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

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

Notes on the genus Dendroceros in West Africa and South Atlantic Islands

Abstract

Following revision of Dendroceros specimens from West Africa and South Atlantic islands, three species are recognized for this geographical area: Dendroceros adglutinatus, D. africanus and finally D. herasii which is described as new. On the other hand, D. crispatus is excluded from Africa while Folioceros fuciformis (Megaceros lacerus, Anthoceros fuciformis) is excluded from Equatorial Guinea. A preliminary key to the five Dendroceros species known to occur in Africa is included.     

Diversity, biomass and distribution pattern of Sargassum beds in the South West lagoon of New Caledonia (South Pacific)

Sargassum (Phaeophyceae, Fucales) is a genus of worldwide distribution recently recognised to proliferate in several regions of the South Pacific. In New Caledonia, species of this genus naturally structure one of the major lagoon habitats but their extent, composition and biomass remain largely unknown. To fill these gaps in our knowledge over large areas, we applied a combination of remote sensing and in situ methods applied to the Neo Caledonian South West Lagoon. Space borne high resolution Landsat (30-m resolution) and Quickbird (2.4-m resolution) images of the Neo Caledonian South West lagoon were used to estimate the spatial extent of the different beds of interest. Species composition was determined for 11 Sargassum beds and seasonal variations were investigated for four representative beds over an 18 month period using quadrats and transects. Relationships between surface cover and biomass were estimated from seasonal field data sets. Finally, four methods requiring variable levels of sampling effort were designed to estimate the total biomass at the scale of each bed, considering (or not) the specific composition, and spatial and temporal variations. Seven Sargassaceae species were identified. Mean surface cover (24.4–51.6%) and total biomass [3.4–1,461.9 t dry matter (DM)] varied widely between beds. Overall, biomass temporal variations were not significantly different, but species-level variations seemed to be bed-specific. The extent of the 11 beds was 9 km²; their total biomass was estimated and compared using each of the four methods, and the most precise method provided an estimate of 2,900 t DM. This study demonstrates a way of characterising Sargassum beds, efficiently and on a large scale, using a combination of remotely sensed and in situ data. These methods should be useful for possible future biomonitoring of Sargassum beds in New Caledonia, and in other areas worldwide.     

Biostratigraphy and histories of upper Miocene - Pliocene Globorotalia, South Atlantic and South West Pacific

The upper Miocene-Pliocene histories of the Globorotalia miozea Plexus and G. crassaformis from the S.W. Pacific and a S. Atlantic site are assessed for their biostratigraphic utility in the mid latitudes of the Southern Hemisphere. As in the S.W. Pacific, Pliocene descendants of G. conoidea (G. pliozea, G. mons) appear in the S. Atlantic near the level at which the G. inflata lineage diverges (as G. puncticulata sphericomiozea). But there is an incomplete history of the latter at the Atlantic site examined, probably due to its northern position. Unlike S.W. Pacific sequences, S. Atlantic strata record a phyletic connection between G. juanai (= G. cibaoensis of authors) and G. crassaformis. The alignment of events in the plexus between the two regions is poor as judged by other foraminiferal datums (extinction of Globoquadrina dehiscens, entries of Globorotalia margaritae and G. crassaformis. As with the G. miozea plexus the records of these taxa probably vary according to latitudinal position.     

The pelagic system in the photic zone of the tropical Atlantic

Summary A multidisciplinary study of the carbon budget in the upper 300 meter of permanently stratified waters was started by two NECTAR-expeditions (North Equatorial Current Trans Atlantic Research) with HMS TYDEMAN in 1977 and 1978 (BAARS, ZIJLSTRA and TIJSSEN, 1979). In 1979 additional measurements were performed during the Gulf of Guinea-expedition with MS TYRO.Primary production in the nutrient depleted mixed layer of the North Equatorial Current estimated from the diurnal cycle in the O₂ concentration (TIJSSEN, 1979) and in POC (POSTMA and ROMMETS, 1979) revealed values 4–10 times higher than the data from the¹⁴C method in the literature: 800–2000 against ca. 200 mg C/m²/day. Moreover,¹⁴C incubations performed in bottle volumes of 4 liter and over 2 hour periods, instead of the recommended 12 hours for oligotrophic waters, gave 5–15 times higher values as incubations in the commonly used 300 ml (or smaller) bottles (GIESKES, KRAAY and BAARS, 1979). In the latter bottles a dramatic decrease of chlorophyll concentrations was observed, suggesting either heavy damage to fragile microflagellates by glasswall contacts and/or insufficient nutrient recycling by the lack of zooplankton in small samples. This then could account for the phenomena of low production and decreasing algal stock in long incubations with small bottles. These results suggest that today's picture of the primary production in the world's oceans (DE VOOIJS, 1979) needs probably a thorough reexamination. On the other hand, experiments in the North Sea and in the Gulf of Guinea did not indicate an effect of bottle size on¹⁴C incorporation and the¹⁴C method gave comparable estimates of primary production as the high precision oxygen method (photometric endpoint detection in the Winkler titration,cf. TIJSSEN, 1979). However, in these waters nutrient concentrations are always clearly above detection levels so that all previous data from somewhat richer areas may have been correct. In relation to the large oligotrophic parts of the oceans a question remains about the fate of the probably high primary production. Is it consumed by the algae themselves by night (POSTMA, 1980), are bacteria and microzooplankton more important consumers than formerly thought, or is the daily ration of zooplankters much higher as expected from extrapolation of filtration experiments in more eutrophic waters? We hope to clarify this topic during the coming NECTAR'82 expedition.Another main objective of the programme concerns the character of the deep chlorophyll maximum in permanently stratified waters. This layer is in the North Equatorial Current located at the depth of the nitratocline and at ca. 1% of the incident light at the surface (SPITZER and WERNAND, 1981). Primary production in the layer seems to be negligible while detailed vertical profiles of zooplankton, obtained with a Longhurst-Hardy Plankton Recorder, revealed no obvious concentrations in the layer. Chlorophyll analysis by thin-layer-chromatography (GIESKES, KRAAY and TIJSSEN, 1978) demonstrated that more than half of the chlorophyll a in the layer consists of an isomer which bleaches rapidly when transferred to higher light levels. The hypothesis was formulated that the chlorophyll maximum layer, at least in this region of the Atlantic, is an accumulation of chlorophyll breakdown products with a quite long turnover time at low light levels. In contrast to these findings, chlorophyll maxima near West-Africa and in the Gulf of Guinea were located at 5–10% light and contained more living algal cells (most of them as small as 1–3 m) than the nutrientpoor mixed layer above it. Primary production profiles had therefore a bimodal shape, with peaks at 30–50% light and at the chlorophyll maximum. Chlorophyll isomers, now analysed with High Pressure Liquid Chromatography, were also in these waters present at nearly all stations and comprised 30–70% of total chlorophyll comparable to the situation in the North Equatorial Current (GIESKES and KRAAY, 1981). It could be shown that these isomers are not involved in primary production, so that well established relationships between chlorophyll, light and primary production, found in oceanographic literature, have to be reconsidered.     

Size-fractionated Primary Production in the South Atlantic and Atlantic Sectors of the Southern Ocean

Results are presented from size fractionated chlorophyll a (Chla) and primary production studies along a transect between Antarcticaand southern Africa during the second South African AntarcticMarine Ecosystem Study (SAAMES II), conducted in late australsummer (January to February) 1993. Total integrated Chl a alongthe transect was highest in the vicinity of the Marginal IceZone (MIZ) and Antarctic Polar Front (APF). At these stations,integrated Chl a biomass was always >25 mg Chl a m^–2and was dominated by microphytoplankton. Although nominal increasesinChl a biomass were also associated with the Subantarctic Front(SAF) and Subtropical Convergence (STC), total Chl a biomassin these regions was dominated by nanophytoplankton. Withinthe inter-frontal regions, total integrated Chl a biomass waslower, generally <25 mg Chl a m^–2, and was always dominatedby nanophytoplankton. An exception was found in the AgulhasReturn Current (ARC) where picophytoplankton dominated. Totaldaily integrated production along the transect ranged between60 and 436 mg C m^–2 day^–1. Elevated production rateswere recorded at stations occupied in the vicinity of the MIZand at all the major oceanic frontal systems. The contributionsof the various size fractions to total daily production displayedthe same spatial pattern as integrated biomass, with microphytoplanktonbeing the most important contributor in areas characterizedby elevated phytoplankton biomass. Outside these regions, nanophytoplanktondominated the total phytoplankton production. Again, an exceptionwas found in the ARC north of the STC where picophytoplanktondominated total production. There, the lowest production alongthe entire transect was recorded, with total daily integratedproduction always <90 mg C m^–2 day^–1. The increasedproduction rates recorded in the MIZ appeared to result fromincreased water column stability as indicated by a shallow mixed-layerdepth. Within the inter-frontal regions, the existence of adeep mixed layer appeared to limit phytoplankton production.Low silicate concentrations in the waters north of the APF mayalso have limited the growth of large microphytoplankton.     

Two new anthiine fishes from the eastern tropical Atlantic

Two anthiine fishes from the eastern tropical Atlantic are described as newHolanthias cyprinoides andAnthias helenensis. The former is distinguished from the other Atlantic species ofHolanthias in having the forked caudal fin with rounded lobes and from the Indo-Pacific species in having no elongated dorsal spines or soft rays. The latter is closely related toAnthias asperilinguis Günther (South America, Atlantic coast), but differs from it in having more pectoral fin rays and more gill rakers. The present investigation onAnthias suggests that AtlanticAnthias is a genus distinct fromPseudanthias of the Indo-Pacific.     

Spatial and temporal assessment of sources contributing to the annual austral spring mid-tropospheric ozone maxima over the tropical South Atlantic

The primary source of the annual austral spring mid‐tropospheric ozone maxima over the tropical South Atlantic has generally been assumed to be biomass burning. However, ozone precursor emissions from biogenic, lightning, and anthropogenic sources in subequatorial Africa before and during the ozone peak are shown to be comparable, if not greater, in magnitude to regional biomass burning production. Moreover, an investigation of the spatial and temporal characteristics of these ozone precursor sources (i.e. vegetative and microbial activity, lightning‐induced generation, and anthropogenic emissions) suggests that these alternative sources can potentially make a substantial contribution to the seasonal ozone peak. This argument is supported by the practical limitations of atmospheric transport available to regionally produced ozone and ozone precursors.     

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 degrees N and 56.6 to 18.5 degrees 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 gamma-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 gamma-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 degrees C and where nitrate was undetectable. In contrast, nonfilamentous nifH sequences were found throughout a broader temperature range, 15 to 30 degrees C, more often in waters with temperature of <26 degrees C, and were sometimes recovered from waters with detectable nitrate concentrations.     

Structural-functional relationships in the pelagic community of the eastern tropical Atlantic Ocean

Sampling was carried out in the eastern tropical Atlantic duringJuly–September, 1987 on four transects at latitudes 6,12, 18 and 24°W, from 6°S to 4°N. A total numberof 81 stations were occupied where physical, chemical and biologicalmeasurements were taken. Temperature, salinity, chlorophylla, phytoplankton, primary production (PP), bacterial production,micro-, meso-, macrozooplankton and mesopelagic fish and squidwere sampled in the upper 120 m layer. The biomass size spectrumof the plankton community was relatively flat within the divergencezone but the slope of the size spectrum increased towards theconvergence zone meaning an increased contribution of the small-sizedorganisms. Higher turnover rates (the ratio of PP to the totalcommunity biomass) were observed in regions where the size spectrasloped more steeply. PP exceeded the community metabolism bythree to five times in regions where the size spectra were sharplysloped. In regions with flat-type spectra (with slopes from-0.2 to 0) a balance between PP and community metabolism wasobserved. The variation coefficient and the dispersion index(the variance to mean ratio) indicated an increase of the spatialvariability of biomass in the sequence: ‘phytoplankton–mesozooplankton–macrozooplankton–micronekton’.At the higher trophic level of the zooplankton community, betweenthe biomass of squid and gelatinous zooplankton, a logarithmiclinear relationship was derived. Squid actively consumed gelatinousorganisms in regions of high concentrations of their prey, althoughlarge copepods, euphausiids, amphipods, flying fish, and myctophidfish also contributed to their diet.     

Pelagic communities of seabirds in the South Atlantic Ocean

This paper presents quantitative analyses at the community level of the spatial dispersion of 44 species of pelagic birds in the South Atlantic Ocean. I have examined the extent to which assemblages of pelagic bird species are influenced by the combined influences of the abundances of other species and the physical structure of the pelagic environment. Results are based upon strip transects collected from oceanographic research vessels during the four consecutive austral summers of 1982–1986.
The distributions of most seabird species were statistically independent of the distributions of other species. Species assemblages, however, did occur and were correlated with the physical structure of the ocean. The study area was divided into four zones, each defined by a distinctive assemblage of birds. Physical oceanography proved to be important in determining the species composition of each zone. These relationships were apparent despite the potentially confounding influence of large numbers of immatures and non-breeding adults.     

Abundance and distribution of major groups of diazotrophic cyanobacteria and their potential contribution to N2 fixation in the tropical Atlantic Ocean

The abundances of six N₂‐fixing cyanobacterial phylotypes were profiled at 22 stations across the tropical Atlantic Ocean during June 2006, and used to model the contribution of the diazotrophs to N₂ fixation. Diazotroph abundances were measured by targeting the nifH gene of Trichodesmium, unicellular groups A, B, C (UCYN‐A, UCYN‐B and UCYN‐C), and diatom‐cyanobiont symbioses Hemiaulus–Richelia, Rhizosolenia–Richelia and Chaetoceros–Calothrix. West to east gradients in temperature, salinity and nutrients [NO₃^‐ + NO₂^‐, PO₄^3?, Si(OH)₄] showed the influence of the Amazon River plume and its effect on the distributions of the diazotrophs. Trichodesmium accounted for more than 93% of all nifH genes detected, dominated the warmer waters of the western Atlantic, and was the only diazotroph detected at the equatorial upwelling station. UCYN‐A was the next most abundant (> 5% of all nifH genes) and dominated the cooler waters of the eastern Atlantic near the Cape Verde Islands. UCYN‐C was found at a single depth (200 m) of high salinity and low temperature and nutrients, whereas UCYN‐B cells were widespread but in very low abundance (6.1 × 10¹ ± 4.6 × 10² gene copies l^?1). The diatom‐cyanobionts were observed primarily in the western Atlantic within or near the high Si(OH)₄ input of the Amazon River plume. Overall, highest diazotroph abundances were observed at the surface and declined with depth, except for some subsurface peaks in Trichodesmium, UCYN‐B and UCYN‐A. Modelled contributions of Trichodesmium, UCYN‐B and UCYN‐A to total N₂ fixation suggested that Trichodesmium had the largest input, except for the potential of UCYN‐A at the Cape Verde Islands.     

Cleaning interactions at the only atoll in the South Atlantic

In marine ecosystems, cleaning is a mutualistic relationship in which so-called cleaners remove ectoparasites, diseased tissue, or mucus from the body of their clients, and thus help to maintain a healthy reef community. In spite of its importance in many marine habitats, this interaction remains poorly understood, particularly at oceanic islands. Here, we present the first comprehensive study of cleaning interactions in a reef fish assemblage at Rocas, the only atoll in the South Atlantic. We recorded 318 cleaning events, in which six fish species, including two endemic ones, and two shrimp species acted as cleaners. The clients serviced by these cleaners were 21 bony fish species, one shark and one sea turtle. The cleaner wrasse Thalassoma noronhanum and the cleaner goby Elacatinus phthirophagus were the cleaners with the greatest number of events and species richness of clients. Additionally, 82% of clients in the cleaning events were non-piscivores, and the abundance of both cleaners and clients positively influenced the number of cleaning events (R² = 0.4; p < 0.001). Our results indicate that Rocas atoll has a high species richness of cleaner species despite its small size and highlight the importance of studies of cleaning symbiosis, even in isolated places with low species richness, for a better comprehension of this association in reefs.     

A comparative study of cleaning activity of two reef fishes at Fernando de Noronha Archipelago,tropical West Atlantic

Cleaner fishes are usually classified as obligate or facultative cleaners according to their diet and the extent to which their nutritional requirements in the different ontogenetic stages are gained from cleaning. While obligate cleaners clean throughout their lives and ingest mainly food taken from the clients’ body surface, facultative cleaners clean only as juveniles and have a broader diet. In addition, some facultative cleaners may experience a relatively higher predation risk, and thus rarely interact with piscivorous fishes. Despite these acknowledged differences, there are very few studies that compare cleaning activity of obligate and facultative cleaners within the same area. Cleaning activity of the obligate cleaner goby Elacatinus cf. randalli and the facultative cleaner wrasse Thalassoma noronhanum were comparatively examined at Fernando de Noronha Archipelago, tropical West Atlantic. The client assemblage attended by the two cleaners differed, as the goby attended a slightly greater diversity of species (22), mostly piscivores and zoobenthivores, and the wrasse attended fewer species (19), mostly planktivores. Chromis multilineata was the most common client species of both cleaners, although body size (which is expected to be positively correlated to clients’ ectoparasite load) of C. multilineata individuals attended by the goby was larger than that of the individuals attended by the wrasse. Despite such differences, T. noronhanum showed a surprisingly species-rich client assemblage when compared with other cleaners of the genus Thalassoma. In addition, the frequency and time spent on cleaning interactions, as well as the number of client species attended per 10-min period, was similar for both cleaner species, which indicate that they have important yet complimentary ecological roles in the reef community at Fernando de Noronha Archipelago.     

Copepod feeding in a tuna fishery area of the tropical Atlantic Ocean

Biomass, feeding and metabolic rates of planktonic copepods were studied in an oligotrophic area of the tropical Atlantic Ocean during an instability wave period (boreal summer) and a stratified period (boreal winter). In summer, zooplankton biomass was higher than in winter, showing a positive effect of the instability wave. Moreover, feeding equilibrated metabolic expenditures of copepods in most cases during the instability. In contrast, in stratified conditions copepods did not equilibrate their metabolic budget. Our results suggest that the microbial loop was the dominant trophic pathway during both periods but with a quicker cycling during the instability.     

Three new species of gnathostomulida from the West Atlantic

Two new species of Gnathostomulida from Bermuda and one from North Carolina are described. All belong to the higher Bursovaginoidae with two occurring in the genus Austrognathia and one in the genus Gnathostomula. Tables of major diagnostic characters showing morphological variability within species and between previously described species in the same genus are included. Clarification in recognition of Gnathostomula jenneri and Gnathostomula brunidens is given under “Remarks on Known Species”.