Biogeography and Potential Exchanges Among the Atlantic Equatorial Belt Cold-Seep Faunas

Like hydrothermal vents along oceanic ridges, cold seeps are patchy and isolated ecosystems along continental margins, extending from bathyal to abyssal depths. The Atlantic Equatorial Belt (AEB), from the Gulf of Mexico to the Gulf of Guinea, was one focus of the Census of Marine Life ChEss (Chemosynthetic Ecosystems) program to study biogeography of seep and vent fauna. We present a review and analysis of collections from five seep regions along the AEB: the Gulf of Mexico where extensive faunal sampling has been conducted from 400 to 3300m, the Barbados accretionary prism, the Blake ridge diapir, and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin. Of the 72 taxa identified at the species level, a total of 9 species or species complexes are identified as amphi-Atlantic. Similarity analyses based on both Bray Curtis and Hellinger distances among 9 faunal collections, and principal component analysis based on presence/absence of megafauna species at these sites, suggest that within the AEB seep megafauna community structure is influenced primarily by depth rather than by geographic distance. Depth segregation is observed between 1000 and 2000m, with the middle slope sites either grouped with those deeper than 2000m or with the shallower sites. The highest level of community similarity was found between the seeps of the Florida escarpment and Congo margin. In the western Atlantic, the highest degree of similarity is observed between the shallowest sites of the Barbados prism and of the Louisiana slope. The high number of amphi-atlantic cold-seep species that do not cluster according to biogeographic regions, and the importance of depth in structuring AEB cold-seep communities are the major conclusions of this study. The hydrothermal vent sites along the Mid Atlantic Ridge (MAR) did not appear as “stepping stones” for dispersal of the AEB seep fauna, however, the south MAR and off axis regions should be further explored to more fully test this hypothesis.     

Distribution and Ecophysiology of Calanoid Copepods in Relation to the Oxygen Minimum Zone in the Eastern Tropical Atlantic

Oxygen minimum zones (OMZs) affect distribution patterns, community structure and metabolic processes of marine organisms. Due to the prominent role of zooplankton, especially copepods, in the marine carbon cycle and the predicted intensification and expansion of OMZs, it is essential to understand the effects of hypoxia on zooplankton distribution and ecophysiology. For this study, calanoid copepods were sampled from different depths (0–1800 m) at eight stations in the eastern tropical Atlantic (3°47′N to 18°S) during three expeditions in 2010 and 2011. Their horizontal and vertical distribution was determined and related to the extent and intensity of the OMZ, which increased from north to south with minimum O₂ concentrations (12.7 µmol kg⁻¹) in the southern Angola Gyre. Calanoid copepod abundance was highest in the northeastern Angola Basin and decreased towards equatorial regions as well as with increasing depth. Maximum copepod biodiversity was observed in the deep waters of the central Angola Basin. Respiration rates and enzyme activities were measured to reveal species-specific physiological adaptations. Enzyme activities of the electron transport system (ETS) and lactate dehydrogenase (LDH) served as proxies for aerobic and anaerobic metabolic activity, respectively. Mass-specific respiration rates and ETS activities decreased with depth of occurrence, consistent with vertical changes in copepod body mass and ambient temperature. Copepods of the families Eucalanidae and Metridinidae dominated within the OMZ. Several of these species showed adaptive characteristics such as lower metabolic rates, additional anaerobic activity and diel vertical migration that enable them to successfully inhabit hypoxic zones.     

The ecology of Saint Paul's Rocks (Equatorial Atlantic)

St Paul's Rocks are a remote group of islets, barely 400 m across, in the equatorial mid-Atlantic (o°55'N 29°21'W). The results of a primarily zoological survey of the Rocks, carried out in September 1979 by the Cambridge Expedition to Saint Paul's Rocks, are reported. In addition, earlier studies of the Rocks' biology are discussed in the light of recent research. The terrestrial fauna is dominated by three species of sea-bird and a land crab. Also reported from on land are a few species of insects, ticks and spiders, an endemic pseudoscorpion, and a centipede; the majority of these species are associated with the seabirds.
The islets are devoid of tracheophytes, bryophytes and lichens.
The marine ecology of the Rocks is discussed with respect to habitat. Five habitats are distinguished in the supralittoral and littoral zones; the fauna and to a limited extent the flora of each is described. Sublittorally the Rocks consist almost entirely of steep escarpments extending to 60 m depth and beyond. The shallow sublittoral (< 60 m deep) is divided into four habitats: (1) Palythoa zone; (2) Caulerpa zone; (3) sub- Cauferpa zone, and (4) areas of unstable substrates. The Palythoa zone occupies a band extending from the base of the littoral to a depth of c . 5m and is dominated by a mat-forming hexacoral, Palythoa caribaeorum. Below this zone to a depth of c . 33 m the rock faces are thickly invested by a species of green-alga, Caulerpa racemosa. At depth this Caulerpa dominated zone is succeeded by an invertebrate dominated community. Unstable substrates are represented by a few pockets of coarse sand and some patches of rubble. The communities in each of these habitats are described. A primarily taxonomic review of the deep-water fauna is also included. Finally, the influence of ocean currents on the zoogeographical relationships of the Rocks' fauna is briefly examined.     

Percolated mantle peridotites along the Romanche Fracture Zone (Equatorial Atlantic ocean)

Pl-peridotites dredged along the Romanche Fracture Zone (FZ) are peculiar rocks. These peridotites derived from weakly deformed upper mantle rocks widespread percolated by melts with gabbroic composition. Mantle peridotite reacted with hot melt percolating through it and trapped melt which crystallised in pockets and in localised conduits at higher levels in the lithosphere. Reaction and crystallisation caused the peculiar microstructures, modal and chemical compositional variations, both in the bulk rocks and in the residual mineral phases. Electron and ion microprobe analyses suggest that percolating melts, characterised by different geochemical signatures, interacted, at deep levels in the lithosphere, with the Romanche peridotites mostly modifying the chemical compositions of Spl and Cpx. These new data show that impregnated mantle peridotites were emplaced all along the Romanche FZ, also in the western Romanche ridge transform intersection, were only fertile Pl-free peridotites were previously described. The cold equatorial upper mantle belt seems to influence strongly the dynamic of melting, melt migration, extraction and its final emplacement on the seafloor. Melts produced in this cold oceanic lithosphere are not able to segregate and join the overlying oceanic crust, but freeze and remain trapped at depth. Nella seduta del 17 novembre 1999.     

Distribution and conservation status of primates in Bioko island,Equatorial Guinea

Ten species of non-human primates are indigenous to Bioko; half of these are endangered and between five and eight are endemic subspecies. Recent data on their status and distribution have been lacking. In 1986, a ten-week survey of the island was carried out to determine the distribution and status of the primates and the natural vegetation, and to evaluate the effects of man on them. This paper presents the results of that survey, gives an update of conservation achievements since 1986, and highlights current concerns. Between 1974 and 1986 it is probable that numbers of all Bioko primates rose as a result of an increase in habitat and of reduced hunting. At the time of the survey there was considerably more natural, undisturbed, vegetation remaining in Bioko tran expected. Much of this vegetation occurs within two large blocks that are of outstanding importance to the conservation of species in tropical Africa, particularly of plants and primates.     

Distribution of chiorophylls and phaeopigments in the Northwestern Atlantic Ocean

During a cruise in the Northwestern Atlantic the distributionsof chlorophylls and phaeopigments were investigated in relationto mesoscale features by high-performance liquid chromatography.Samples were collected from hydrographic Stations and from twoContinuous underway sampling systems fished simultaneously atthe surface (2 m) and at deeper (30–90 m) depths. Highestpigment concentrations were associated with the transitionalzones between Slope Water, the Gulf Stream and Northern SargassoSea. Chlorophyll b occurred in 47% of the samples analyzed andthe dominant chlorophyll a degradation product was phaeophorbidea. Chlorophyllide a was usually found at low concentrationsand phaeophytin a levels were undetectable. Concentrations ofthe various chlorophylls and their degradation products wereused to infer Changes in taxonomic composition, phytoplanktonbiomass and zooplankton grazing pressure across several frontalzones.     

Competition for polymers among heterotrophic bacteria, isolated from particles of the Equatorial Atlantic.

Three heterotrophic bacterial strains, isolated from organic particles of the upper water column of the Equatorial Atlantic, taken during a cruise on the R/V METEOR (1997), were investigated concerning their physiological and phylogenetic properties using classic microbiological and modern molecular-biological methods. All isolates are gram-negative rods able to use polymers such as cellulose, chitin or starch as sole carbon source. The phylogeny of these isolates was investigated by fluorescence in situ hybridization (FISH) and 16S rDNA sequencing. The three isolated strains belong to the Cytophaga/Flavobacteria, gamma-Proteobacteria (Marinobacter sp.), and alpha-Proteobacteria (Sulfitobacter pontiacus). In order to study succession during growth on polymers naturally occurring in marine habitats, FISH was used as a new approach to detect cells from different phylogenetic clusters in the course of a single growth experiment. Mixed cultures consisting of the isolated strains in equal amounts were incubated with cellulose, chitin or starch. Isolate 4301-10/2, a member of the gamma-Proteobacteria, dominated in mixed cultures growing on cellulose, chitin, or starch after only 10 days, with 55, 60, and 95%, respectively, of cells hybridizing with 4,6-diamidino-2-phenylindole (DAPI).     

Stable isotopic study of Oligocene-Miocene sediments from DSDP Site 354, Equatorial Atlantic

The oxygen- and carbon-isotope compositions of planktic and benthic foraminifera and calcareous nannofossils from Middle Oligocene-Early Miocene Equatorial Atlantic sediments (DSDP Site 354) indicate two important paleoceanographic changes, in the Late Oligocene (foraminiferal Zone P.21) and in the Early Miocene (foraminiferal Zone N.5). The first change, reflected by a δ¹⁸O increase of 1.45‰ inGlobigerina venezuelana, affected only intermediate pelagic and not surface, deep or bottom waters. The second change affected surface and intermediate waters, whereas deep and bottom waters showed only minor fluctuations. In the case of the former the isotope effect of the moderate ice accumulation on the Antarctic continent is amplified in the Equatorial Atlantic by changes in the circulation pattern. The latter paleoceanographic change, reflected by a significant increase in¹⁸O in both planktic and benthic forms (about 1.0‰ and 0.5‰, respectively), may have been caused by ice volume increase and temperature decrease. Both oxygen- and carbon-isotope compositions indicate a marked depth-habitat stratification for planktic foraminifera and calcareous nannofossils. Three different dwelling groups are recognized: shallowGlobigerinoides, Globoquadrina dehiscens, Globorotalia mayeri and nannofossils; intermediateGlobigerina venezuelana; and deepCatapsydrax dissimilis. The comparison of foraminifera and calcareous nannofossils suggests that the isotopic compositions of nannofossils are generally controlled by the same parameters which control the isotopic composition of shallow-dwelling foraminifera, but the former are more enriched in¹⁸O.     

Seasonal and interannual variability of chlorophyll a and primary production in the Equatorial Atlantic: in situ and remote sensing observations

The seasonal variability of phytoplankton in the EquatorialAtlantic was analysed using Sea-viewing Wide Field-of-view Sensor(SeaWiFS)-derived chlorophyll a (Chl a) concentration data from1998 to 2001, together with in situ Chl a and primary productiondata obtained during seven cruises carried out between 1995and 2000. Monthly averaged SeaWiFS Chl a distributions werein agreement with previous observations in the Equatorial Atlantic,showing marked differences between 10° W in the EasternTropical Atlantic (ETRA) and 25° W in the Western TropicalAtlantic (WTRA) provinces (Longhurst et al. 1995. J. PlanktonRes., 17, 1245–1271). The seasonal cycle of SeaWiFS-derivedChl a concentration calculated for 0–10° S, 0–20°W (ETRA) is consistent with in situ Chl a measurements, withvalues ranging from 0.16 mg m^–3, from February to April,to 0.52 mg m^–3 in August. Lower variability was observedin 10° N–10° S, 20–30° W (WTRA) whereminimum and maximum concentrations occurred in April (0.15 mgm^–3) and in August (0.24 mg m^–3), respectively.A significant empirical relationship between depth-integratedprimary production and in situ measured sea surface Chl a wasfound for ETRA, allowing us to estimate the seasonal cycle ofdepth-integrated primary production from SeaWiFS-derived Chla. As for Chl a, this model was verified in a small area ofthe Eastern Equatorial Atlantic (0–10° S, 0–20°W), although in this instance it was not completely able todescribe the magnitude and temporal variability of in situ primaryproduction measurements. The annual euphotic depth-integratedprimary production rate estimated for ETRA by our empiricalmodel was 1.4 Gt C year^–1, which represents 16% of theopen ocean primary production estimated for the whole AtlanticOcean.     

The size distribution of phytoplankton and participate organic matter in the Equatorial Atlantic Ocean: importance of ultraseston and consequences

The size fractionation of paniculate matter (<200, <35,<3 and <1 µm) has been measured in the EquatorialAtlantic Ocean at different stations. Chlorophyll a, phaeophytin,particulate carbon, nitrogen and phosphorus have been analysed.Primary production by ¹⁴CO₂ uptake was also measured with prescreeningtechnique. It appears from this study, that the pariculate matter has avery small size: 40–60% of the chlorophyll passed through1 µm Nucleopore filter, and 75–90% of the paniculatecarbon and nitrogen passed through 3 µm Nucleopore filterin offshore waters. From the atomic ratio C/N, C/P and C/chla, and primary productionvalues, the <3 µm fraction would be mainly constitutedby inactive photosynthetic organisms or partides of detritus.The 3–35 µm fraction, in contrast, would be principallyactive phytoplankton.     

Occurrence of the ocean triggerfish in the Eastern Atlantic (Canary Islands)

The occurrence, habitat and abundance are reported for the population of ocean triggerfish Canthidermis sufflamen (Balistidae) at El Hierro, Canary Islands. This is the first record from the Eastern Atlantic.     

Distribution of capsulated bacterioplankton in the North Atlantic and North Sea

In laboratory experiments, bacterioplankton were incubated under different nutrient conditions, and the percentage of bacteria exhibiting a polysaccharidic capsule (capsulated bacteria) and that of CTC (cyanotetrazolium chloride)-positive and therefore metabolically highly active bacteria were determined. In these seawater cultures amended with nutrients more than 95% of the CTC-positive cells exhibited a capsule. During two cruises, one to the North Atlantic and one to the North Sea, we investigated the distribution of capsulated bacteria throughout the water column. Capsulated bacteria were generally more abundant in eutrophic surface waters than in deeper layers or more oligotrophic regions. In the upper 100 m of the North Atlantic, about 6–14% of the total bacterioplankton community was capsulated, while in the layers below 100 m depth, 97% of the bacteria lacked a visible capsule. The percentage of capsulated bacteria correlated with bacterial abundance and production, and chlorophyll a concentration. Also, the bioavailability of DOC (dissolved organic carbon), estimated by the ratio between bacterial production and DOC concentration, significantly correlated with the percentage of capsulated bacteria. In the North Sea, the contribution of capsulated bacteria to the total number of bacteria decreased from the surface (3 m depth) to the near-bottom (25–35 m) layers from 20% to 14% capsulated bacteria. In the nearshore area of the North Sea, about 27% of the bacteria exhibited a capsule. Overall, a pronounced decrease in the contribution of capsulated bacteria to the total bacterial abundance was detectable from the eutrophic coastal environment to the open North Atlantic. Using this epifluorescence-based technique to enumerate capsulated bacterioplankton thus allowed us to routinely assess the number of capsulated bacteria even in the oceanic water column. Based on the data obtained in this study we conclude that almost all metabolically highly active bacteria exhibit a capsule, but also some of the metabolically less active cells express a polysaccharide capsule detectable with this method.     

Diazotrophic Diversity and Distribution in the Tropical and Subtropical Atlantic Ocean

To understand the structure of marine diazotrophic communities in the tropical and subtropical Atlantic Ocean, the molecular diversity of the nifH gene was studied by nested PCR amplification using degenerate primers, followed by cloning and sequencing. Sequences of nifH genes were amplified from environmental DNA samples collected during three cruises (November-December 2000, March 2002, and October-November 2002) covering an area between 0 to 28.3°N and 56.6 to 18.5°W. A total of 170 unique sequences were recovered from 18 stations and 23 depths. Samples from the November-December 2000 cruise contained both unicellular and filamentous cyanobacterial nifH phylotypes, as well as γ-proteobacterial and cluster III sequences, so far only reported in the Pacific Ocean. In contrast, samples from the March 2002 cruise contained only phylotypes related to the uncultured group A unicellular cyanobacteria. The October-November 2002 cruise contained both filamentous and unicellular cyanobacterial and γ-proteobacterial sequences. Several sequences were identical at the nucleotide level to previously described environmental sequences from the Pacific Ocean, including group A sequences. The data suggest a community shift from filamentous cyanobacteria in surface waters to unicellular cyanobacteria and/or heterotrophic bacteria in deeper waters. With one exception, filamentous cyanobacterial nifH sequences were present within temperatures ranging between 26.5 and 30°C and where nitrate was undetectable. In contrast, nonfilamentous nifH sequences were found throughout a broader temperature range, 15 to 30°C, more often in waters with temperature of <26°C, and were sometimes recovered from waters with detectable nitrate concentrations.     

Apparently contradictory routes in the expansion of two fish species in the Eastern Atlantic

One specimen of the boreal Clupea harengus (Clupeidae) was caught southerly, in the Gulf of Cádiz (southern Spain), whereas a specimen of the subtropical Epinephelus aeneus (Serranidae) was caught northerly, in Galician waters (north-west of Spain). These catches represent a new southernmost record in the eastern Atlantic for C. harengus and the second northernmost for E. aeneus. The co-occurrence of warm and cold affinity fishes in temperate latitudes is unusual, but the existence of anomalies in the temperature of the Atlantic waters could be behind these findings.     

Abyssal Megafaunal Necrophages: Latitudinal Differences in the Eastern North Atlantic Ocean

A pop-up time-lapse camera system (BATHYSNACK) was used to study megafaunal organisms attracted to bait at three abyssal localities in the eastern North Atlantic Ocean, a northern site on the Porcupine Abyssal Plain and two sub-tropical sites on the Madeira Abyssal Plain. Fish and decapod crustaceans appeared on 47% of photographs taken during deployments of up to seven days duration. The fishes Coryphaenoides (Nematonurus) armatus and Pachycara bulbiceps and the decapod Munidopsis spp. dominated appearances at the northern locality whereas over 95% of records at the two subtropical localities were of the decapod Plesiopenaeus armatus. Cyclical variations in necrophage abundance did not correlate with tidal cycles of near-bottom currents, although high P. armatus numbers were inversely related to current speed. Most of the frequently recorded entities showed contagious intraspecific distributions. In contrast, many of the interspecies relationships were non-random, with co-occurrences lower than expected, probably as a result of physical disturbance or potential predation. Population densities based on times of first arrival were calculated. Assemblages dominated by fish at higher latitudes and crustaceans at lower latitudes are discussed in terms of organic carbon supply to the deep sea and pelagic community structures.     

Isolation and identification of root associated diazotrophs

Diazotrophs have been isolated from the rhizosphere or roots of plants by many workers. To recognize a certain diazotroph as the most abundant bacterium at a certain site or as the principal agent responsible for N₂-fixation is much more difficult. It is probable that many diazotrophs, including possibly the most efficient ones, have not been identified yet. The use of proper selective media which simulate the environment of the various diazotrophsin situ has led to the discovery of 10 new root-associated diazotrophs, three of them during 1986/1987 (Azospirillum halopraeferans, Herbaspirillum seropedicae and the recently proposedAcetobacter diazotrophicus). The importance of using a variety of carbon substrates in the growth media with pH indicators, and the use of N-free semi-solid media, is discussed. Recognition of plant-bacteria interactions requires, in addition to the identification of the bacteria, the demonstration of effects of the plant on the bacteria and of the bacteria on the plant. Confirmation of the identity of diazotrophs responsible for response of plants to inoculation must be made in experiments with strains labelled with antibiotic resistance or other markers. If establishment of the inoculated strain is demonstrated in plants grown in¹⁵N-labelled soil, the¹⁵N enrichment of the plants will reveal if any observed responses in N yield are due to N₂-fixation or increased soil/fertilizer-N uptake.