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The shrimp Rimicaris exoculata is the most conspicuous component of vent communities developing around hydrothermal fluid emissions below 2000 m on the northern Mid-Atlantic Ridge (nMAR). Its high genetic connectivity suggests a remarkable ability to produce dispersing larval stages. However, so far brooding females have been rarely observed and reproduction remained enigmatic. Spatially complex population structures related to the heterogeneity of local habitat conditions are described for many vent species, this information being fundamental to gain a better understanding of their life history. Here our aim was to assess such complexity along with reproductive development in R. exoculata populations within two vent fields, TAG and Snake Pit (3620m and 3470m depth respectively). We compared samples collected in January–February 2014 among visually distinct assemblages with different degrees of exposure to vent fluids. Dense aggregations located near active venting included mostly females and immature individuals, while inactive peripheries harbored low density assemblages of large males. Small juveniles gathering around low temperature diffusions belonged to another species, Rimicaris chacei. One third of the sexually mature females were ovigerous at the two vent fields during our sampling period, with lower fecundities and egg sizes in the TAG population. Overall, the observed shrimp distribution patterns were consistent across both vent fields, although a high degree of heterogeneity in population structure was observed locally within dense aggregations, probably reflecting micro-scale variations in environmental conditions. Our results thus highlight spatially complex population structures where R. exoculata females brood eggs within dense aggregations exposed to vent fluids, while peripheral inactive areas may be important mating grounds for adults. In addition, we suggest temporal variability in reproductive activity, increasing in the winter season, which que

The shrimp Rimicaris exoculata is the most conspicuous component of vent communities developing around hydrothermal fluid emissions below 2000 m on the northern Mid-Atlantic Ridge (nMAR). Its high genetic connectivity suggests a remarkable ability to produce dispersing larval stages. However, so far brooding females have been rarely observed and reproduction remained enigmatic. Spatially complex population structures related to the heterogeneity of local habitat conditions are described for many vent species, this information being fundamental to gain a better understanding of their life history. Here our aim was to assess such complexity along with reproductive development in R. exoculata populations within two vent fields, TAG and Snake Pit (3620m and 3470m depth respectively). We compared samples collected in January–February 2014 among visually distinct assemblages with different degrees of exposure to vent fluids. Dense aggregations located near active venting included mostly females and immature individuals, while inactive peripheries harbored low density assemblages of large males. Small juveniles gathering around low temperature diffusions belonged to another species, Rimicaris chacei. One third of the sexually mature females were ovigerous at the two vent fields during our sampling period, with lower fecundities and egg sizes in the TAG population. Overall, the observed shrimp distribution patterns were consistent across both vent fields, although a high degree of heterogeneity in population structure was observed locally within dense aggregations, probably reflecting micro-scale variations in environmental conditions. Our results thus highlight spatially complex population structures where R. exoculata females brood eggs within dense aggregations exposed to vent fluids, while peripheral inactive areas may be important mating grounds for adults. In addition, we suggest temporal variability in reproductive activity, increasing in the winter season, which que

Whilst the fauna inhabiting hydrothermal vent structures in the Atlantic Ocean is reasonably well known, less is understood about the spatial distributions of the fauna in relation to abiotic and biotic factors. In this study, a major active hydrothermal edifice (Eiffel Tower, at 1690 m depth) on the Lucky Strike vent field (Mid-Atlantic Ridge (MAR)) was investigated. Video transects were carried out by ROV Victor 6000 and complete image coverage was acquired. Four distinct assemblages, ranging from dense larger-sized Bathymodiolus mussel beds to smaller-sized mussel clumps and alvinocaridid shrimps, and two types of substrata were defined based on high definition photographs and video imagery. To evaluate spatial variation, faunal distribution was mapped in three dimensions. A high degree of patchiness characterizes this 11 m high sulfide structure. The differences observed in assemblage and substratum distribution were related to habitat characteristics (fluid exits, depth and structure orientation). Gradients in community structure were observed, which coincided with an increasing distance from the fluid exits. A biological zonation model for the Eiffel Tower edifice was created in which faunal composition and distribution can be visually explained by the presence/absence of fluid exits.

Numerous attempts have been made over the last thirty years to estimate fluid flow rates at hydrothermal vents, either at the exit of black smoker chimneys or within diffuse flow areas. In this study, we combine two methods to accurately estimate fluid flow velocities at diffuse flow areas. While the first method uses a hot film anemometer that performs high-frequency measurements, the second allows a relatively rapid assessment of fluid flow velocity through video imagery and provides in situ data to calibrate the sensor. Measurements of flow velocities on hydrothermal diffuse flow areas were obtained on the Mid-Atlantic Ridge (MAR). They range from 1.1 to 4.9 mm/s at the substratum level, in low-temperature (4.5-16.4 degrees C) diffuse flow areas from the Tour Eiffel sulfide edifice. A strong correlation was observed between fluid flow velocities and temperature, supporting the possible use of temperature as a proxy to estimate the flow rates in diffuse flow areas where such a simple linear flow/temperature relation is shown to dominate.

Trophic relationships in Bathymodiolus azoricus mussel bed communities on the Tour Eiffel hydrothermal edifice (Lucky Strike) were assessed using delta C-13 and delta N-15 signatures from 14 hydrothermal species. The nutritional basis of B. azoricus was also investigated with delta S-34. Faunal samples and environmental data (temperature, pH, total dissolved sulfide, iron and copper concentrations) were collected from 12 different locations on the edifice. Chemical conditions varied between microhabitats, and were all correlated to temperature. Carbon and nitrogen isotopic results revealed the presence of two, apparently independent, trophic groups. The first was composed of symbiont-bearing fauna (B. azoricus and their associated polychaetes Branchipolynoe seepensis), while the second enclosed heterotrophic fauna (bacterivores, cletritivores, scavengers, predators). A majority of mussels displayed delta C-13 values ranging from -27 parts per thousand to -34 parts per thousand, supporting thiotrophy as the dominant nutritional pathway at Tour Eiffel, with methanotrophy and filter feeding emerging as secondary strategies. This result was corroborated by delta S-34 signatures. However, higher delta C-13 values in larger mussels suggested that, as they grow, B. azoricus mussels rely more heavily on their methanotrophic enclosymbionts. Significant spatial variability in isotopic signatures for single faunal species was observed at the scale of the edifice for three species (B. azoricus, B. seepensis, Amathys lutzi), and environmental conditions explained variation in isotopic signatures for one-third of the species. This confirms the hypothesis raised by several authors on the role of hydrothermal fluids on the trophic network at small spatial scales. We suggest that vent fluid characteristics, by influencing microbial production, are key factors in the variation of local carbon sources at vents.

The objective of this study was to deter-mine the concentrations of different fractions of dissolved copper (after filtration at 0.45 mu m) along the cold part of the hydrothermal fluid-seawater mixing zone on the Tour Eiffel edifice (MAR). Dissolved copper was analyzed by stripping chronopotentiometry (SCP) after chromatographic C-18 extraction. Levels of total dissolved copper (0.03 to 5.15 mu M) are much higher than those reported for deep-sea oceanic waters but in accordance with data previously obtained in this area. Speciation measurements show that the hydrophobic organic fraction (C18Cu) is very low (2 +/- 1%). Dissolved copper is present mainly as inorganic and hydrophilic organic complexes (nonC(18)CU). The distribution of copper along the pH gradient shows the same pattern for each fraction. Copper concentrations increase from pH 5.6 to 6.5 and then remain relatively constant at pH>6.5. Concentrations of oxygen and total sulphides demonstrate that the copper anomaly corresponds to the transition between suboxic and oxic waters. The increase of dissolved copper should correspond to the oxidative redissolution of copper sulphide particles formed in the vicinity of the fluid exit. The presence of such a secondary dissolved copper source, associated with the accumulation of metal sulphide particles, could play a significant role in the distribution of fauna in the different habitats available at vents.

Whilst the fauna inhabiting hydrothermal vent structures in the Atlantic Ocean is reasonably well known, less is understood about the spatial distributions of the fauna in relation to abiotic and biotic factors. In this study, a major active hydrothermal edifice (Eiffel Tower, at 1690 m depth) on the Lucky Strike vent field (Mid-Atlantic Ridge (MAR)) was investigated. Video transects were carried out by ROV Victor 6000 and complete image coverage was acquired. Four distinct assemblages, ranging from dense larger-sized Bathymodiolus mussel beds to smaller-sized mussel clumps and alvinocaridid shrimps, and two types of substrata were defined based on high definition photographs and video imagery. To evaluate spatial variation, faunal distribution was mapped in three dimensions. A high degree of patchiness characterizes this 11 m high sulfide structure. The differences observed in assemblage and substratum distribution were related to habitat characteristics (fluid exits, depth and structure orientation). Gradients in community structure were observed, which coincided with an increasing distance from the fluid exits. A biological zonation model for the Eiffel Tower edifice was created in which faunal composition and distribution can be visually explained by the presence/absence of fluid exits.

Trophic relationships in Bathymodiolus azoricus mussel bed communities on the Tour Eiffel hydrothermal edifice (Lucky Strike) were assessed using delta C-13 and delta N-15 signatures from 14 hydrothermal species. The nutritional basis of B. azoricus was also investigated with delta S-34. Faunal samples and environmental data (temperature, pH, total dissolved sulfide, iron and copper concentrations) were collected from 12 different locations on the edifice. Chemical conditions varied between microhabitats, and were all correlated to temperature. Carbon and nitrogen isotopic results revealed the presence of two, apparently independent, trophic groups. The first was composed of symbiont-bearing fauna (B. azoricus and their associated polychaetes Branchipolynoe seepensis), while the second enclosed heterotrophic fauna (bacterivores, cletritivores, scavengers, predators). A majority of mussels displayed delta C-13 values ranging from -27 parts per thousand to -34 parts per thousand, supporting thiotrophy as the dominant nutritional pathway at Tour Eiffel, with methanotrophy and filter feeding emerging as secondary strategies. This result was corroborated by delta S-34 signatures. However, higher delta C-13 values in larger mussels suggested that, as they grow, B. azoricus mussels rely more heavily on their methanotrophic enclosymbionts. Significant spatial variability in isotopic signatures for single faunal species was observed at the scale of the edifice for three species (B. azoricus, B. seepensis, Amathys lutzi), and environmental conditions explained variation in isotopic signatures for one-third of the species. This confirms the hypothesis raised by several authors on the role of hydrothermal fluids on the trophic network at small spatial scales. We suggest that vent fluid characteristics, by influencing microbial production, are key factors in the variation of local carbon sources at vents.

Numerous attempts have been made over the last thirty years to estimate fluid flow rates at hydrothermal vents, either at the exit of black smoker chimneys or within diffuse flow areas. In this study, we combine two methods to accurately estimate fluid flow velocities at diffuse flow areas. While the first method uses a hot film anemometer that performs high-frequency measurements, the second allows a relatively rapid assessment of fluid flow velocity through video imagery and provides in situ data to calibrate the sensor. Measurements of flow velocities on hydrothermal diffuse flow areas were obtained on the Mid-Atlantic Ridge (MAR). They range from 1.1 to 4.9 mm/s at the substratum level, in low-temperature (4.5-16.4 degrees C) diffuse flow areas from the Tour Eiffel sulfide edifice. A strong correlation was observed between fluid flow velocities and temperature, supporting the possible use of temperature as a proxy to estimate the flow rates in diffuse flow areas where such a simple linear flow/temperature relation is shown to dominate.

The objective of this study was to deter-mine the concentrations of different fractions of dissolved copper (after filtration at 0.45 mu m) along the cold part of the hydrothermal fluid-seawater mixing zone on the Tour Eiffel edifice (MAR). Dissolved copper was analyzed by stripping chronopotentiometry (SCP) after chromatographic C-18 extraction. Levels of total dissolved copper (0.03 to 5.15 mu M) are much higher than those reported for deep-sea oceanic waters but in accordance with data previously obtained in this area. Speciation measurements show that the hydrophobic organic fraction (C18Cu) is very low (2 +/- 1%). Dissolved copper is present mainly as inorganic and hydrophilic organic complexes (nonC(18)CU). The distribution of copper along the pH gradient shows the same pattern for each fraction. Copper concentrations increase from pH 5.6 to 6.5 and then remain relatively constant at pH>6.5. Concentrations of oxygen and total sulphides demonstrate that the copper anomaly corresponds to the transition between suboxic and oxic waters. The increase of dissolved copper should correspond to the oxidative redissolution of copper sulphide particles formed in the vicinity of the fluid exit. The presence of such a secondary dissolved copper source, associated with the accumulation of metal sulphide particles, could play a significant role in the distribution of fauna in the different habitats available at vents.

Whilst the fauna inhabiting hydrothermal vent structures in the Atlantic Ocean is reasonably well known, less is understood about the spatial distributions of the fauna in relation to abiotic and biotic factors. In this study, a major active hydrothermal edifice (Eiffel Tower, at 1690 m depth) on the Lucky Strike vent field (Mid-Atlantic Ridge (MAR)) was investigated. Video transects were carried out by ROV Victor 6000 and complete image coverage was acquired. Four distinct assemblages, ranging from dense larger-sized Bathymodiolus mussel beds to smaller-sized mussel clumps and alvinocaridid shrimps, and two types of substrata were defined based on high definition photographs and video imagery. To evaluate spatial variation, faunal distribution was mapped in three dimensions. A high degree of patchiness characterizes this 11 m high sulfide structure. The differences observed in assemblage and substratum distribution were related to habitat characteristics (fluid exits, depth and structure orientation). Gradients in community structure were observed, which coincided with an increasing distance from the fluid exits. A biological zonation model for the Eiffel Tower edifice was created in which faunal composition and distribution can be visually explained by the presence/absence of fluid exits.

Trophic relationships in Bathymodiolus azoricus mussel bed communities on the Tour Eiffel hydrothermal edifice (Lucky Strike) were assessed using delta C-13 and delta N-15 signatures from 14 hydrothermal species. The nutritional basis of B. azoricus was also investigated with delta S-34. Faunal samples and environmental data (temperature, pH, total dissolved sulfide, iron and copper concentrations) were collected from 12 different locations on the edifice. Chemical conditions varied between microhabitats, and were all correlated to temperature. Carbon and nitrogen isotopic results revealed the presence of two, apparently independent, trophic groups. The first was composed of symbiont-bearing fauna (B. azoricus and their associated polychaetes Branchipolynoe seepensis), while the second enclosed heterotrophic fauna (bacterivores, cletritivores, scavengers, predators). A majority of mussels displayed delta C-13 values ranging from -27 parts per thousand to -34 parts per thousand, supporting thiotrophy as the dominant nutritional pathway at Tour Eiffel, with methanotrophy and filter feeding emerging as secondary strategies. This result was corroborated by delta S-34 signatures. However, higher delta C-13 values in larger mussels suggested that, as they grow, B. azoricus mussels rely more heavily on their methanotrophic enclosymbionts. Significant spatial variability in isotopic signatures for single faunal species was observed at the scale of the edifice for three species (B. azoricus, B. seepensis, Amathys lutzi), and environmental conditions explained variation in isotopic signatures for one-third of the species. This confirms the hypothesis raised by several authors on the role of hydrothermal fluids on the trophic network at small spatial scales. We suggest that vent fluid characteristics, by influencing microbial production, are key factors in the variation of local carbon sources at vents.

The objective of this study was to deter-mine the concentrations of different fractions of dissolved copper (after filtration at 0.45 mu m) along the cold part of the hydrothermal fluid-seawater mixing zone on the Tour Eiffel edifice (MAR). Dissolved copper was analyzed by stripping chronopotentiometry (SCP) after chromatographic C-18 extraction. Levels of total dissolved copper (0.03 to 5.15 mu M) are much higher than those reported for deep-sea oceanic waters but in accordance with data previously obtained in this area. Speciation measurements show that the hydrophobic organic fraction (C18Cu) is very low (2 +/- 1%). Dissolved copper is present mainly as inorganic and hydrophilic organic complexes (nonC(18)CU). The distribution of copper along the pH gradient shows the same pattern for each fraction. Copper concentrations increase from pH 5.6 to 6.5 and then remain relatively constant at pH>6.5. Concentrations of oxygen and total sulphides demonstrate that the copper anomaly corresponds to the transition between suboxic and oxic waters. The increase of dissolved copper should correspond to the oxidative redissolution of copper sulphide particles formed in the vicinity of the fluid exit. The presence of such a secondary dissolved copper source, associated with the accumulation of metal sulphide particles, could play a significant role in the distribution of fauna in the different habitats available at vents.

Numerous attempts have been made over the last thirty years to estimate fluid flow rates at hydrothermal vents, either at the exit of black smoker chimneys or within diffuse flow areas. In this study, we combine two methods to accurately estimate fluid flow velocities at diffuse flow areas. While the first method uses a hot film anemometer that performs high-frequency measurements, the second allows a relatively rapid assessment of fluid flow velocity through video imagery and provides in situ data to calibrate the sensor. Measurements of flow velocities on hydrothermal diffuse flow areas were obtained on the Mid-Atlantic Ridge (MAR). They range from 1.1 to 4.9 mm/s at the substratum level, in low-temperature (4.5-16.4 degrees C) diffuse flow areas from the Tour Eiffel sulfide edifice. A strong correlation was observed between fluid flow velocities and temperature, supporting the possible use of temperature as a proxy to estimate the flow rates in diffuse flow areas where such a simple linear flow/temperature relation is shown to dominate.

The deep-sea hydrothermal vent mussel Bathymodiolus azoricus has been the subject of several studies aimed at understanding the physiological adaptations that vent animals have developed in order to cope with the particular physical and chemical conditions of hydrothermal environments. In spite of reports describing successful procedures to maintain vent mussels under laboratory conditions at atmospheric pressure, few studies have described the mussel's physiological state after a long period in aquaria. In the present study, we investigate changes in mucocytes and hemocytes in B. azoricus over the course of several months after deep-sea retrieval. The visualization of granules of mucopolysaccharide or glycoprotein was made possible through their inherent auto-fluorescent property and the Alcian blue-Periodic Acid Schiff staining method. The density and distribution of droplets of mucus-like granules was observed at the ventral end of lamellae during acclimatization period. The mucus-like granules were greatly reduced after 3 months and nearly absent after 6 months of aquarium conditions. Additionally, we examined the depletion of endosymbiont bacteria from gill tissues, which typically occurs within a few weeks in sea water under laboratory conditions. The physiological state of B. azoricus after 6 months of acclimatization was also examined by means of phagocytosis assays using hemocytes. Hemocytes from mussels held in aquaria up to 6 months were still capable of phagocytosis but to a lesser extent when Compared to the number of ingested yeast particles per phagocytic hemocytes from freshly collected vent mussels. We suggest that the changes in gill mucopolysaccharides and hemocyte glycoproteins, the endosymbiont abundance in gill tissues and phagocytosis are useful health criteria to assess long term maintenance of B. azoricus in aquaria. Furthermore, the laboratory set up to which vent mussels were acclimatized is an applicable system to study physiological reac

The deep-sea hydrothermal vent mussel Bathymodiolus azoricus has been the subject of several studies aimed at understanding the physiological adaptations that vent animals have developed in order to cope with the particular physical and chemical conditions of hydrothermal environments. In spite of reports describing successful procedures to maintain vent mussels under laboratory conditions at atmospheric pressure, few studies have described the mussel's physiological state after a long period in aquaria. In the present study, we investigate changes in mucocytes and hemocytes in B. azoricus over the course of several months after deep-sea retrieval. The visualization of granules of mucopolysaccharide or glycoprotein was made possible through their inherent auto-fluorescent property and the Alcian blue-Periodic Acid Schiff staining method. The density and distribution of droplets of mucus-like granules was observed at the ventral end of lamellae during acclimatization period. The mucus-like granules were greatly reduced after 3 months and nearly absent after 6 months of aquarium conditions. Additionally, we examined the depletion of endosymbiont bacteria from gill tissues, which typically occurs within a few weeks in sea water under laboratory conditions. The physiological state of B. azoricus after 6 months of acclimatization was also examined by means of phagocytosis assays using hemocytes. Hemocytes from mussels held in aquaria up to 6 months were still capable of phagocytosis but to a lesser extent when Compared to the number of ingested yeast particles per phagocytic hemocytes from freshly collected vent mussels. We suggest that the changes in gill mucopolysaccharides and hemocyte glycoproteins, the endosymbiont abundance in gill tissues and phagocytosis are useful health criteria to assess long term maintenance of B. azoricus in aquaria. Furthermore, the laboratory set up to which vent mussels were acclimatized is an applicable system to study physiological reac

The deep-sea hydrothermal vent mussel Bathymodiolus azoricus has been the subject of several studies aimed at understanding the physiological adaptations that vent animals have developed in order to cope with the particular physical and chemical conditions of hydrothermal environments. In spite of reports describing successful procedures to maintain vent mussels under laboratory conditions at atmospheric pressure, few studies have described the mussel's physiological state after a long period in aquaria. In the present study, we investigate changes in mucocytes and hemocytes in B. azoricus over the course of several months after deep-sea retrieval. The visualization of granules of mucopolysaccharide or glycoprotein was made possible through their inherent auto-fluorescent property and the Alcian blue-Periodic Acid Schiff staining method. The density and distribution of droplets of mucus-like granules was observed at the ventral end of lamellae during acclimatization period. The mucus-like granules were greatly reduced after 3 months and nearly absent after 6 months of aquarium conditions. Additionally, we examined the depletion of endosymbiont bacteria from gill tissues, which typically occurs within a few weeks in sea water under laboratory conditions. The physiological state of B. azoricus after 6 months of acclimatization was also examined by means of phagocytosis assays using hemocytes. Hemocytes from mussels held in aquaria up to 6 months were still capable of phagocytosis but to a lesser extent when Compared to the number of ingested yeast particles per phagocytic hemocytes from freshly collected vent mussels. We suggest that the changes in gill mucopolysaccharides and hemocyte glycoproteins, the endosymbiont abundance in gill tissues and phagocytosis are useful health criteria to assess long term maintenance of B. azoricus in aquaria. Furthermore, the laboratory set up to which vent mussels were acclimatized is an applicable system to study physiological reac

The poorly known ferromanganese nodule fauna is a widespread hard substratum community in the deep sea that will be considerably impacted by large-scale nodule mining operations. The objective of this study was to analyze the spatial distribution of the fauna attached to nodules in the Clarion-Clipperton Fracture Zone at two scales; a regional scale that includes the east (14 degrees N, 130 degrees W) and the west (9 degrees N, 150 degrees W) zones and a local scale in which different geological facies (A, B, C and west) are recognizable. The fauna associated with 235 nodules was quantitatively described: 104 nodules from the east zone (15 of facies A, 50 of facies B and 39 of facies C) and 131 nodules from the west zone. Percent cover was used to quantify the extent of colonization at the time of sampling, for 42 species out of the 62 live species observed. Fauna covered up to 18% of exposed nodule surface with an average of about 3%. While species richness increased with exposed nodule surface, both at the regional and at the facies scales (except for facies A), total species density decreased (again except for facies A). When all nodules were included in the statistical analysis, there was no relation between faunal cover and exposed nodule surface. Nevertheless, faunal cover did decrease with exposed nodule surface for the east zone in general and for both facies B and C in particular. Species distributions among facies were significantly different but explained only a very small portion of the variance (similar to 5%). We identified two groups of associated species: a first group of two species and a second group of six species. The other species (34) were independently distributed.. suggesting that species interactions play only a minor role in the spatial distribution of nodule fauna. The flux of particulate organic carbon to the bottom is the only major environmental factor considered to vary between the two zones within this study. We conclude that the high

Manganese nodules provide an important habitat for sessile benthic organisms, many of them novel foraminifera, in the abyssal Pacific. The surface texture and microhabitat heterogeneity of nodules may play a significant role in structuring these communities. We analyzed the distribution of foraminiferal species on 20 nodules collected using either a USNEL box corer or the Nautile submersible sampling tool at a 5000-m-deep site in the Tropical North Pacific. The nodules had an upper region characterized by two microhabitats, namely raised surfaces and intervening depressed surfaces. The more or less vertical sides of the nodules were regarded as a third microhabitat. The upper region had a predominantly smooth texture while the sides had a rough texture. We tested whether the percent cover of each of the 34 species examined was distributed evenly over the three microhabitats and the two surface textures (smooth vs rough). More than half (20 out of 34) of species covered a greater area on smooth surfaces than on rough surfaces, while one-third of the species (12 out of 34) occurred more on raised microhabitats than on depressed surfaces and nodule sides. These two results are closely interrelated, since 78% of the raised surfaces were characterized by a smooth texture. The nodule-encrusting species were feeding either on suspended particles or on particles deposited on the nodule surfaces. Because most species had a mat-like or domed morphology, we suspect that nodule surface feeding was more prevalent. Current flow, which transports propagules and particulate food, may be the main environmental factor explaining the observed faunal distributions, as it is in the case of other hard-substrate assemblages.

The poorly known ferromanganese nodule fauna is a widespread hard substratum community in the deep sea that will be considerably impacted by large-scale nodule mining operations. The objective of this study was to analyze the spatial distribution of the fauna attached to nodules in the Clarion-Clipperton Fracture Zone at two scales; a regional scale that includes the east (14 degrees N, 130 degrees W) and the west (9 degrees N, 150 degrees W) zones and a local scale in which different geological facies (A, B, C and west) are recognizable. The fauna associated with 235 nodules was quantitatively described: 104 nodules from the east zone (15 of facies A, 50 of facies B and 39 of facies C) and 131 nodules from the west zone. Percent cover was used to quantify the extent of colonization at the time of sampling, for 42 species out of the 62 live species observed. Fauna covered up to 18% of exposed nodule surface with an average of about 3%. While species richness increased with exposed nodule surface, both at the regional and at the facies scales (except for facies A), total species density decreased (again except for facies A). When all nodules were included in the statistical analysis, there was no relation between faunal cover and exposed nodule surface. Nevertheless, faunal cover did decrease with exposed nodule surface for the east zone in general and for both facies B and C in particular. Species distributions among facies were significantly different but explained only a very small portion of the variance (similar to 5%). We identified two groups of associated species: a first group of two species and a second group of six species. The other species (34) were independently distributed.. suggesting that species interactions play only a minor role in the spatial distribution of nodule fauna. The flux of particulate organic carbon to the bottom is the only major environmental factor considered to vary between the two zones within this study. We conclude that the high

Manganese nodules provide an important habitat for sessile benthic organisms, many of them novel foraminifera, in the abyssal Pacific. The surface texture and microhabitat heterogeneity of nodules may play a significant role in structuring these communities. We analyzed the distribution of foraminiferal species on 20 nodules collected using either a USNEL box corer or the Nautile submersible sampling tool at a 5000-m-deep site in the Tropical North Pacific. The nodules had an upper region characterized by two microhabitats, namely raised surfaces and intervening depressed surfaces. The more or less vertical sides of the nodules were regarded as a third microhabitat. The upper region had a predominantly smooth texture while the sides had a rough texture. We tested whether the percent cover of each of the 34 species examined was distributed evenly over the three microhabitats and the two surface textures (smooth vs rough). More than half (20 out of 34) of species covered a greater area on smooth surfaces than on rough surfaces, while one-third of the species (12 out of 34) occurred more on raised microhabitats than on depressed surfaces and nodule sides. These two results are closely interrelated, since 78% of the raised surfaces were characterized by a smooth texture. The nodule-encrusting species were feeding either on suspended particles or on particles deposited on the nodule surfaces. Because most species had a mat-like or domed morphology, we suspect that nodule surface feeding was more prevalent. Current flow, which transports propagules and particulate food, may be the main environmental factor explaining the observed faunal distributions, as it is in the case of other hard-substrate assemblages.

Manganese nodules provide an important habitat for sessile benthic organisms, many of them novel foraminifera, in the abyssal Pacific. The surface texture and microhabitat heterogeneity of nodules may play a significant role in structuring these communities. We analyzed the distribution of foraminiferal species on 20 nodules collected using either a USNEL box corer or the Nautile submersible sampling tool at a 5000-m-deep site in the Tropical North Pacific. The nodules had an upper region characterized by two microhabitats, namely raised surfaces and intervening depressed surfaces. The more or less vertical sides of the nodules were regarded as a third microhabitat. The upper region had a predominantly smooth texture while the sides had a rough texture. We tested whether the percent cover of each of the 34 species examined was distributed evenly over the three microhabitats and the two surface textures (smooth vs rough). More than half (20 out of 34) of species covered a greater area on smooth surfaces than on rough surfaces, while one-third of the species (12 out of 34) occurred more on raised microhabitats than on depressed surfaces and nodule sides. These two results are closely interrelated, since 78% of the raised surfaces were characterized by a smooth texture. The nodule-encrusting species were feeding either on suspended particles or on particles deposited on the nodule surfaces. Because most species had a mat-like or domed morphology, we suspect that nodule surface feeding was more prevalent. Current flow, which transports propagules and particulate food, may be the main environmental factor explaining the observed faunal distributions, as it is in the case of other hard-substrate assemblages.

The poorly known ferromanganese nodule fauna is a widespread hard substratum community in the deep sea that will be considerably impacted by large-scale nodule mining operations. The objective of this study was to analyze the spatial distribution of the fauna attached to nodules in the Clarion-Clipperton Fracture Zone at two scales; a regional scale that includes the east (14 degrees N, 130 degrees W) and the west (9 degrees N, 150 degrees W) zones and a local scale in which different geological facies (A, B, C and west) are recognizable. The fauna associated with 235 nodules was quantitatively described: 104 nodules from the east zone (15 of facies A, 50 of facies B and 39 of facies C) and 131 nodules from the west zone. Percent cover was used to quantify the extent of colonization at the time of sampling, for 42 species out of the 62 live species observed. Fauna covered up to 18% of exposed nodule surface with an average of about 3%. While species richness increased with exposed nodule surface, both at the regional and at the facies scales (except for facies A), total species density decreased (again except for facies A). When all nodules were included in the statistical analysis, there was no relation between faunal cover and exposed nodule surface. Nevertheless, faunal cover did decrease with exposed nodule surface for the east zone in general and for both facies B and C in particular. Species distributions among facies were significantly different but explained only a very small portion of the variance (similar to 5%). We identified two groups of associated species: a first group of two species and a second group of six species. The other species (34) were independently distributed.. suggesting that species interactions play only a minor role in the spatial distribution of nodule fauna. The flux of particulate organic carbon to the bottom is the only major environmental factor considered to vary between the two zones within this study. We conclude that the high