The Biogeography of Tropical Phytoplankton Species in the Pacific Ocean

The ranges of 10 tropical Pacific phytoplankton species are shown in the maps. Five types of these ranges are discussed. Except for broadly tropical species some species with much narrower ranges are characterized. The bases of ranges and the expatriation areas are given. The bases of ranges of all the species studied fall into similar temperature and salinity limits. Most of the species inhabit a wide range of nutrients concentration, but Climacodium frauenfeldianum and Hemiaulus hauckii are only found at low phosphate and silicate concentrations. These peculiarities determine the geography of ranges of these two species. Among the species investigated only Ceratium gravidum belongs to the shade flora.     

Phytoplankton Biogeography and Community Stability in the Ocean

Background

Despite enormous environmental variability linked to glacial/interglacial climates of the Pleistocene, we have recently shown that marine diatom communities evolved slowly through gradual changes over the past 1.5 million years. Identifying the causes of this ecological stability is key for understanding the mechanisms that control the tempo and mode of community evolution.

Methodology/Principal Findings

If community assembly were controlled by local environmental selection rather than dispersal, environmental perturbations would change community composition, yet, this could revert once environmental conditions returned to previous-like states. We analyzed phytoplankton community composition across >10⁴ km latitudinal transects in the Atlantic Ocean and show that local environmental selection of broadly dispersed species primarily controls community structure. Consistent with these results, three independent fossil records of marine diatoms over the past 250,000 years show cycles of community departure and recovery tightly synchronized with the temporal variations in Earth''s climate.

Conclusions/Significance

Changes in habitat conditions dramatically alter community structure, yet, we conclude that the high dispersal of marine planktonic microbes erases the legacy of past environmental conditions, thereby decreasing the tempo of community evolution.     

Phytoplankton rate processes in the oligotrophic waters of the central North Pacific Ocean

The central North Pacific is one of the more oligotrophic regionsof the world oceans. There the particulate organic nitrogen:cabonratio of surface waters is variable and less than the Redfieldratio of 16N:106C by atoms. The phytoplankton P/B ratio basedupon both C and N assimilation rate varied directly with theparticulate matter PON:POC ratio as did the productivity index[mg C (mg chl a)^–1h^–1]. At steady state the doublingtime of the phytoplankton, the turnover time of the limitingnutrient supplied via herbivore grazing, and the time for herbivoresto filter a unit volume of water would be equivalent. They appearto be of the order of 5–9 days based on present methodologyand straightforward interpretation of its results. The rate measurements involved incubation of water samples forseveral hours in bottles. In the central N. Pacific the valueswere similar using bottles of different sizes. Addition of chelatorsdid not enhance the rates implying no poisoning of the planktonby heavy metal contaminants. The observed specific activitiesof ¹⁴C and ¹⁵N of the particulate matter in the rate measurementsare inconsistent with the notion of an active, rapidly growingand recycling microplankton food web within the incubation bottlesand support the idea that phytoplankton are growing slowly.     

Phytoplankton photosynthesis in the subsurface chlorophyll-maximum layer of the tropical North Pacific Ocean

Maximum quantum yield (φ_m^B) and maximum photosynthetic rate (P_m^B) of light-saturation curves of phytoplankton photosynthesis were determined for nannoplankton (< 20 μm) and netplankton (>20 μm) from the subsurface chlorophyll-maximum layer at 14 stations in the tropical North Pacific Ocean in the spring of 1976. The maximum quantum yield $\text{(φ}{}_{\mathrm{m}}{}^{\mathrm{B}}\text{±}\text{s.e.}\text{)}$ was significantly higher for nannoplankton (0.056 ± 0.006 moles CO₂·Einstein^?1 absorbed) than netplankton (0.039 ± 0.002 moles CO₂·Einstein^?1 absorbed). The importance of nannoplankton in the maximum photosynthetic rate (P_m^B) appears to be less consistent. At least 60% of the theoretical maximum quantum yield (0.12 moles CO₂·Einstein^?1 absorbed) was probably incorporated into the particulate fraction at the subsurface chlorophyll-maximum layer.     

Vertical distribution and phylogenetic composition of bacteria in the Eastern Tropical North Pacific Ocean

The vertical community structure of bacteria along a depth profile in the Eastern Tropical North Pacific Ocean (13 degrees N, 104 degrees W) was studied by flow cytometry measurement and 16S rRNA gene clone libraries analysis. Picoeukaryotes and Synechococcus peaked at 30 m and decreased sharply below 50 m, while Prochlorococcus peaked at both 30 and 100 m layers and disappeared below 200 m. Heterotrophic bacteria peaked above shallow thermocline and decreased along the depth profile. Sequences of total 322 clones from four clone libraries (10, 100, 1000, and 3000 m) clustered into nine major lineages. gamma-Proteobacteria dominated all the depths and occupied almost the whole bacterial community at the 3000 m. alpha-Proteobacteria was abundant throughout the water column except near the sea bottom, and delta-Proteobacteria peaked at the 1000 m depth. Cyanobacteria were primarily limited to the photic zone, and the genetic diversity of Prochlorococcus showed a good correlation with niche adaptation. The appearance of the Cytophaga-Flexibacter-Bacteroides (CFB) group did not show a clear relationship with depth. Actinobacteria were found both in the photic zone and in deep water. Planctomyetes, Acidobacteria, and Verrucomicrobia were present as minor groups and more dominant in the deeper layers of water.     

Anaerobic Respiration on Tellurate and Other Metalloids in Bacteria from Hydrothermal Vent Fields in the Eastern Pacific Ocean

This paper reports the discovery of anaerobic respiration on tellurate by bacteria isolated from deep ocean (1,543 to 1,791 m) hydrothermal vent worms. The first evidence for selenite- and vanadate-respiring bacteria from deep ocean hydrothermal vents is also presented. Enumeration of the anaerobic metal(loid)-resistant microbial community associated with hydrothermal vent animals indicates that a greater proportion of the bacterial community associated with certain vent fauna resists and reduces metal(loid)s anaerobically than aerobically, suggesting that anaerobic metal(loid) respiration might be an important process in bacteria that are symbiotic with vent fauna. Isolates from Axial Volcano and Explorer Ridge were tested for their ability to reduce tellurate, selenite, metavanadate, or orthovanadate in the absence of alternate electron acceptors. In the presence of metal(loid)s, strains showed an ability to grow and produce ATP, whereas in the absence of metal(loid)s, no growth or ATP production was observed. The protonophore carbonyl cyanide m-chlorophenylhydrazone depressed metal(loid) reduction. Anaerobic tellurate respiration will be a significant component in describing biogeochemical cycling of Te at hydrothermal vents.     

Biogeography of the Tropical Eastern Pacific: distribution and phylogeny of chaenopsid fishes

The Tropical Eastern Pacific Biogeographic Region (TEP) is delimited by steep thermal gradients to the north and south, by a wide expanse of open ocean (the East Pacific Barrier) to the west, and by the Central American land mass to the east. Four provinces within the TEP have been recognized based on the distribution of rocky shore fishes and marine invertebrates: the Cortez, Mexican, Panamic, and Galápagos Provinces. For rocky shore fishes, hypothesized barriers between these provinces are areas lacking rocky outcroppings, specifically the Central American Gap between the Panamic and Mexican Provinces, the Sinaloan Gap between the Mexican and Cortez Provinces, and the Pelagic Gap between the mainland and the Islas Galápagos. The occurrence of 33 chaenopsid fish species within these provinces, as well as other oceanic islands or archipelagos in the TEP (Isla de Malpelo, Isla del Coco, and Islas Revillagigedo) were tallied based on literature records and observations of museum specimens. Chaenopsid distributions within the TEP support these hypothesized provinces and their intervening gaps. Twenty‐one species (64% of the TEP chaenopsid fauna) are restricted to a single mainland province or one of the oceanic islands or archipelagos. Of the mainland provinces, the Cortez and Panamic exhibit similar levels of endemism (50%), but the Mexican Province has only one endemic (10%). Of the remaining chaenopsids in the Mexican Province, three are widespread, occurring in all three mainland provinces, four are shared only with the Cortez Province, and two are shared only with the Panamic Province. Within the TEP, the Pelagic Gap is the most effective (crossed by only 3 of 33 species adjacent to it), followed by the Central American Gap (crossed by 5 of 21 species), and the Sinaloan Gap (crossed by 7 of 17 species). Only one species, Chaenopsis alepidota, which is found off southern California and in the Cortez Province, crosses a barrier delimiting the TEP. Species‐level phylogenetic hypotheses for the Chaenopsidae imply exclusively allopatric speciation for these fishes in the TEP. Of the barriers delimiting the TEP, the most important in the recent evolution of chaenopsids is the Isthmian Barrier which is implicated in six speciation events. Within the TEP, the Central American Gap and Sinaloan Gap are each implicated in three speciation events, while the Pelagic Gap is implicated in three speciation events of island endemics from mainland populations and one inter‐island speciation event.     

Biogeography of Southern Ocean prokaryotes: a comparison of the Indian and Pacific sectors

We investigated the Southern Ocean (SO) prokaryote community structure via zero-radius operational taxonomic unit (zOTU) libraries generated from 16S rRNA gene sequencing of 223 full water column profiles. Samples reveal the prokaryote diversity trend between discrete water masses across multiple depths and latitudes in Indian (71–99°E, summer) and Pacific (170–174°W, autumn-winter) sectors of the SO. At higher taxonomic levels (phylum-family) we observed water masses to harbour distinct communities across both sectors, but observed sectorial variations at lower taxonomic levels (genus-zOTU) and relative abundance shifts for key taxa such as Flavobacteria, SAR324/Marinimicrobia, Nitrosopumilus and Nitrosopelagicus at both epi- and bathy-abyssopelagic water masses. Common surface bacteria were abundant in several deep-water masses and vice-versa suggesting connectivity between surface and deep-water microbial assemblages. Bacteria from same-sector Antarctic Bottom Water samples showed patchy, high beta-diversity which did not correlate well with measured environmental parameters or geographical distance. Unconventional depth distribution patterns were observed for key archaeal groups: Crenarchaeota was found across all depths in the water column and persistent high relative abundances of common epipelagic archaeon Nitrosopelagicus was observed in deep-water masses. Our findings reveal substantial regional variability of SO prokaryote assemblages that we argue should be considered in wide-scale SO ecosystem microbial modelling.     

High Fungal Diversity and Abundance Recovered in the Deep-Sea Sediments of the Pacific Ocean

Knowledge about the presence and ecological significance of bacteria and archaea in the deep-sea environments has been well recognized, but the eukaryotic microorganisms, such as fungi, have rarely been reported. The present study investigated the composition and abundance of fungal community in the deep-sea sediments of the Pacific Ocean. In this study, a total of 1,947 internal transcribed spacer (ITS) regions of fungal rRNA gene clones were recovered from five sediment samples at the Pacific Ocean (water depths ranging from 5,017 to 6,986 m) using three different PCR primer sets. There were 16, 17, and 15 different operational taxonomic units (OTUs) identified from fungal-universal, Ascomycota-, and Basidiomycota-specific clone libraries, respectively. Majority of the recovered sequences belonged to diverse phylotypes of Ascomycota (25 phylotypes) and Basidiomycota (18 phylotypes). The multiple primer approach totally recovered 27 phylotypes which showed low similarities (≤97 %) with available fungal sequences in the GenBank, suggesting possible new fungal taxa occurring in the deep-sea environments or belonging to taxa not represented in the GenBank. Our results also recovered high fungal LSU rRNA gene copy numbers (3.52?×?10⁶ to 5.23?×?10⁷copies/g wet sediment) from the Pacific Ocean sediment samples, suggesting that the fungi might be involved in important ecological functions in the deep-sea environments.     

Distribution and movements of fin whales in the North Pacific Ocean

• 1 We summarize fin whale Balaenoptera physalus catch statistics, sighting data, mark recoveries and acoustics data. The annual cycle of most populations of fin whales had been thought to entail regular migrations between high‐latitude summer feeding grounds and lower‐latitude winter grounds. Here we present evidence of more complex and varied movement patterns.
• 2 During summer, fin whales range from the Chukchi Sea south to 35 °N on the Sanriku coast of Honshu, to the Subarctic Boundary (ca. 42 °N) in the western and central Pacific, and to 32 °N off the coast of California. Catches show concentrations in seven areas which we refer to as ‘grounds’, representing productive feeding areas.
• 3 During winter months, whales have been documented over a wide area from 60 °N south to 23 °N. Coastal whalers took them regularly in all winter months around Korea and Japan and they have been seen regularly in winter off southern California and northern Baja California. There are also numerous fin whale sightings and acoustic detections north of 40 °N during winter months. Calves are born during the winter, but there is little evidence for distinct calving areas.
• 4 Whales implanted with Discovery‐type marks were killed in whaling operations, and location data from 198 marked whales demonstrate local site fidelity, consistent movements within and between the main summer grounds and long migrations from low‐latitude winter grounds to high‐latitude summer grounds.
• 5 The distributional data agree with immunogenetic and marking findings which suggest that the migratory population segregates into at least two demes with separate winter mating grounds: a western ground off the coast of Asia and an eastern one off the American coast. Members of the two demes probably mingle in the Bering Sea/Aleutian Islands area.
• 6 Prior research had suggested that there were at least two non‐migratory stocks of fin whale: one in the East China Sea and another in the Gulf of California. There is equivocal evidence for the existence of additional non‐migratory groups in the Sanriku‐Hokkaido area off Japan and possibly the northern Sea of Japan, but this is based on small sample sizes.

     

Bacterioplankton nutrient metabolism in the Eastern Tropical North Pacific

Bacterioplankton nutrient metabolism in the Eastern Tropical North Pacific (ETNP) was assessed using specific activities of intracellular nitrogen (N) assimilation enzymes and hydrolytic ectoenzymes during amendment experiments, mesocosms, and diel studies of in situ rates. Glutamine synthetase (GS) and assimilatory nitrate reductase (ANR) were used to investigate N bioavailability, alkaline phosphatase (AP) to assess phosphorous (P) bioavailability and β-glucosidase (β-Glu) to detect shifts in the use of labile dissolved organic carbon (DOC). Conditions regulating activity of each enzyme were tested using incubations of < 0.6 mm size-fractionated seawater amended with different combinations of N, P, and DOC as glucose. Overall, N-deficiency was indicated by pronounced growth stimulation and repression of GS and ANR activity in incubations amended with dissolved free amino acid and ammonium. Phosphate and glucose amendments produced little or no growth stimulation, but did influence activity of all enzymes measured. Enzyme activities of bacterioplankton in mesocosms of whole plankton indicated enhanced N-deficiency and glucoside hydrolysis when the plankton community was released from any P-deficiency. Spatially, enzyme activity of bacterioplankton during two diel studies (at one slope and one open-ocean station) suggested greater N-deficiency at surface depths than within the chlorophyll maximum where activity of AP and b-Glu was often greatest. There was also greater GS and ANR activity at the open-ocean station, which had lower concentrations of dissolved inorganic N (DIN) relative to soluble reactive P (SRP), than along the continental slope of Mexico. These data suggest that bacterioplankton in surface waters of the ETNP require a large flux of DOC to drive N-deficiency; whereas, bacterioplankton deeper in the chlorophyll maximum depend on hydrolysis of complex DOC and DOP to meet their carbon demand in the presence of elevated nutrients with a low DIN:SRP ratio.     

Egg brooding by deep-sea octopuses in the North Pacific Ocean

Videotapes made from the submersible Alvin on Baby Bare, a 2600-m-deep North Pacific basalt outcrop, and at two other deep-sea localities document that octopuses of the genera Graneledone and Benthoctopus attach their eggs to hard substrate and apparently brood them through development. The behavior of brooding females was generally similar to that of shallow-water octopuses, but the genera showed apparent differences. In addition to the high density of brooding females observed at Baby Bare, which may relate to the increased availability of exposed hard substrates for egg attachment and of prey, females are suggested to increasingly associate with hard substrates as they mature. The biology of Baby Bare may seem unduly unique because the outcrop is isolated on a sedimented plain and is among the few exposures of hard substrate other than hydrothermal vents that have been explored by submersible. On the sediment-covered ocean floor, the availability of hard substrate may strongly affect the distribution of brooding octopuses. The size and shape of boreholes in 19 of over 400 thyasirid clam shells collected from Baby Bare support the hypothesis that octopuses had preyed upon the clams.     

Spatial Segregation in Eastern North Pacific Skate Assemblages

Skates (Rajiformes: Rajoidei) are common mesopredators in marine benthic communities. The spatial associations of individual species and the structure of assemblages are of considerable importance for effective monitoring and management of exploited skate populations. This study investigated the spatial associations of eastern North Pacific (ENP) skates in continental shelf and upper continental slope waters of two regions: central California and the western Gulf of Alaska. Long-term survey data were analyzed using GIS/spatial analysis techniques and regression models to determine distribution (by depth, temperature, and latitude/longitude) and relative abundance of the dominant species in each region. Submersible video data were incorporated for California to facilitate habitat association analysis. We addressed three main questions: 1) Are there regions of differential importance to skates?, 2) Are ENP skate assemblages spatially segregated?, and 3) When skates co-occur, do they differ in size? Skate populations were highly clustered in both regions, on scales of 10s of kilometers; however, high-density regions (i.e., hot spots) were segregated among species. Skate densities and frequencies of occurrence were substantially lower in Alaska as compared to California. Although skates are generally found on soft sediment habitats, Raja rhina exhibited the strongest association with mixed substrates, and R. stellulata catches were greatest on rocky reefs. Size segregation was evident in regions where species overlapped substantially in geographic and depth distribution (e.g., R. rhina and Bathyraja kincaidii off California; B. aleutica and B. interrupta in the Gulf of Alaska). Spatial niche differentiation in skates appears to be more pronounced than previously reported.     

Aphotic N2 Fixation in the Eastern Tropical South Pacific Ocean

We examined rates of N₂ fixation from the surface to 2000 m depth in the Eastern Tropical South Pacific (ETSP) during El Niño (2010) and La Niña (2011). Replicated vertical profiles performed under oxygen-free conditions show that N₂ fixation takes place both in euphotic and aphotic waters, with rates reaching 155 to 509 µmol N m⁻² d⁻¹ in 2010 and 24±14 to 118±87 µmol N m⁻² d⁻¹ in 2011. In the aphotic layers, volumetric N₂ fixation rates were relatively low (<1.00 nmol N L⁻¹ d⁻¹), but when integrated over the whole aphotic layer, they accounted for 87–90% of total rates (euphotic+aphotic) for the two cruises. Phylogenetic studies performed in microcosms experiments confirm the presence of diazotrophs in the deep waters of the Oxygen Minimum Zone (OMZ), which were comprised of non-cyanobacterial diazotrophs affiliated with nifH clusters 1K (predominantly comprised of α-proteobacteria), 1G (predominantly comprised of γ-proteobacteria), and 3 (sulfate reducing genera of the δ-proteobacteria and Clostridium spp., Vibrio spp.). Organic and inorganic nutrient addition bioassays revealed that amino acids significantly stimulated N₂ fixation in the core of the OMZ at all stations tested and as did simple carbohydrates at stations located nearest the coast of Peru/Chile. The episodic supply of these substrates from upper layers are hypothesized to explain the observed variability of N₂ fixation in the ETSP.     

Plankton Biogeography of the Indian Ocean

Knowledge of plankton biogeography for the Indian and adjacent seas is necessary for an understanding of the regional characteristics and changes in the plankton composition. Some of the plankters will serve as good biogeographical indicator species. Some phytoplankters and tintinnids among zooplankton are promising for use as such labels for the presence of different waters in the Porto Novo (Coromandel coast, Bay of Bengal) region. A biogeographical classification of the local phytoplankton and peculiarities in the distribution of some rare tintinnids occurring there are discussed.     

Age and growth of albacore Thunnus alalunga in the North Pacific Ocean

The age and growth of North Pacific albacore Thunnus alalunga were investigated using obliquely sectioned sagittal otoliths from samples of 126 females and 148 males. Otolith edge analysis indicated that the identified annulus in a sagittal otolith is primarily formed during the period from September to February. The assessments of the fish age at first annulus formation indicated that the first annulus represents an age of <1 year. This study presents an age estimate (0·75 years) for the formation of the first annulus. The oldest fish ages observed in this study were 10 years for females and 14 years for males. The von Bertalanffy growth parameters of females estimated were L(∞) = 103·5 cm in fork length (L(F) ), K = 0·340 year(-1) and t(0) = -0·53 years, and the parameters of males were L(∞) = 114·0 cm, K = 0·253 year(-1) and t(0) = -1·01 years. Sexual size dimorphism between males and females seemed to occur after reaching sexual maturity. The coefficients of the power function for expressing the L(F) -mass relationship obtained from sex-pooled data were a = 2·964 × 10(-5) and b = 2·928.     

Biogeography of Photosynthetic Light-Harvesting Genes in Marine Phytoplankton

Background

Photosynthetic light-harvesting proteins are the mechanism by which energy enters the marine ecosystem. The dominant prokaryotic photoautotrophs are the cyanobacterial genera Prochlorococcus and Synechococcus that are defined by two distinct light-harvesting systems, chlorophyll-bound protein complexes or phycobilin-bound protein complexes, respectively. Here, we use the Global Ocean Sampling (GOS) Project as a unique and powerful tool to analyze the environmental diversity of photosynthetic light-harvesting genes in relation to available metadata including geographical location and physical and chemical environmental parameters.

Methods

All light-harvesting gene fragments and their metadata were obtained from the GOS database, aligned using ClustalX and classified phylogenetically. Each sequence has a name indicative of its geographic location; subsequent biogeographical analysis was performed by correlating light-harvesting gene budgets for each GOS station with surface chlorophyll concentration.

Conclusion/Significance

Using the GOS data, we have mapped the biogeography of light-harvesting genes in marine cyanobacteria on ocean-basin scales and show that an environmental gradient exists in which chlorophyll concentration is correlated to diversity of light-harvesting systems. Three functionally distinct types of light-harvesting genes are defined: (1) the phycobilisome (PBS) genes of Synechococcus; (2) the pcb genes of Prochlorococcus; and (3) the iron-stress-induced (isiA) genes present in some marine Synechococcus. At low chlorophyll concentrations, where nutrients are limited, the Pcb-type light-harvesting system shows greater genetic diversity; whereas at high chlorophyll concentrations, where nutrients are abundant, the PBS-type light-harvesting system shows higher genetic diversity. We interpret this as an environmental selection of specific photosynthetic strategy. Importantly, the unique light-harvesting system isiA is found in the iron-limited, high-nutrient low-chlorophyll region of the equatorial Pacific. This observation demonstrates the ecological importance of isiA genes in enabling marine Synechococcus to acclimate to iron limitation and suggests that the presence of this gene can be a natural biomarker for iron limitation in oceanic environments.