Ecological assessment data of the marine flora and fauna of Point Lookout in 2014 (PLEA)

In 2014, UniDive (The University of Queensland Underwater Club) conducted an ecological assessment of the Point Lookout Dive sites for comparison with similar surveys conducted in 2001. Involvement in the project was voluntary. Members of UniDive who were marine experts conducted training for other club members who had no, or limited, experience in identifying marine organisms and mapping habitats. Since the 2001 detailed baseline study, no similar seasonal survey has been conducted. The 2014 data is particularly important given that numerous changes have taken place in relation to the management of, and potential impacts on, these reef sites. In 2009, Moreton Bay Marine Park was re-zoned, and Flat Rock was converted to a marine national park zone (Green zone) with no fishing or anchoring. In 2012, four permanent moorings were installed at Flat Rock. Additionally, the entire area was exposed to the potential effects of the 2011 and 2013 Queensland floods, including flood plumes which carried large quantities of sediment into Moreton Bay and surrounding waters. The population of South East Queensland has increased from 2.49 million in 2001 to 3.18 million in 2011 (BITRE, 2013). This rapidly expanding coastal population has increased the frequency and intensity of both commercial and recreational activities around Point Lookout dive sites (EPA 2008). Methodology used for the PLEA project was based on the 2001 survey protocols, Reef Check Australia protocols and Coral Watch methods. This hybrid methodology was used to monitor substrate and benthos, invertebrates, fish, and reef health impacts. Additional analyses were conducted with georeferenced photo transects. The PLEA marine surveys were conducted over six weekends in 2014 totaling 535 dives and 376 hours underwater. Two training weekends (February and March) were attended by 44 divers, whilst biological surveys were conducted on seasonal weekends (February, May, July and October). Three reefs were surveyed, with two semi-permanent transects at Flat Rock, two at Shag Rock, and one at Manta Ray Bommie. Each transect was sampled once every survey weekend, with the transect tapes deployed at a depth of 10 m below chart datum. Fish populations were assessed using a visual census along 3 x 20 m transects. Each transect was 5 m wide (2.5 m either side of the transect tape), 5 m high and 20 m in length. Fish families and species were chosen that are commonly targeted by recreational or commercial fishers, or targeted by aquarium collectors, and that were easily identified by their body shape. Rare or otherwise unusual species were also recorded. Target invertebrate populations were assessed using visual census along 3 x 20 m transects. Each transect was 5 m wide (2.5 m either side of the transect tape) and 20 m in length. The diver surveying invertebrates conducted a 'U-shaped' search pattern, covering 2.5 m on either side of the transect tape. Target impacts were assessed using a visual census along the 3 x 20 m transects. Each transect was 5 m wide (2.5 m either side of the transect tape) and 20 m in length. The transect was surveyed via a 'U-shaped' search pattern, covering 2.5 m on either side of the transect tape. Substrate surveys were conducted using the point sampling method, enabling percentage cover of substrate types and benthic organisms to be calculated. The substrate or benthos under the transect line was identified at 0.5m intervals, with a 5m gap between each of the three 20m segments. Categories recorded included various growth forms of hard and soft coral, key species/growth forms of algae, other living organisms (i.e. sponges), recently killed coral, and, non-living substrate types (i.e. bare rock, sand, rubble, silt/clay).

Eocene to Quaternary benthic foraminifers from the Izu-Bonin Arc

The benthic foraminifer fauna at Sumisu Rift Sites 790 and 791 indicates that a deep open-ocean (>2300 m) or a basin with open-ocean access existed between 1.1 and 0.7 Ma at the time of the initiation of rifting. The appearance of a low- to medium-oxygen fauna (1600-2300 m) between 0.7 and 0.5 Ma suggests that the open-ocean access may have been terminated at this time because of the development of volcanoes and rift flank uplifts around the basin. The occurrence of low-oxygen faunas at 0.03 Ma suggests a secondary closing of the basin. The lower bathyal benthic faunas from lower Pliocene sediments of rift margin Site 788 suggest about 0.6-1.6 km of total basement uplift. This uplift may have led to the formation of the major hiatus between 2.3 and <0.3 Ma. The faunal changes of benthic foraminifers at Sites 792 and 793 in the forearc basin document a shallowing water depth from below the carbonate compensation depth (CCD) (about 3.5 km) in the late early Oligocene to the present depths of 1800 and 2975 m, respectively. These data suggest about 1 km of total basement uplift in the inner part of the forearc basin (Site 792) and about 0.6 km total basement subsidence in the central part of the forearc basin (Site 793) since about 31 Ma. The former uplift led to a thinner sediment accumulation (800 m) and the latter subsidence to a thicker sediment accumulation (1400 m) at these sites. Faunal changes of benthic foraminifers observed in Sites 782 and 786 sequences drilled at the outer-arc high document a deepening water depth from 1.3 to 2.1 km in late Eocene to the present depth of about 3 km. These data suggest about 1.1-1.9 and 1.3-2.1 km of total basement subsidence at Sites 786 and 782, respectively. These results indicate total basement uplift in the inner part of the Bonin arc-trench system since late Oligocene and total basement subsidence in the outer part of the system since late Eocene. The last occurrence (LO) of Stilostomella spp. and Pleurostomella spp. and the first occurrence (F0) of Bulimina aculeata d'Orbigny occurred consistently at 0.7 Ma at all three arc proximal sites (790,791, and 792). This fact is taken to suggest a change of water mass, from one originating from the central part of the ocean to that originating from ocean-margin areas at that time.

Stable Isotope composition of particulate organic matter and fauna in a deep Mediterranean cold-water coral bank

Cold-water corals (CWC) are frequently reported from deep sites with locally accelerated currents that enhance seabed food particle supply. Moreover, zooplankton likely account for ecologically important prey items, but their contribution to CWC diet remains unquantified. We investigated the benthic food web structure of the recently discovered Santa Maria di Leuca (SML) CWC province (300 to 1100 m depth) located in the oligotrophic northern Ionian Sea. We analyzed stable isotopes (delta13C and delta15N) of the main consumers (including ubiquitous CWC species) exhibiting different feeding strategies, zooplankton, suspended particulate organic matter (POM) and sedimented organic matter (SOM). Zooplankton and POM were collected 3 m above the coral colonies in order to assess their relative contributions to CWC diet. The delta15N of the scleractinians Desmophyllum dianthus, Madrepora oculata and Lophelia pertusa and the gorgonian Paramuricea cf. macrospinawere consistent with a diet mainly composed of zooplankton. The antipatharian Leiopathes glaberrima was more 15N- depletedthan other cnidarians, suggesting a lower contribution of zooplankton to its diet. Our delta13C data clearly indicate that the benthic food web of SML is exclusively fuelled by carbon of phytoplanktonic origin. Nevertheless, consumers feeding at the water sediment interface were more 13C-enriched than consumers feeding above the bottom (i.e. living corals and their epifauna). This pattern suggests that carbon is assimilated via 2 trophic pathways: relatively fresh phytoplanktonic production for 13C-depleted consumers and more decayed organic matter for 13C-enriched consumers. When the delta13C values of consumers were corrected for the influence of lipids (which are significantly 13C-depleted relative to other tissue components), our conclusions remained unchanged, except in the case of L. glaberrima which could assimilate a mixture of zooplankton and resuspended decayed organic matter.

Sediment geochemistry, foraminiferal fauna parameter, census counts and Varimax PC-factor loadings and scores of foraminiferal assemblages of sediment core GeoB12615-4

Analogous to West- and North Africa, East Africa experienced more humid conditions between approximately 12 to 5 kyr BP, relative to today. While timing and extension of wet phases in the North and West are well constrained, this is not the case for the East African Humid Period. Here we present a record of benthic foraminiferal assemblages and sediment elemental compositions of a sediment core from the East African continental slope, in order to provide insight into the regional shallow Indian Ocean paleoceanography and East African climate history of the last 40 kyr. During glacial times, the dominance of a benthic foraminiferal assemblage characterized by Bulimina aculeata, suggests enhanced surface productivity and sustained flux of organic carbon to the sea floor. During Heinrich Stadial 1 (H1), the Nuttallides rugosus Assemblage indicates oligotrophic bottom water conditions and therefore implies a stronger flow of southern-sourced AAIW to the study site. During the East African Humid Period, the Saidovina karreriana Assemblage in combination with sedimentary C/N and Fe/Ca ratios suggest higher river runoff to the Indian Ocean, and hence more humid conditions in East Africa. Between 8.5 and 8.1 kyr, contemporaneous to the globally documented 8.2 kyr Event, a severe reduction in river deposits implies more arid conditions on the continent. Comparison of our marine data with terrestrial studies suggests that additional moisture from the Atlantic Ocean, delivered by an eastward migration of the Congo Air Boundary during that time period, could have contributed to East African rainfall. Since approximately 9 kyr, the gaining influence of the Millettiana millettii Assemblage indicates a redevelopment of the East African fringe reefs.

Benthic foraminifera of the central Indian Ocean

Late Oligocene to late Pliocene vertical water-mass stratification along depth traverses in the northern Indian Ocean is depicted in this paper by benthic foraminifer index faunas. During most of this time, benthic faunas indicate well-oxygenated, bottom-water conditions at all depths except under the southern Indian upwelling and in the Pliocene in the southern Arabian Sea. Faunas suggest the initiation of lower oxygen conditions at intermediate depths in the northern Indian Ocean beginning in Oligocene Zone P21a. Lower oxygen conditions intensified during primary productivity pulses, possibly related to increased upwelling vigor, in the latest Oligocene and throughout most of the late middle through late Miocene. During times of elevated primary production, there may be more oxygen flux into sedimentary pore waters and the shallow infaunal habitat may become more oxygenated. One criterion for locating the source of "new" water masses is vertical homogeneity of benthic foraminifer indexes for well-oxygenated water masses from intermediate through abyssal depths. In the northern Mascarene Basin, this type of faunal homogeneity with depth corroborates the proposal that the northern Indian Ocean was an area of sinking well-oxygenated waters through most of the Miocene before Zone N17. Oxygenated, possibly "new" intermediate-water masses in the low- to middle-latitude Mascarene and Central Indian basins first developed in the late Oligocene. These well-oxygenated waters were probably more fertile than the Antarctic Intermediate Waters (AAIW) that cover intermediate depths in these areas today. Production of intermediate waters more similar to modern AAIW is indicated by the sparse benthic population of epifaunal rotaloid species in the northern Mascarene Basin during middle Miocene Zone N9 and from early through late Pliocene time. Deep-water characteristics are more difficult to interpret because of the extensive redeposition at the deeper sites. Redeposited intermediate, rather than shallow, water fossils and erosion from north to south in the Mascarene Basin are incompatible with the sluggish circulation from south to north through the western Indian Ocean basins today. Such erosion could result from the vigorous sinking of an intermediate-depth water mass of northern origin. Before late Oligocene Zone P22, benthic faunas indicate a twofold subdivision of the troposphere, with the boundary between upper and lower well-oxygenated water masses located from 2500-3000 mbsl. No characteristic bottom-water fauna developed before the end of late Oligocene Zone P22. Deep and abyssal benthic indexes suggest the development of water masses similar to those of the present day in the latest Miocene. Faunas containing deep-water benthic indexes, including the uvigerinids, suggestive of a water mass similar to modern Indian Deep Water (IDW), appeared during the late Miocene in the northern Mascarene and Central Indian basins. In the early Pliocene, this deep-water fauna was found only in the Central Indian Basin, whereas a fauna typical of modern Antarctic Bottom Water (AABW) spread through deep waters at 2800 mbsl in the Mascarene Basin. By late Pliocene Zone N21, however, deep-water faunas similar to their modern analogs were developed in both the eastern and western basins. Abyssal faunas, studied only in the Mascarene Basin, show more or less similarity to those under modern AABW. Bottom-water faunas containing Nuttallides umbonifera or Epistominella exiguua were first differentiated at the end of Zone P22, then appeared episodically during the early Miocene. These AABW-type faunas reappeared and migrated updepth into deep waters during the glacial episodes at the end of the Miocene and at the beginning of the Pliocene. By late Pliocene Zone N21, however, a bottom-water fauna similar to that under eastern Indian Bottom Water (IBW) developed in the Mascarene Basin. Modern bottom-water characteristics of the Mascarene Basin must have developed after ZoneN21.

Benthic foraminifera, stable isotope record and sedimentology of Holocene sediments in the Skagerrak

A high-resolution multi-proxy study of core MD99-2286 reveals a highly variable hydrographic environment in the Skagerrak from 9300 cal. yr BP to the present. The study includes foraminiferal faunas, stable isotopes and sedimentary parameters, as well as temperature and salinity reconstructions of a ca. 29 m long radiocarbon-dated core record. The multivariate technique fuzzy c-means was applied to the foraminiferal counts, and it was extremely valuable in defining subtle heterogeneities in the foraminiferal fauna data corresponding to hydrographic changes. The major mid-Holocene (Littorina) transgression, led to flooding of large former land areas in the North Sea, the opening of the English Channel and Danish straits and initiation of the modern circulation system. This is reflected by fluctuating C/N values and an explosive bloom of Hyalinea balthica. A slight indication of ameliorated conditions between 8000-5750 cal. yr BP is related to the Holocene Thermal Maximum. A subsequent increase in fresh water/Baltic water influence between 5750-4350 cal. yr BP is reflected by dominance of Bulimina marginata and depleted d18O-values. The Neoglacial cooling (after 4350 cal. yr BP) is seen in the Skagerrak as enhanced turbidity, increasing TOC-values and short-term changes in an overall Cassidulina laevigata dominated fauna suggesting a prevailing influence of Atlantic waters. This is in agreement with increased strength of westerly winds, as recorded for this period. The last 2000 years were also dominated by Atlantic Water conditions with generally abundant nutrient supply. However, during warm periods, particularly the Medieval Warm Period and the modern warming, the area was subject to a restriction in the supply of nutrients and/or the nutrient supply had a more refractory character.

Upper Maestrichtian to Eocene benthic foraminifera in ODP Leg 121 holes

Late Maestrichtian to late Eocene bathyal benthic foraminiferal faunas at Sites 752,753, and 754 on Broken Ridge in the eastern Indian Ocean were analyzed as to their stratigraphic distribution of species to clarify the relation between faunal turnovers and paleoceanographic changes. Based on Q-mode factor analysis, eight varimax assemblages were distinguished: the Stensioina beccariiformis assemblage in the upper Maestrichtian to upper Paleocene; the Cibicidoides hyphalus assemblage in the upper Maestrichtian; the Cibicidoides cf. pseudoperlucidus assemblage in the upper Paleocene; the Anomalinoides capitatusldanicus assemblage in the uppermost Paleocene to lower Eocene; the Cibicidoides subspiratus assemblage in the lower Eocene; the Nuttallides truempyi assemblage in the lower and middle Eocene; the Osangularia sp. 1 - Hanzawaia ammophila assemblage in the upper Eocene; and the Lenticulina spp. assemblage in the uppermost Eocene, Oligocene, and lower Miocene. The presence of the Osangularia sp. 1 - Hanzawaia ammophila assemblage is related to the shallowing episode on Broken Ridge (upper bathyal), as a result of the rifting event that occurred in the middle Eocene. The most distinct faunal change (the disappearance of about 37% of the species) occurred between the S. beccariiformis assemblage and the A. capitatusldanicus assemblage, at the end of the upper Paleocene. A. capitatusldanicus, Lenticulina spp., and varied forms of Cibicidoides replaced the Velasco-type fauna at this time. The timing of this event is well correlated with the known age at South Atlantic sites (Thomas, 1990 doi:10.2973/odp.proc.sr.113.123.1990; Kennett and Stott, 1990 doi:10.2973/odp.proc.sr.113.188.1990; Katz and Miller, 1990 doi:10.2973/odp.proc.sr.114.147.1991). The primary cause of the extinction of the Stensioina beccariiformis assemblage is elusive, but may have resulted from the cessation of deep-water formation in the Antarctic (Katz and Miller, 1990), and subsequent arrival of warm saline deep water (Thomas, 1990; Kennett and Stott, 1990). Another possibility may be a weakened influence of high-salinity water formed at the low latitudes such as the Tethys Sea. The extinction event corresponds to the change from higher delta13C values in benthic foraminifers to lower ones. An interpretation of delta13C values is that the eastern Indian deep water, characterized by young and nutrient-depleted water, became old water which was devoid of a supply of new water during the latest Paleocene to early Eocene. Prior to this benthic event, signals of related faunal change were detected in the following short periods: early and late Paleocene, near the boundary of nannofossil Zone CP4, and Zone CP5 of the late Paleocene at Site 752. Among common taxa in the upper Maestrichtian, only seven species disappeared or became extinct at the Cretaceous/ Tertiary boundary at Site 752. The benthic foraminiferal population did not change for up to 2 m above the boundary, in contrast to the rapid decrease of the plankt onic foraminiferal population at the boundary. A decrease in the number of benthic foraminifers occurs after that level, corresponding to an interval of decreased numbers of planktonic foraminifers and higher abundance of volcanic ash. Reduced species diversity (H') suggests a secondary effect attributable to the dissolution of foraminiferal tests. The different responses of planktonic and benthic foraminifers to the event just above the boundary suggest that the Cretaceous/Tertiary event was a surface event as also suggested by Thomas (1990). In addition, a positive shift of delta13C in benthic foraminifers after the event indicates nutrient-depleted bottom water at Site 752.

Neogene benthic foraminifers from the Kerguelen Plateau

Benthic foraminifers were studied quantitatively in 120 lower Miocene through upper Pleistocene samples from Ocean Drilling Program Site 747 (Central Kerguelen Plateau) and Sites 748 and 751 (Southern Kerguelen Plateau). These sites are situated on an 450-km-long, north-south transect between 54°49'S and 58°26'S at present water depths between 1696 and 1288 m. Principal component analysis on the census data of the most abundant 92 taxa helped to identify 8 benthic foraminifer assemblages. These benthic foraminifer assemblages were compared with Holocene faunas from southern high latitudes to reconstruct paleoenvironmental conditions. Middle lower Miocene sediments are characterized by a Uvigerina hispidocostata assemblage, indicating high paleoproductivity and/or not well-ventilated bottom water. From late early to late middle Miocene time, the Southern Kerguelen Plateau was bathed by a young, well-oxygenated, and carbonate-aggressive water mass, as indicated by a Nuttallides umbonifer-dominated benthic foraminifer assemblage. During late middle Miocene time, an Astrononion pusillum assemblage took over for only about 1 m.y., probably indicating the first injection of an aged water mass, similar to the North Atlantic Deep Water (NADW), into a developing circumpolar current system. Around the middle to late Miocene boundary, the fauna again became dominated by N. umbonifer. After the last appearance of N. umbonifer, reestablishment of the A. pusillum assemblage from the early late through at least the late late Miocene, indicated the established influence of a NADW-like water mass. The latest Miocene through middle late Pliocene benthic foraminifer assemblage was characterized by Epistominella exigua and strong carbonate dissolution, indicating very high biosiliceous production, and this in turn may indicate the formation and paleoposition of an Antarctic Polar Frontal Zone. From the late late Pliocene, a Trifarina angulosa assemblage (indicative today of sandy substrate and vigorous bottom currents) strongly dominated the fauna up to the late Pleistocene, when Bulimina aculeata (indicative today of calm sedimentation with high organic matter fluxes) became an important and partly dominating constituent of the fauna. This is interpreted as the faunal response to the decreased winnowing force (bottom current velocities) of the Antarctic Circumpolar Current during periods of global climatic amelioration and raised sea level.

Response of benthic foraminifera to ocean acidification in their natural sediment environment: a long-term culturing experiment

Calcifying foraminifera are expected to be endangered by ocean acidification; however, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment with pCO2-enriched seawater of 430, 907, 1865 and 3247 µatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Omega calc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size-frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 µm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 µm (pCO2 from 430 to 1865 µatm) to 258 µm (pCO2 3247 µatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g/m**2/a, whereas dead empty tests accumulated a rate of 0.27 g /m**2/a. Although Omega calc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 µatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

Benthic foraminifera in Northwest Indian Ocean surface sediments

The relationship between the distribution of benthic foraminifera and sediment type and depositional environment in the Arabian Sea is discussed. The benthic foraminiferal fauna were sampled in nineteen Recent surface sediment samples, and geochemical variables of the sediment of the same samples were measured. The water depths for the box core samples varies from 440 to 4040 m. A total of 103 species and six species-complexes were identified. The geochemical properties were found to correspond well to the sediment type and depositional environment and six different sediment/depositional environment types could be distinguished. Analysis of the benthic foraminiferal fauna reveals specific faunal assemblages that are closely related to these sediment/depositional environment types.

Benthic foraminifers in Mesozoic and Cenozoic sediments of the southwestern Atlantic

Benthic foraminiferal assemblages in Mesozoic and Cenozoic sediments were studied at Sites 511, 512, 513, and 514 drilled during Leg 71 in the southwestern Atlantic on the Maurice Ewing Bank and in the Argentine Basin. Benthic foraminifers in almost all stratigraphic subdivisions of Sites 511 and 512 reflect the gradual subsidence of the Falkland Plateau from shelf depths in the Barremian-Albian, when a semiclosed basin with restricted circulation of water masses and anaerobic conditions existed, to lower bathyal depths in the Late Cretaceous and Cenozoic, with an abrupt acceleration at the boundary of Lower and Upper Cretaceous. The composition, distribution, and preservation of Late Cretaceous assemblages of benthic foraminifers suggest considerable fluctuations of the foraminiferal lysocline and the CCD. This is evidenced by dissolution facies and foraminiferal assemblages in which agglutinated and resistant calcareous forms predominated during high stands of the CCD and by calcareous facies in which rich assemblages of calcareous species predominated during low stands. The highest position of the CCD on the Plateau (less than 1500-2000 m) was in the late Cenomanian, Turonian, and Coniacian. In the Santonian and Campanian the CCD was at depths below 1500-2000 meters. At the end of the Campanian the CCD shifted again to depths comparable with those of Cenomanian and Turonian time. In the latest Campanian and the Maestrichtian the CCD was low and nanno-foraminiferal oozes with a rich assemblage of benthic foraminifers accumulated. Foraminiferal assemblages at Sites 513 and 514 in the Argentine Basin also testify to oceanic subsidence from lower bathyal depths in the Oligocene to abyssal ones at present. This process was complicated by the influence of geographical migrations of the Polar Front caused by extensions of the ice sheet in the Antarctic after the opening of the Drake Passage during the Oligocene. In Mesozoic and Cenozoic deposits of the Falkland Plateau and the Argentine Basin seven assemblages of benthic foraminifers were distinguished by age: early-middle Albian, middle-late Albian, Late Cretaceous (including four groups), middle Eocene, late Eocene-early Miocene, middle-late Miocene, and Pliocene-Quaternary. The Albian assemblages contain many species common to the foraminiferal fauna of the Austral Biogeographical Province. The Late Cretaceous assemblage contains, along with Austral species, species common to foraminifers of North America, Western Europe, the Russian platform, and the south of the U.S.S.R. Deep-sea cosmopolitan species prevail in Cenozoic assemblages.

Distribution of benthic foraminifera in surface sediments on the continental margin off North-West Africa

The benthic foraminiferal populations along three traverses across the Northwest African continental margin were analyzed on the base of ca. 60 surface sediment samples. Depth ranges of 213 species were established and the main trends of vertical distribution are compared with those known from adjacent regions. Main faunal breaks occure at 100/200 m and 1000/1500 m depth of water. Some species show latitudinal distribution boundaries and the same applies to population density (standing stock), reflecting the regional distribution of nutrients supply by river discharge and upwelling processes. - High proportions of Bolivina test at the lower slope indicate extended downslope transport.