keratoisis sp. : Watling et al., 2011) to thousands of years (black coral Leiopathes sp. Roark et al., 2009, Carreiro-Silva et al., 2013).
5-Although age estimates for sponge species are scarce, they suggest multi-centennial age spans, e.g., 220 and 440 years (Leys and Lauzon, 1998; Fallon et al., 2010), whereas some sponge reefs are estimated to be up to 9,000 years old (e.g., Krautter et al., 2001).
1- The presence of a mid-ocean ridge with a truncated water column disrupts the general oceanographic circulation, potentially creating regions of high biomass that may arise from topographic influences on water circulation (St Laurent and Thurnherr 2007) upwelling nutrient-rich deep water as well as concentrating biomass over summits creating mid ocean regions of high productivity (Priede et al., 2013).
2-In the vent biotopes, there is local primary production of energy and organic molecules by chemolithoautotrophic bacteria (Synnes, 2007; Le Bris et al., 2016).
3-. Hydrothermal vents are involved in the biogeochemical cycling and elemental transformation of carbon, sulfur, and nitrogen (Petersen et al., 2011; Lilley et al., 1995; Sievert and Vetrini, 2012) and contribute to the huge diversity of deep-sea organisms and habitats.
4- This ecosystem enhances trophic and structural complexity relative to the surrounding deep sea and provides the setting for complex trophic interactions (e.g., Colaço et al., 2007; Portail et al., 2017). The chemosynthetic productivity from vents is exchanged with the nearby deep-sea environments, providing labile organic resources to benthic and pelagic ecosystems that otherwise have limited availability of food (Levin et al., 2016).
5-Organic matter produced at vent complexes, with metals such as iron or copper released from vents with organic ligands (Bennett et al., 2008; Hoffman et al., 2018), is spread with the buoyant plume, contributing to the global ocean micronutrient budgets (Tagliabue et al., 2010; Resing et al., 2015).
6- The hydrothermal fluids are rich in iron (Charlou et al,2010 Le Bris et al, 2019). Recent assessments of these iron sources indicate their significance for deep-water budgets at oceanic scales and underscore the possibility for fertilizing surface waters through vertical mixing in particular regional settings (Tagliabue et al., 2010) and supporting long-range organic carbon transport to abyssal oceanic areas (German et al., 2015).
7-Both cold-water coral communities and sponge grounds are important for global biogeochemical cycles and the ocean’s benthic pelagic coupling loop, being responsible for nearly 30 per cent of the coupling between organic matter produced at the ocean surface and the seafloor (Cathalot et al., 2015). They represent hotspots of ecosystem functioning, processing substantial amounts of organic matter (White et al., 2012; Cathalot et al., 2015), and release nutrients back into the surrounding water (Van Oevelen et al., 2009; Cathalot et al., 2015) that become available to associated fauna, thereby potentially increasing overall biodiversity and biological productivity of these habitats.
1-Fauna associated with vents are characterized by a high degree of specialization and relatively high productivity and species abundances compared with the surrounding deep sea. However, slow-spreading ridges, such as the MAR, that are present in the area, present the highest species diversity found at vent communities (Dubilier et al., 2008; Bernardino et al., 2012).
2-The adjacent bathyal and abyssal areas are characterized by low biomass and high diversity. During recent years, new exploration led to new discoveries. Around 60 different habitats are identified by the European Nature Information System (EUNIS) (Tempera et al., 2013).
3-Cold-water coral reefs, gardens and sponge grounds support and enhance a highly diverse community, comprising faunal biomass that is orders of magnitude above that of the surrounding seafloor (Henry and Roberts, 2007; Roberts et al., 2008; Lindsay et al, 2013). The composition of megafauna significantly differed between sponge grounds and non-sponge grounds and between different sponge morphologies (Lindsay et al., 2013).
Overall the naturalness of the described area is classified as high, as it is located in a relatively remote area.
- Mapes and Figures of Ridge South of the Azores.docx (/api/v2013/documents/E0A2A005-93E6-CC17-49F5-276B146A1F0E/attachments/613211/Mapes%20and%20Figures%20of%20Ridge%20South%20of%20the%20Azores.docx)
