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? Geographical distribution of Lamellibrachia species: L. anaximandri n. sp. (), L. barhami Webb, 1969 (), L. columna Southward, 1991 (), L. juni Miura & Kojima, 2006 (), L. luymesi Van der Land & N?rrevang, 1975 (), L. satsuma Miura, Tsukahara & Hashimoto, 1997 (), L. victori Ma??-Garz?n & Montero, 1985 () and unnamed species ().
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A new species of lamellibrachiid vestimentiferan, Lamellibrachia anaximandri n. sp., has been found in the Eastern Mediterranean, close to cold seeps of fluid carrying dissolved methane and sources of sulfide in superficial sediments. It occurs at about 1100 to 2100 m depth, on some of the mud volcanoes on the Anaximander Mountains, south of Turkey...
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... The genera Lamellibrachia Webb, 1969 andEscarpia Jones, 1985 are considered basal to the vestimentiferan radiation (Li et al. 2015) and although certain vestimentiferan species are endemic to a particular chemosynthetic setting (e.g. Riftia pachyptila Jones, 1981 and Ridgeia piscesae Jones, 1985 are only found at hydrothermal vents), Lamellibrachia and Escarpia exhibit greater flexibility, occurring within deep-sea cold seeps (Gardiner and Hourdez 2003;Andersen et al. 2004;Miglietta et al. 2010), at whale falls (Feldman et al. 1998), other organic falls (Hughes and Crawford 2008;Southward et al. 2011) and hydrothermal vents (Southward 1991;Plouviez et al. 2015). Such habitat flexibility is likely to have been key in enabling vestimentiferans to spread throughout the world's oceans. ...
... Invertebrate Systematics single specimen a very long, brown root was present. The longest tube was measured at 1309 mm that is still shorter than those of the species of the genus with the largest tubes, L. barhami Webb, 1969 and L. anaximandri with tubes of 800-1546 and 200-1530 mm respectively (Webb 1969;Southward 1991;Southward et al. 2011). The tubes from the VDVF appear to still be growing with no obvious reduction in growth in the larger specimens ( Supplementary Fig. S2), however the sample size is relatively small. ...
... Maximum width of tubes ranging from 4.2 to 11.5 mm at the apical ends, minimum width of tubes ~0.6 mm in posterior root region. Tubes generally straight at anterior 1/3-1/2 of length, Superscript numbers in the species column are citations of the following references: 1, Webb (1969); 2, Jones (1985); 3, Southward et al. (2011);4, Southward (1991); 5, Gardiner and Hourdez (2003); 6, Miura and Kojima (2006); 7, Kobayashi et al. (2015); 8, Miura et al. (1997); 9, Mañé-Garzón and Montero (1986); 10, van der Land and Nørrevang (1975);11, van der Land and Nørrevang (1977); 12, Southward, unpubl. data;13, McCowin and Rouse (2018); 14, Woodside et al. (1997); 15, Woodside et al. (1998) www.publish.csiro.au/is ...
The vestimentiferan tubeworm genera Lamellibrachia and Escarpia inhabit deep-sea chemosynthesis-based ecosystems, such as seeps, hydrothermal vents and organic falls, and have wide distributions across the Pacific, Atlantic and Indian Oceans. In 2010–2012 during initial explorations of hydrothermal vents of the Mid-Cayman Spreading Centre (MCSC), both genera were found to co-occur at the Von Damm Vent Field (VDVF), a site characterised by diffuse flow, therefore resembling a ‘hydrothermal seep’. Here, we erect two new vestimentiferan tubeworm species from the VDVF, Lamellibrachia judigobini sp. nov. and Escarpia tritentaculata sp. nov. Lamellibrachia judigobini sp. nov. differs genetically and morphologically from other Lamellibrachia species, and has a range that extends across the Gulf of Mexico, MCSC, off Trinidad and Tobago, and Barbados, and also across both vents and seeps and 964–3304-m water depth. Escarpia tritentaculata sp. nov. is distinguished from other Escarpia species primarily based on morphology and is known only from vents of the MCSC at 2300-m depth. This study highlights the incredible habitat flexibility of a single Lamellibrachia species and the genus Escarpia, and historic biogeographic connections to the eastern Pacific for L. judigobini sp. nov. and the eastern Atlantic for E. tritentaculata sp. nov. ZooBank: urn:lsid:zoobank.org:pub:D9F72BD4-FDE1-4C0A-B84B-A08D06F2A981
... First data on Mediterranean cold-seep communities were collected (Salas and Woodside 2002;Olu-Le Roy et al. 2004;Werne et al. 2004;Duperron et al. 2009;Brissac et al. 2011). Siboglinid polychaetes belonging to the clades Obturata (Lamellibrachia anaximandri, Southward et al. 2011) and Frenulata (Siboglinum sp.) have also been observed. Later in 2012, a new Spionid polychaete was discovered in deep-sea sediments of the Amsterdam mud volcano site by the E/V Nautilus in the same area (Blake and Ramey-Balci 2020). ...
Deep sea ecosystems are home to many unique habitats. Considering the Mediterranean Sea, extreme environments are clustered mainly in the eastern basin such as the Nile Deep Sea Fan, Finike (Anaximander) and Eratosthenes Seamounts, Olimpi mud volcano field, Calabrian Arc and Marmara deep fault systems. This study was conducted in the Finike Seamounts, to assess the ichthyofauna of the area with a depth of down to 2200 m. Samples of larval, juvenile and adult phases indicated the presence of 20 fish species belonging to 13 families in and around the Finike Seamounts area. The collected individuals consisted of 422 adults, 6 juveniles and 31 larvae. Most of the individuals belonged to Cyclothone braueri, all of them being adults.
... Thus, cold seep vestimentiferans-representatives of the genera Lamellibrachia, Seepiophila and Escarpiareach high numbers around hydrocarbon seeps in the Gulf of Mexico and on the slope off Louisiana [55][56][57][58][59][60][61][62][63]. This pattern is also valid for siboglinids of the subfamilies Frenulata and Monilifera [16,31,48,[64][65][66][67][68][69][70][71][72][73][74][75][76][77]. ...
Frenulate siboglinids are a characteristic component of communities living in various reducing environments, including sites with hydrocarbon seeps. High concentrations of hydrocarbons in the sediments of the Arctic basin seas, including the Barents Sea, suggest the presence of a rich siboglinid fauna there. This reflects the fact that microbiological oxidation of methane occurs under reducing conditions, generating high concentrations of hydrogen sulfide in the sediment. This hydrogen sulfide acts as an energy source for the sulfide-oxidizing symbionts of siboglinids. Here we report on the findings of the frenulate siboglinid species Nereilinum murmanicum made between 1993 and 2020 in the Barents Sea. These data significantly expand the range of this species and yield new information on its habitat distribution. The depth range of N. murmanicum was 75-375 m. The species was most abundant from 200 to 350 m and was associated with temperatures below 3 • C and salinities from 34.42 to 35.07. Most of the findings (43 locations or 74%) fall on areas highly promising for oil and gas production. Twenty-eight locations (48%) are associated with areas of known oil deposits, 22 locations (37%) with explored areas of gas hydrate deposits. N. murmanicum was also found near the largest gas fields in the Barents Sea, namely Shtokman, Ludlovskoye and Ledovoye.
... Yet, the hosts can also be 'promiscuous' and establish partnerships with more than one symbiont lineage. Two distinct symbiont phylotypes were found in tubeworms from the Lamellibrachia genus: amplicon and Sanger sequencing, as well as fluorescence in situ hybridization, revealed that Lamellibrachia barhami from the cold seeps in the Eastern Pacific, as well as Lamellibrachia anaximandri, which is endemic to the Mediterranean Sea (Southward et al., 2011), often host more than one symbiont phylotype, based on the dissimilarities in the 16S rRNA genes of the symbionts (Rubin-Blum et al., 2014;Zimmermann et al., 2014;Breusing et al., 2020a). On one hand, this is unusual, especially since the competition among symbionts may be unfavourable for the fitness of the host (Frank, 1996). ...
The symbioses between the vestimentiferan tubeworms and their chemosynthetic partners (Gammaproteobacteria, Chromatiales and Sedimenticolaceae) hallmark the success of these organisms in hydrothermal vent and hydrocarbon seep deep‐sea habitats. The fidelity of these associations varies, as both the hosts and the symbionts can be loose in partner choice. Some tubeworms may host distinct symbiont phylotypes, which often co‐occur in a single host individual. To better understand the genetic basis for the promiscuity of tubeworm symbioses, we assembled and investigated metagenome‐assembled genomes of two symbiont phylotypes (species, based on the average nucleotide identity < 95%) in Lamellibrachia anaximandri, a vestimentiferan endemic to the Mediterranean Sea, in individuals collected from Palinuro hydrothermal vents (Italy) and hydrocarbon seeps (Eratosthenes seamount and Palmahim disturbance). Using comparative genomics, we show that mainly mobilome and genes involved in defence mechanisms distinguish the symbiont genotypes. While many central metabolic functions are conserved in the tubeworm symbionts, nitrate respiration (Nar, Nap and Nas proteins) is modular, yet this modularity is not linked to phylotype, but rather to geographic location, potentially implying adaptation to the local environment. Our results hint that variation in a single moonlighting protein may be responsible for the fidelity of these symbioses.
... Yet, the hosts can also be 'promiscuous' and establish partnerships with more than one symbiont lineage. Two distinct symbiont phylotypes were found in tubeworms from the Lamellibrachia genus: amplicon and Sanger sequencing, as well as fluorescence in situ hybridization (FISH), revealed that L. barhami from the cold seeps in the Eastern Pacific, as well as L. anaximandri, which is endemic to the Mediterranean Sea (Southward et al., 2011), often host more than one symbiont phylotype, based on the dissimilarities in the 16S rRNA genes of the symbionts (Rubin-Blum et al., 2014;Zimmermann et al., 2014;Breusing, Franke, et al., 2020). On one hand, this is unusual, especially since the competition among symbionts may be unfavorable for the fitness of the host (Frank, 1996). ...
The symbioses between the vestimentiferan tubeworms and their chemosynthetic partners (Gammaproteobacteria, Chromatiales, Sedimenticolaceae) hallmark the success of these organisms in hydrothermal vent and hydrocarbon seep deep-sea habitats. The fidelity of these associations varies, as both the hosts and the symbionts can be loose in partner choice. Some tubeworms may host distinct symbiont phylotypes, which often co-occur in a single host individual. To better understand the genetic basis for the promiscuity of tubeworm symbioses, we curated and investigated metagenome-assembled genomes of two symbiont phylotypes (species, based on the average nucleotide identity <95%) in Lamellibrachia anaximandri, a vestimentiferan endemic to the Mediterranean Sea, in individuals collected from Palinuro hydrothermal vents (Italy) and hydrocarbon seeps (Eratosthenes seamount and Palmahim disturbance). Using comparative genomics, we show that mainly mobilome and defense mechanism-related features distinguish the symbiont genotypes. While many central metabolic functions are conserved in the tubeworm symbionts, nitrate respiration (Nar, Nap and Nas proteins) is modular, yet this modularity is not linked to speciation, but rather to local adaptation. Our results hint that variation in a single moonlighting protein may be responsible for the host-symbiont fidelity.
... Active seeping was evident at several mud volcanoes including Amsterdam, Thessaloniki, and Athina, where methane bubbles were observed escaping the seafloor through fractured carbonate crust (Raineault et al. 2013). These areas were host to abundant Siboglinidae tubeworm populations, Lamellibrachia anaximandri Southward, Andersen & Hourdez, 2011. Temperature of the actively seeping fluid was the same as the ambient seawater temperature, 14°C (Raineault et al. 2013). ...
A new spionid polychaete was discovered in deep-sea sediments in the eastern Mediterranean Sea during an expedition by the Ocean Exploration Trust. Specimens were collected by the E/V Nautilus in August 2012 off Turkey, at a depth of 2216 m on the Anaximander Seamount at the Amsterdam mud volcano site. Cores were taken from sediments covered with microbial mats. The new species belongs to the Pygospiopsis-Atherospio Group, which has unusual neuropodial hooks, modified neurosetae in some anterior setigers, and branchiae in middle body segments that are broad, flattened, and fused to the dorsal lamellae. The new species is assigned to a new genus and species, Aciculaspio anaximanderi n. gen., n. sp., and is unusual in having a reduced setiger 1 lacking notosetae; well-developed pre- and postsetal lamellae that encompass the neurosetae and notosetae; notopodial lamellae free from the branchiae in anterior setigers that become fused and flattened in middle and posterior segments; unidentate hooded hooks in both noto- and neuropodia; neuropodial spines in setigers 4–10; and a pygidium with three anal cirri. Aciculaspio anaximanderi n. gen., n. sp. is the first species in the Atherospio-Pygospiopsis Group collected from a deep-water cold seep habitat.
... There are a few possibilities of how frenulates' tubes can receive sulfide: through the opening at the posterior, directly through the tube wall or through the posterior end of the body (opistosome) protruding from the posterior end of the tube (Southward et al., 1986). Distribution locations of frenulates and moniliferans often coincide with regions of methane seepage (Flügel and Callsen-Cencic, 1992;Dando et al., 2008;Gebruk et al., 2003;Sahling et al., 2005;Niemann et al., 2006;Sauter et al., 2006;Lösekann et al., 2008;Dubilier et al., 2008;Sommer et al., 2009;Southward et al., 2011;Aquilina et al., 2013;Savvichev et al., 2018;Åström et al., 2018;Rimskaya-Korsakova et al., 2020;Sen et al., 2020;Baranov et al., 2020;Vedenin et al., 2020). ...
Siboglinids are a characteristic feature of reducing environments. More than 75% of all siboglinids were found in the Sea of Okhotsk at a depth of less than 400 m, while some species are known to inhabit the abyssal depth in other regions. Among the six species of siboglinids encountered in the Sea of Okhotsk, only two are widespread: Siboglinum caulleryi and Oligobrachia dogieli. A significant number of all findings belong to the area where, according to geological data, the methane concentration varies between 0.22 and 4.46*10 9 nmol/kg. There is a vast territory in the central part of the Sea of Okhotsk that is not inhabited by siboglinids and is characterized by minimum methane concentration values. Thus, data on the Sea of Okhotsk indicate that siboglinids are related to sites of methane seepage.
... It would be necessary to establish even more uncommon and fractional taxa to give the phylogenetic definition for the frenulate orders, in particular. Thus, it is currently inadvisable to renounce the multilevel taxonomic system hitherto existing within Pogonophora, and therefore the older classification and family names are used here (Ivanov, 1963;Southward et al., 2002Southward et al., , 2005Southward et al., , 2011; note comments by Bartolomaeus et al., 2005). ...
A new species of Pogonophora obtained from one station at a depth of 25 m from near the Dikson Island in the Kara Sea is described. Galathealinum karaense sp. nov. is one of the largest pogonophorans, the first known representative of the rare genus Galathealinum Kirkegaard, 1956 in the Eurasian part of the Arctic Ocean and a highly unusual finding for the desalted shallow of the Yenisey Gulf. Several characters occurring in the new species are rare or unique among the congeners: under-developed, hardly discernible frills on the tube segments, extremely thin felted fibres in the external layer of the tube, and very faintly separated papillae in the anterior part of the trunk. Morphological characters useful in distinguishing species within the genus Galathealinum are defined and summarised in a table. Diagnosis of the genus Galathealinum is emended and supplemented by new characters. Additionally, three taxonomic keys are provided to the species of Galathealinum and to the known species of the Arctic pogonophorans using either animals or their empty tubes only, with the brief zoogeographical information on each Arctic species.
... However, they are usually smaller than their Atlantic counterparts such as the small Idas modioliformis or the vesicomyid Isorropodon perplexum (Olu-Le Roy et al., 2004;von Cosel and Salas, 2001). The Siboglinidae tubeworm Lamellibrachia anaximandri (Southward et al., 2011) colonize several of these cold seep sites. Among symbiont-bearing taxa, only bivalves, including the same or closely related species to those of the Eastern Mediterranean, have been reported from the Sea of Marmara (SoM) (Ritt et al., 2010(Ritt et al., , 2012b. ...
... It is worth noting that although the long tubeworms seen in the Çinarcik basin near seepages which may be symbiont-bearing Siboglinidae tubeworms, also named pogonophorans or vestimentiferans, they have never been sampled in the SoM even if they colonize cold seeps in the Eastern Mediterranean from the South of Crete and Turkey (Olu-Le Roy et al., 2004) and from the Nile Deep Sea Fan (Ritt et al., 2011) with the species Lamellibrachia anaximandri (Southward et al., 2011). ...
The Sea of Marmara hosts part of the North Anatolian Fault as an active submarine strike-slip fault. This area has suffered numerous earthquakes and presents a major seismic risk. Although the Sea of Marmara has been studied for many years, the link between geological morphostructures, the nature of fluids and biological communities is still rarely described. During the Marsite cruise (November 2014), dives with Remotely Operated Vehicle (ROV) VICTOR 6000 focused on detailed seafloor explorations of four different areas: the Central and Western highs and the Tekirdağ and Çinarcik basins. Based on 130 h of in situ videos, high-resolution seafloor mapping of seeps was conducted, emphasizing their significant geological and biological diversity from one seeping site to another, from one basin/high to another. Gas bubbles (CH4, CO2), shimmering water (brine, marine and fresh water) and oil, escape from the seafloor into the water column with low to strong fluxes. Black patches of reduced sediments, authigenic carbonate crusts and chimneys compose the seep environments with various types of bacterial mats and chemosynthetic fauna. Several venting sites discovered during previous cruises are still active 7–12 years later. The seeps are mostly, but not only, focalized along the Main Marmara Fault (MMF), at the southern border of the Tekirdağ Basin and along the Western High. Fluid emission is also occurring at secondary faults and at their intersection with the MMF. Our study emphasizes the location of seeps at the foot of slopes, gully outlets and crossroads. Sedimentary features, such as mass wastings, stratigraphic discontinuities or canyons, also interact with fluid emissions. The observed fauna is dominated by Bathymodiolinae, Vesicomyidae, Lucinidae-like empty shells and tubiculous worms resembling Ampharetidae polychatea. Most of the symbiont-bearing taxa encountered and previously sampled in the Marmara Sea, are characterized by thiotrophic symbioses. Vesicomyids and Idas sp. mussels are present at gas seeps, but also in areas where crude oil escapes from the seafloor. Moreover, other taxa unusually encountered at cold seeps such as large-sized amphipod and vagile worms were observed in the Çinarcik Basin. Idas-like mussels were observed in the western part of the Sea of Marmara, in the Tekirdağ Basin and possibly on the Western High active seep sites. There, the sampled fluids had high methane content (reaching 65 μmol/l) but not as high as on the Central High (363 μmol/l) and Çinarcik Basin (228 μmol/l) where no mussels were observed in the video records. Bottom waters oxygen levels in the Sea of Marmara showed a west to east decreasing gradient (57–8.5 μmol/l). These oxygen conditions, which fall under the limit of Oxygen Minimum Zones (OMZ <20 μm/l) in the eastern part, may impact benthic fauna and explain the absence of symbiotrophic bivalves at cold seep sites of the Çinarcik Basin, whereas densely aggregated amphipods, likely more tolerant to oxygen stress were observed in the seepage area. Finally, no specific fauna was observed near the CO2-rich seep sites. First observations suggest that seep fauna composition in the Sea of Marmara does not seem to be strongly influenced by the nature (e.g., oil, gas bubbling, brines) of fluid venting through seeps. The seep environments are highly variable and characterized by distinctive geological morphostructures. They sustain typical Mediterranean cold seep fauna, but also unusual communities likely related to the interaction of seeps with hypoxic conditions.
... Its claws are consistently present and volumetrically dominant in the >2 mm fraction of the box-cores (Fig. 3), exhibiting a variety of sizes, which indicate the species goes through a full life cycle with several moulting stages at the locality. Several clusters of living chemosymbiotic tubeworms Lamellibrachia anaximandri Southward et al. (2011) have been observed by ROV at seeps upon carbonates at WA2a and WA3 (Fig. 4). ...
... Class Polychaeta Grube, 1850Order Sabellida Family Siboglinidae Caullery, 1914Genus Lamellibrachia Webb, 1969 Lamellibrachia anaximandri Southward et al. (2011). The species was first described from the Anaximander Mountains off the southwest shore off Turkey (Southward et al., 2011) and found to be closely related to Lamellibrachia luymesi from the Gulf of Mexico. ...
... Class Polychaeta Grube, 1850Order Sabellida Family Siboglinidae Caullery, 1914Genus Lamellibrachia Webb, 1969 Lamellibrachia anaximandri Southward et al. (2011). The species was first described from the Anaximander Mountains off the southwest shore off Turkey (Southward et al., 2011) and found to be closely related to Lamellibrachia luymesi from the Gulf of Mexico. Tube worms belonging to the genus Lamellibrachia were located in various mud volcanos and cold seeps in the Mediterranean sea typically 1100-2800 m deep (Olu-Le Roy et al., 2004;Duperron et al., 2009;Hil� ario et al., 2011;Carey et al., 2012;Thiel et al., 2012). ...
Chemosymbiotic micro- and macro-fauna related to cold-seep sites were recovered in the Palmahim Disturbance (PD), offshore Israel, during EU EUROFLEETS2 SEMSEEP Cruise, by box-coring and Remotely Operated Vehicle (ROV) dives. No live macrofauna was identified in the collected sediments, with the exception of the seep-related crustacean Calliax lobata (de Gaillande and Lagardère, 1966). Numerous Calliax claws testify the past colonization of these soft bottoms by several generations of this ghost shrimp. After sediment sieving on 1 mm, we identified gastropods belonging to the families Trochidae, Eucyclidae, unassigned Seguenzioidea (genus Anekes), Rissoidae, Elachisinidae, Raphitomidae, Mangeliidae, Architectonicidae, Orbitestellidae, and Acteonidae. The identified bivalves belong to the families Nuculidae, Yoldiidae, Mytilidae, Lucinidae, Thyasiridae, Semelidae, Kelliellidae, Vesicomyidae, Xylophagidae, and Cuspidariidae. A seep-related group of chemosymbiotic molluscs was detected, including: Taranis moerchii (Malm, 1861), Lurifax vitreus Warén and Bouchet, 2001, Idas ghisottii Warén and Carrozza, 1990, Lucinoma kazani Salas and Woodside, 2002, Thyasira biplicata (Philippi, 1836), Isorropodon perplexum Sturany, 1896, and the newly described Vesicomyid species Waisiuconcha corsellii n. sp., that represents also the first record of the genus Waisiuconcha in recent Mediterranean sediments.
The ROV dives recorded local patches of several m² of seafloor covered by dead shells of L. kazani, with a density of up to about 200 loose shells per square meter. The potential occurrence of seep-related foraminifera, among low-oxygen tolerant species, was explored by comparison with previously sampled adjacent localities, and lead to the identification of Chilostomella oolina, Globobulimina affinis and G. pseudospinescens as potential foraminiferal seep indicators in the southeastern Mediterranean Sea. The absence of live, seep-related fauna in surface sediments in the PD, where seepage has been confirmed, suggests intermittent activity and a pause or decline of the investigated seeps.
