Marine Micropaleontology 58 (2006) 103 – 113

www.elsevier.com/locate/marmicro

Foraminiferal response to the Northeast Monsoon in the

western and southern Arabian Sea
Petra Heinz *, Christoph Hemleben
Institute of Geosiences, Tübingen University, Sigwartstr. 10, 72076 Tübingen, Germany
Received 15 March 2005; received in revised form 16 September 2005; accepted 13 October 2005

Abstract

Sediments from the western and southern part of the Arabian Sea were collected periodically in the spring intermonsoon
between March and May 1997 and additionally at the end of the Northeast Monsoon in February 1998. Assemblages of Rose
Bengal stained, living deep-sea benthic foraminifera, their densities, vertical distribution pattern, and diversity were analysed
after the Northeast Monsoon and short-time changes were recorded. In the western Arabian Sea, foraminiferal numbers
increased steadily between March and the beginning of May, especially in the smaller size classes (30–63 Am, 63–125 Am).
At the same time, the deepening of the foraminiferal living horizon, variable diversity and rapid variations between dominant
foraminiferal communities were observed. We interpret these observations as the time-dependent response of benthic forami-
nifera to enhanced organic carbon fluxes during and after the Northeast Monsoon. In the southern Arabian Sea, constant low
foraminiferal abundances during time, no distinctive change in the vertical distribution, reduced diversity, and more stable
foraminiferal communities were noticed, which indicates no or little influence of the Northeast Monsoon to benthic foraminifera
in this region.
D 2005 Elsevier B.V. All rights reserved.

Keywords: deep-sea; Arabian Sea; recent benthic foraminifera; meiobenthos; Northeast monsoon/intermonsoon

1. Introduction proxies for deep- and bottom-water circulation and new

Foraminifera are a very successful evolutionary
group of marine organisms. They live in all oceanic
environments and show broad ecological adaptability.
The majority of foraminiferal species build a solid test
of CaCO3 with a high fossilisation potential. Analyses
of stable isotopes in their fossil tests are often used as
tools in paleoceanographic studies. Stable carbon iso-
topes (y13C) in benthic foraminifera can be utilised as
* Corresponding author. Fax: +49 7071 29 57 66.
E-mail address: petra.heinz@uni-tuebingen.de (P. Heinz).
0377-8398/$ - see front matter D 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.marmicro.2005.10.001
production (e.g. Shackleton, 1977; Duplessy et al.,
1984; Curry et al., 1988; Mackensen et al., 1994;
Sarnthein et al., 1994; Mackensen et al., 2001) or
have been proposed as a paleoproxy for the location
and degree of methane venting (Kennett et al., 2000;
Rathburn et al., 2000). Additionally, assemblages, spe-
cies composition, abundance and distributional patterns
of fossil benthic foraminifera are important for biostra-
tigraphy and the reconstruction of a paleoenvironment.
Benthic foraminifera mainly depend on environmental
conditions, such as food supply and oxygen distribution
(e.g. Jorissen et al., 1995; Van der Zwaan et al., 1999).
Organic carbon supply regulates population dynamics
104 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113
and reproduction cycles. Benthic foraminifera are
strongly influenced by surface ocean productivity and
organic carbon flux to the seabed (e.g. Gooday, 1988;
Altenbach and Sarnthein, 1989; Altenbach et al., 1999)
and may provide a better proxy for estimating past
fluxes than bulk sediment organic carbon (Austin and
Evans, 2000). In laboratory experiments, the addition of
organic material resulted in a rise of biomass, a higher
number of food vacuoles (Altenbach, 1992; Linke et al.,
1995) and increased foraminiferal densities (Heinz et
al., 2001; 2002). Investigations on recent foraminifera
and their response to changing environmental condi-
tions therefore provide data of living populations as a
basis for the interpretation of paleoecology. Areas,
which exhibit regional and/or seasonal environmental
variations, can be of interest and ecological relation-
ships seen in these modern faunas may be applicable to
the fossil record.
One of these interesting areas is the Arabian Sea,
which is strongly influenced by the monsoonal system
that triggers seasonal reversals in the surface circula-
tion. Upwelling, mixing and entrainment generate nu-
trient fluxes into the euphotic zone (e.g. McCreary et
al., 1996), which leads to one of the highest produc-
tion rates in the world. Seasonal and regional changes
of surface water productivity and the variability of
organic carbon fluxes make this study site ideal to
investigate the impact of trophic gradients on the
abyssal benthos. Assemblages of living deep-sea ben-
thic foraminifera were affected by these gradients
Fig. 1. Station map of the Arabian Sea.
(Kurbjeweit et al., 2000; Heinz and Hemleben,
2003). The Northeast Monsoon (with strong winds
from November to February) leads to an increased
carbon flux particularly in the northern and eastern
regions of the Arabian Sea (Qasim, 1982; Madhupra-
tap et al., 1996; Smith et al., 1998; Lee et al., 1998;
Wiggert et al., 2000), which results in raised forami-
niferal densities in this area (Heinz and Hemleben,
2003). Size distribution and species composition indi-
cated the response to enhanced organic carbon supply
in spring 1997. Foraminifera in the southern Arabian
Sea were not influenced significantly by the monsoon-
al system (Kurbjeweit et al., 2000; Heinz and Hemle-
ben, 2003). The impact of the monsoonal system on
benthic foraminiferal assemblages has therefore been
demonstrated to a certain extent. However, we still do
not know much about a detailed chronology and am-
plitude of this benthic response in the deep-sea. The
observation of reproductive events or temporal faunal
assemblage changes needs samples in high resolution,
which can provide useful information for interpreta-
tions of the fossil record.
In the present study, we present a more detailed
record of foraminiferal response to the Northeast Mon-
soon. For this, sediments from the western and southern
parts of the Arabian Sea were collected monthly in the
intermonsoon period from March to Mai 1997 and
additionally at the end of the Northeast Monsoon in
February 1998. Assemblages of living deep-sea benthic
foraminifera were analysed and short-time changes
 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113 105

were recorded. Furthermore, these data were compared

with results from 1995 (Kurbjeweit et al., 2000; Heinz
and Hemleben, 2003).

2. Materials and methods

2.1. Study area and sample processing

Multicorer sediment samples containing living ben-

thic foraminifera were collected during different cruises
with the German R/V SONNE in March to May 1997
(spring intermonsoon) and February 1998 (end of
Northeast Monsoon) in the Arabian Sea. Collection
sites were located in the western (WAST = station
Western Arabian Sea Sediment Trap) and southern
(SAST = station Southern Arabian Sea Sediment Trap)
Arabian Sea (Fig. 1). A list of sampling data is given
in Table 1. WAST is close to the upwelling region off
Oman. The existence of a steep mountain slope divides
this site into the abyssal station WAST and the bathyal
WAST-Top (= WAST-T), located 65 km west of WAST.
Only one core was analysed for each sampling date.
Core diameter was 95 mm. All cores were cut at 0.5-
cm intervals for the first 2 to 3 cm (~ 36 cm3 sediment)
and then in 1-cm slices down to a depth of 5 cm (~ 71
cm3 sediment). Each layer was preserved separately
with ethanol and stained with Rose Bengal (1 g/l eth-
anol) for at least 14 days to distinguish living from
dead foraminifera. The sediment was washed over a
30-Am mesh screen. The residue was dried and frac- Fig. 2. Abundance of Rose Bengal stained living benthic foraminifera
tionated into different size classes: 30–63, 63–125, in the first 5 cm of the sediment, divided in three different size classes
N 125 Am. Sample splits (1 / 8–1 / 1) were picked, (**Kurbjeweit et al., 2000; *Heinz and Hemleben, 2003).
depending on the density of stained individuals. To
minimise wrongly identified living foraminifera, speci- plasm were counted. Numbers in each slice were re-
mens were individually wetted with water and only ferred to 10 cm3.
foraminifera containing well-stained dark red proto-
2.2. Species identification
Table 1
Sampling sites and dates (3–4 May 97 = 5/97a, 25 May 97 = 5/97b)
The literature used for the identification of species
was described in Kurbjeweit et al. (2000) in detail.
Station Date Position Depth (m) Cruise
Original data on the foraminiferal assemblages are
WAST-T 27-Mar-97 16810.54VN 59846.03VE 1909 So 117 available from the Arabian Sea database: http://
03-Apr-97 16810.38VN 59846.00VE 1916 So 118
04-May-97a 16810.50VN 59846.00VE 1920 So 118
www.pangaea.de/PangaVista?query=bigset. Please note
25-May-97 16810.49VN 59846,00VE 1915 So 119 that the species mentioned in this paper as Fursenkoina
06-Feb-98 16810.46VN 59846.00VE 1915 So 129 texturata was listed in the Arabian Sea database under
WAST 26-Mar-97 16813.71VN 60810.30VE 4032 So 117 the older name Virgulina texturata.
03-Apr-97 16813.00VN 60816:00VE 4044 So 118
03-May-97a 16813.00VN 60816.00VE 4045 So 118
03-Feb-98 16813.08VN 60816.00VE 4041 So 129
2.3. Statistical analysis
SAST 08-Mar-97 10809.65VN 65800.18VE 4420 So 117
11-Apr-97a 10801.98VN 65800.00VE 4426 So 118 Diversity H(S) was determined according to the
11-Feb-98 10802.00VN 65800.00VE 4427 So 129 Shannon–Wiener Index (Shannon and Weaver,
a
Heinz and Hemleben (2003). 1963). S is the number of observed species. Equita-
106 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113
Fig. 3. Vertical distribution of living foraminifera in the upper 5 cm.
bility was calculated as e H(S)/S (Buzas and Gibson,
1969). When species are totally equally distributed,
the ratio reaches its maximum value of 1. Three size-
standardised fragments of tubular agglutinated forms
such as Rhizammina algaeformis were used to define
a single specimen to gain a semiquantitative number
(Kurbjeweit et al., 2000). Q-mode principal compo-
nent analysis was carried out with SYSTAT 5.2.1.
Only species constituting more than 0.5% in one of
the samples were used as single species. All other
species were grouped in the residual. Factor loadings
of 0.5 and higher were considered as significant for
the Q-mode principal component analysis (Backhaus
et al., 1989).
3. Results
Abundance of Rose Bengal stained living benthic
foraminifera found in the upper 5 cm of the sediment is
shown in Fig. 2. WAST and SAST were not sampled as
frequently as WAST-T. Published data of 1995 (Kurb-
jeweit et al., 2000) and April (SAST) to beginning of
May 1997 (WAST-T, WAST, Heinz and Hemleben,
2003) were also integrated in this figure for a better
comparison between long- and short-time scale sam-
pling. After the Northeast Monsoon in spring 1997,
densities increased steadily between March and the
beginning of May from 16 to 109 foraminifera/50
cm3 at WAST-T and from 11 to 170 foraminifera/50
cm3 at WAST. At the end of May 1997, numbers
decreased again to 74 foraminifera/50 cm3 at WAST-
T (WAST was not sampled). Accounts in February
1998 were slightly higher as in March 1997 (33 fora-
minifera/50 cm3 at WAST-T and 23 foraminifera/50
cm3 at WAST). But maximum densities were still
recorded in 1995 (Fig. 2; Kurbjeweit et al., 2000).
Constant low abundances during time were observed
at SAST (10 to 17 foraminifera/50 cm3 in spring 1997
and 1998). The increase of foraminifera during April to
May 1997 at the western stations was largely caused by
rising densities of smaller specimens, especially in the
63–125 Am fraction.
A comparison of the vertical distribution of the total
living assemblages down to 5-cm sediment depth is
given in Fig. 3. Most stations showed a typical distri-
bution with a maximum in the first cm and decreasing
 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113 107

and only low numbers of foraminifera were found

Fig. 4. Diversity H(S) (x) and equitability (o) at the different stations
in spring 1997 and 1998.
numbers with increasing depths. Only few foraminifera
were counted below 2 cm directly after the Northeast
Monsoon in March 1997 and February 1998. At
WAST-T and WAST, the upper maximum strongly
increased between March and the beginning of May
1997. At the same time, higher densities were recorded
also in deeper sediment layers. At WAST-T, this was
observed especially in May 1997. At the abyssal
WAST, it started already in April 1997. No distinctive
change in the vertical distribution was found at SAST,
below 2 cm.
Diversity and equitability data are given in Fig. 4.
Highest diversity was observed at WAST-T, with a
Shannon–Wiener Index between 2.73 and 3.49 and
an increasing diversity between April and the end of
May 1997. At the abyssal WAST, diversity was
recorded between 2.37 and 3.18, but no general
trend can be observed, and the maximum was found
in February 1998. SAST showed reduced diversity,
between 2.17 and 2.69. Equitability varied at the
different stations and sampling dates between 0.32
and 0.70.
A principal component analysis was performed to
compare short-time development of faunal assem-
blages at the three stations. It showed that foraminifera
were characterized by five principal component com-
munities (= PCCs), which explained 85.41% of vari-
ance of the live assemblages. Species with a score of
z 6.0 were considered to be dominant, between 3.0
and 5.9 as associated. Variance percentage, species
composition and PCC scores are given in Table 2. A
comparison of the dominant communities is demon-
strated in Fig. 5. A high time-dependent variation
between important faunas was observed at the stations
WAST-T and WAST. In March 1997, a community
which was dominated by Hippocrepina indivisa, asso-
ciated by Rhizammina algaeformis (= PCC 2), was
found at WAST-T. One week later, at the beginning
of April, it was characterized by a Reophax scorpiurus
assemblage, accompanied by Lagenammina difflugifor-
mis (= PCC 1). Both samples in May recorded the
dominance of Epistominella pussila, together with
Epistominella exigua and Fursenkoina texturata
(= PCC 3). In February 1998, an Astrorhiza sp. group
could be described (= PCC 5). At WAST, characteristic
communities changed from PCC 1 in March 1997 to
PCC 2 in April 1 week later and to a mixture of PCC 1
and 2 in May. An additional fauna arose in February
1998, dominated by Reophax horrida, which was
again associated by Lagenammina difflugiformis.
SAST was influenced by only two communities,

Table 2
Principal component communities of living benthic foraminifera in the western and southern Arabian Sea
PCC Variance (%) Dominant species Score Associated species Score
1 28.94 Reophax scorpiurus 9.11 Lagenammina difflugiformis 3.54
2 24.46 Hippocrepina indivisa 9.44 Rhizammina algaeformis 3.03
3 15.16 Epistominella pusilla 6.97 Epistominella exigua 3.42
Fursenkoina texturata 3.29
4 8.51 Reophax horrida 8.05 Lagenammina difflugiformis 3.79
5 8.34 Astrorhiza sp. 9.3 – –
108 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113
Fig. 5. Principal component communities of living foraminifera dom-
inating at the different sites in spring 1997 and 1998.
PCC 2 in March 1997 and PCC 1 in April 1997 and
February 1998.
4. Discussion
4.1. General response to the Northeast Monsoon
Living benthic foraminifera in the western Arabian
Sea quickly increased abundances between end of
March and May 1997. We interpret this rise as a
response to improved food availability caused by en-
hanced organic carbon fluxes during and after the
Northeast Monsoon. This monsoon has considerable
impact on the productivity in the western Arabian Sea
(Pfannkuche and Lochte, 2000; Smith, 2001a,b). Inten-
sive studies about the oceanic response to monsoonal
forcing, mixed-layer processes, export fluxes, and in-
terannual variations were done in the Arabian Sea
(Smith, 1998, 1999, 2000, 2001a,b, 2002; Burkill,
1999 and articles cited therein). A time lag of several
months was observed between the beginning of the
meteorological monsoon and biological responses. A
coupling of biological processes in surface waters and
rapid export flux of organic carbon into the deep were
shown (Marra et al., 1998; Dickey et al., 1998). Carbon
fluxes peaked at the end of the Northeast Monsoon
(Honjo et al., 1999). Following settling velocities of
90–330 m/day in the Arabian Sea (Berelson, 2002),
particles will need 6–21 days to reach the bathyal
WAST-T and 12–45 days to arrive at the abyssal
WAST. Phytodetrital deposition during February to
April was indicated by high amounts of chlorophyll a
(= chl a) and chloroplastic pigment equivalents at
WAST (Pfannkuche et al., 2000). In April 1997, fresh
phytodetritus on the sea floor, high porewater fluxes
and enhanced biological activity implied a recent sed-
imentation pulse of particulate organic matter at WAST
(Pfannkuche and Lochte, 2000). Potential respiratory
activity determinations of bacteria and meiofauna
(ETSA measurements) and bacterial degradation
(FDA analysis) showed highest activities during April
1997, compared to other seasons and years (Pfannkuche
et al., 2000). DNA concentrations as a proxy of bio-
mass for small-sized biota (bacteria, protists and small
meiofauna) showed increased values. High concentra-
tions of organic carbon in the upper sediment were
measured after the Northeast Monsoon 1997, compared
to other investigated stations (Böttcher et al., 2000).
Biotic and abiotic data used for the analysis of phyto-
detritus deposition are summarized in Table 3.
4.2. Foraminiferal densities in the western Arabian Sea
Food supply is one of the main environmental fac-
tors that influence foraminifera. Higher densities were
recorded in areas or times of higher organic carbon
fluxes. Experimental laboratory and in situ studies on
deep-sea foraminifera confirmed a response to simulat-
ed phytoplankton pulses (Heinz et al., 2001; Moodley
et al., 2002; Witte et al., 2003; Kitazato et al., 2003;
Nomaki et al., 2005a,b). In the northwestern Arabian
Sea, benthic foraminiferal communities were influenced
 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113 109

Table 3
Summary of biotic and abiotic data used for the analysis of phytodetritus deposition
WAST SAST Reference
2
POC-flux (mg m day 1) 7.4 1.00 Rixen et al. (2000)
2
Flux chl a (Ag m day 1) 6.05 0.39 Rixen et al. (2000)

Apr-97 Feb-98 Apr-97 Feb-98

chl a (Ag 2 cm 3) 0.4700 0.5081 0.0761 0.0490 Pfannkuche et al. (2000)
CPE (Ag 2 cm 3) 7.926 7.801 1.281 1.086 Pfannkuche et al. (2000)
ETSA (Al O2 h 1 2 cm 3) 0.5370 0.5285 0.2555 0.0830 Pfannkuche et al. (2000)
FDA (nmol h 1 2 cm 3) 24.40 13.83 15.74 10.68 Pfannkuche et al. (2000)
DNA (Ag 2 cm 3) 29.86 46.40 15.64 18.28 Pfannkuche et al. (2000)
J nitrate + nitrite (mmol m 2 day 1
) 0.28 0.49 0.12 0.15 Grandel et al. (2000)
J phosphate (Amol m 2 day 1) 51 14 22 10 Grandel et al. (2000)
J silicic acid (mmol m 2 day 1
) 0.9 0.9 0.2 0.3 Grandel et al. (2000)
TOC (% dwt.) 1.3–1.6 – 0.25–0.35 – Böttcher et al. (2000)
POC-flux = flux of particulate organic carbon measured in sediment traps 500 m above sea floor; chl a = chlorophyll a; CPE = chloroplastic pigment
equivalents; ETSA= electronic transport system activity; FDA= FDA turnover bacterial ectoencymatic activity; DNA= DNA concentration; 2
cm 3 = average value integrated over the upper 0–2 cm layer of the sediment; J nitrate + nitrite/phosphate/silicic acid = diffusive benthic fluxes of
nutrients at the sediment water interface; TOC = total organic carbon in % dry weight in the upper 0–2 cm sediment.

by enhanced monsoonal carbon fluxes (Kurbjeweit et
al., 2000; Heinz and Hemleben, 2003). Abundance
correlates with the carbon content in the sediment and
sediment-bound chloroplastic pigment equivalents
(Kurbjeweit et al., 2000). In the present study, increased
densities occurred between the end of March and the
beginning of April 1997, with a presumable delay about
2–5 weeks after enhanced organic supply. This re-
sponse time is in accordance to other field and labora-
tory investigations. Fontanier et al. (2003) found strong
increase of most opportunistic taxa 4–6 weeks after
elevated chl a concentrations at an open-slope station
in the Bay of Biscay. Four weeks after spring bloom
sedimentation, foraminiferal biomass increased signifi-
cantly (Altenbach, 1985). In the laboratory, increased
numbers were recorded 21 days (Heinz et al., 2002) and
19 days (Ernst and van der Zwaan, 2004) after feeding.
Both western stations recorded increasing numbers
(3–7) at the beginning of April, 1 week after the
March sampling. Fresh phytodetritus can initiate strong
foraminiferal reproduction, as it was observed for op-
portunistic species (Gooday and Lambshead, 1989;
Gooday and Turley, 1990). Rise continued the next
4–5 weeks. We observed 7–15 times higher densities
at the beginning of May. Especially the two smaller size
fractions became more important, indicating triggered
reproduction. Three weeks later, the decrease of fora-
minifera at WAST-T indicated diminished food supply.
Synchronous temporal changes of foraminiferal
amounts in different water depths were also described
from bathyal stations in the Bay of Biscay (Fontanier et
al., 2003). The comparison of total densities between
WAST and WAST-T (Fig. 2) showed that in most
investigated seasons, values at WAST-T were lower.
Higher densities may indicate higher levels of organic
material in the abyssal plain, due to an accumulation by
lateral and vertical transport and resuspension process-
es. Elevated total organic carbon values were observed
at WAST, compared to WAST-T (Böttcher et al., 2000).
Additionally, no regular decrease in particle flux at
WAST was observed in trap samples at 1000 and
3000 m water depth (Haake et al., 1993).
Foraminiferal numbers showed interannual variation
in the western Arabian Sea. Stable abundances were
observed at the end of the Northeast Monsoon (March
1997 and February 1998), which indicates a compara-
ble situation before the response to elevated particle
fluxes. But records of March and October 1995 out-
numbered them strongly, and reflect an earlier or more
pronounced Northeast and Southwest Monsoon. Inter-
annual variation in surface chl a concentrations and
particle flux rates were observed (Haake et al., 1993;
Wiggert et al., 2002). Highest sediment chl a concen-
trations and chl a/pheopigment ratios were found at
WAST in March 1995, compared to spring seasons of
1997 and 1998 (Pfannkuche et al., 2000).
4.3. Foraminiferal densities in the southern Arabian
Sea
The southern station SAST, located in the open sea,
was not strongly influenced by monsoon forcing, but
still bears some signs of monsoonal influences. No
significant differences in organic carbon flux content
but in mass flux were observed with season, although in
much reduced amplitudes (Honjo et al., 1999). The
110 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113
Northeast Monsoon did not cause elevated primary
productivity (Pfannkuche and Lochte, 2000). Reduced
flux rates, a low total organic carbon content and low
chl a and chloroplastic pigment equivalent concentra-
tions in the sediment were determined at SAST (Lee et
al., 1998; Böttcher et al., 2000; Pfannkuche et al., 2000,
compare Table 3). Diminished foraminiferal and macro-
faunal densities and biomass as well as lower microbial
activities mirrored the reduced biological response to
such oligotrophic conditions (Kurbjeweit et al., 2000;
Witte, 2000; Boetius et al., 2000; Heinz and Hemleben,
2003). The present study confirmed constant low fora-
miniferal abundances at SAST throughout the year and
between years due to continuous small food supply.
4.4. Spatial variability of foraminifera
Spatial variability and heterogeneous distribution
surely complicates interpretation of temporal foraminif-
eral change within the study area. Only one sediment
core was analysed for each time point and station, and
small-scale patchiness cannot be excluded. Small-scale
patch structures were noted for benthic foraminifera in
the abyssal realm (Bernstein et al., 1978; Bernstein and
Meador, 1979), probably due to habitat selection, re-
productive patterns and small-scale variability of food
supply. Current activity and microtopography may in-
duce patchy distribution of phytodetritus at the sea floor
(Thiel et al., 1988/89). But in the western Arabian Sea,
primary production and organic matter supply record
high seasonality, which will have an impact on the
benthic organism in this area. Simultaneous increasing
densities of mainly small foraminifera in the same range
at both western stations and stable abundances in the
south convince us that the response to the Northeast
Monsoon was recorded in spring 1997.
4.5. Vertical distribution after the Northeast Monsoon
Vertical distribution patterns of total assemblages
changed during spring in the western stations as a
response to enhanced food availability (Fig. 3). At the
end of the Northwest Monsoon (March 1997 and Feb-
ruary 1998), very few foraminifera were observed
below 2 cm depth at WAST-T and WAST. A distinct
maximum in the surface sediment layers and increasing
abundances in the depth formed during the following
weeks in spring 1997. These distribution patterns mir-
ror the dependence of foraminifera on the flux of labile
easily consumable organic matter. Many foraminiferal
species use fresh organic matter deposits (Kitazato et
al., 2003; Nomaki et al., 2005a) and concentrate at first
at the sediment surface because of limited food supply
in deeper layers. Time-dependent transport of food
material downward enables them to disperse vertical
occurrence. This response is in agreement with the
TROX-model (= Trophic–Oxygen–Microhabitat–Reac-
tion, after Jorissen et al., 1995, Fontanier et al.,
2002), which explains benthic foraminiferal microhabi-
tats as a function of organic flux and benthic ecosystem
oxygenation. At SAST, no significant deepening of the
foraminiferal living horizon was observed due to con-
stant oligotrophic conditions.
4.6. Faunal response after the Northeast Monsoon
Diversity was high but variable during 1997 and
1998 (Fig. 4). Only WAST-T recorded an increasing
trend in time. Foraminifera were characterized by five
PCCs (Table 2 and Fig. 5). Time-dependent variability
in flux masses and flux composition may cause faunal
changes at one station. In spring 1997, all stations were
mainly dominated by two assemblages: a Reophax scor-
piurus community, accompanied by Lagenammina dif-
flugiformis (= PCC 1), and a Hippocrepina indivisa
community, associated by Rhizammina algaeformis
(= PCC 2). PCC 2 in the present comparison now com-
bined two communities of a regional and interannual
comparison between 1995 and 1997 (Heinz and Hem-
leben, 2003). Reophax scorpiurus and L. difflugiformis
were both important infaunal taxa in the west and south
of the Arabian Sea and can be considered as background
assemblages, indicating oligotrophic to mesotrophic and
well oxygenated conditions. They are not opportunistic
species, although some Reophax taxa were more abun-
dant in sediments with recorded phytodetritus deposits
(Gooday, 1996) or showed opportunistic behavior, like
Reophax guttiferus (Fontanier et al., 2003). Reophax
scorpiurus and other Reophax species observed in the
Arabian Sea are probably detritus feeders (Jones and
Charnock, 1985; Kaminski et al., 1988) and seem to
benefit from elevated fluxes. PCC 2 combined Hippo-
crepina indivisa and Rhizammina algaeformis. Hippo-
crepina indivisa is probably opportunistic. In laboratory
experiments, Hippocrepina sp. densities increased after
a phytodetrital event (Heinz et al., 2001, 2002). Kurbje-
weit et al. (2000) showed that the sessile tubular R.
algaeformis is positively correlated with high carbon
content and silica. Rhizammina species were described
as typical constituents of eutrophic assemblages in the
Cretaceous (Holbourn and Kaminski, 1997). We can
interpret PCC 2 as an indicator for slightly increased
organic-rich sediments, which requires higher amounts
of labile organic carbon than PCC 1.
 P. Heinz, C. Hemleben / Marine Micropaleontology 58 (2006) 103–113
At WAST, PCC 1 characterized the sediment in
March 1997, displaced by PCC 2 in April. This corre-
sponded well with the interpretation of total abun-
dances, indicating a foraminiferal response to
phytodetritus deposit after the Northeast Monsoon in
April. PCC 2 continued dominating until May, where
PCC 1 was observed additionally and the fluxes were
probably reduced again. At WAST-T and SAST, com-
munities were observed in reversed order, indicating
somewhat higher organic carbon contents in March,
compared to April. For SAST, this would mean that
the Northeast Monsoon influenced this station in March
1997 to a certain degree, at least in variation of faunal
control. For WAST-T, the change from PCC 2 to PCC 1
but increasing foraminiferal densities is difficult to
explain. Here, another faunal change was observed in
May 1997, when an Epistominella pussilla assemblage
(associated with Epistominella exigua and Fursenkoina
texturata) (= PCC 3) was described. Again, two differ-
ent communities of the comparison between 1995 and
1997 (Heinz and Hemleben, 2003) were now combined
in the present comparison. These taxa exhibit strongly
opportunistic taxa. Epistominella reproduce very quick-
ly after the arrival of fresh food material (Gooday and
Turley, 1990; Gooday and Lambshead, 1989; Heinz et
al., 2001, 2002; Fontanier et al., 2003; Ernst and van
der Zwaan, 2004). Fursenkoina species were reported
to be opportunistic and associated with phytodetritus
and enhanced organic flux (Gooday, 1993; Ohga and
Kitazato, 1997). In recent and fossil studies, they be
found tolerant to low oxic and dysoxic conditions and
occurred in pre- and post-sapropel assemblages (Bern-
hard and Sen Gupta, 1999; Jorissen, 1999; Holbourn et
al., 2001). The dominance of PCC 3 in May 1997 at
WAST-T and high abundances strongly indicated the
response of opportunistic foraminiferal species to the
Northeast Monsoon. We can speculate that a slight
reaction was already observed in March by PCC 2
which was then replaced by PCC 3 in May. In-between,
background assemblages like PCC 1 could also be able
to profit from higher trophic conditions. But patchy
distribution of PCC 1 and PCC 2 may pretend faunal
changes. Additionally, spatial variability of surface and
export production make interpretations of biota re-
sponse in the Arabian Sea difficult, especially at
WAST (Pfannkuche et al., 2000).
Both western stations recorded increasing densities
in the smaller size fractions in spring 1997, despite
different faunal changes. We assume that mainly repro-
duction was responsible. PCC 3 was only observed at
WAST-T, not at WAST. Divergences of eutrophic com-
munities between both stations were probably caused
111
by different water depths. Water depth surely affects the
amount of degradation of the sinking phytodetritus and
thus will alter the quality of food. Additionally, varia-
tions in the sediment caused by different bottom cur-
rents, oxygen content, and temperature, may play a
role. WAST-T showed more coarse sediments with
higher calcium carbonate contents (Böttcher et al.,
2000). Bottom water temperature was higher, but oxy-
gen concentration in the bottom water was lower than at
WAST (Grandel et al., 2000).
High annual variability was recorded in the western
Arabian Sea. In February 1998, WAST was mainly
described by Reophax horrida, which is again associ-
ated with Lagenammina difflugiformis (PCC 4), where-
as WAST-T was dominated by PCC 5. PCC 4 is very
similar to PCC 1 and is probably indicating comparable
trophic conditions, just as PCC 5, characterized by
Astrorhiza sp., a genus of sessile tubular forms that
feature passive suspension feeding (Cedhagen, 1988)
and carnivorism (Buchanan and Hedley, 1960).
Acknowledgements
Thanks to Wiebke Ruschmeier and Christina
Schwarz for assistance during sample collection and
processing and crew of F. S. bSonneQ for good collab-
oration during the cruises. We greatly appreciate the
helpful comments of Alexander Altenbach and Olaf
Pfannkuche. This work received financial support
from the Bundesministerium für Bildung und For-
schung (BMBF), BIGSET programme (Biogeochem-
ische Stoff-und Energietransporte in der Tiefsee), No.
03F0177C, which is gratefully acknowledged.
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