Marine Micropaleontology 51 (2004) 345 – 371
www.elsevier.com/locate/marmicro
Distribution of diatoms, coccolithophores and planktic foraminifers
along a trophic gradient during SW monsoon in the Arabian Sea
Ralf Schiebel a,*,1, Alexandra Zeltner b, Ute F. Treppke a, Joanna J. Waniek c,
Jörg Bollmann d, Tim Rixen e, Christoph Hemleben a
a
Institute of Geosciences und Palaontologie, Tübingen University, Sigwartstrasse 10, D-72076 Tübingen, Germany
b
Wilonstrasse 19, 72072 Tübingen, Germany
c
Southampton Oceanography Centre, SOC, European Way / Empress Dock, Southampton SO14 3ZH, UK
d
Department of Earth Sciences, Swiss Federal Institute of Technology, ETHZ, Sonneggstrasse 5, 8092 Zurich, Switzerland
e
Zentrum für Marine Tropenökologie, Bremen University, ZMT, Fahrenheitstr. 6, 28359 Bremen, Germany
Received 1 September 2003; received in revised form 19 January 2004; accepted 14 February 2004
Abstract
The distribution of diatoms, coccolithophores and planktic foraminifers mirrored the hydrographic and trophic conditions of
the surface ocean (0 – 100 m) across the upwelling area off the Oman coast to the central Arabian Sea during May/June 1997
and July/August 1995. The number of diatoms was increased in waters with local temperature minimum and enhanced nutrient
concentration (nitrate, phosphate, silicate) caused by upwelling. Vegetative cells of Chaetoceros dominated the diatom
assemblage in the coastal upwelling area. Towards the more nutrient depleted and stratified surface waters to the southeast, the
number of diatoms decreased, coccolithophore and planktic foraminiferal numbers increased, and floral and faunal composition
changed. In particular, the transition between the eutrophic upwelling region off Oman and the oligotrophic central Arabian Sea
was marked by moderate nutrient concentration, and high coccolithophore and foraminifer numbers. Florisphaera profunda,
previously often referred as a ‘lower-photic-zone-species’, was frequent in water depths as shallow as 20 m, and at high nutrient
concentration up to 14 Amol NO3 l 1 and 1.2 Amol PO4 l 1. To the oligotrophic southeast of the divergence, cell densities of
coccolithophores declined and Umbellosphaera irregularis prevailed throughout the water column down to 100 m depth. In
general, total coccolithophore numbers were limited by nutrient threshold concentration, with low numbers ( < 10 103 cells
l 1) at high [NO3] and [PO4], and high numbers (>70 103 cells l 1) at low [NO3] and [PO4]. The components of the complex
microplankton succession, diatoms, coccolithophores and planktic foraminifers (and possibly others), should ideally be used as
a combined paleoceanographic proxy. Consequently, models on plankton ecology should be resolved at least for the seasonality,
to account for the bias of paleoceanographic transfer calculations.
D 2004 Elsevier B.V. All rights reserved.
Keywords: coccoliths; diatom flora; foraminifera; nutrients; oceanography; paleoceanography
* Corresponding author. Tel.: +41-1-6323676; fax: +41-1-6321080.
E-mail address: [email protected] (R. Schiebel).
1
Current address: Department of Earth Sciences, Swiss Federal Institute of Technology, ETHZ, Sonneggstrasse 5, 8092 Zürich,
Switzerland.
0377-8398/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.marmicro.2004.02.001
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R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
1. Introduction
The Indian Ocean Dipole is one of the most
prominent climate systems on Earth today, and is
connected to other climatic patterns such as the North
Atlantic Oscillation (NAO), El Niño Southern Oscillation (ENSO) and the Indonesian to Australian region
via tele-connections (e.g., Ashok et al., 2003). The
Indian Monsoon affects the precipitation on land as
well as the hydrography and the biogeochemistry of
the Indian Ocean (e.g., Ittekkot and Nair, 1993;
Fleitmann et al., 2003). Distribution and the standing
stock of marine planktic organisms, including diatoms, coccolithophores and planktic foraminifers, is
related to the monsoonal oscillation, which varied
over the historic past and over geologic time scales
(e.g., Gupta et al., 2003; Ivanova et al., 2003).
The fossil remains of microorganisms, for example
skeletons and alkenones, are used as paleoceanographic tools although microplankton dynamics and
trophic conditions are not yet fully understood. In this
study, we therefore focus on the effect of the southwest monsoon (SWM) on the ecology of living
diatoms, coccolithophores and planktic foraminifers,
and on the differential population dynamics among
these three microplankton groups.
Different phytoplankton groups co-occur according to ecological demand on trophic (nutrients, food),
physical (e.g., light, mixing, temperature) or biological factors (e.g., competition, predation) (cf. Smayda,
1986). In particular, we focus on the effect of
hydrography on nutrient concentration in the upper
100 m of the water column and production of
autotrophic organisms (diatoms and coccolithophores). This is pursued by quasi-synoptic investigations on the spatial and temporal succession of
diatoms, coccolithophores and planktic foraminifers
along a productivity gradient from the coastal
upwelling off Oman (eutrophic) to the stratified
(oligotrophic) central Arabian Sea.
1.1. Previous studies on diatoms, coccolithophores
and planktic foraminifers in the Arabian Sea
Diatoms of the Indian Ocean were studied by
Cleve (1901), Karsten (1907), Taylor (1967), Sournia
(1968), Hendey (1970), Thorrington-Smith (1970)
and Mathur and Singh (1993). Certain diatom taxa
in coastal waters off India were studied for their
standing crop, unfortunately without being discussed
for their ecology (Prasad and Nair, 1960; Durairatnam, 1964). Simonsen (1974) gave a broad and
detailed overview of species in plankton samples from
net hauls taken during the International Indian Ocean
Expedition (IIOE) in 1964/1965, but only little information on diatom ecology was presented. Diatoms of
the Arabian Sea have not yet been investigated in a
temporal and spatial resolution that would allow an
ecological interpretation.
The taxonomic composition of coccolithophore
assemblages from the Arabian Sea and northern
Indian Ocean was investigated by Norris (1965,
1971, 1983, 1984, 1985), Kleijne et al. (1989) and
Kleijne (1991, 1992, 1993). Bernard and Lecal (1960)
reported the distribution of extant coccolithophores in
a general phytoplankton study. Martini and Müller
(1972) and Guptha et al. (1995) analyzed the coccolithophore assemblage along transect at 65jE during
the late SWM. Sediment trap studies from the northeastern Arabian Sea (Andruleit et al., 2000), and from
the Somalia upwelling region (Broerse et al., 2000)
revealed a strong relationship between the monsoon
and the seasonal flux of coccolithophores. A first
comparison between monsoon induced changes of
the living coccolithophore assemblage in the northwestern Arabian Sea, compared to Holocene and
Quaternary assemblages, was carried out by Woellner
et al. (1988). Holocene sediment assemblages in the
Arabian Sea are dominated by Gephyrocapsa oceanica (Martini and Müller, 1972; Guptha, 1985;
Houghton and Guptha, 1991; Andruleit and Rogalla,
2002). Potential environmental control of coccolithophores in surface waters off Pakistan was investigated
by Andruleit et al. (2003).
Planktic foraminifers were investigated for their
general distribution, ecology and fossil record in the
Indian Ocean by Bé and Tolderlund (1971), Bé and
Hutson (1977) and Guptha et al. (1994). Seasonal
changes of the planktic foraminiferal fauna and stable
isotope composition over the course of the monsoons
are described by Kroon and Ganssen (1988) and
Curry et al. (1992). Kroon (1988) and Ivanova et al.
(1998) reported Globigerinita glutinata, Neogloboquadrina dutertrei and Globorotalia menardii as upwelling indicators in the Arabian Sea. While
Globigerina bulloides occurs mainly during the late
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
SWM (September, after Kroon and Ganssen, 1988),
N. dutertrei is more frequent during the early SWM,
in June and July, along with decreased sea surface
temperatures, which is again indicative of upwelling.
2. Materials and methods
Water and microplankton was collected from the
upper 100 m of the water column in the Arabian Sea.
During R/V METEOR cruise 32/5 (M32/5), July 14 to
August 14, 1995, nine stations along a transect perpendicular to the coast of Oman towards 14jN/65jE
(Central Arabian Sea Station, CAST) and at 10jN/
65jE (Southern Arabian Sea Station, SAST) were
sampled (Table 1, Fig. 1). During R/V SONNE cruise
119 (SO119), May 12 to June 10, 1997, nine stations
were sampled along the coast of the Oman and at the
Northern, Western, Central and Southern Arabian Sea
Stations (NAST, WAST, CAST and SAST).
To obtain diatoms and coccolithophores, water
was sampled with 5-liters Niskin bottles at 10, 20,
40, 60, 75 and 100 m water depth. The Niskin bottles
were attached to and synchronized with an opening–
closing-net. Water samples were treated according to
347
the method of Kleijne (1991). Coccolithophores were
filtered onto a 50-mm diameter, regenerated cellulose
membrane filter with a pore size of 0.45 Am (SARTORIUS) using a vacuum pump. The vacuum was
adjusted to 200 mbar and the actual filtration area was
1249 mm2. All filters were air dried for 12 h, stored
in plastic petri dishes and were kept dry in closed
boxes with silica gel. For the analysis on a scanning
electron microscope (SEM), circular pieces (area of
133 mm2) were punched out of the center of the filter
membrane and mounted onto aluminium stubs using
double-sided adhesive tape. Subsequently, samples
were coated with 20 nm of gold/palladium, and
colloidal silver suspension was put on the border of
the membrane to provide optimal conductivity of the
sample. The stubs were examined on a Cambridge
Stereoscan S250 SEM at 15– 20 kV.
Diatoms and coccolithophores were counted on
the same microscope stub. Diatoms were counted
from eight sites sampled during M32/5 and SO119
(Table 1) at a magnification of 1000 . Diatom
valves were counted in an area of 15 – 275 SEM
screens, randomly distributed on the filter. In general,
diatoms were preserved as single valves and complete
cells. Counting units were defined according to
Table 1
List of sampling locations
Cruise
SO119
M32/5
a
b
Date
16.05.97
20.05.97
22.05.97
24.05.97
26.05.97
31.05.97
01.06.97
02.06.97
03.06.97
24.07.95
27.07.95
30.07.95
01.08.95
04.08.95
04.08.95
06.08.95
07.08.95
10.08.95
Sta. #a
NAST-3
CAST-5
SAST-6
WAST-7
12
28
31
36
37
404
414
423
430
438
440
444
446
460
Lat. (jN)a
19j56.9V
14j27.5V
10j00.7V
16j12.1V
17j14.5V
17j18.6V
17j45.9V
19j02.1V
20j33.9V
09j58.8V
14j26.8V
16j01.9V
17j06.8V
18j31.8V
18j36.7V
18j13.7V
17j41.2V
16j13.5V
Long. (jE)a
65j49.5V
64j35.1V
64j59.9V
60j18.5V
58j31.3V
58j23.6V
59j04.2V
58j48.9V
60j03.1V
65j01.1V
65j00.9V
62j00.8V
60j00.8V
57j20.5V
57j28.5V
58j13.2V
58j53.2V
61j28.3V
Sta. = station, Lat. = latitude, Long. = longitude, MN = multinet number.
D = diatoms, Cocco = coccolithophores, plF = planktic foraminifers.
MN #a
1272
1278
1281
1284
1288
1295
1298
1301
1303
971
974
977
979
981
982
986
988
990
Db
Sample depth (m)
Coccob
–
–
–
–
–
–
–
20
–
–
–
60
10
10
20
20
20
–
20/60/100
20/60/100
20/60/100
20/60/100
60/100
20/60/100
20/60
20/60/100
20/60/100
20/60/100
20/60/100
20/60/100
60/100
40/75
10/40/75
60/100
20/60/100
20/60/100
plFb
0 – 100
0 – 100
0 – 100
–
0 – 100
–
–
0 – 100
–
0 – 100
–
0 – 100
–
–
–
–
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R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Fig. 1. Location of sampling sites during R/V Sonne cruise 119 and R/V Meteor cruise 32/5. Primary production (after Antoine et al., 1996,
g C m 2 year 1) is given in italic numbers (dashed lines). Coastal upwelling off Oman causes highest primary production. Lowest biological
productivity occurs in oligotrophic waters at the southeast of the study area. The Findlater Jet Axis is indicated by the arrow (cf. Rixen et al.,
2000).
Schrader and Gersonde (1978). The abundance of
diatoms is given as valves per liter and the relative
amount of taxa was calculated (see Appendix A).
Coccolithophores were counted on 500 randomly
chosen fields (49.5038.25 Am = 1893.38 Am2) at
2000 magnification along transects over the
complete circular sub-sample of the membrane filter.
If possible, at least 200 individuals (complete coccospheres) per sample were analyzed. Unequivocally
collapsed coccospheres were considered complete.
The number of coccolithophores is given in cells
per liter (Appendices B and C). Morphometric measurement of gephyrocapsids was carried out according
to Bollmann (1997), by analyzing 50 specimens per
sample on the SEM.
Planktic foraminifers were sampled in five 20-m
depth intervals between the ocean surface water and
100 m water depth (Appendices D and E) with a
multiple-opening-closing-net (100-Am mesh size).
Samples were fixed on board by a 4% formaldehyde
solution buffered with hexamin at pH 8.2. In the
laboratory, planktic foraminiferal tests were picked,
dried, sieved into size classes of >100 –125 – 150–
200 – 250 – 315 and >315 Am then counted on a
species level. Cytoplasm bearing tests (‘living specimens’) were counted separately from empty tests
(‘dead specimens’). The taxonomy of planktic foraminifers used here follows Hemleben et al. (1989). On
average, more than 90 % of specimens are classified
as ‘living’ (up to 99.25%). Therefore, in the following
we refer to the total numbers of specimens (sum of
‘living’ and ‘dead’).
During cruise M32/5, temperature and salinity
were measured with a Neil Brown Mark III CTD with
fluorometer and oxygen sensor, and during cruise
SO119 a SBE 4-Conductivity Sensor was used.
Nutrients (dissolved nitrate [NO3], phosphate [PO4]
and silicate [SiO4]) were analyzed with a Continuous
Flow Analyzer according to Grasshoff et al. (1988).
Samples were taken from the rosette sampler, along
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
with CTD measurements, in polyethylene bottles and
analyzed within 24 h after collection. Chlorophyll a
analysis (M32/5) was performed according to Herbland et al. (1985). The data is available from http://
www.pangaea.de/home/rschiebel/.
3. On the hydrography of the modern Arabian Sea
Detailed knowledge of the hydrography is crucial
for the understanding of the ecology of diatom, cocco-
349
lithophore and planktic foraminiferal species, which, in
turn, will allow for detailed paleoceanographic interpretation. Hydrographic circulation, trophic condition
and biological productivity of the Arabian Sea are
largely affected by the seasonal oscillation of the
monsoon winds. Northeastern winds prevail between
November and March (northeast monsoon, NEM), and
southwestern winds prevail from May to October
(SWM). Different heat capacity of land and sea causes
barometric pressure differences and strong winds,
which result in the low-level atmospheric Findlater
Fig. 2. Temperature (jC) and salinity of the upper 100 m along the Sonne 119 (panels a and b) and Meteor 32/5 transect (panels c and d). The
cross-hatched square at the lower left corner of panel c and d indicates the sea floor. For the exact position of the stations (numbers on top of
panel a), see Fig. 1 and Table 1. Note the different latitudinal extension of the upper (a, b) and lower (c, d) panels.
350
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Jet, parallel to the Somalian coast during the SWM
(Findlater, 1971) (Fig. 1). As a result, surface currents
turn towards the northeast, surface waters are displaced
away from the coast, and strong upwelling occurs at the
coast off Somalia and Oman. Two different upwelling
modes have been suggested: (1) Ekman driven upwelling caused by the wind stress curl, between 15jN and
22jN up to 30 km off the coast of Oman (Wyrtki, 1973;
Bauer et al., 1991), and also at the open ocean along the
Findlater-Jet axis (Smith and Bottero, 1977; Swallow et
al., 1983; Manghnani et al., 1998; Rixen et al., 2000)
(Fig. 1). (2) Quasi-stationary filaments are tied off from
the coastal upwelling area. Filaments are tongues of
cold and nutrient rich surface water (visible from
satellite images) associated to upwelling, and reach
far beyond the coast (cf. Weaks, 1983; Washburn et al.,
1991; Brink and Cowles, 1991; Brock et al., 1992;
Waniek, 1997). Depending on the intensity of the
SWM winds, filaments are features that are supplied
by coastal upwelling and often triggered by topography
(Strub et al., 1991).
3.1. Environmental conditions during sampling
In contrast to warm surface waters during May/
June 1997, comparatively cold surface waters during
July/August 1995 were observed, indicating strong
upwelling to the north of 15jN (Fig. 2). Concentration
of silicate, nitrate and phosphate in the coastal upwelling area off Oman increased with water depth (Fig. 3)
and was negatively related to water temperature (Appendices A –E) (cf. Tomczak, 1984). Concentration of
chlorophyll (>0.82 Ag l 1) was high throughout the
upper water column close to the Oman coast and
indicates high primary production (Appendix A,
M32/5, Stations 438 and 444). At Stations 423 and
460, at the southern margin of the upwelling area
(Fig. 1), updoming of isotherms from below 180 m
water depth possibly indicates a filament (cf. Waniek
et al., 1996). At the oligotrophic site in the open
Arabian Sea (Station 404), a Deep Chlorophyll
Maximum (DCM) indicates stratification of the
upper water column, nutrient concentrations were
Fig. 3. Concentration (Amol l 1) of (a) silicate, (b) phosphate and (c) nitrate in the upper 100 m of the water column during July/August 1995.
Sample positions are indicated by dots.
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
decreased, and nitrate was only detected below 90 m
water depth (Von Bröckel et al., 1996).
The SWM in 1997 started on May 24, indicated by
a sudden shift of wind direction from 315j to 225 –
270j, and increased wind speed from 3 to 9 m s 1
(Bange et al., 1999). The sea surface temperature
(SST) remained high between 29 and 30 jC at the
open ocean Stations 6 to 31 (Fig. 2). Close to the
Oman coast, at Stations 36 and 37, slight upwelling
was indicated by decreased SST at 18 –20jN (Fig. 2).
To the southeast of the coastal upwelling area, updoming of isotherms was recorded, which possibly indicates diverging currents and slight upwelling caused
by the atmospheric Findlater Jet (Stations 5, 7, 12 and
28). To the southeast of the divergence, in the oligotrophic open Arabian Sea, surface waters were stratified between 10jN and 14jN (Fig. 2).
4. Results
4.1. Diatom distribution
The highest numbers of diatom valves off the
Oman coast occurred closest to land and numbers
351
decreased towards the open ocean at the beginning of
August 1995 (Fig. 4). In the coastal area, vegetative
cells of Chaetoceros with more than 80% of the total
assemblage were most abundant. In general, lowest
species richness occurred along with high valve
concentration in the upwelling area close to the Oman
coast and the highest species richness occurred along
with decreased nutrient concentration (Appendix A).
Most Chaetoceros species in the coastal area belong
to the group of hyalochaetes. In addition, Thalassionema nitzschioides var. nitzschioides was also predominant, and Stephanopyxis turris occurred as a
typical species of this assemblage (Appendix A).
Further from the coast, absolute and relative abundances of Pseudo-nitzschia spp. (mainly P. delicatissima and P. pseudodelicatissima) and species of the
oceanic Nitzschia bicapitata-group increased. Highest
numbers of N. bicapitata species, as well as taxa
related to the Palgiotropis-group, occurred at Station
446. In the open Arabian Sea (Station 423), the
diatom assemblage was still characterized by the N.
bicapitata-group, Chaetoceros and Pseudo-nitzschia.
In May/June 1997, the number of diatom specimens
off the Oman coast (Station 37) resembled that at
Stations 440 and 444, in July/August 1995. Virtually,
Fig. 4. Distribution of the diatom taxa (a) Chaetoceros spp., (b) Nitzschia bicapitata-group, (c) Thalassionema nitzschioides and (d) Pseudonitzschia spp. along a transect from the Oman coast (left) towards the central Arabian Sea (right), given in percent of total diatoms. The total
number of diatoms (valves 104 l 1) is given as bold line in panel a. The break between 1995 (Stations 438 – 423) and 1997 (Station 37) is
indicated by the dashed vertical line.
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R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
no freshwater diatoms were observed during May/
June 1997 (Appendix A). During May/June 1995,
diatom valve numbers were positively related to
nutrient concentration (N = 6, linear regression: SiO4,
r = 0.68, P = 14.0%; NO3, r = 0.80, P = 5.6%; PO4,
r = 0.81, P = 5.2 %) (Appendix A).
4.2. Coccolithophore distribution
In total, 83 coccolithophore taxa were recorded
in the Arabian Sea during the analyzed time
periods. The highest species richness (48 taxa)
was recorded at Station 31 during June 1997,
and the overall diversity increased towards the
oligotrophic open ocean during both sampling
campaigns (Appendices B and C). However, the
diversity varied significantly between the two
cruises with 80 taxa during May/June 1997 and
40 taxa during July/August 1995. Standing stock
increased from the upwelling area towards the
oligotrophic open ocean, although in different
modes during both sampling periods. Total cell
Fig. 5. Coccolithophore cell numbers are negatively related to nitrate [NO3] and phosphate [PO4] concentration (N = 46 for both [NO3] and
[PO4]). Below lower threshold of 4 Amol NO3 and 0.35 Amol PO4 l 1 more than 70 103 coccolithophore cells l 1 occurred, and above the
upper threshold of 14 Amol NO3 and >1.25 Amol PO4 l 1 cell densities are lower than 10 103 cells l 1. Nutrient thresholds are similar for
total coccolithophore cell numbers (panel a) and for distinct species (panels b to g), although cell numbers ( 103 cells l 1) vary between
species.
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
density varied between 0 and 112 103 cells l 1,
and in the oligotrophic central Arabian Sea cell
densities were lower during May/June 1997, than
during July/August 1995.
Standing stocks were negatively related to NO3
and PO4 concentrations. In the oligotrophic Arabian Sea, high cell densities (>70 103 cells l 1)
occur at < 4 Amol NO3 l 1 and < 0.35 Amol PO4
l 1. At a concentration below 14 Amol NO3 l 1
and below 1.25 Amol PO4 l 1 cell densities
exceeded 10 103 cells l 1 (Fig. 5a). At these
levels of NO3 and PO4 concentration, cell numbers
of the total coccolithophore flora (Fig. 5a) and
of all frequent species (Fig. 5b – g) changed
distinctly.
During July/August 1995, G. oceanica was the
most abundant species (Fig. 6b) with up to
51 103 cells l 1. The joint second most abundant
species were Emiliania huxleyi and the ‘lower
photic zone species’ Florisphaera profunda with
up to 22 103 cells l 1 (Appendix C). Oolithotus
antillarum reached up to 20 103 cells l 1 at the
open ocean Station 404 (Fig. 1, Appendix C). In
contrast, during May/June 1997, F. profunda was
the most abundant species with up to 31 103 cells
l 1 at the Stations 3 and 7, and at the stratified
353
oligotrophic central Arabian Sea Station 6, Umbellosphaera irregularis was most abundant (Fig. 7c
and d). The high cell concentrations of up to
45 103 cells l 1 of O. antillarum (Station 5, 20
m) and Calcidiscus leptoporus (Station 7, 20 m;
‘type A’ after Kleijne, 1993; ‘small morphotype’
after Knappertsbusch et al., 1997) during May/June
1997, are remarkable, as both species usually show
much lower cell densities. G. oceanica shows a
general decrease of the proportion of its ‘larger
morphotype’ (after Bollmann, 1997) from the upwelling area (Stations 37 and 438; diameter of 4.3 –
4.5 Am) towards the oligotrophic open ocean (Stations 7 and 404) where the ‘equatorial morphotype’
of G. oceanica (diameter of 3.5 – 3.9 Am) was
dominant.
4.3. Planktic foraminiferal distribution
During the beginning of the SWM in May/June
1997, high average planktic foraminiferal abundance
was restricted to the upper 40 m of the water
column (Fig. 8) with maximum numbers at Station
7. Medium numbers were recorded at the open
Arabian Sea (Stations 5 and 6) and low numbers
occurred at the two sites closest to the Oman coast
Fig. 6. (a) Total coccolithophore cell densities ( 103 cells l 1) and (b) relative floral part of G. oceanica from the upwelling area off Oman
(left) to the central Arabian Sea (right) during July/August 1995 (see Appendix C).
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Fig. 7. (a) Total coccolithophore cell densities ( 103 cells l 1) and (b, c, d) relative floral part of frequent species from the upwelling area off
Oman (left) to the central Arabian Sea (right) during May/June 1997 (see Appendix B). Along the transect, from the north to the south, three
hydrographic regimes can be distinguished: upwelling (Stations 37, 3, 36 and 31), an area of diverging currents (Stations 28, 12, 7 and 5) and
stratified waters in the central Arabian Sea (Station 6).
(Appendix D: Stations 28 and 37). The most frequent species were Globigerinoides ruber at Station
7 and Globigerinoides sacculifer at Station 6, increasing in numbers from the north towards the
south. High relative frequency of G. ruber and G.
sacculifer was recorded only south of the area of
coastal upwelling (Fig. 8). In the area of the coastal
upwelling (Station 37) G. bulloides was most frequent, constituting up to 60% of the assemblage in
the uppermost 60 m (Fig. 8) and Neogloboquadrina
dutertrei also occurred in high numbers. South of
the upwelling area G. bulloides and N. dutertrei
were rare. Globigerinoides glutinata was rare at
the northern Station 37 and numbers only slightly
increased at Stations 28, 6 and 5 (Fig. 1, Appendix
D). In 1995, numbers of planktic foraminifers were
much higher than in 1997 and the main depth
habitat was broadened to 80 m instead of 40 m in
1997 (Appendices D and E). The species composition was different comparing both years and, in
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
355
Fig. 8. Distribution of (a) total planktic foraminifers and (b, c, d) frequent species in the upper 100 m of the water column (see Appendix D),
from the upwelling area off Oman (left) to the central Arabian Sea (right), during May/June 1997, indicates three different hydrographic and
trophic regimes (on top of panel a).
addition to G. ruber and G. sacculifer, high numbers of Globoturborotalita tenella and Globigerina
falconensis occurred at Stations 5 and 7 during
July/August 1995 (Appendix E).
4.4. Succession of diatoms, coccolithophores and
planktic foraminifers along the trophic gradient
Along transect sampled in the northern to central
Arabian Sea during early (May/June 1997) to mid
SWM (July/August 1995), a gradient from high to
low nutrients was observed (Fig. 3). In general, diatom
valve numbers were positively related to nutrient
concentration and negatively related to coccolithophore cell numbers (Fig. 9). The general distribution
of planktic foraminifers (Fig. 8) did not correlate to a
high degree to both the diatom and the coccolithophore
distribution (cf. Appendices A, C and E). However,
highest planktic foraminiferal abundance occurred in
the area of divergence where mesotrophic conditions
were recorded, and which resembles the distribution of
coccolithophores during May/June 1997 (Figs. 7 and
8). Lowest frequency of planktic foraminifers occurred
along with high nutrient concentration under upwelling
conditions and was negatively related to the distribution of diatoms (Figs. 8 and 9).
356
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
5. Discussion
5.1. Specific diatom flora in different upwelling
regimes
Diatoms are the dominant phytoplankton group in
upwelled coastal waters during SWM off Oman, in
terms of valve concentration in surface waters, carbon
cycling and mass flux (cf. Garrison et al., 2000). The
abundance of diatoms is closely related to the availability of nutrients (Fig. 9). The dominating diatom
species were Chaetoceros spp. and Thalassionema
nitzschioides during both investigated years 1995 and
1997 (cf. Brock et al., 1991). The Chaetoceros species
that occurred along with upwelling off Oman are
hyalochaete taxa, which are characteristic of the high
productive coastal regime (Pitcher et al., 1991), and cooccurring T. nitzschioides, which is related to coastal
waters and high nutrient levels (Abrantes, 1988;
Treppke et al., 1996) and to enhanced biological
productivity caused by river runoff (e.g., off southwest
Africa; Van Iperen et al., 1987). Both Chaetoceros and
T. nitzschioides indicated coastal upwelling and high
primary productivity during the sampling campaigns
M32/5 and SO119. Outside the upwelling area, diatom
frequency decreased by two orders of magnitude and
the species composition changed (Fig. 4). Enhanced
biological productivity towards the central Arabian Sea
was indicated by Nitzschia bicapitata, a diatom species, which is possibly related to upwelled open-oceanic waters (cf. Lange et al., 1994) or to filaments,
propagating from the Oman upwelling. The cooccurrence of more coastal Pseudo-nitzschia spp. and Chaetoceros spp. with the more open marine N. bicapitata
points towards mixing of different nutrient rich water
masses (cf. Cupp, 1943; Hasle, 1965; Pitcher et al.,
1991, Lange et al., 1994; Treppke et al., 1996).
5.2. Coccolithophores, nutrient concentration and
surface water stratification
Coccolithophore cell densities (up to 107 103 l 1)
and species richness in the Arabian Sea are comparable
to cell densities in the Atlantic at Bermuda (BATS) and
in the Pacific Ocean near Hawaii (HOT) (cf. Haidar and
Thierstein, 2001; Cortés et al., 2001). The significant
differences in coccolithophore cell densities and diversity in the Arabian Sea, comparing May/June 1997 and
July/August 1995, may be related either to the seasonal
succession or to interannual variations in the strength of
Fig. 9. Diatom (open bars) and coccolithophore (black bars) abundance are negatively related (r = 0.69, N = 6) along transect of varying
nutrient concentration (exemplified by [NO3], bold line; linear regression: r = 0.80, N = 6) in the northern Arabian Sea. High nutrient
concentration occurs in the upwelling area off Oman (NW of transect, left side) and in the central Arabian Sea nutrient concentration is low (SE
of transect, right side). The break between 1995 (Stations 438 – 423) and 1997 (Station 37) is indicated by the dashed line.
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
the monsoonal upwelling. Seasonal variation in cell
densities was also reported from the time series Stations
BATS and HOT, and, in addition, significant interannual variability was observed at HOT (Haidar and
Thierstein, 2001; Cortés et al., 2001).
The general increase of cell density and species
richness towards the oligotrophic open ocean supports
the assumption that coccolithophores are adapted to
low to medium nutrient concentration (Fig. 5), low
turbulence and stratified surface waters (cf. Winter,
1985; Mitchell-Innes and Winter, 1987; Kleijne, 1993).
From the complete Arabian Sea data set on coccolithophore cell density, [NO3] and [PO4], there is striking
evidence that coccolithophore cell density does not
follow nutrient concentration in some negative linear or
exponential way, but is limited by thresholds of nitrate
and phosphate (Fig. 5). This assumption is true for the
total coccolithophore flora and for at least the most
frequent species observed in the study presented here.
Complete ecological preferences of coccolithophores, however, are difficult to infer from our data
set because of missing data on, for example, light and
turbulence, and due to the complex relations between
environmental parameters and differential biologic
prerequisites of species. Unfortunately, cell densities
of more than 106 cells l 1, as reported from neritic
environments of the North Atlantic (e.g., Birkenes and
Braarud, 1952; Burkill et al., 2002), were neither
observed during the study presented here, nor at BATS
and HOT, and, therefore, the upper end of cell density
and maybe a lower threshold of nutrient concentration
could be missing here (Fig. 5).
The dominance of Gephyrocapsa oceanica in the
studied area confirms the results obtained in Holocene
sediments (Martini and Müller, 1972; Guptha, 1985;
Houghton and Guptha, 1991; Andruleit and Rogalla,
2002) and supports the general assumption that G.
oceanica morphotypes are adapted to upwelling, neritic settings, and equatorial open-ocean warm-water
conditions (Bollmann, 1997 and references therein).
Our morphometric data show that the ‘larger morphotype’ of G. oceanica is positively related to upwelling
and neritic settings, whereas the ‘equatorial morphotype’ appears to be adapted to the open ocean warm
water conditions. G. oceanica ‘equatorial’ is negatively
related to the concentration of nitrate and exhibits much
higher cell densities than G. oceanica ‘larger’. These
findings confirm the ecological model of these mor-
357
photypes that was previously inferred from Holocene
sediments (Bollmann, 1997).
In contrast to the observation that the lower-photiczone-species Florisphaera profunda occurs mainly in
subsurface waters of 75 – 200 m depth (Okada and
Honjo, 1973; Reid, 1980; Cortés et al., 2001; Haidar
and Thierstein, 2001), F. profunda occurred in high cell
numbers in waters of 20 –60 m depth during May/June
1997 (Fig. 7, cf. Andruleit et al., 2003). The occurrence
of F. profunda throughout the upper water column
supports the assumption that F. profunda has rather
an affinity to enhanced nutrient concentration (Fig. 5)
and low light intensity than for a specific water depth
(cf. Ahagon et al., 1993; Cortés et al., 2001; Haidar and
Thierstein, 2001). F. profunda was most frequent in the
area of divergence between 13jN and 17jN (Fig. 7,
Appendix B), where updoming isotherms indicated
nutrient entrainment into surface waters from below,
and stimulated primary productivity at depths of suitable light intensity. However, updoming isotherms may
also indicate upward transportation of coccolithophores from below, and not their normal habitat. Along
with F. profunda, also Gladiolithus flabellatus, Calcidiscus leptoporus and Oolithotus antillarum were
frequent close to the nutricline.
Calcidiscus leptoporus is attributed to oligotrophic
open marine conditions (Blasco et al., 1980; MitchellInnes and Winter, 1987; Giraudeau, 1992; C̆epek,
1996). However, there are at least three different
morphotypes of C. leptoporus with specific environmental preferences, out of which the small morphotype
is present here. The small morphotype (type A after
Kleijne, 1993; type small after Knappertsbusch et al.,
1997) appears to prefer water temperature higher than
26 jC, which is confirmed by our observations. Umbellosphaera irregularis was the most characteristic species, dominating the flora in warm, stratified, and
oligotrophic waters (Fig. 7d), confirming earlier observation of Kleijne (1993), Haidar and Thierstein (2001)
and Cortés et al. (2001).
5.3. Planktic foraminifers and phytoplankton prey
Enhanced availability of nutrients and prey
(particularly diatoms) in the upwelling area off Oman,
at Station 37, was mirrored by high numbers of the
planktic foraminiferal species Globigerina bulloides
and Neogloboquadrina dutertrei (cf. Kroon, 1988;
358
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Ivanova et al., 1998), while the total number of planktic
foraminifers decreased (Fig. 8, Appendix D). G. bulloides is reported as an upwelling species (e.g., Kroon
and Ganssen, 1988; Hemleben et al., 1989; Brock et al.,
1992; Naidu and Malmgren, 1996; Kiefer, 1999),
indicating highest levels of primary production (Berger
et al., 1988), and high concentration of chlorophyll a
(Appendix E). Chlorophyll a is linearly correlated to
the concentration of autotrophs, the autotrophs serving
as prey for larger zooplankton including planktic
foraminifers (Hemleben et al., 1989; Garrison et al.,
2000). We found high faunal parts of G. bulloides
synchronous to high numbers of diatoms, during the
early ‘upwelling phytoplankton stage’, not only during
the ending ‘upwelling zooplankton stage’ (cf. Kroon
and Ganssen, 1988). Therefore, we suggest that G.
bulloides increases production at local upwelling cells
not only during a final stage of upwelling but during
the whole upwelling season. Enhanced numbers of G.
bulloides always occurred together with an increase of
numbers of N. dutertrei. Although G. glutinata is
specialized on a diatom diet (Hemleben et al., 1989),
G. glutinata was rare in the upwelling region where
diatoms were frequent (Appendices D and E) Fig. 4a).
G. glutinata is not as opportunistic as G. bulloides and
N. dutertrei, and is more adapted to nutrient entrainment at nutricline depth, and therefore deep production
of diatoms (cf. Schiebel et al., 2001).
To the south of the upwelling region off Oman, in
May/June 1997, Globigerinita bulloides and Neogloboquadrina dutertrei were rare and the fauna
was dominated by subtropical to tropical species
Globigerinoides ruber and Globigerinoides sacculifer
(cf. Conan and Brummer, 2000). According to a
higher salinity (Fig. 2), G. ruber dominated the fauna
at Station 7 (Fig. 9b), while G. sacculifer dominated
at Station 6 where the upper ocean salinity was lower
( < 35.5) than at Station 7 (>36.4) (cf. Bé and
Tolderlund, 1971; Bé and Hutson, 1977; Shenoi et
al., 1993). Along with G. ruber, Globoturborotalita
tenella was frequent in 1995 and may favour similar
temperature and salinity conditions as G. ruber in the
Arabian Sea.
5.4. Microplankton succession in the Arabian Sea
During SW monsoon in the Arabian Sea, the
succession of diatoms, coccolithophores and
planktic foraminifers in 1995 and 1997 resembled
the hydrography and followed a gradient in nutrient concentration (Fig. 10). In the Arabian Sea,
three microplankton associations were identified
and assigned to different hydrographic situations
(Fig. 10).
(A)
(B)
(C)
In the upwelling area off Oman, turbulent
mixing caused high nutrient concentration at
the sea surface. At Station 438 in the central
upwelling area (Fig. 1), Chaetoceros spp.
and Thalassionema nitzschioides diatoms
occurred in high numbers, and among low
coccolithophore numbers the ‘larger morphotype’ of Gephyrocapsa oceanica was most
frequent. To the northeast of the upwelling
center, at Station 37, though still under
upwelling conditions and with a species
composition similar to Station 438, diatom
numbers were reduced and coccolithophore
numbers were still low. Therefore, we
assume that coccolithophores are not actively
replaced by diatoms but by environmental
conditions, which, however, needs further
investigation. The planktic foraminiferal fauna under coastal upwelling conditions was
dominated by the opportunistic species
Globigerina bulloides.
Updoming of isotherms and entrainment of
nutrients into surface waters from below was
caused by strong Findlater Jet winds to the
southeast of the coastal upwelling area. Diatoms occurred in much lower numbers than
under upwelling conditions and the diatom
flora was characterized by Nitzschia bicapitata.
Coccolithophore and planktic foraminifer
numbers were very high. The most frequent
coccolithophore species was Florisphaera profunda at nutricline depths. The foraminiferal
fauna was characterized by Globigerinoides
ruber; Globigerinoides sacculifer was the
second most frequent species.
In the open Arabian Sea, the water column
was well stratified and nutrient concentration
in surface waters was low. Diatoms occurred
in low numbers and coccolithophores were
very abundant. The coccolithore flora was
characterized by Umbellosphaera irregularis
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
359
Fig. 10. Model of the diatom, coccolithophore and planktic foraminiferal succession (A, B, C), according to hydrographic zonation and
trophic condition along transect (. – .) from the Oman coast to the central Arabian Sea during the southwest monsoon. A = coastal
upwelling, turbulent mixing, eutrophic; B = strong Findlater Jet, updoming isotherms and isohalines; C = stratified water column,
oligotrophic.
throughout the upper 100 m of the water
column. Globigerinoides sacculifer was the
most frequent species in the upper 40 m,
below 40 water depth Globigerinoides ruber
dominated the planktic foraminiferal fauna.
5.5. Paleoceanographic implications of modern
microplankton ecology in the Arabian Sea
Mechanistical understanding of modern diatom,
coccolithophore and planktic foraminiferal ecology
is essential for reconstructing paleoceanography and
paleoclimatology using their fossil remains (cf.
Fischer and Wefer, 1999; Henderson, 2002).
Planktic foraminifers are widely used to reconstruct
water temperature and ancient current systems by
transfer calculations, stable isotopes and element
ratios (e.g., Pflaumann et al., 1996; Schulz et al.,
2002; Rohling et al., 2004). Although diatoms are
suited for transfer calculations of marine paleotemperature in high latitudes (Zielinski et al.,
1998), they could not successfully be applied to
the low latitudes until now. However, diatoms add
information on the plankton succession and seem to
compete with coccolithophores for nutrients (Fig. 9)
(cf. Margalef, 1978). In addition, diatoms are part
of the foraminiferal diet (Hemleben et al., 1989).
Coccolithophores are the assumed main producer
of alkenones that are applied as paleo-thermometer
although their ecology is not fully understood to
date (e.g., Mix et al., 2000; Bard, 2001; Niebler et
al., 2003). Modern coccolithophores are negatively
related to nutrient concentration and, therefore,
during upwelling, alkenones are produced only to
a minor amount. Highest coccolithophore abundance occurs in regions with low nutrient concentration ( < 3 Amol NO3 l 1 and < 0.3 Amol PO4 l 1)
and possibly also during low-productive seasons,
namely non-upwelling periods (cf. Niebler et al.,
2003). Therefore, alkenones could not unequivocally
be applied to reconstruct water temperature during
upwelling.
Under a paleoclimatological perspective, upwelling regions in general, and the Arabian Sea in
particular, are of main interest, because a major
part of the global release of greenhouse gases, CO2,
N2O and CH4, takes place in these areas (Körtzinger et al., 1997; Patra et al., 1999; Bange et al.,
360
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
1999; Bange, 2000; Sarma et al., 2003). The
greenhouse gases are relevant to global climate
change. In turn, global climate change does trigger
productivity in the Arabian Sea through monsoonal
activity and its teleconnection to other major systems of the Earth’s climate, as, for example, ENSO
and NAO (Ivanova et al., 2003; Zahn, 2003).
Combining coccolithophore data with information on planktic foraminifers and diatoms will add
information on oceanographic settings with enhanced nutrient concentration (mesotrophic to eutrophic regions and high-productive seasons).
Seasonality is of particular importance to understand teleconnection of the climatic dipoles like
ENSO and the Indian Monsoon, because all of
these dipoles are seasonal features.
6. Conclusion
Different planktic foraminiferal, coccolithophore
and diatom species are adapted to differential scales
in trophic condition and hydrography. During SWM
in 1995 and 1997, the distribution of total coccolithophores and planktic foraminifers was positively
related, and both were negatively related to the
abundance of diatoms.
High valve numbers of diatoms, in particular
Chaetoceros spp. and Thalassionema nitzschioides,
were related to low water temperature and high
nutrient concentration caused by upwelling driven
by SWM winds in the northern Arabian Sea in
1995 (Fig. 10). To the southeast of the upwelling
area, diatom frequency decreased by two orders of
magnitude, and the species composition changed to
a dominance of N. bicapitata. The diatom assemblage further offshore points towards mixing of
different water masses and to the existence of
filaments.
Total planktic foraminiferal numbers were lower
in the upwelling area off Oman than in lower
productive waters to the southeast. In contrast,
Globigerinita bulloides and Neogloboquadrina
dutertrei numbers were increased along with upwelling and enhanced chlorophyll concentration.
Low productive waters were dominated by subtropical to tropical species Globigerinoides ruber and
Globigerinoides sacculifer.
According to nutrient concentration, coccolithophore cell numbers as well as diversity was low in
the upwelling area close to the Oman coast. Towards the open ocean, coccolithophore cell concentration increased along with decreasing nutrient
concentration and increasing stratification of the upper water column. Coccolithophore cell number is
possibly limited by thresholds in nutrient concentration. High standing stocks occur at low nitrate and
phosphate concentration. Below the lower threshold of
4 Amol NO3 and 0.35 Amol PO4 l 1 more than
70 103 coccolithophore cells l 1 occurred, and
above the upper threshold of 14 Amol NO3 and >1.25
Amol PO4 l 1 cell densities are lower than 10 103
cells l 1. The nutrient thresholds are similar for total
coccolithophore numbers and for the abundance of
single species.
The complex microplankton succession must be
understood in detail before each of its components,
diatoms, coccoliths and planktic foraminifers (and
others), can unequivocally be used as a paleoceanographic proxy. Models on plankton ecology
should be resolved at least for the seasonality
and for the hydrography on mesoscale, to account
for the ecological bias of, for example, transfer
function and alkenone derived paleotemperature
estimate.
Acknowledgements
The masters and crews of R/V METEOR
cruise 32/5 and R/V SONNE cruise 119 are
gratefully acknowledged for their technical assistance. We thank B. Hiller, M. Bayer and S. Flaiz
for support with sampling and preparation of
water and net samples, and T. Mitzka (Dept. of
Marine Planktology, IfM, Kiel University) and R.
Heuermann (Physical Institute, Oldenburg University) who provided CTD data. Nutrient data was
gratefully provided by P. Fritsche, I. Kriest and K.
Nachtigall (Meteor 32/5, Dept. of Marine Planktology, IfM, Kiel University, German JGOFS).
This project was funded by the German Ministry
of Technology and Science (BMBF), JGOFS-Indic
grant number 03F0183E, and by the German
Science Foundation (DFG), grant number He 697/
29 to CH.
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
361
Appendix A . Diatom species (103 valves l 1). Taxa are arranged according to absolute frequency
Cruise
SO119
Station
Water Depth (m)
Number of counted cells
Analyzed water volume (ml)
Counted area of filter (1250 mm2)
Filtered water (1)
Temperature (jC)
Salinity
Total Chl a (Ag l 1)
NO3 (Amol l 1)
PO4 (Amol l 1)
SiO4 (Amol l 1)
Taxa
37
20
194
0.53
0.24
2.80
25.99
36.25
n.d.a
0.15
0.10
0.27
Chaetoceros Hyalochaete
Thalassionema nitzschioides
Nitzschia bicapitata-group
Thalassiosira spp.
Stephanopyxis turris
Pseudo-nitzschia spp.
Tropidoneis-group
Eucampia spp.
Thalassiosira delicatula
Rhizosolenia spp.
Proboscia alata
Chaetoceros Phaeoceros
Nitzschia spp.
Corethron sp.
Nitzschia sicula
Pleurosigma spp.
Fragilariopsis pseudonana
Leptocylindrus mediterraneus
Dactyliosolen spp.
Bacteriastrum spp.
Thalassiosira partheneia/T. oceanica
Thalassiosira lineata
Rhizosolenia setigera
Cerataulina pelagica
Thalassionema nitzschioides var. parva
Chaetoceros resting spore
Stauroneis spp.
Navicula spp.
Fragilaria spp.
Asteromphalus sarcophagus
Porosira denticulata
Thalassiosira subtilis
Eunotia spp.
Roperia tesselata
Asteromphalus heptactis
Rhizosolenia af. antennata f. semispina
Coscinodiscus spp.
Thalassiothrix spp.
Climacodium frauenfeldianum
Pseudohimantidium pacificum
Azpeitia spp.
Centrale, other
Pennale, other
Total
144.4
100.4
–
16.9
–
1.9
3.8
9.4
13.1
5.6
5.6
–
–
–
–
–
–
3.8
3.8
1.9
–
–
1.9
–
–
1.9
1.9
–
–
–
–
–
–
–
–
–
–
–
–
–
–
30.0
16.9
363.0
a
n.d. = not determined.
M32-5
438
10
509
0.34
0.11
4.00
20.51
35.70
0.82
16.91
1.54
6.50
1303.6
178.6
–
–
32.7
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
1514.9
M32-5
M32-5
M32-5
M32-5
M32-5
M32-5
440
10
315
0.93
0.67
1.75
20.49
35.69
n.d.a
n.d.a
n.d.a
n.d.a
444
20
370
2.52
1.26
2.50
25.99
36.03
1.10
10.95
0.99
5.81
446
20
166
2.52
1.05
3.00
26.05
36.03
0.55
4.39
0.54
3.20
430
60
224
6.16
1.93
4.00
26.61
36.20
0.36
1.60
0.39
1.49
460
20
71
2.47
0.77
4.00
25.21
35.94
0.43
8.03
0.73
3.41
423
20
114
3.14
0.98
4.00
25.59
36.07
0.34
0.00
0.00
0.50
154.7
135.9
1.1
33.3
3.2
–
–
2.1
–
2.1
1.1
–
–
–
1.6
1.1
–
–
–
–
–
–
–
–
–
–
–
–
1.1
–
–
–
–
–
–
–
–
–
–
–
–
1.1
–
338.3
42.1
11.9
15.5
19.4
–
22.6
0.8
2.8
–
3.2
1.6
4.8
4.0
6.3
1.2
2.8
1.2
0.4
–
0.8
–
–
–
1.6
1.2
–
–
–
–
0.4
0.4
–
–
0.4
0.4
0.4
0.4
–
–
–
–
–
0.4
146.8
–
0.8
41.3
2.0
–
0.6
10.7
–
3.9
1.1
14.0
0.8
–
2.4
0.5
0.2
2.0
0.6
13.0
1.2
–
1.0
1.6
–
3.5
1.8
12.1
4.8
–
1.3
1.0
–
–
–
–
–
–
–
0.4
2.8
–
–
–
–
–
0.4
–
0.4
–
–
0.4
–
–
–
–
0.4
–
–
–
–
–
–
–
–
–
5.8
65.9
1.0
–
2.3
1.1
–
0.6
–
0.2
0.8
0.8
0.5
1.5
0.2
–
0.6
0.2
–
–
–
–
0.2
0.5
0.3
–
–
–
–
–
0.2
0.3
–
0.2
0.6
1.4
36.3
0.8
–
–
–
–
0.8
–
0.4
–
–
–
–
1.2
–
–
0.4
–
–
–
–
0.4
–
–
–
–
–
–
–
0.2
–
–
–
2.0
3.0
28.8
0.3
–
1.6
–
1.6
1.0
0.6
–
–
1.0
2.6
1.0
–
–
–
–
–
1.3
–
–
–
0.3
–
–
–
–
–
–
0.3
–
0.3
–
36.2
F. profunda var.
elongata
U. irregularis
O. antillarum
G. oceanica
C. leptoporus
E. huxleyi var.
huxleyi
G. flabellatus
R. xiphos
Ophiaster spp.
A. robusta
C. murrayi
F. profunda
with spine
S. nodosa
Syracosphaera spp.
M. adriaticus
U. hulburtiana
U. tenuis
D. tubifera
Corisphaera sp.
S. pulchra
M. elegans
H. triarcha
R. clavigera var.
stylifera
Gephyrocapsa sp.
S. dilatata
C. rigidus
S. prolongata
P. maximus
A. pinnigera
S. noroitica
A. quattrospina
S. lamina
37
20
18
37
60
42
37
100
7
2.55
3.03
3.79
1.136 0.946 0.946
2.80
25.99
36.25
n.da
0.15
0.10
0.27
4.00
24.65
36.37
n.da
1.96
0.50
1.60
36
20
51
) recorded during SO119. Taxa are arranged according to absolute frequency. Taxonomy refers to Jordan and Green (1994)
36
60
89
31
20
115
31
60
186
31
100
113
3.79
3.79
3.79
3.79
4.55
0.946
1.136 0.946 0.946 0.946
5.00
5.00
5.00
22.80 27.73 24.36
36.26 36.22 36.10
n.da
0.089 0.149
7.90
0.36
7.70
1.20
0.64
0.63
5.41
0.73
3.00
28
60
75
28
100
159
12
20
55
12
60
115
12
100
63
7
20
382
7
60
226
7
100
122
3
20
149
3
60
105
3
100
8
5.30
3.79
4.55
3.79
3.48
3.79
3.79
3.79
4.55
3.79
4.55
1.136
0.946
0.946
0.946
0.946
0.946
1.136
0.946
1.136 1.325 0.946
5.00
5.00
23.73 26.66
36.41 36.20
n.da
0.196
13.71 3.68
1.11
0.34
5.30
1.02
5.00
4.60
24.51 29.09
36.41 36.44
0.066 0.060
9.17
1.66
0.94
0.03
3.90
0.24
5.00
26.61
36.24
0.147
1.50
0.14
1.05
5.00
22.95
36.16
0.092
10.21
0.74
8.70
5.00
29.77
36.50
0.071
1.12
0.04
0.00
5.00
24.55
36.04
0.322
4.11
0.34
1.46
5.00
23.41
36.16
0.117
12.84
0.63
5.38
5.00
28.84
36.42
0.077
0.56
0.24
0.70
5.00
24.80
36.39
n.da
7.63
0.76
2.20
14.00
–
21.13
–
–
2.11
10.30
31.48
21.66
14.75
16.60 1.58
10.30 13.47
–
0.26
1.58 6.60
–
–
–
8.45
0.53
3.17
2.91
0.26
0.26
3.96
–
–
–
2.42
0.26
8.19
0.53
–
–
4.59
–
0.86
–
0.86
2.64
0.79
3.17
–
6.08
–
2.64
7.66
–
0.53
–
13.21
14.00
44.11
0.79
–
1.10
1.10
0.88
–
–
0.26
1.06
0.53
–
–
2.42
3.30
0.88
1.32
–
0.38
0.57
0.19
0.19
1.13 0.26
–
–
–
–
–
–
–
–
0.75 –
5.00
29.14
36.41
n.da
0.01
0.15
0.42
5.00
24.98
36.26
n.da
2.70
0.34
1.20
–
–
–
1.32 0.26
–
15.19
–
0.39
5.50
–
0.79
0.33 –
3.63 0.26
4.62 1.32
–
–
0.66 –
3.70
0.26
1.85
0.26
5.55
–
0.44
0.88
–
0.66
–
–
–
–
–
–
0.66
–
–
0.99
–
–
–
–
–
–
–
–
–
–
–
0.26
0.53
–
–
–
–
–
–
–
–
10.83
0.26
–
0.53
–
–
2.64
0.26
0.26
3.70
–
3.96
–
1.06
1.06
–
–
–
1.54
0.44
–
0.22
–
5.28
0.53
1.32
1.58
–
0.79
–
3.16
–
0.29
–
–
–
1.85
5.02
0.26
3.43
–
0.26
1.06
–
–
0.26
–
–
–
0.53
2.91
–
0.53
6.16
–
1.54
2.20
–
2.64
6.87
0.26
–
0.26
–
0.53
0.88
–
–
1.76
–
5.94
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
0.22
–
–
–
–
0.22
–
–
–
–
–
–
1.06
0.79
0.26
–
–
1.32
–
0.53
0.53
1.32
–
–
1.58
0.53
0.53
0.53
–
0.53
–
1.06
–
–
–
–
–
–
–
–
–
–
0.44
0.22
–
–
–
0.22
0.22
0.44
–
0.88
–
0.26
–
–
–
–
–
–
0.26
–
–
–
–
–
–
0.57
1.15
0.29
–
–
1.44
–
0.53
0.26
0.53
–
–
–
–
0.26
0.79
–
–
–
–
0.26
–
–
–
–
–
–
–
–
9.51
0.26
0.26
0.53
–
–
–
0.26
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
0.22
–
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.79
–
2.11
0.26
0.26
0.26
–
–
0.53
–
–
–
–
0.26
–
–
–
–
–
–
–
0.44
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.53
0.26
0.79
0.26
–
–
–
0.79
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
1.58
–
–
0.26
–
–
–
–
–
–
–
–
0.66
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.88
–
–
–
–
–
–
–
–
–
5.00
22.19
36.13
n.da
11.97
1.43
7.30
5
20
249
5
60
44
5
100
26
6
20
164
6
60
173
6
100
74
3.79
3.79
3.79
3.79
3.79
3.79
0.946
0.946
0.946
0.946
0.946
0.946
5.00
5.00
29.89 28.36
35.43 36.08
0.047 0.086
0.24
2.46
0.56
0.22
0.69
1.20
5.00
24.67
35.99
0.096
14.51
0.91
6.40
5.00
30.24
34.95
0.035
0.18
0.17
0.83
5.00
28.25
35.66
0.088
0.24
0.24
1.05
5.00
25.81
35.88
0.114
4.30
0.52
2.44
9.24
0.26
–
–
–
–
–
45.17
0.53
–
0.26
–
0.79
0.26
–
0.26
2.91
–
0.53
–
–
20.60
–
0.53
–
3.70
28.26
–
–
–
–
5.02
–
1.85
–
–
–
0.53
3.17
3.17
0.26
0.79
–
–
1.06
0.26
3.96
–
–
–
–
–
–
–
–
1.85
0.26
–
1.85
–
–
0.26
1.06
–
–
–
–
0.79
0.79
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
0.26
–
–
–
2.11
0.79
1.06
0.79
–
0.26
0.79
0.26
–
0.79
–
4.23
1.06
2.11
–
–
1.58
0.53
–
–
0.53
–
1.06
–
1.32
2.64
1.06
0.26
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
0.79
–
–
0.53
–
0.26
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
0.26
–
0.53
–
–
–
0.53
–
1.32
1.32
–
–
–
–
–
0.26
–
–
0.26
–
1.06
0.26
–
–
0.53
–
0.26
0.26
–
–
–
–
–
–
–
–
–
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
SO119, Station
Water depth (m)
Number of counted
specimens
Analyzed water
volume (ml)
Counted filter area
(mm2)
Filtered Water (l)
Temperature (jC)
Salinity
Total Chl a (Agl 1)
NO3 (Amol l 1)
PO4 (Amol l 1)
SiO4 (Amol l 1)
Taxa
1
362
Appendix B. Coccolithophore species (103 cells l
a
–
–
–
–
–
–
–
0.33
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.44
1.32
–
–
–
–
–
0.26
–
–
0.26
–
0.79
0.26
0.26
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
–
0.22
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.29
0.29
–
–
–
0.26
1.06
–
–
0.53
–
–
1.06
–
–
–
–
–
–
0.53
–
–
–
0.26
–
–
–
0.44
0.44
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.53
1.06
0.26
–
–
0.53
–
–
–
–
0.26
–
–
0.26
–
0.79
–
–
–
–
–
–
–
–
–
0.26
–
0.26
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
0.33
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.79
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
0.26
–
–
–
–
–
–
–
0.26
0.53
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
0.26
–
–
–
0.79
–
–
–
0.26
0.26
–
0.26
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
0.22
0.44
–
–
–
0.22
–
0.22
–
0.88
0.22
–
0.22
–
–
–
0.22
0.22
0.22
–
–
0.44
–
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
0.22
0.22
–
–
–
–
–
–
–
–
0.26
–
–
–
0.53
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
0.29
–
–
–
–
–
–
–
0.57
–
–
0.29
–
–
0.29
0.29
–
–
–
–
0.29
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
0.26
–
–
–
0.26
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.22
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.53
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
0.53
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
0.53
–
–
0.26
–
–
–
–
–
–
0.26
–
0.26
–
–
–
–
–
–
–
0.53
0.26
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
0.53
–
0.79
–
–
–
–
–
–
–
–
–
–
0.53
–
0.26
–
0.26
–
–
–
0.26
–
–
–
–
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
0.26
0.79
0.53
–
–
–
–
–
0.26
0.53
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.26
–
–
–
–
–
0.26
–
–
–
1.06
–
–
–
–
–
–
–
0.26
–
–
–
–
–
0.26
–
–
0.26
–
0.53
–
0.26
–
–
–
0.26
–
–
–
–
–
–
–
–
–
–
–
0.22
–
16.3
0.53
–
42.0
0.44
–
49.7
0.26
–
32.2
–
–
6.9
0.26
0.26
43.3
–
0.79
45.7
0.66 –
–
–
13.5 1.8
–
–
13.5
–
–
19.6
–
–
30.4
–
–
49.1
–
–
29.8
–
–
15.8
–
–
30.4
–
–
16.6
–
–
100.9
–
–
32.8
–
–
19.8
–
–
2.1
–
–
65.8
–
–
11.6
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
H. carteri var. carteri
–
P. lepida
–
G. corolla
–
A. brasiliensis
–
S. ossa
–
C. mediterranea
–
S. marginaporata
–
F. profunda var.
–
profunda
S. epigrosa
–
H. carteri var. hyalina
–
S. halldalii
–
C. gracilis
–
H. corn‘ifera
–
S. molischii
–
U. sibogae var. sibogae
–
C. multipora
–
S. orbiculus
0.39
C. papillifera
–
S. rotula
–
R. sessilis
–
C. oblonga
–
S. ampliora
–
S. pirus
–
P. poritectus
–
C. acuelata
–
A. periperforata
–
A. unicornis
–
S. anthos
–
C. cucullata
–
S. sp. II cf. S. epigrosa
–
P. galapagensis
–
C. diconstricta
–
O. fragilis var. fragilis
–
P. magnaghii
–
Syracolithus sp.
–
S. dilatata var. I
–
C. valliformis
–
S. exigua
–
A. fragaria
–
S. confusus
–
Z. hellenica
–
P. flabellifera
–
Acanthoica sp.
–
A. cidaris
–
A. spatula
–
T. latericioides
–
S. histrica
–
S. adenensis
–
Heterococcolithophore
–
spp.
Holococcolithophore spp. –
Indet spec.
–
Total
7.1
–
–
19.5
n.d. = not determined.
363
364
M32/5, Station
Depth (m)
Number of counted
specimens
Analyzed water
volume (ml)
Counted filter area
(mm2)
Filtered water (1)
Temperature (jC)
Salinity
Total Chl a (Ag l 1)
NO3 (Amol l 1)
PO4 (Amol l 1)
SiO4 (Amol l 1)
Taxa
G. oceanica
E. huxleyi var.
huxleyi
O. antillarum
F. profunda var.
elongata
C. murrayi
A. robusta
U. hulburtiana
C. leptoporus
C. mediterranea
Ophiaster spp.
C. rigidus
A. brasiliensis
G. flabellatus
H. carteri var.
carteri
Gephyrocapsa sp.
M. adriaticus
438
40
15
438
75
7
440
10
3
440
40
0
440
75
2
3.03
1.33
1.52
1.52
2.27
0.947 0.947 0.947 0.947 0.947
2.00
20.24
35.69
n.da
n.da
n.da
n.da
3.00
18.76
35.68
n.da
n.da
n.da
n.da
444
60
21
444
100
20
446
20
81
446
60
35
446
100
15
430
60
191
430
100
24
460
20
19
460
60
169
460
100
5
423
20
182
423
60
38
423
100
0
414
20
166
414
60
241
414
100
94
404
20
325
404
60
339
404
100
266
3.03
3.03
2.27
3.03
3.03
3.03
3.03
2.12
3.06
3.03
3.03
3.03
3.03
2.65
2.62
3.03
3.03
3.03
3.03
0.947
0.947
0.947
0.947
0.947
0.947
0.947
0.661
0.954
0.947
0.947
0.947
0.947
0.947
0.818
0.947
0.947
0.947
0.947
4.00
26
36.03
0.038
17.35
1.39
9.61
4.00
22.64
36.14
0.038
21.42
1.73
12.52
3.00
26.05
36.03
0.547
4.39
0.54
3.20
4.00
25.95
36.02
0.523
4.47
0.49
3.09
4.00
21.34
35.83
0.022
17.93
1.34
7.61
4.00
26.61
36.2
0.36
1.6
0.39
1.49
4.00
24.71
36.16
0.038
9.94
0.86
3.50
4.00
25.21
35.94
0.43
8.03
0.73
3.41
4.00
23.81
35.98
0.48
13.33
0.99
6.21
4.00
20.94
36.01
0.01
22.77
1.64
12.20
4.00
25.59
36.07
0.34
0.00
0.00
0.50
4.00
24.93
36.13
0.34
9.25
0.87
3.89
4.00
21.87
36.09
0.01
24.11
1.95
10.97
3.50
27.42
36.49
0.60
0.00
0.29
0.48
4.00
27.30
36.47
0.60
0.00
0.29
0.48
4.00
24.68
36.29
0.11
8.86
0.85
2.31
4.00
27.90
36.55
0.21
0.00
0.18
0.01
4.00
27.88
36.57
0.43
0.00
0.19
0.11
4.00
27.78
36.55
0.07
0.74
0.28
0.45
4.00
19.75
35.74
0.384
21.48
1.69
n.da
1.75
18.58
35.7
0.139
24.13
2.07
n.da
2.00
20.49
35.69
n.da
n.da
n.da
n.da
3.6
–
2.3
–
1.5
–
–
–
0.4
–
3.3
0.7
4.0
1.0
14.5
7.9
5.0
1.7
2.3
–
24.1
13.9
2.6
1.3
3.3
3.3
10.8
4.3
0.7
0.3
22.8
14.2
5.0
1.0
–
–
29.8
15.1
51.6
20.3
1.7
0.3
48.8
22.1
51.2
22.1
42.3
20.8
–
0.3
–
–
–
–
–
–
–
–
0.3
1.3
0.3
0.3
0.9
–
–
–
0.3
1.7
4.3
–
1.7
1.0
–
–
3.3
18.0
–
0.7
1.7
–
0.3
4.0
–
–
4.5
–
3.8
0.8
1.7
21.8
19.8
–
20.5
–
16.2
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.8
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.4
–
–
–
–
–
–
–
–
–
1.3
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
0.7
–
–
2.2
–
4.8
1.3
–
–
–
–
2.2
–
0.3
–
3.3
0.3
–
–
–
–
0.7
–
0.7
–
–
–
–
–
–
–
–
0.7
4.6
0.3
3.6
0.7
3.0
3.0
2.3
–
–
–
–
–
–
–
–
–
–
1.0
–
0.5
–
–
0.9
–
0.9
–
–
–
–
0.7
7.2
0.3
1.0
–
2.6
1.0
–
–
1.3
–
–
–
–
–
–
–
–
–
–
2.0
6.6
–
0.7
7.3
0.7
0.7
1.0
–
1.3
–
0.3
–
0.7
0.7
–
0.3
–
–
0.3
–
–
–
–
–
–
–
–
–
–
4.5
1.1
2.3
–
–
–
–
1.1
–
–
3.4
1.1
7.3
–
–
–
–
–
–
–
–
0.3
0.3
–
–
–
–
–
4.3
–
5.9
0.7
3.0
–
–
1.0
2.0
0.7
–
–
9.2
–
2.3
–
0.7
0.3
1.3
1.7
–
–
3.3
1.3
2.0
–
–
0.3
–
1.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
0.3
–
–
–
0.3
–
–
–
–
–
1.5
1.1
2.3
0.8
–
–
–
1.3
–
0.7
–
–
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Appendix C. Coccolithophore species (103 cells l 1) recorded during M32/5. Taxa are arranged according to absolute frequency. Taxonomy refers to Jordan and Green (1994)
a
n.d. = not determined.
–
–
–
–
–
–
1.8
–
–
0.3
0.3
–
0.7
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
1.0
1.0
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.8
–
–
–
–
–
0.4
0.4
–
0.4
–
–
–
0.4
–
–
–
–
–
–
–
–
–
0.3
–
–
0.3
–
–
0.3
–
–
–
0.7
0.3
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.7
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
0.3
–
0.3
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
0.3
0.7
–
–
–
–
–
–
–
–
–
–
0.3
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.3
0.3
0.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
2.3
–
2.3
–
0.0
–
0.9
–
6.9
–
6.6
–
35.7
–
11.6
–
5.0
0.3
63.1
–
7.9
–
9.0
0.3
55.4
–
1.7
–
60.1
–
12.5
–
0.0
–
62.6
–
92.1
–
31.0
–
107.1
–
111.9
–
87.8
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
U. sibogae var.
–
foliosa
M elegans
–
S. nodosa
–
A. quattrospina
–
P. lepida
–
T. quadrilaminata
–
A. unicornis
–
G. ericsonii
–
U. sibogae var.
–
sibogae
S. exigua
–
S. halldalii
–
Syracosphaera spp.
–
F. profunda var.
0.7
profunda
S. pirus
–
O. fragilis var.
–
fragilis
H. carteri var.
–
hyalina
S. lamina
–
S. orbiculus
–
S. adenensis
–
S. marginaporata
–
S. rotula
–
U. irregularis
–
R. xiphos
–
A. periperforata
–
Holococcolithophore –
spp.
Indet spec.
–
Total
5.0
365
366
Appendix D. Planktic foraminiferal species >125 Am (numbers m
5
0 – 20
357
29.92
35.42
0.051
0.00
0.90
0.65
Globigerinoides sacculifer
46.2
Globigerinoides ruber
6.8
Globigerina bulloides
–
Globigerinita glutinata
6.8
Neogloboquadrina dutertrei
2.4
Globigerina falconensis
–
Globoturborotalita tenella
–
Globorotalia menardii
–
Globoturborotalita rubescens
–
Globigerinella siphonifera
3.6
Globigerinita minuta
1.2
Pulleniatina obliquiloculata
–
Globigerinella calida
–
Globigerinoides conglobatus
–
Hastigerina pelagica
0.4
Globigerinella adamsi
–
Tenuitella parkerae
–
Globoquadrina conglomerata 2.4
Orbulina universa
–
Dentagloborotalia anfracta
–
Globigerinita uvula
–
Neogloboquadrina incompta
–
Globorotaloides hexagonus
–
Globorotalia scitula
–
Turborotalita quinqueloba
–
Globorotalia inflata
–
Globorotalia theyeri
–
Globorotalia tumida
–
Indet spec.
1.6
Total
71.4
a
n.d. = not determined.
) recorded during SO 119. Species are arranged according to absolute frequency
5
20 – 40
318
29.89
35.43
0.044
0.43
0.31
0.73
5
40 – 60
86
28.74
36.04
0.059
1.41
0.14
0.78
5
60 – 80
131
27.35
36.05
0.119
3.74
0.33
1.50
5
80 – 100
129
26.08
36.17
0.130
9.40
0.65
4.30
6
0 – 20
458
30.23
34.79
0.027
0.20
0.13
0.83
6
20 – 40
236
29.91
35.44
0.041
0.16
0.20
0.83
6
40 – 60
86
28.87
35.64
0.065
0.21
0.22
0.83
6
60 – 80
109
27.43
35.67
0.120
0.29
0.27
1.10
6
80 – 100
122
26.06
35.64
0.145
1.88
0.39
1.80
7
0 – 20
758
29.92
36.51
0.051
1.27
0.01
0.00
7
20 – 40
525
29.51
36.44
0.091
0.97
0.07
0.00
38.6
4.6
0.4
6.4
3.2
–
1.4
0.6
–
3.6
–
–
0.6
0.2
0.2
–
–
2.4
–
–
–
–
0.2
–
–
–
–
–
1.2
63.6
6.2
1.0
–
–
2.4
–
0.2
1.2
0.2
3.6
–
–
–
–
0.2
1.8
–
–
–
–
–
–
0.2
–
–
–
–
–
0.2
17.2
6.2
1.8
–
1.6
2.4
–
0.8
2.6
–
7.4
–
–
–
–
0.8
1.8
–
0.4
0.2
–
–
–
–
–
–
–
–
–
0.2
26.2
4.6
2.4
0.2
1.8
1.6
–
2.8
0.4
0.6
8.8
0.6
–
–
–
1.0
0.2
–
0.2
0.2
–
–
–
–
–
–
–
–
–
0.4
25.8
58.2
15.6
–
6.8
2.2
–
2.8
0.8
–
2.2
0.8
–
–
–
–
0.6
–
1.0
–
–
–
–
0.6
–
–
–
–
–
–
91.6
26.6
10.8
0.2
2.2
0.8
–
1.2
1.2
–
0.4
1.6
–
–
–
–
0.6
–
0.2
1.0
–
–
–
0.2
–
–
–
0.2
–
–
47.2
3.0
5.8
0.6
0.6
–
–
1.0
0.8
–
0.8
0.2
–
–
–
–
0.4
–
–
2.6
–
–
–
1.2
–
–
–
–
–
0.2
17.2
1.0
10.0
0.4
0.6
–
–
2.2
1.8
–
0.2
1.4
–
–
–
–
1.4
–
0.2
0.8
–
–
–
1.8
–
–
–
–
–
–
21.8
1.4
6.2
0.4
2.8
0.2
–
5.0
1.2
0.8
0.2
0.6
–
0.8
2.0
0.2
1.4
–
–
0.2
–
–
–
0.6
–
–
–
–
–
0.4
24.4
35.4
7.8
88.6 81.2
–
0.2
1.6
–
1.6
0.2
–
–
16.8
9.0
0.2
–
–
0.2
0.8
1.0
–
–
–
–
–
0.2
5.2
4.8
–
–
0.2
–
–
–
–
–
1.0
–
–
–
–
–
–
0.2
–
–
–
–
–
–
–
–
–
–
–
–
0.2
0.2
151.6 105.0
7
40 – 60
144
25.87
36.11
0.395
2.11
0.27
0.53
28
0 – 20
242
29.96
36.49
0.046
1.99
0.18
0.80
28
20 – 40
242
29.89
36.48
0.076
1.76
0.12
1.04
28
40 – 60
173
27.95
36.24
0.150
2.88
0.24
1.00
28
60 – 80
86
25.41
36.18
0.156
5.68
0.52
1.40
28
80 – 100
51
24.65
36.37
0.156
8.41
0.52
2.83
37
0 – 20
213
28.07
36.26
0.075
0.33
0.10
0.25
37
20 – 40
75
27.6
36.23
n.da
0.00
0.10
0.32
37
40 – 60
40
25.07
36.14
n.da
0.05
0.34
0.74
37
60 – 80
24
23.72
36.06
n.da
3.67
0.65
2.50
37
80 – 100
85
23.02
36.13
n.da
6.61
1.00
4.60
4.6
13.0
0.4
0.4
–
–
9.2
–
0.2
0.4
0.2
–
–
0.4
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
28.8
20.2
17.0
0.4
6.6
2.6
–
0.4
–
–
0.2
–
–
–
–
–
–
–
–
0.4
–
–
–
–
–
–
–
–
–
0.6
48.4
22.4
16.2
0.2
6.8
0.8
0.4
–
0.6
–
–
–
–
0.8
–
0.2
–
–
–
–
–
–
–
–
–
–
–
–
–
–
48.4
11.2
14.8
–
1.4
1.8
–
–
0.4
–
0.2
–
–
–
0.8
0.2
–
–
–
0.2
–
–
–
–
3.4
–
–
–
–
0.2
34.6
6.6
3.6
0.2
0.8
2.8
–
0.4
0.4
–
0.4
–
–
0.4
0.2
–
–
–
–
1.0
–
–
–
–
0.4
–
–
–
–
–
17.2
0.2
2.8
–
0.2
2.8
–
–
1.4
–
–
–
0.2
1.0
–
–
–
–
–
0.8
–
–
–
–
0.8
–
–
–
–
–
10.2
1.2
1.4
14.6
2.2
18.2
–
0.2
–
–
0.2
–
–
3.0
–
–
0.4
–
–
–
–
–
–
–
–
–
–
–
–
1.2
42.6
–
–
10.2
1.0
2.4
–
–
–
–
–
–
–
1.0
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.4
15.0
–
0.4
4.0
0.2
1.6
–
–
–
–
0.4
–
–
1.0
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.2
0.2
8.0
–
0.4
1.0
0.4
0.8
–
–
–
–
0.4
–
0.2
1.4
–
–
0.2
–
–
–
–
–
–
–
–
–
–
–
–
–
4.8
0.6
0.4
4.4
1.8
1.8
0.4
0.6
–
1.6
1.2
0.8
0.2
2.8
–
–
0.2
–
–
–
–
–
–
–
–
–
–
–
–
0.2
17.0
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
SO 119, Station
Water depth (m)
Number of counted tests
Temperature (jC)
Salinity
Total Chl a (Ag l 1)
NO3 (Amol l 1)
PO4 (Amol l 1)
SiO4 (Amol l 1)
Species
3
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
367
Appendix E. Planktic foraminiferal species >125 Am (numbers m 3) recorded during M32/5. Species are
M 32/5, Station
Water depth (m)
Number of counted tests
Temperature (jC)
Salinity
Total Chl a (Ag l 1)
NO3 (Amol l 1)
PO4 (Amol l 1)
SiO4 (Amol l 1)
Species
414
0 – 20
1215
27.42
36.49
0.672
0.00
0.29
0.49
414
20 – 40
891
27.36
36.47
0.552
0.00
0.29
0.37
414
40 – 60
908
27.31
36.48
0.576
0.00
0.29
0.40
414
60 – 80
1166
27.29
36.47
0.576
4.42
0.57
1.23
414
80 – 100
542
24.68
36.29
0.187
8.86
0.85
2.31
430
0 – 20
1732
26.65
36.22
0.336
1.45
0.38
1.50
430
20 – 40
882
26.66
36.22
0.360
1.39
0.38
1.49
430
40 – 60
1344
26.62
36.21
0.360
1.50
0.39
1.47
430
60 – 80
2165
26.61
36.20
0.228
3.42
0.51
1.94
430
80 – 100
471
24.71
36.16
0.067
7.59
0.75
2.95
Globigerinoides sacculifer
Globigerinoides ruber
Globigerina bulloides
Globigerinita glutinata
Neogloboquadrina dutertrei
Globigerina falconensis
Globoturborotalita tenella
Globorotalia menardii
Globoturborotalita rubescens
Globigerinella siphonifera
Globigerinita minuta
Pulleniatina obliquiloculata
Globigerinella calida
Globigerinoides conglobatus
Hastigerina pelagica
Globigerinella adamsi
Tenuitella parkerae
Globoquadrina conglomerata
Orbulina universa
Dentagloborotalia anfracta
Globigerinita uvula
Neogloboquadrina incompta
Globorotaloides hexagonus
Globorotalia scitula
Turborotalita quinqueloba
Globorotalia inflata
Globorotalia theyeri
Globorotalia tumida
Indet spec.
Total
18.4
15.2
34.2
29.4
33.4
25.2
34.0
6.8
16.0
4.4
13.2
7.2
1.6
–
–
–
–
–
–
0.8
1.6
–
–
–
1.6
–
–
–
–
243.0
21.4
17.6
17.0
14.2
25.2
24.4
24.0
2.8
7.2
3.0
6.4
12.8
2.0
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0.2
178.2
12.8
13.4
29.0
19.4
22.2
19.6
27.0
3.6
10.4
4.0
4.8
9.8
2.8
–
2.2
–
–
–
–
–
–
–
–
–
–
0.2
–
–
0.4
181.6
21.2
17.8
36.4
17.2
14.4
51.2
23.6
7.4
14.4
7.2
12.8
8.0
0.8
–
–
–
–
–
–
0.8
–
–
–
–
–
–
–
–
–
233.2
10.4
10.4
15.6
8.4
12.6
8.0
14.0
3.0
8.8
1.2
8.8
2.4
2.4
–
–
–
–
2.4
–
–
–
–
–
–
–
–
–
–
–
108.4
32.6
29.2
28.8
53.6
40.4
55.2
12.8
39.4
12.8
18.6
8.0
11.6
–
–
1.6
–
–
0.2
–
1.6
–
–
–
–
–
–
–
–
–
346.4
19.8
10.8
26.6
21.4
19.0
23.0
5.6
18.6
7.2
7.2
3.2
5.2
–
–
2.0
–
2.4
–
–
–
1.6
2.4
–
–
–
–
–
–
0.4
176.4
25.2
22.0
74.0
33.0
21.0
17.6
24.0
18.2
9.6
8.8
9.6
4.6
–
–
–
0.4
0.8
–
–
–
–
–
–
–
–
–
–
–
–
268.8
40.6
43.8
83.8
58.8
44.4
28.8
13.6
50.6
16.8
19.8
4.8
11.4
–
–
3.6
2.0
5.6
–
–
0.8
2.4
–
–
–
–
–
–
–
1.4
433.0
5.2
7.6
14.4
11.4
7.2
5.6
3.8
9.6
10.0
4.6
0.6
2.2
2.2
–
0.4
–
1.0
–
0.2
3.6
1.4
2.2
–
–
0.2
–
–
–
0.8
94.2
1
n.d. = not determined.
References
Abrantes, F., 1988. Diatom assemblages as upwelling indicators in
surface sediments off Portugal. Marine Geology 85, 15 – 39.
Ahagon, N., Tanaka, Y., Ujiie, H., 1993. Florisphaera profunda, a
possible nannoplankton indicator of late Quaternary changes in
sea-water turbidity at the northwestern margin of the Pacific.
Marine Micropaleontology 22, 255 – 273.
Andruleit, H., Rogalla, U., 2002. Coccolithophores in surface
sediments of the Arabian Sea in relation to environmental gradients in surface waters. Marine Geology 186,
505 – 526.
Andruleit, H.A., von Rad, U., Bruns, A., Ittekkot, V., 2000. Coccolithophore fluxes from sediment traps in the northeastern Arabian Sea off Pakistan. Marine Micropaleontology 38 (3 – 4),
285 – 308.
Andruleit, H., Stäger, S., Rogalla, U., Čepek, P., 2003. Living
coccolithophores in the northern Arabian Sea: ecological tol-
368
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
erances and environmental control. Marine Micropaleontology
49, 157 – 181.
Antoine, D., André, J.M., Morel, A., 1996. Oceanic primary production: 2. Estimation at global scale from satellite (Coastal
Zone Color Scanner) chlorophyll. Global Biogeochemical
Cycles 10 (1), 57 – 69.
Ashok, K., Guan, Z., Yamagata, T., 2003. Influence of the Indian
Ocean Dipole on the Australian winter rainfall. Geophysical
Research Letters 30 (15), 1821 (doi: 10.1029/2003GL017926).
Bange, H.W., 2000. Global change—it’s not a gas. Nature 408,
301 – 302.
Bange, H.W., Breves, W., Mitzka, T., Lendt, R., Petuhov, K., Hupe,
A., Rapsomanikis, S., Andreae, M.O., Reuter, R., Zeitzschel, B.,
Ittekkot, V., 1999. The surface distribution of nutrients, chlorophyll and trace gases (CO2, NO2, CH4) in the upwelling area of
the northwestern Arabian Sea during the SW monsoon 1997. In:
Schlünz, B., Wefer, G.Bericht über den 7. JGOFS-Workshop.
Berichte aus dem Fachbereich Geowissenschaften der Universität Bremen, vol. 131, pp. 12 – 13.
Bard, E., 2001. Comparison of alkenones estimate with other paleotemperature proxies. Geochemistry Geophysics Geosystems 2
(2000GC000050), 1 – 12.
Bauer, S., Hitchcock, G.L., Olson, D.B., 1991. Influence of monsoonally-forced Ekman dynamics upon surface layer depth and
plankton biomass distribution in the Arabian Sea. Deep-Sea
Research 18 (5), 531 – 553.
Bé, A.W.H., Hutson, W.H., 1977. Ecology of planktonic foraminifera and biogeographic patterns of life and fossil assemblages in
the Indian Ocean. Micropaleontology 23 (4), 369 – 414.
Bé, A.W.H., Tolderlund, D.S., 1971. Distribution and ecology of
living planktonic foraminifera in surface waters of the Atlantic and Indian Oceans. In: Funnel, B.M., Riedel, W.R.Micropaleontology of the Oceans. Cambridge Univ. Press,
Cambridge, pp. 105 – 149.
Berger, W.H., Fischer, K., Lai, C., Wu, G., 1988. Ocean carbon
flux: global maps of primary production and export production.
In: Agegioan, C.R. (Ed.), Biogeochemical Cycling and Fluxes
between the Deep Euphotic Zone and Other Oceanic Realms,
NOAA National Undersea Research Program, Research Reports
88-1, pp. 131 – 176.
Bernard, F., Lecal, J., 1960. Plancton unicellulaire recolte dans
l’ocean Indien par le Charcot (1950) et le Norsel (1955 – 56).
Bulletin de l’Institut Oceanographique 1166, 1 – 59.
Birkenes, E., Braarud, T., 1952. Phytoplankton in the Oslo Fjord
during a ‘Coccolithus huxleyi-summer’. Avhandlinger - Det
Norske Videnskaps-Akademi i Oslo 1952/2, 1 – 23.
Blasco, D., Estrada, M., Jones, B., 1980. Relationship between the
phytoplankton distribution and composition and the hydrography in the northwest African upwelling region near Cabo Corbeiro. Deep-Sea Research 27A, 799 – 821.
Bollmann, J., 1997. Morphology and biogeography of Gephyrocapsa coccoliths in Holocene sediments. Marine Micropaleontology 29 (3 – 4), 319 – 350.
Brink, K.H., Cowles, T.J., 1991. The coastal transition program.
Journal of Geophysical Research 96 (C8), 14637 – 14647.
Brock, J.C., McClain, Ch.R., Luther, M.A., Hay, W.W., 1991. The
phytoplankton bloom in the northwestern Arabian Sea during
the southwest monsoon of 1979. Journal of Geophysical Research 96 (C11), 20623 – 20642.
Brock, J.C., McClain, Ch.R., Hay, W.W., 1992. A southwest monsoon hydrographic climatology for the northwestern Arabian
Sea. Journal of Geophysical Research 97 (6), 9455 – 9465.
Broerse, A.T.C., Brummer, G.J.A., Van Hinte, J.E., 2000. Coccolithophore export production in response to monsoonal upwelling
off Somalia (northwestern Indian Ocean). Deep-Sea Research II
47 (9 – 11), 2179 – 2205.
Burkill, P.H., Archer, S.D., Robinson, C., Nightingale, P.D.,
Groom, S.B., Tarran, G.A., Zubkov, M.V., 2002. Dimethyl sulphide biogeochemistry within a coccolithophore bloom (DISCO): an overview. Deep-Sea Research II 49, 2863 – 2885.
Čepek, M., 1996. Zeitliche und räumliche Variationen von Coccolithophoriden-Gemeinschaften im subtropischen Ost-Atlantik:
Untersuchungen an Plankton, Sinkstoffen und Sedimenten.
Berichte aus dem Fachbereich Geowissenschaften der Universität Bremen 86 (160 pp.).
Cleve, P.T., 1901. Plankton from the Indian Ocean and the Malay
Archipelago. Kungliga Svenska Vetenskapsakademiens Handlingar 35 (5), 1 – 58.
Conan, S.M.-H., Brummer, G.-J., 2000. Fluxes of planktic Foraminifera in response to monsoonal upwelling on the Somalia Basin
margin. Deep-Sea Research II 47 (9 – 10), 2207 – 2227.
Cortés, M.Y., Bollmann, J., Thierstein, H.R., 2001. Coccolithophore ecology at the HOT station ALOHA, Hawaii. Deep-Sea
Research II 48, 1957 – 1981.
Cupp, E.E., 1943. Marine Plankton Diatoms of the West Coast of
North America. University of California Press, Berkeley and Los
Angelos; Reprint 1977, Koeltz Science Publishers, Koenigstein.
Curry, W.B., Ostermann, D.R., Guptha, M.V.S., Ittekkot, V., 1992.
Foraminiferal production and monsoonal upwelling in the Arabian Sea: evidence from sediment traps. In: Summerhays, C.P.,
Prell, W.S., Emeis, K.C.Upwelling Systems: Evolution Since
the Early Miocene. Special Publication - Geological Society,
vol. 64, pp. 93 – 106.
Durairatnam, M., 1964. Some planktonic diatoms from the Indian
Ocean. Bulletin of the Fisheries Research Station, Ceylon 17
(2), 159 – 168.
Findlater, J., 1971. Mean monthly air flow at low levels over the
western Indian Ocean. Geophysical Memoires 115, 53.
Fischer, G., Wefer, G., 1999. Use of Proxies in Paleoceanography.
Examples from the South Atlantic. Springer-Verlag, New York.
735 pp.
Fleitmann, D., Burns, S.J., Mudelsee, M., Neff, U., Kramers, J.,
Mangini, A., Matter, A., 2003. Holocene forcing of the Indian
Monsoon recorded in a stalagmite from southern Oman. Science
300, 1737 – 1739.
Garrison, D.L., Gowing, M.M., Hughes, M.P., Campbell, L., Caron,
D.A., Dennett, M.R., Shalapyonok, A., Olson, R.J., Landry,
M.R., Brown, S.L., Liu, H.-B., Azam, F., Steward, G.F.,
Ducklow, H.W., Smith, D.C., 2000. Microbial food web structure
in the Arabian Sea: a US JGOFS study. Deep-Sea Research II 47,
1387 – 1422.
Giraudeau, J., 1992. Distribution of recent nannofossils beneath the
Benguela system: southwest African continental margin. Marine
Geology 108, 219 – 237.
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Grasshoff, K., Erhardt, M., Kremling, K., 1988. Methods of Seawater Analysis. Verlag Chemie, Weinheim. 419 pp.
Gupta, A.K., Anderson, D.M., Overpeck, J.T., 2003. Abrupt changes
in the Asian southwest monsoon during the Holocene and their
links to the North Atlantic Ocean. Nature 421 (23), 354 – 357.
Guptha, M.V.S., 1985. Distribution of calcareous nannoplankton
from the sediments of the northwestern continental shelf of
India. Journal of the Geological Society of India 26, 267 – 274.
Guptha, M.V.S., Rahul, M., Muralinath, S., 1994. Living planktonic
foraminifera during the late summer monsoon period in the
Arabian Sea. Marine Geology 120, 365 – 371.
Guptha, M.V.S., Mohan, R., Muralinath, A.S., 1995. Living coccolithophorids from the Arabian Sea. Rivista Italiana di Paleontologia e Stratigrafia 100 (4), 551 – 574.
Haidar, A.T., Thierstein, H.R., 2001. Coccolithophore dynamics off
Bermuda (N. Atlantic). Deep-Sea Research II 48, 1925 – 1956.
Hasle, G.R., 1965. Nitzschia and Fragilariopsis species studied in
the light and electron microscope: II. The group Pseudonitzschia. Det Norske Videndskaps-Akademi i Oslo. I, Matematisk-Naturvitenskapelig Klasse. Ny Serie 18, 1 – 45.
Hemleben, Ch., Spindler, M., Anderson, O.R., 1989. Modern
Planktonic Foraminifera. Springer Verlag, Berlin. 363 pp.
Henderson, G.M., 2002. New oceanic proxies for paleoclimate.
Earth and Planetary Science Letters 203, 1 – 13.
Hendey, N.I., 1970. Some littoral diatoms of Kuwait. Nova Hedwigia. Beihefte 31, 101 – 167.
Herbland, A., Le Bouteller, A., Raimbault, P., 1985. Size structure
of phytoplankton biomass in the equatorial Atlantic Ocean.
Deep-Sea Research 32, 819 – 836.
Houghton, S.D., Guptha, S.M.V., 1991. Monsoonal and fertility
controls on recent marginal sea and continental shelf coccolith
assemblages from the western Pacific and northern Indian
oceans. Marine Geology 97 (3 – 4), 251 – 259.
Ittekkot, V., Nair, R.R., 1993. Monsoon biogeochemistry. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 76, 1 – 192.
Ivanova, E.M., Conan, S.M.-H., Peeters, F.J.C., Troelstra, S.R.,
1998. Living Neogloboquadrina pachyderma sin and its distribution in the sediments from Oman and Somalia upwelling
areas. Marine Micropaleontology 36, 91 – 107.
Ivanova, E., Schiebel, R., Singh, A.D., Schmiedl, G., Niebler,
H.-S., Hemleben, Ch., 2003. Primary production in the Arabian Sea during the last 135,000 years. Palaeogeography,
Palaeoclimatology, Palaeoecology 197, 61 – 82.
Jordan, R.W., Green, J.C., 1994. A check-list of the extant haptophyta of the world. Journal of the Marine Biological Association
of the United Kingdom 75, 769 – 814.
Karsten, G., 1907. Das indische Phytoplankton nach dem Material
der deutschen Tiefsee-Expedition 1898 – 1899. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition 2, 223 – 544.
Kiefer, A. 1999. Reaktionen planktischer Foraminiferen auf trophische Unterschiede in der produktiven Zone des Arabischen Meeres. Tübingen University, Diploma thesis, 1 – 44.
[unpublished].
Kleijne, A., 1991. Holococcolithophorids from the Indian Ocean,
Red Sea, Mediterranean Sea and North Atlantic Ocean. Marine
Micropaleontology 17 (1 – 2), 1 – 76.
369
Kleijne, A., 1992. Extant Rhabdosphaeraceae (coccolithophorids,
class Prymnesiophyceae) from the Indian Ocean, Red Sea, Mediterranean Sea and North Atlantic Ocean. Scripta Geologica 65,
1 – 100.
Kleijne, A. 1993. Morphology, taxonomy and distribution of extant
coccolithophorids (calcareous nannoplankton) PhD thesis, Vrjie
Univ., 321pp.
Kleijne, A., Kroon, D., Zevenboom, W., 1989. Phytoplankton and
foraminiferal frequencies in northern Indian Ocean and Red Sea
waters. Netherlands Journal of Sea Research 24, 531 – 539.
Knappertsbusch, M., Cortes, M.Y., Thierstein, H.R., 1997. Morphologic variability of the coccolithophorid Calcidiscus leptoporus
in the plankton, surface sediments and from the Early Pleistocene. Marine Micropaleontology 30, 293 – 317.
Körtzinger, A., Duinker, J.C., Mintrop, L., 1997. Strong CO2 emissions from the Arabian Sea during South-West Monsoon. Geophysical Research Letters 24 (14), 1763 – 1766.
Kroon, D., 1988. Distribution of extant planktic foraminiferal
assemblages in Red Sea and northern Indian Ocean surface
waters. In: Brummer, G.-J.A., Kroon, D. (Eds.), Planktonic Foraminifers as Tracers of Ocean Climate History, pp. 229 – 267.
Kroon, D., Ganssen, G., 1988. Northern Indian Ocean upwelling
cells and the stable isotope composition of living planktic foraminifera. In: Brummer, G.J.A., Kroon, D.Planktonic Foraminifers as Tracers of Ocean Climate History, pp. 299 – 317.
Lange, C.B., Treppke, U.F., Fischer, G., 1994. Seasonal diatom
fluxes in the Guinea Basin and their relationship to trade winds,
hydrography and upwelling events. Deep-Sea Research 41 (5/6),
859 – 878.
Manghnani, V., Morrison, J.M., Hopkins, T.S., Böhm, E., 1998.
Advection of upwelled waters in the form of plumes off Oman
during Southwest Monsoon. Deep-Sea Research II 45 (10/11),
2027 – 2052.
Margalef, R., 1978. Life-forms of phytoplankton as survival alternatives in an unstable environment. Oceanologica Acta 1 (4),
493 – 509.
Martini, E., Müller, C., 1972. Nannoplankton aus dem nördlichen
Arabischen Meer. ‘‘Meteor’’ Forschungsergebnisse, Reihe C 10,
63 – 74.
Mathur, K., Singh, K., 1993. Diatom flora from the Arabian Sea.
Geoscience Journal 14 (1/2), 67 – 84.
Mitchell-Innes, B.A., Winter, A., 1987. Coccolithophores: a major
phytoplankton component in mature upwelled waters off the
Cape Peninsula, South Africa in March, 1983. Marine Biology
95, 25 – 30.
Mix, A.C., Bard, E., Eglinton, G., Keigwin, L.D., Ravelo, A.C.,
Rosenthal, Y., 2000. Alkenones and multiproxy strategies in
paleoceanographic studies. Geochemistry Geophysics Geosystem 1 (2000GC000056), 1 – 22.
Naidu, P.D., Malmgren, B.A., 1996. Relationship between late Quaternary upwelling history and coiling properties of Neogloboquadrina pachyderma and Globigerina bulloides in the Arabian
Sea. Journal of Foraminiferal Research 26 (1), 64 – 70.
Niebler, H.-S., Arz, H.W., Donner, B., Mulitza, S., Pätzold, J.,
Wefer, G., 2003. Sea surface temperatures in the equatorial
and South Atlantic Ocean during the Last Glacial Maximum
(23 – 19 ka). Paleoceanography 18 (3), 14-1 – 14-12.
370
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
Norris, R.E., 1965. Living cells of Ceratolithus cristatus (Coccolithophorineae). Archiv für Protistenkunde 108, 19 – 24.
Norris, R.E., 1971. Extant calcareous nannoplankton from the Indian Ocean. In: Farinacci, A.Proceedings of the II Planktonic
Conference, Roma 1970, vol. 2. Editioni Tecnoscienza, Rome,
pp. 899 – 909.
Norris, R.E., 1983. The family position of Papposphaera Tangen
and Pappomonas Manton and Oates (Prymnesiophyceae) with
record from the Indian Ocean. Phycologia 22 (2), 161 – 169.
Norris, R.E., 1984. Indian Ocean nannoplankton: I. Rhabdosphaeraceae (Prymnesiophyceae) with review of extant taxa. Journal of
Phycology 20, 27 – 41.
Norris, R.E., 1985. Indian Ocean Nannoplankton: II. Holococcolithophorids (Calyptrosphaeraceae, Prymnesiophyceae) with a review of extant genera. Journal of Phycology 21, 619 – 641.
Okada, H., Honjo, S., 1973. The distribution of oceanic coccolithophorids in the Pacific. Deep-Sea Research 20, 355 – 374.
Patra, P.K., Lal, S., Venkataramani, S., de Sousa, S.N., Sarma,
V.V.S.S., Sardesai, S., 1999. Seasonal and spatial variability in
N2O distribution in the Arabian Sea. Deep-Sea Research. Part I 46
(3), 529 – 543.
Pflaumann, U., Duprat, J., Pujol, C., Labeyrie, L.D., 1996. SIMMAX: a modern analog technique to deduce Atlantic sea surface
temperatures from planktonic foraminifera in deep-sea sediments. Paleoceanography 11 (1), 15 – 35.
Pitcher, G.C., Walker, D.R., Mitchell-Innes, B.A., Moloney, C.L.,
1991. Short-term variability during an anchor station study in
the southern Benguela upwelling system: phytoplankton dynamics. Progress in Oceanography 28, 39 – 64.
Prasad, R.R., Nair, P.V.R., 1960. Observations on the distribution
and occurrence of diatoms in the inshore waters of the Gulf of
Mannar and Palk Bay. Indian Journal of Fisheries 7, 49 – 68.
Reid, F.M.H., 1980. Coccolithophorids of the North Pacific Central
Gyre with notes on their vertical and seasonal distribution. Micropaleontology 26, 151 – 176.
Rixen, T., Haake, B., Ittekkot, V., 2000. Sedimentation in the western Arabian Sea the role of coastal and open-ocean upwelling.
Deep-Sea Research II 47 (9 – 11), 2155 – 2178.
Rohling, E.J., Sprovieri, M., Cane, T., Casford, J.S.L., Cooke,
S., Bouloubassi, I., Emeis, K.C., Schiebel, R., Rogerson, M.,
Hayes, A., Jorissen, F.J., Kroon, D., 2004. Reconstructing past
planktic foraminiferal habitats using stable isotope data: a case
history for Mediterranean sapropel S5. Marine Micropaleontology 50, 89 – 123.
Sarma, V.V.S.S., Swathi, P.S., Dileep Kumar, M., Prasannakumar,
S., Bhattathiri, P.M.A., Madhupratap, M., Ramaswamy, V.,
Sarin, M.M., Gauns, M., Ramaiah, N., Sardessai, S., de Sousa,
S.N., 2003. Carbon budget in the eastern and central Arabian Sea:
an Indian JGOFS synthesis. Global Biogeochemical Cycles 17
(4), 13-1 – 13-13.
Schiebel, R., Waniek, J., Bork, M., Hemleben, Ch., 2001. Planktic
foraminiferal production stimulated by chlorophyll redistribution and entrainment of nutrients. Deep-Sea Research I 48,
721 – 740.
Schrader, H., Gersonde, R., 1978. Diatoms and silicoflagellates. In:
Zachariasse, W.J., et al.Micropaleontological counting methods
and techniques—an exercise on an eight metres section of the
lower Pliocene of Capo Rosello, Sicily. Utrecht Micropaleontological Bulletins, vol. 17, pp. 129 – 176.
Schulz, H., von Rad, U., Ittekkot, V., 2002. Planktic foraminifera, particle flux and oceanic productivity off Pakistan, NE
Arabian Sea: modern analogues and application to the paleoclimatic record. In: Clift, P.D., Kroon, D., Gaedicke, C.,
Craig, J.The Tectonic and Climatic Evolution of the Arabian
Sea Region. The Geological Society of London, vol. 195,
pp. 499 – 516.
Shenoi, S.S.C., Shetye, S.R., Gouveia, A.D., Michael, G.S., 1993.
Saliniy extrema in the Arabian Sea. In: Ittekkot, V., Nair,
R.R.Monsoon Biogeochemistry. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg, vol.
76, pp. 37 – 49.
Simonsen, R., 1974. The diatom plankton of the Indian ocean expedition of R/V ‘‘Meteor’’ 1964 – 1965. ‘‘Meteor’’ Forschungsergebnisse, Reihe D 19, 1 – 107.
Smayda, T.J., 1986. Phytoplankton species succession. In: Morris,
I.The Physiological Ecology of Phytoplankton. Studies in Ecology, vol. 7, pp. 493 – 570.
Smith, R.L., Bottero, J.S., 1977. On upwelling in the Arabian Sea.
In: Angel, M.V.A voyage of Discovery. Deep-Sea Research,
Supplement, pp. 291 – 304.
Sournia, A., 1968. Diatomées planctoniques du Canal de Mozambique et de l’ile Maurice. Mémoire O.R.S.T.O.M. 31, 1 – 120.
Strub, P.T., Kosro, P.M., Huyer, A., 1991. The nature of the cold
filaments in the California current system. Journal of Geophysical Research 96 (8), 14743 – 14768.
Swallow, J.C., Molinari, R.L., Bruce, J.G., Brown, O.B., Evans,
R.H., 1983. Development of near-surface flow pattern and water
mass-distribution in the Somali Basin in response to the southwest monsoon of 1979. Journal of Physical Oceanography 13 (8),
1398 – 1415.
Taylor, F.J.R., 1967. Phytoplankton of the south western Indian
Ocean. Nova Hedwigia. Beihefte 12, 433 – 476.
Thorrington-Smith, M., 1970. Some new and little-known planktonic diatoms from the west Indian Ocean. Nova Hedwigia.
Beihefte 31, 815 – 835.
Tomczak Jr., M., 1984. Verbreitung und Vermischung der Zentralwassermassen in den Tropengebieten der Ozeane. 2: Indischer
und Pazifischer Ozean. Oceanologica Acta 7 (3), 271 – 288.
Treppke, U.F., Lange, C.B., Donner, B., Fischer, G., Ruhland, G.,
Wefer, G., 1996. Diatom and silicoflagellate fluxes at the Walvis
Ridge: an environment influenced by coastal upwelling in the
Benguela System. Journal of Marine Research 54, 991 – 1016.
Van Iperen, J.M., van Weering, T.C.E., Jansen, J.H.F., van Benekom, A.J., 1987. Diatoms in surface sediments of the Zaire
deep-sea fan (SE Atlantic Ocean) and their relation to overlying
water masses. Netherlands Journal of Marine Research 21,
203 – 217.
Von Bröckel, K., Reinecke, C., Molis, M., Fritsche, P., Kriest, I.,
Nachtigal, K., 1996. Phytoplankton and nutrients. In: Schott, F.,
Pollehne, F., Quadfasel, D., Stramma, L., Wiesner, M., Zeitzschel, B.METEOR-Berichte, Arabian Sea 1995, Cruise No. 32,
JGOFS Results of Leg 32/5, Hamburg University, pp. 96 – 108.
Waniek, J., 1997. Auftriebserscheinungen in der Arabischen See
während des SW-Monsunes. In: Giese, M., Wefer, G.Bericht
R. Schiebel et al. / Marine Micropaleontology 51 (2004) 345–371
über den 5. JGOFS-Workshop. Berichte aus dem Fachbereich
Geowissenschaften der Universität Bremen, vol. 89, p. 60.
Waniek, J., Mitzka, T., Prien, R., Schartau, M., 1996. The cold
water structure (17 jN, 62 jE) and the upwelling region at the
coast off the Arabian Peninsula. In: Schott, F., Pollehne, F., Quadfasel, D., Stramma, L., Wiesner, M., Zeitzschel, B.METEORBerichte, Arabian Sea 1995, Cruise No. 32, JGOFS Results of
Leg 32/5, Hamburg University, pp. 91 – 93.
Washburn, L., Kadko, D.C., Jones, B.H., Hayward, T., Kosro, P.M.,
Stanton, T.P., Ramp, S., Cowles, T., 1991. Water mass subduction
and the transport of phytoplankton in a coastal upwelling system.
Journal of Geophysical Research 96 (C8), 14927 – 14945.
Weaks, M.L., 1983. Satellite image of the month: upwelling in the
Gulf of Oman. Oceanographic Monthly Summary 4 (10), 15.
Winter, A., 1985. Distribution of living coccolithophores in the
371
California current system, Southern California Borderland. Marine Micropaleontology 9, 385 – 393.
Woellner, R.A., Blackwelder, P.L., Peterson, L.C., Lynn, M.J.,
1988. Monsoonal upwelling in the NW Indian Ocean: modern
and late Quaternary coccolithophorid assemblages. Abstracts
with Programs - Geological Society of America 20, A252.
Wyrtki, K., 1973. Physical oceanography of the Indian Ocean. In:
Zeitzschel, B.Ecological Studies: Analysis and Synthesis, vol. 3.
Springer-Verlag, New York, pp. 18 – 36.
Zahn, R., 2003. Global change: monsoon linkages. Nature 421,
324 – 325.
Zielinski, U., Gersonde, R., Sieger, R., Fütter, D., 1998. Quaternary
surface water temperature estimations: calibration of a diatom
transfer function for the Southern Ocean. Paleoceanography 13
(4), 365 – 383.