Spring phytoplankton assemblages in the Southern Ocean between Australia and Antarctica

Variations of phytoplankton assemblages were studied in November–December 2001, in surface waters of the Southern Ocean along a transect between the Sub-Antarctic Zone (SAZ) and the Seasonal Ice Zone (SIZ; 46.9°–64.9°S; 142°–143°E; CLIVAR-SR3 cruise). Two regions had characteristic but different phytoplankton assemblages. Nanoflagellates(<20 μm) and pico-plankton (∼2 μm) occurred in similar concentrations along the transect, but were dominant in the SAZ, Sub-Antarctic Front (SAF), Polar Front Zone (PFZ) and the Inter-Polar Front Zone (IPFZ), (46.9°–56.9°S). Along the entire transect their average cell numbers in the upper 70 m of water column, varied from 3 × 10⁵ to 1.1 × 10⁶ cells l⁻¹. Larger cells (>20 μm), diatoms and dinoflagellates, were more abundant in the Antarctic Zone-South (AZ-S) and the SIZ, (60.9°–64.9°S). In AZ-S and SIZ diatoms ranged between 2.7 × 10⁵ and 1.2 × 10⁶ cells l⁻¹, dinoflagellates from 3.1 × 10⁴ to 1.02 × 10⁵ cells l⁻¹. A diatom bloom was in progress in the AZ-S showing a peak of 1.8 × 10⁶ cells l⁻¹. Diatoms were dominated by Pseudo-nitzschia spp., Fragilariopsis spp., and Chaetoceros spp. Pseudo-nitzschia spp. outnumbered other diatoms in the AZ-S. Fragilaropsis spp. were most numerous in the SIZ. Dinoflagellates contained autotrophs (e.g. Prorocentrum) and heterotrophs (Gyrodinium/Gymnodinium, Protoperidinium). Diatoms and dinoflagellates contributed most to the cellular carbon: 11–25 and 17–124 μg C l⁻¹, respectively. Small cells dominated in the northern region characterized by the lowest N-uptake and new production of the transect. Larger diatom cells were prevalent in the southern area with higher values of N-uptake and new production. Diatom and nanoflagellate cellular carbon contents were highly correlated with one another, with primary production, and productivity related parameters. They contributed up to 75% to the total autotrophic C biomass. Diatom carbon content was significantly correlated to nitrate uptake and particle export, but not to ammonium uptake, while flagellate carbon was well correlated to ammonium uptake, but not to export. Diatoms have contributed highly to particle export along the latitudinal transect, while flagellates played a minor role in the export.     

Sea surface distribution of coccolithophores in the eastern Pacific sector of the Southern Ocean (Bellingshausen and Amundsen Seas) during the late austral summer of 2001

Horizontal distributions of coccolithophores were observed in sea surface water samples collected on the RV Polarstern between 27 February and 10 April, 2001, in the Pacific sector of the Southern Ocean (Bellingshausen and Amundsen Seas). These samples were analyzed to gain information about the distribution of coccolithophores in relation to the oceanic fronts of the Southern Ocean. A total of fifteen species of coccolithophores were identified, showing cell abundances of up to 67 × 10³ cells/l down to 63°S. Emiliania huxleyi was the most abundant taxon, always accounting for more than 85% of the assemblage. The second most abundant species was Calcidiscus leptoporus, with values lower than 7%. Cell density increases significantly in both the Subantarctic and Polar Fronts (155 and 151 × 10³ cells/l, respectively), decreasing abruptly in the intervening Polar Frontal Zone and to the south of the Polar Front. Although temperature at high latitudes is the main factor controlling the biogeographical distribution of coccolithophores, at the regional level (Southern Ocean) the frontal systems, and consequently nutrient distribution, play a crucial role.     

Effects of Asian dust input on eukaryotic phytoplankton community structure in the open areas in the Northwestern Pacific Ocean

To study the effects of aerosol particulates originated from Asian dust on the growth of eukaryotic phytoplankton in the oligotrophic open ocean, we conducted deck-board incubation experiments in the oligotrophic region of Northwestern Pacific Ocean (NWPO). Our results showed that when dust was added at a concentration of 2?mg/L (Dust-2), the NO₃^–N concentration increased by 3.2 fold, and chlorophyll a (chl a) concentration of nano- and micro-eukaryotic phytoplankton increased, while that of pico-eukaryotic phytoplankton did not change significantly. The microphytoplankton abundance increased but the species number decreased in Dust-2. Community structure of eukaryotic microphytoplankton also changed after dust addition. The abundance of diatoms in Dust-2 (23,072 cells/L) was 4.0 fold of that in the control (5750 cells/L), and 2.4 fold of that in Dust-1 (0.2?mg/L of dust addition) (9425 cells/L) at the 7th day of incubation. Abundance of dinoflagellates decreased in Dust-2, being 42.2% that of the control at the 7th day of the incubation. Effects of dust addition on the growth of phytoplankton differed among the dominant genera: growth of Pseudo-nitzschia and Chaetoceros were promoted while that of Prorocentrum was inhibited, and growth of Thalassiosira, Heterocapsa and Scrippsiella was not influenced significantly. The growth of nano- and pico-eukaryotic phytoplankton was promoted, with the cell abundance in Dust-2 2.4 fold of that in control. Our results indicated that Asian dust-originated aerosol particulates could provide nutrients to the oligotrophic NWPO, increase the marine productivity in the area, and alter the eukaryotic phytoplankton community structure.     

Dispersal and speciation of skinks among archipelagos in the tropical Pacific Ocean

Summary We examined the potential effects of geography on the distribution and speciation of skinks on tropical Pacific archipelagos. The entire tropical Pacific skink fauna was divided into continental (found also in continental areas), Pacific (endemic to the study area but found within more than one archipelago) and endemic (found within only one archipelago) species categories. The number and proportion of skinks within each species category were determined for each of the 27 archipelagos in the study area. Nine geographic variables reflecting archipelago size, isolation and elevation were estimated for each archipelago. Principal components analysis was used to reduce the nine variables to three uncorrelated composite variables that were interpreted as representing archipelago size, isolation and elevation. Numbers and proportions of skinks in each category within an archipelago were related to the composite geographic variables using multiple linear regression analysis. Archipelago size and isolation were important predictors of both skink diversity and endemism. Results were then compared to diversity and endemism of birds within the study area. Skinks showed an archipelago-wide level of endemism similar to that of birds. On an archipelago by archipelago basis, however, large differences between birds and skinks were evident. In particular, the New Caledonia skink fauna was much more endemic than that of birds. The bird faunas of Hawaii and the Marquesas were nearly completely endemic, while no endemic skinks occurred in these two archipelagos. These differences presumably reflect the relative dispersal powers of skinks and birds and, consequently, rates of colonization and speciation. Differences may also be due partly to morphological conservatism among isolated skink populations and the occurrence of cryptic species that have not yet been identified as separate species. The discovery of such cryptic species, however, is unlikely to increase the endemic skink fauna of Hawaii and other distant archipelagos to a level commensurate with that of birds. Differences in endemism between skinks and birds may also be due to unknown local ecological interactions.     

Effects of nutrients and dissolved organic matter on the response of phytoplankton to ultraviolet radiation: experimental comparison in spring versus summer

The effects of nutrients and dissolved organic matter (DOM) on the response of phytoplankton community structure to ultraviolet radiation (UVR) was studied using natural phytoplankton assemblages from Lake Giles (Northeastern Pennsylvania), a temperate, oligotrophic, highly UVR-transparent lake. Microcosm experiments were conducted in 1-l bags in the spring and summer. A factorial design was used, with two UVR treatments (ambient and reduced), two nutrient treatments (control with no nutrients added, and nitrogen and phosphorus addition together), and two DOM treatments (control of 1 mg l⁻¹ and doubled). In April, UVR affected the overall phytoplankton community structure, causing a shift in the dominant species. Significant interactive effects of UVR × nutrients and UVR × DOM were found on total phytoplankton biovolumes. In July, all taxa responded positively to the N + P addition, and were affected differentially by the UVR treatments. The initial communities varied in April and July, but Synura sp. and Chroomonas sp. were present in both seasons. Synura sp. responded positively to the addition of DOM in April and the reduction of UVR in July. Chroomonas sp. responded positively to the reduction of UVR in April and the addition of nutrients in July. The differential sensitivity of these two species suggests that changing environmental factors between spring and summer promoted differences in the relative importance of UVR in changing phytoplankton community structure. Handling editor: Luigi Naselli-Flores     

In situ and remote-sensed chlorophyll fluorescence as indicator of the physiological state of phytoplankton near the Isles Kerguelen (Southern Ocean)

Shipboard and remote-sensed Chlorophyll fluorescence were determined in the natural phytoplankton assemblage above the iron-enriched Kerguelen Plateau and the adjacent high-nutrient, low-Chlorophyll open Southern Ocean. The variance between fluorescence yield and photosynthetic efficiency was determined in combination with Chlorophyll a concentrations, irradiance and phytoplankton species distribution. A co-variance between the fluorescence measurements would allow the refinement of remote-sensing primary production algorithms. Distinct differences were found in photosynthetic efficiency and water-leaving fluorescence, with relatively high values for the Kerguelen Plateau and low values in the open ocean, reflecting the differences in Chlorophyll a concentrations. The co-variance of the fluorescence properties suggested that remote-sensed fluorescence measurements could be used to infer differences in the physiological state of the phytoplankton, hence primary production. Fluorescence yield, however, did not show the differences in the research area, most likely due to the low signal and the diurnal variation in water-leaving fluorescence.     

Production of viruses during a spring phytoplankton bloom in the South Pacific Ocean near of New Zealand

Lagrangian studies of virus activity in pelagic environments over extended temporal scales are rare. To address this, viruses and bacteria were examined during the course of a natural phytoplankton bloom in the pelagic South Pacific Ocean east of New Zealand. Daily samples were collected in a mesoscale eddy from year days 263-278 (September 19th-October 4th, 2008). The productive bloom transitioned from a diatom to a pico- and nanoplankton-dominated system, resulting in chlorophyll a concentrations up to 2.43?μg?L(-1) . Virus abundances fluctuated c.?10-fold (1.8?×?10(10) -1.3?×?10(11) L(-1) ) over 16?days. The production rates of virus particles were high compared with those reported in other marine systems, ranging from 1.4?×?10(10) to 2.1?×?10(11) L(-1) day(-1) . Our observations suggest viruses contributed significantly to the mortality of bacteria throughout the bloom, with 19-216% of the bacterial standing stock being lysed daily. This mortality released nutrient elements (N, Fe) that likely helped sustain the bloom through the sampling period. Parametric analyses found significant correlations with both biotic (e.g. potential host abundances) and abiotic parameters (e.g. nutrient concentrations, temperature). These observations demonstrate that viruses may be critical in the extended maintenance of regeneration-driven biological production.     

New approach in studies of microalgal cell lysis

A new approach to the studies of the microalgal cell lysis by utilizing a combination of two complementary methods is presented. Delayed fluorescence (DF) is a measure of the living algal biomass, detecting only cells with active photosynthesis. Thermal lens spectrometry (TLS) detects the total pigment amount released from lysed cells. Both methods select for photosynthetic organisms, reducing possible background from other sources (e.g. heterotrophic bacteria, zooplankton, and abiotic substances). The DF/TLS method was tested with a laboratory Skeletonema costatum culture exposed to a geometric dilution series of the lysing factor poly- APS. The exposure resulted in similar EC₅0 values for DF intensity, TLS and dissolved esterase activity of 0.8±0.2, 1.77±0.35, and 1.25±0.1 mg poly-APS l⁻¹, respectively. The combined DF/TLS method enabled a rapid evaluation of the living vs. dead cells without any sample pretreatment or manipulation.     

Extinction and replacement in the Indo-West Pacific Ocean

Aim To provide evidence suggesting the existence of a dynamic system of extinction and replacement. Location The Indo-West Pacific Ocean. Methods Utilization of species distribution patterns produced by detailed systematic works. Results The distribution patterns appear to suggest a sequence of events that is consistent with the centre of origin hypothesis. The East Indies centre is considered to operate according to the centrifugal speciation model. Main conclusions Disjunct patterns appear to have originated in the East Indies and spread out from there. In cases where the vacated area is occupied by a sibling species, competitive (sympatric) speciation may have occurred. Over time the extinction pattern will gradually extend outward until it is played out among small populations far from their centre of origin     

Differences of the protozoan biomass and grazing during spring and summer in the Indian sector of the Southern Ocean

The dynamics of protozoa were investigated during two cruises in the Indian sector of the Southern Ocean: the early spring ANTARES 3 cruise (28 September to 8 November 1995) and the late summer ANTARES 2 cruise (6 February to 8 March 1994). Biomass and feeding activity of protozoa were measured as well as the biomass of their potential prey – bacteria and phototrophic flagellates – along the 62°E meridian. The sampling grid extended from the Polar Frontal region to the Coastal and Continental Shelf Zone in late summer and to the ice edge in spring, crossing the Antarctic Divergence. Protozoan biomass, although low in absolute terms, contributed 30% and 20% to the total microbial biomass (bacteria, phytoplankton and protozoa) in early spring and late summer, respectively. Nanoprotozoa dominated the total protozoan biomass. The geographical and seasonal distribution of protozoan biomass was correlated with that of phototrophic flagellates. However, bacterial and phototrophic flagellate biomass were inversely correlated. Phototrophic flagellates dominated in the Sea Ice Zone whereas bacteria were predominant at the end of summer in the Polar Frontal region and Coastal and Continental Shelf Zone. Furthermore, bacteria were the most important component of the microbial community (57% of the total microbial biomass) in late summer. Phototrophic flagellates were ingested by both nano-and microprotozoa. In contrast, bacteria were only ingested by nanoprotozoa. Protozoa controlled up to 90% of the daily bacterial production over the period examined. The spring daily protozoan ingestion controlled more than 100% of daily phototrophic flagellate production. This control was less strong at the end of summer when protozoan grazing controlled 42% of the daily phototrophic flagellate production. Accepted: 30 October 1999     

Late Holocene variability in pelagic fish scales and dinoflagellate cysts along the west coast of Vancouver Island, NE Pacific Ocean

Fish stocks and dinoflagellates are essential components of the marine food chain. Sediment cores from a predominantly anoxic basin in Effingham Inlet, Vancouver Island, British Columbia, archive a late Holocene (500–5300 years BP) record of paleoproductivity in the North American Coastal Upwelling Domain (CUD). We present evidence that late Holocene changes in the dinoflagellate cyst assemblages, sedimentary record, and fish stocks in the northeastern Pacific Ocean fluctuated, at least partially, in accordance with regional and global climate cycles.Principal components analysis (PCA), and trend, wavelet and spectral analyses were used to identify relationships, cycles and trends in sediment grey-scale values, and the abundances of fish scales and dinoflagellate cysts on centennial to millennial time scales. Most observed cycles fluctuated in intensity over time, particularly following transition of the regional climate to a higher rainfall phase that impacted coastal oceanic dynamics 3400 ± 150 years ago. Correlation of the marine paleoproductivity records observed in Effingham Inlet with solar influenced climate proxy cycles observed in the North Atlantic region indicates that solar forcing at different scales might have influenced the climate in the northeast Pacific as well. In particular an 1100- to 1400-year cycle in regional climate is well represented in the fish productivity proxy and sedimentological record. It was also observed that colder water, high-productivity, Selenopemphix nephroides and anchovy-dominated “Anchovy Regime” ecosystems alternate with warmer water, herring-dominated “Herring Regime” ecosystems at millennial time scales. The fish scale record preserved in Effingham Inlet indicates that the NE Pacific is now in transition from an ‘anchovy-’ to a ‘herring’-dominated regime.     

Light utilization efficiency of size-fractionated phytoplankton in the subarctic Pacific, spring and summer 1999: high efficiency of large-sized diatom

Higher light utilization efficiency of the >10 µm fractionwas observed in the western subarctic Pacific in spring. Thehigher efficiency was possibly due to large-sized diatoms.     

Description of phytoplankton and nutrient in spring in the western North Atlantic Ocean

Abundance of diatoms in the northern Sargasso Sea in springhas been observed in a 4-year period between 1981 and 1985 andthen again in 1987. A theory is presented that if stratificationblocks get-through of deep nutrients to the surface, as in thesouthern Sargasso, then in-situ cycling via non-diatoms andammonia dominates. The theory is that if in-situ cycling doesnot dominate, as northward, then stratification does not blockget-through of deep nutrients to the surface, which, in combinationwith the marked growth capacity of diatoms, produce an abundanceof cells. This abundance consumes nitrate, phosphate and silicateat the surface and ultimately this nutrient-depleted water arrivesin the southern Sargasso via the gyre circulation. Thus thenutrient-depleted water of the western Atlantic gyre is consideredto be primarily achieved only by diatoms, in spring and in thenorthern Sargasso. A transect during April 13–16, 1985,north from the Virgin Islands along 63°–64°W showsa great change in the diatom species plus Emiliania huxleyifrom negligible concentrations in the southern half of the sectionto large concentrations in the northern half. This unevennessof distribution can be rephrased by indicating that these speciesare narrow-niched with regard to nutrient availability. Theother species, coccolithophores and dinoflagellates, are evenlydistributed at low concentrations south-north (though some diminishslightly) and so they are broad-niched with respect to nutrient.High species diversity depends on broadness of nutrient nicheand equilibrium conditions of hydrographically stable water(i), implying that when non-equilibrium conditions occur (ii),when niche breadth is less (iii) or both (iv), then diversityis less marked, (i) is confirmed by broad-niched species southward,(ii) is confirmed by broad-niched species northward, (iii) isconfirmed by narrow-niched species southward and (iv) by narrow-nichedspecies northward.     

New species and new records of Callionymus from the Pacific Ocean (Teleostei: Callionymidae)

Recently obtained material of Callionymus from the Pacific Ocean includes: Callionymus marquesensis sp. n. (Marquesas Islands), characterized within the C. variegatus-group of the subgenus Calliurichthys by its preopercular spine formula of with an upcurved main tip and a bony keel between the points on the dorsal margin, and a spotted dorsal fin without filaments which is higher than the 1st D₂ ray in both sexes; C. pleurostictus Fricke, 1982, record from New Britain and Solomon Islands, with a discussion of competition with C. delicatulus; C.simplicicornis Valenciennes, 1837, record from Society Islands and Marquesas Islands, with a discussion of competition with C. marquesensis; C. obscurus sp. n. (Ambon, Indonesia) which is characterized within the subgenus Spini-capitichthys by its preopercular spine formula with an upcurved main tip and very small ventral serrae, its weakly armed head, and its 7 anal fin rays.     

Isolation by distance and vicariance drive genetic structure of a coral reef fish in the Pacific Ocean

We studied the genetic diversity of a coral reef fish species to investigate the origin of the differentiation. A total of 727 Acanthurus triostegus collected from 15 locations throughout the Pacific were analyzed for 20 polymorphic loci. The genetic structure showed limited internal disequilibrium within each population; 3.7% of the loci showed significant Hardy-Weinberg disequilibrium, mostly associated with Adh*, and we subsequently removed this locus from further analysis of geographic pattern. The genetic structure of A. triostegus throughout the tropical Pacific Ocean revealed a strong geographic pattern. Overall, there was significant population differentiation (multilocus F(ST) = 0.199), which was geographically structured according to bootstraps of neighbor-joining analysis on Nei's unbiased genetic distances and AMOVA analysis. The genetic structure revealed five geographic groups in the Pacific Ocean: western Pacific (Guam, Philippines, Palau, and Great Barrier Reef); central Pacific (Solomons, New Caledonia, and Fiji); and three groups made up of the eastern populations, namely Hawaiian Archipelago (north), Marquesas (equatorial), and southern French Polynesia (south) that incorporates Clipperton Island located in the northeastern Pacific. In addition, heterozygosity values were found to be geographically structured with higher values grouped within Polynesian and Clipperton populations, which exhibited lower population size. Finally, the genetic differentiation (F(ST)) was significantly correlated with geographic distance when populations from the Hawaiian and Marquesas archipelagos were separated from all the other locations. These results show that patterns of differentiation vary within the same species according to the spatial scale, with one group probably issued from vicariance, whereas the other followed a pattern of isolation by distance. The geographic pattern for A. triostegus emphasizes the diversity of the evolutionary processes that lead to the present genetic structure with some being more influential in certain areas or according to a particular spatial scale.     

Determination of the cell lytic properties of amphiphilic inhibitors of the cytosolic phospholipase A2 against human platelets by measuring the liberation of serotonin with high-performance liquid chromatography and fluorescence detection

A procedure for the determination of the influence of detergents on the cell integrity of human platelets by measuring the release of serotonin with high-performance liquid chromatography and fluorescence detection is presented. After exposure of the platelets to the test compounds the cells and cell fragments, respectively, are centrifuged off. In the supernatants the liberated serotonin is determined directly without a further sample clean up. The amphiphilic inhibitors of cytosolic phospholipase A2 (cPLA2), arachidonyltrifluoromethyl ketone (AACOCF3), palmityltrifluoromethyl ketone (PACOCF3) and methyl arachidonylfluorophosphonate (MAFP), and the polyoxyethylene detergent Brij 58 were investigated for their cell lytic properties with this method. All compounds lysed the platelets liberating serotonin at a concentration of 33 microM. AACOCF, and Brij 58 even caused cell lysis at lower concentrations.     

Distribution and identification of red yeasts in deep-sea environments around the northwest Pacific Ocean

We isolated 99 yeast strains, including 40 red yeasts, from benthic animals and sediments collected from the deep-sea floor in various areas in the northwest Pacific Ocean. Comparing the yeast isolates from animals and sediments collected from shallow locations, the proportion of red yeasts differed considerably, comprising 81.5% and 10.6% of the isolates from animals and sediments, respectively. All of the red yeast isolates belonged to the genera Rhodotorula and Sporobolomyces. On the basis of morphological and physiological characteristics, the isolates were identified as R. aurantiaca, R. glutinis, R. minuta and R. mucilaginosa of the genus Rhodotorula, and S. salmonicolor and S. shibatanus of the genus Sporobolomyces. Only R. glutinis and R. mucilaginosa were isolated from sediments. All of the others were isolated from animal sources. Phylogenetic analyses based on internal transcribed spacer (ITS) regions and 5.8S rRNA gene sequences allowed us to establish the precise taxonomic placement of each of the isolates and thereby investigate the intraspecific relationships among the isolates. Twenty-two strains identified as members of R. glutinis, which showed a wide distribution in the deep-sea, and five isolates identified as R. minuta, which were isolated only from benthic animals, showed substantial heterogeneity within the species. The isolates phenotypically identified as Sporobolomyces species and R. mucilaginosa phylogenetically occupied the placements corresponding to these species. Some strains assigned to known species on the basis of phenotypic features should be regarded as new species as suggested by the results of molecular analysis.     

Preliminary study of summer phytoplankton in coastal waters of the Gulf of Taranto (Ionian Sea)

Abstract

Phytoplankton abundances and species composition of coastal waters of the Gulf of Taranto (Ionian Sea) were investigated in July 1995. Abundances were lower than those of other Mediterranean coastal areas. Higher densities values were found in the central part of the studied area. In surface waters, small phytoflagellates (<10μm) and dinoflagellates, such as Heterocapsa niei and Oxytoxum variabile, were the dominant species. Diatoms, mainly belonging to the genus Chaetoceros, were also present.     

Planktonic foraminifera from the eastern Indian Ocean: distribution and ecology in relation to the Western Pacific Warm Pool (WPWP)

Faunal assemblages, principal component (PCA), canonical correspondence (CCA), and factor analysis are applied to planktonic foraminifera from 57 core-top samples from the eastern Indian Ocean. The foraminiferal lysocline occurs at 2400 m north of 15°S where carbonate dissolution is induced by the Java upwelling system, and occurs deeper south of 15°S where carbonate dissolution is characteristic of the oligotrophic regions in the Indian Ocean. Dissolution effects, the February standing stock at the time of collection of the plankton-tow material, and different production rates explain the different foraminiferal assemblages found between plankton-tow and core-top samples. Core-top samples are differentiated by PCA into four groups — Upwelling, Western Pacific Warm Pool (WPWP), Transitional, and Southern — that are related to environmental variables (temperature, salinity and nutrients); all environmental variables follow a strong latitudinal component as indicated by the CCA analysis. Similarly, three assemblages are recognized by factor analysis: Factor 1 (dominated by Globigerinoides sacculifer, G. ruber, Globigerinita glutinata and Globorotalia cultrata), factor 2 (dominated by Globigerina bulloides and Globorotalia inflata) and factor 3 (dominated by Neogloboquadrina dutertrei) explain more than 92% of the variance, and are related to sea-surface temperature, thermocline depth and nutrient levels. The seasonal influence of the Java upwelling system supplies nutrients, phyto- and zooplankton to the oligotrophic eastern Indian Ocean (factor 1). South of 24°S, a deep chlorophyll maximum, a deep euphotic zone, a deep thermocline, SSTs below 22°C, and brief upwelling pulses seem to explain factors 2 and 3. The ratio of G. sacculifer and N. dutertrei, two mutually excluding species, appears to indicate the southern boundary of the WPWP. This ratio is applied to core Fr10/95-11 to demonstrate past shifts of the southern boundary of the WPWP.