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Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234 – 245

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A major change in monsoon-driven productivity in the tropical Indian Ocean

during ca 1.2–0.9 Myr: Foraminiferal faunal and stable isotope data
Anil K. Gupta ⁎, M. Sundar Raj, K. Mohan, Soma De
Department of Geology & Geophysics, Indian Institute of Technology, Kharagpur – 721 302, India
Received 16 July 2007; received in revised form 4 January 2008; accepted 7 January 2008

Abstract

Tropical climate is variable on astronomical time scale, driving changes in surface and deep-sea fauna during the Pliocene–Pleistocene. To
understand these changes in the tropical Indian Ocean over the past 2.36 Myr, we quantitatively analyzed deep-sea benthic foraminifera and
selected planktic foraminifera from N 125 μm size fraction from Deep Sea Drilling Project Site 219. The data from Site 219 was combined with
published foraminiferal and isotope data from Site 214, eastern Indian Ocean to determine the nature of changes. Factor and cluster analyses of the
28 highest-ranked species distinguished four biofacies, characterizing distinct deep-sea environmental settings. These biofacies have been named
after their most dominant species such as Stilostomella lepidula–Pleurostomella alternans (Sl–Pa), Nuttallides umbonifer–Globocassidulina
subglobosa (Nu–Gs), Oridorsalis umbonatus–Gavelinopsis lobatulus (Ou–Gl) and Epistominella exigua–Uvigerina hispido-costata (Ee–Uh)
biofacies. Biofacies Sl–Pa ranges from ~ 2.36 to 0.55 Myr, biofacies Nu–Gs ranges from ~ 1.9 to 0.65 Myr, biofacies Ou–Gl ranges from ~ 1 to
0.35 Myr and biofacies Ee–Uh ranges from 1.1 to 0.25 Myr. The proxy record indicates fluctuating tropical environmental conditions such as
oxygenation, surface productivity and organic food supply. These changes appear to have been driven by changes in monsoonal wind intensity
related to glacial–interglacial cycles. A shift at ~ 1.2–0.9 Myr is observed in both the faunal and isotope records at Site 219, indicating a major
increase in monsoon-induced productivity. This coincides with increased amplitude of glacial cycles, which appear to have influenced low latitude
monsoonal climate as well as deep-sea conditions in the tropical Indian Ocean.
© 2008 Elsevier B.V. All rights reserved.

Keywords: DSDP 214 and 219; Tropical Indian Ocean; Faunal and isotope data

1. Introduction This climatic transition is often known as the Mid-Pleistocene

Earth's climate has evolved from a state of relative warmth
during the mid-Pliocene, with major ice sheets restricted to
Antarctica, to a cooler world during the late Pleistocene marked
with extensive bipolar ice sheets and increased pole to equator
temperature gradients (Raymo, 1997; Ravelo et al., 2004).
These changes are believed to have been driven by changes in
Earth's orbital parameters with a switch from 41-Kyr mode to a
100-Kyr rhythm after 900 Kyr BP (Raymo, 1997; Lisiecki and
Raymo, 2005). Changes in polar ice volume brought significant
changes in the tropics (Sirocko et al., 1999; Gupta et al., 2001).
⁎ Corresponding author. Tel.: +91 3222 283368; fax: +91 3222 282700.
E-mail address: anilg@gg.iitkgp.ernet.in (A.K. Gupta).
0031-0182/$ - see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2008.01.012
Transition (MPT) (e.g., Raymo et al., 1997) or Mid-Pleistocene
Revolution (MPR) (e.g., Hayward, 2001; Xu et al., 2005), and is
well-documented in numerous proxy records (Mix et al., 1995;
Raymo et al., 1997; Gupta et al., 2001; Hayward, 2001; Gupta
and Dhingra, 2004; Xu et al., 2005). At ~ 900 Kyr BP,
the mixed-layer thickness reduced and thermocline shoaled in
the eastern Indian Ocean, and a 100-Kyr component began to
dominate in the Indian monsoon variability (Gupta et al., 2001;
Gupta and Dhingra, 2004). Elongated benthic foraminifera
(including Stilostomella lepidula) suffered widespread extinc-
tion across this climatic transition (Weinholz and Lutze, 1989;
Gupta, 1993; Hayward, 2001; Kawagata et al., 2005, 2006).
Numerous studies have recently been undertaken to under-
stand paleoclimatic and paleoceanographic evolution of the
Indian Ocean during the Plio-Pleistocene and the Holocene
 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245 235

Fig. 1. Location map of Deep Sea Drilling Program (DSDP) Sites 214 and 219 in the tropical Indian Ocean. Also shown are deep-ocean currents (You, 2000). AABW =
Antarctic Bottom Water, NADW = North Atlantic Deep Water, CPDW = Circumpolar Deep Water.

using faunal and geochemical data (Gupta and Srinivasan, 2. Site location: materials and methods
1990; Kroon et al., 1991; Gupta et al., 2003). Most of these
studies have focused on high-productivity areas of the Arabian DSDP Site 214 is located in the eastern Indian Ocean on the
Sea such as the northwestern and northeastern Arabian Sea, Ninetyeast Ridge (11o20.21′S, 88o43.08′E; present water depth
where surface productivity changes are driven by seasonal 1665 m). This site is presently located beneath a hydrological
reversals in the monsoon winds (e.g., Reichart et al., 1998; Jung
et al., 2001; Schmiedl and Leuschner, 2005; Schmiedl and
Mackensen, 2006). The southeastern Arabian Sea is also an
important region where low salinity surface currents from the
Bay of Bengal mix with those of the Arabian Sea.
Changes in organic flux to the seafloor due to variations in
surface productivity modulate deep-sea faunal composition
(e.g. Jorissen et al., 1995; Gupta and Thomas, 1999). The
amount of organic flux to the seafloor not only depends on
surface production but also on the nature of deep-sea column.
Well-oxygenated deep-sea circulation may cause reminera-
lization of organic carbon resulting in little organic material
reaching the seafloor (Schmiedl and Mackensen, 2006). To
understand if the changes in the surface and deep-water column
of the tropical Indian Ocean were driven by the Indian Ocean
climate (monsoon) and deep-sea circulation, we analyzed a
2.36 Myr record of deep-sea benthic foraminifera from Deep
Sea Drilling Project (DSDP) Site 219 from the southeastern
Arabian Sea and combined it with published planktic and
benthic records from Site 214, eastern Indian Ocean (Fig. 1).
The sediment accumulation rate is low to moderate at Sites 214
(0.91 cm/Kyr) and 219 (1.03 cm/Kyr) (Fig. 2). Fig. 2. Numerical age versus depth at Site 219 (Gupta and Thomas, 1999).
236 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245

Table 1
VARIMAX rotated factor scores of eight significant factors at DSDP Site 219
Species name Factor1 Factor2 Factor3 Factor4 Factor5 Factor6 Factor7 Factor8
Astrononian umbilicatulum − 0.27678 −0.54435 0.21522 0.12411 0.24152 − 0.09983 − 0.21964 −0.03865
Bolivina pseudoplicata − 0.09604 0.63124 0.14031 0.28713 0.33582 0.01515 0.26011 0.1557
Bolivina pusilla − 0.18667 0.1082 −0.06034 − 0.35088d 0.54077 0.29692 − 0.10807 0.06134
Bulimina aculeata 0.62835 0.34862 0.16157 − 0.0122 − 0.0527 − 0.00905 0.07341 −0.38974c
Bulimina alazanensis 0.14442 0.2046 0.16591 − 0.12433 − 0.33426 0.43545 0.50979 −0.03206
Bulimina striata 0.68119 −0.10787 0.29073 0.03259 − 0.21666 − 0.30203 0.00952 0.2498
Cassidulina carinata − 0.40159a 0.67055 0.24401 0.34092 − 0.07719 0.12485 0.03409 0.05668
Cibicides bradyi 0.41532 −0.50645 0.13069 0.18526 0.17863 0.15353 0.27073 0.0007
Cibicides wuellerstorfi 0.63842 0.02374 0.28183 − 0.08202 0.02421 0.22133 − 0.30961 −0.03629
Discopulvinulina bertheloti − 0.19432 0.73936 0.32118 0.35622 0.14037 0.19348 0.01261 0.05632
Eggerella bradyi 0.5745 −0.06103 0.14104 0.29633 − 0.08572 − 0.14829 − 0.04954 0.48948
Ehrenbergina carinata − 0.45695a −0.27574 0.1506 − 0.19893 0.14098 − 0.17699 0.26047 0.27035
Epistominella exigua 0.29132 0.06991 0.08212 − 0.48336d 0.59049 − 0.0765 0.14786 0.0468
Evolvocassidulina bradyii 0.46502 0.31869 −0.27943 − 0.06257 0.00804 − 0.36928 − 0.07362 −0.03243
Gavelinopsis lobatulus − 0.37221a −0.26279 0.29207 0.12792 0.34095 − 0.10859 − 0.15582 −0.40186c
Globocassidulina pacifica 0.56338 −0.2219 −0.07875 0.43029 0.02424 0.10288 − 0.25646 0.37342
Globocassidulina subglobosa 0.08172 0.0121 −0.65278b 0.33966 0.2013 0.31311 − 0.01501 −0.04683
Gyroidinoides cibaoensis 0.47499 0.20755 −0.4199b 0.00346 0.02495 − 0.29373 0.12424 0.15028
Hoglandulina elegans 0.51997 −0.42207 0.01022 0.47281 0.24827 0.07963 0.06066 −0.18306
Melonis barleeanum 0.32735 0.03988 0.66648 − 0.24837 − 0.11669 − 0.13728 − 0.15883 −0.09138
Nuttallides umbonifer 0.02991 0.06983 −0.68766b − 0.14073 0.20614 0.07212 − 0.05179 0.13001
Oridorsalis umbonatus 0.03694 0.08683 −0.41545b 0.28318 − 0.23448 − 0.25462 − 0.05876 −0.51602c
Osangularia culter 0.19279 −0.47841 0.10761 0.14219 − 0.06176 0.07296 0.60514 −0.16739
Pleurostomella alternans − 0.66316a −0.0648 0.07837 0.06686 0.05568 − 0.18797 0.01202 0.16325
Pullenia bulloides − 0.04405 −0.28612 0.01745 − 0.09139 − 0.28655 0.68298 − 0.3178 0.08143
Stilostomella lepidula − 0.76712a 0.03078 0.14501 0.22505 − 0.18706 − 0.14994 − 0.2423 −0.0025
Uvigerina hispido-costata 0.60918 0.27717 −0.07434 − 0.38523d − 0.17286 0.07584 − 0.17436 −0.09885
Uvigerina proboscidea − 0.57463a −0.2155 −0.28727b − 0.2008 − 0.4666 − 0.02655 0.15959 0.15555
% variance 27.73 15.18 11.50 8.44 7.66 6.0 5.28 5.02
a
Biofacies Sl–Pa, bBiofacies Nu–Gs, cBiofacies Ou–Gl, dBiofacies Ee–Uh.

front in the core of the Indonesian Throughflow (Du et al.,
2005), and is under the influence of the South Equatorial
Current (SEC), which intensifies during northern summer when
southeast trade winds are stronger and Indian summer monsoon
begins to strengthen (Fig. 1; Tchernia, 1980). Site 214 is bathed
by the North Indian Deep Water (NIDW) with a dissolved
oxygen content of ~ 5.8 ml/L (GEOSECS, 1983), which is a
mixture of Antarctic Bottom Water (AABW), North Atlantic
Deep Water (NADW) and deep water of the northern Indian
Ocean origin (Tchernia, 1980). DSDP Site 219 was drilled
during Leg 23 on the crest of the Laccadive-Chagos Ridge,
southeastern Arabian Sea (9°01.75′N, 72°52.67′E; water depth
1764 m). At present, this site is bathed by the NIDW with
dissolved oxygen content of ~ 4.8 ml/L (GEOSECS, 1983), and
lies within an area where surface production is higher due to
monsoon-induced upwelling (Gupta and Thomas, 1999;
Shankar et al., 2002).
Seventy one core samples of 10 cm3 volume from Site 219
were analyzed from a sediment sequence extending back in time
to ~ 2.36 Myr. Samples were soaked in water with baking soda
for 8–12 h, and washed over a 63 µm size sieve. The washed
samples were dried in an electric oven at ~ 50 °C and transferred
into glass vials. Hard sediment samples were treated with 2–3
drops of 2% hydrogen peroxide. We applied the age model for
Site 219 based on planktic foraminifer datums suggested by
Gupta and Thomas (1999). The sediment accumulation rate at
Site 219 shows a major increase at ~ 1.6 Myr and thereafter is
constant (Fig. 2). The interpolated numerical ages are updated to
the Berggren et al. (1995) time scale, with an average time
resolution of 33 Kyr per sample.
We generated benthic foraminiferal census data from an
aliquot of ~ 300 specimens in the N 125 μm size fraction (census
data are made available online as Table 1s). This size fraction
allows us to compare our results with recent results from the
Atlantic and Indian Oceans (Mackensen et al., 1993, 1995;
Schmiedl et al., 1997; Gupta and Thomas, 2003; Gupta et al.,
2004). Factor and cluster analyses were performed on relative
abundance data of the 28 highest-ranked species (Table 1;
supplementary Table 1s), which are present in at least three
samples with a percentage of 2% or more in at least one sample.
The analysis performs standardization of the data and calcula-
tion of covariance matrix that helps remove noise and reduce the

Fig. 3. Dendogram based on Q-mode cluster analysis of 71 samples from DSDP Site 219 using Ward's Minimum Variance method. Four clusters have been
identified on the basis of the number of clusters versus semi-partial R2. Each cluster corresponds to a biofacies named after the most dominant species within each
cluster.
A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245 237
238 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245

Table 2
Environmental preferences of characteristic benthic foraminiferal species defining different biofacies at Site 219
Species Interpretation
B. pusilla High organic carbon flux (Thomas and Gooday, 1996; Gupta and Satapathy, 2000)
B. aculeata No relation with food flux (Hermelin and Shimmield, 1990)
Calm depositional regimes with clayey and organic-rich sediments (Mackensen et al., 1993)
Interglacial, high productivity (Almogi-Labin et al., 2000)
C. carinata High food supply (Gupta and Thomas, 1999)
C. wuellerstorfi Raised epibenthic, suspension feeder, high energy (Lutze and Thiel, 1989)
Oligotrophic (Linke and Lutze, 1993)
Seasonal food supply (Loubere and Fariduddin, 1999)
AABW (Corliss, 1979, 1983)
E. carinata High organic food (Gupta and Satapathy, 2000)
Low-oxygen, high organic carbon (Nomura, 1991)
E. exigua Epibenthic, cosmopolitan, abyssal, opportunistic, phytodetritus feeder (Gooday, 1988; Gooday and Turley, 1990)
Seasonal food fluxes (Smart et al., 1994; Loubere and Fariduddin, 1999)
G. lobatulus Pulsed food supply (Gooday,1994)
Cool waters, partially pulsed food supply (Gupta and Thomas, 1999)
G. subglobosa Warmer AABW (Corliss, 1979)
Circumpolar Deep Water (Schnitker, 1980)
Phytodetritus feeder (Gooday, 1994)
Oligotrophic (Mackensen et al., 1995)
Low-productivity associated with NADW (Fariduddin and Loubere, 1997)
Strong bottom currents, elevated, oligotrophic (Mackensen et al., 1995; Schmiedl et al., 1997)
G. cibaoensis Low-oxygen (Gupta and Thomas, 1999)
N. umbonifera AABW (Streeter 1973)
Corrosive bottom water (Bremer & Lohmann, 1982)
Oligotrophic (Gooday, 1994)
O. umbonatus AABW (Corliss, 1979)
High carbonate saturation, motile (Miao and Thunell, 1993)
High oxygen, low organic carbon (Mackensen et al., 1995)
Cosmopolitan, both oligotrophic and eutrophic (Schmiedl 1995; Schmiedl and Mackensen, 1997)
P. alternans Abyssal species, characteristic of variable organic flux (Gupta, 1994)
S. lepidula Low-oxygen, organic rich (Boersma, 1990)
High productivity (Gupta, 1993)
Low productivity (Kaiho, 1999)
U. hispido-costata High organic carbon (Altenbach and Sarnthein, 1989; Rathburn and Corliss, 1994)
U. proboscidea Shallow infaunal, high organic carbon, variable oxygenation (Lutze and Coulbourn, 1984)
High, year-round productivity (Gupta and Srinivasan, 1992; Gupta, 1997; Jannink et al., 1998; Gupta and Thomas, 1999;
Gupta et al., 2001, 2004; Murgese and De Deckker, 2005)
Low seasonality (Ohkushi et al., 2000)

data to more meaningful variables. The R-mode factor analysis
identifies the species associations and Q-mode cluster analysis
discerns the sample groups.
R-mode Principal Component Analysis (PCA) was performed
on the correlation matrix followed by orthogonal VARIMAX
rotation using the SAS software package (SAS Inc., 1988). Based
on a scree plot (x–y plot) of eigenvalues versus the number of
factors and screening of the VARIMAX factor scores and species
associations, 8 factors were considered that account for 86.8% of
the total variance (Table 1). Factors that do not show relevant
species associations due to their low factor scores and rare spe-
cies occurrence, were not considered to constitute assemblages.
Q-mode cluster analysis using Ward's Minimum Variance meth-
od was run on a covariance matrix to identify sample groups
belonging to each biofacies (Fig. 3). Four clusters representing
four biofacies were identified on the basis of a plot of semi-partial
R2 values versus the number of clusters. We used information on
environments of modern benthic foraminifera (Table 2) to
interpret different biofacies at Site 219 (Table 3, Figs. 4 and 5).
We calculated Globigerina bulloides percentages from Site
219 and combined them with published percent U. proboscidea,
Globigerinoides sacculifer, Globorotalia menardii (complex)
and Gs. sacculifer stable isotope data from eastern Indian
Ocean Site 214 (Gupta and Dhingra, 2004; Fig. 7). G. bulloides
is a well tested monsoon proxy used in several paleomonsoonal
studies (Kroon et al., 1991; Overpeck et al., 1996; Gupta et al.,
2003).
3. Results and environmental preferences of benthic biofacies
Multivariate analysis was performed on reduced data of
benthic foraminifera from Site 219 to reduce noise from the data
set, which is induced by postmortem taphonomic changes.
The analysis involves standardization of the data and calcula-
tion of covariance matrix that reduces the aliasing. Given below
are details of each biofacies and their inferred environments
(Table 3). Each biofacies is named after the most dominant
species present (Table 4).
 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245 239

Table 3 characteristic species of this biofacies are O. umbonatus,

Benthic foraminiferal biofacies and their interpreted environments at DSDP Site G. lobatulus and Bulimina aculeata, indicating low to interme-
219, southeastern Arabian Sea
diate organic carbon flux and high seasonality (Tables 2 and 3).
Biofacies Environment
Sl–Pa (Factor 1 negative scores) High and sustained organic flux 3.4. Biofacies Epistominella exigua–Uvigerina hispido-costata
Stilostomella lepidula (− 0.76712) (Ee–Uh)
Pleurostomella alternans (− 0.66316)
Uvigerina proboscidea (− 0.57463)
Ehrenbergina carinata (− 0.45695) Epistominella exigua, Uvigerina hispido-costata and Boli-
Cassidulina carinata (− 0.40159) vina pusilla are the characteristic species of biofacies Ee–Uh,
Gavelinopsis lobatulus (−0.37221) with high negative Factor 4 scores (Figs. 4 and 5; Tables 1
Nu–Gs (Factor 3 negative scores) Low and variable flux of and 3). This biofacies is present from ~ 1.1 to 0.25 Myr and is
Nuttallides umbonifer (− 0.68766) organic matter, high seasonality,
suggestive of intermediate to high organic flux, high seasonality
Globocassidulina subglobosa (− 0.65278) well-oxygenated and cold
Gyroidinoides cibaoensis (− 0.4199) deep waters and well-oxygenated conditions (Tables 2 and 3).
Oridorsalis umbonatus (− 0.41545)
Ou–Gl (Factor 8 negative scores) Low to intermediate organic 4. Discussion and conclusions
Oridorsalis umbonatus (− 0.51602) carbon flux, high seasonality
Gavelinopsis lobatulus (−0.40186)
Deep-sea benthic foraminifera have been used to understand
Bulimina aculeata (− 0.38974)
Ee–Uh (Factor 4 negative scores) Intermediate to high organic flux, changes in deep-water conditions driven by climate forcing
Epistominella exigua (−0.48336) high seasonality, well-oxygenated during the Pliocene and Pleistocene (Corliss et al., 1986;
Uvigerina hispido-costata (− 0.38523) conditions Thomas et al., 1995; Schmiedl and Mackensen, 1997; Gupta
Bolivina pusilla (− 0.35088) et al., 2001). Several studies have shown the relationship be-
tween benthic faunal composition, productivity of the overlying
3.1. Biofacies Stilostomella lepidula–Pleurostomella alternans waters and organic flux to the seafloor (Lutze and Coulbourn,
(Sl–Pa) 1984; Miao and Thunell, 1993; Smart et al., 1994; Thomas and
Gooday, 1996; Gupta et al., 2004; Smart et al., 2007). Others
Biofacies Sl–Pa is defined by species with a high negative score suggested oxygen and food supply are the main factors con-
on Factor 1, in the ~2.36 to 0.55 Myr interval (Figs. 4 and 5). The trolling the spatial and in-sediment distribution of benthic fora-
characteristic species of this biofacies are Stilostomella lepidula, minifera (Hermelin and Shimmield, 1990; Bernhard, 1992;
Pleurostomella alternans, Uvigerina proboscidea, Ehrenbergina Jorissen et al., 1995; Schmiedl et al., 1997; Gupta and Thomas,
carinata, Cassidulina carinata and Gavelinopsis lobatulus, 1999; den Dulk et al., 2000). This group explains seasonal
indicating high and sustained organic flux (Tables 2 and 3). fluctuations in primary production (Thomas et al., 1995;
Thomas and Gooday, 1996; Schmiedl et al., 1997, 2000; den
3.2. Biofacies Nuttallides umbonifer–Globocassidulina Dulk et al., 2000; Wollenburg and Kuhnt, 2000; Gupta and
subglobosa (Nu–Gs) Thomas, 2003). Gooday (1994), Thomas et al. (1995) and
Thomas and Gooday (1996) classified phytodetritus species
Characteristic species of this biofacies are Nuttallides (like E. exigua) and mobile, infaunal species (like species of
umbonifer, Globocassidulina subglobosa, Gyroidinoides Melonis and Uvigerina) as indicators of predominantly sea-
cibaoensis and Oridorsalis umbonatus, with Factor 3 negative sonally pulsed versus continuous primary production. Thus
scores. This biofacies is present from ~ 1.9 to 0.65 Myr, benthic foraminifera are considered useful for estimating paleo-
indicating low and variable flux of organic matter (oligotrophic), fluxes in high-productivity areas and they are also more resis-
high seasonality, well-oxygenated and cold deep waters (Figs. 4 tant to diagenetic change compared to planktic foraminifera.
and 5, Tables 2 and 3). It has been suggested by Gupta et al. However, in oligotrophic areas deep-sea oxygenation plays an
(2006) that N. umbonifer is out-competed by species of Buli- important role in controlling benthic foraminifera over variable
mina and Uvigerina when the food supply is high and sustained, time scales. Gupta and Thomas (1999) suggested that changes
unless the waters become corrosive to CaCO3. Under conditions in oxygenation are linked partially to productivity and partially
of relatively low, seasonal flux of food N. umbonifer may be to changes in deep-water ventilation.
swamped by the opportunistic species such as E. exigua. Nut- Wind driven coastal and open-ocean surface productivity
tallides umbonifer thus dominates those regions where other influences organic carbon flux and oxygenation of deep waters
calcareous benthic foraminiferal taxa cannot grow optimally, controlling benthic populations in the Arabian Sea. In the
either as a result of very low food supply (extreme oligotrophy), northern part of the Indian Ocean, the wind regimes follow
or a high carbonate corrosivity (Gupta et al., 2006). seasonal change in circulation producing widespread upwelling
controlling surface productivity near Site 219 (Fig. 1). The in
3.3. Biofacies Oridorsalis umbonatus–Gavelinopsis lobatulus situ (0–50 m) primary production above Sites 214 and 219 is
(Ou–Gl) b20 mg C m− 3 h− 1 during June–September, when the summer
monsoon and trade winds are stronger. This is less than one
This biofacies is defined by species with negative Factor 8 fifth that of the northwestern Arabian Sea primary production
scores, across the ~ 1 to 0.35 Myr interval (Figs. 4 and 5). The (Krey, 1973). The primary production reduces significantly to
240 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245

Fig. 4. Percent distribution of benthic foraminiferal species which are most dominant in different biofacies (panels a–d) at DSDP Site 219. Each colour shade
represents a particular species as given at the bottom of the figure. Open arrows mark the shift in the foraminiferal faunal and stable isotope record.

b 0.50 mg C m− 3 h− 1 above Site 219 and b0.10 mg C m− 3 h− 1
above Site 214 during December–March when variable and
weak winter monsoon winds blow across the Indian Ocean
(Krey, 1973; Tchernia, 1980; Schott and McCreary, 2001).
Because of the location of Sites 214 and 219 in oligotrophic
areas with higher dissolved oxygen content during the studied
interval, the changes in benthic foraminiferal population might
have been linked to the supply of organic food.
At Sites 214 and 219, the faunal and isotope data show a shift
during ~ 1.2–0.9 Myr very close to the Mid-Pleistocene
Transition (MPT), coinciding with the transition from 41-Kyr
to 100-Kyr periodicities, and an increased intensity of ice age
cycles in the Northern Hemisphere (Raymo et al., 1997). This
also corresponds to the Stilostomella extinction observed in
different ocean basins (Gupta, 1993; Gupta et al., 2001;
Hayward, 2001; Kawagata et al., 2005, 2006). During this
time, the East Asian monsoon system strengthened and contrast
between the summer and winter monsoon wind regime
increased (Xiao and An, 1999). Thus variability in the East
Asian monsoon increased (Qiang et al., 2001).
At Site 219, U. proboscidea was a dominant benthic species
from 2.36 to 1.2 Myr and G. bulloides was rare (Fig. 7). The
 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245 241

Fig. 5. Cumulative percentages of benthic foraminiferal species dominant in different biofacies at DSDP Site 219. Open arrow marks a shift benthic foraminiferal
fauna, coinciding with the strengthening of the Indian monsoon.

benthic biofaces Sl–Pa and Nu–Gs alternated during this time crease in % Gr. menardii complex (Fig. 7). This higher
(Fig. 6). This indicates that the surface productivity was low surface productivity could have been linked to strong trade
(rare G. bulloides) above Site 219 during this time but there was winds, which would have intensified during cold intervals,
an increased lateral advection of nutrient-rich deep water thus causing intense open-ocean upwelling and high surface
(NIDW) from the oxygen minimum zone (OMZ) of the productivity. Site 214 is located beneath the core of the
northwestern Arabian Sea to the southeastern Arabian Sea.
Dickens and Owen (1999) suggested lateral expansion of OMZ
waters to the eastern and southeastern parts of the Indian Ocean Table 4
in the late Neogene. In the northwestern Arabian Sea, the Relationship between biofacies factor scores of each species and its average
abundance in respective clusters
summer monsoon winds cause widespread upwelling and high
surface productivity and thus increased organic flux to the Biofacies Factor 1-ve Factor scores Cluster IV
(average %)
seafloor, intensifying oxygen minimum zone between 500 and
1500 m water depths. The faunal and isotope trends at Site 214 Sl–Pa Stilostomella lepidula − 0.76712 15.92655633
Pleurostomella alternans − 0.66316 2.282477444
show similar conditions (Fig. 7). At Site 214, the mixed-layer
Uvigerina proboscidea − 0.57463 19.06312037
was thick (abundant Gs. sacculifer) and thermocline was deep Ehrenbergina carinata − 0.45695 1.806140548
(rare Gr. menardii complex), indicating low surface productiv- Cassidulina carinata − 0.40159 8.077220475
ity in the eastern Indian Ocean. Gavelinopsis lobatulus − 0.37221 3.76770311
After 1.0 Myr, at Site 219 the benthic biofacies are domi-
Nu–Gs Factor 3-ve Factor scores Cluster III
nated by Ou–Gl and Ee–Uh while U. proboscidea decreased
(average %)
in abundance, indicating low to intermediate organic carbon Nuttallides umbonifer − 0.68766 2.40671494
flux to the seafloor and high seasonality — conditions typical Globocassidulina subglobosa − 0.65278 4.369466469
of winter monsoon season. The percent G. bulloides increased Gyroidinoides cibaoensis − 0.4199 2.418400106
after 1.0 Myr, yet its abundance is not significant enough Oridorsalis umbonatus − 0.41545 4.100768771
Uvigerina proboscidea − 0.28727 30.41905914
to attribute it to summer monsoon but to winter monsoon
strengthening. We suggest strengthening of the winter mon- Ou–Gl Factor 8-ve Factor scores Cluster II
soon since 1.0 Myr, driven by the onset of ice age cyclicity. It (average %)
has been observed by Overpeck et al. (1996) that G. bulloides Oridorsalis umbonatus − 0.51602 4.257120018
population remains at b 4–5% during winter monsoon sea- Gavelinopsis lobatulus − 0.40186 1.792069343
Bulimina aculeata − 0.38974 10.77563706
son and increases to N30% during summer monsoon season.
Fontugne and Duplessy (1986) suggested that during cold Ee–Uh Factor 4-ve Factor scores Cluster I
intervals, the winter monsoon strengthens and summer mon- (average %)
soon weakens. In the eastern Indian Ocean, the productivity Epistominella exigua − 0.48336 12.30504722
has increased during the past 1.0 Myr as suggested by step- Uvigerina hispido-costata − 0.38523 7.116840533
Bolivina pusilla − 0.35088 1.355918378
wise decrease in δ13 C values and % Gs. sacculifer and in-
242 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245

Fig. 6. Distribution of benthic foraminiferal biofacies in time along with accumulative percentages of their key species at DSDP Site 219. Also plotted are percent
planktic foraminifer Globigerina bulloides and benthic foraminifer Uvigerina proboscidea (top panel). The shades mark the intervals during which a particular
biofacies dominates.

Indonesian Throughflow (ITF) which has its maximum flow
in summer and minimum in winter (Gordon, 1986; Gordon
and Fine, 1996). We do not link the isotopic and faunal
changes at Site 214 during the last 1.0 Myr to the ITF flow
because during glacial times the sill depths between the
Pacific and the Indonesian Sea shoal, thus restricting the
movement of water between the two regions.
At Sites 214 and 219, oxygen does not appear to have been
a limiting factor since these sites lie in a well-oxygenated
(N3.8 ml/L) environment in the present-day abyssal setting and
 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245 243

Fig. 7. Percent Uvigerina proboscidea and Globigerina bulloides from Site 219 (panel e), plotted with percent U. proboscidea (panel d), mixed-layer species
Globigerinoides sacculifer and thermocline species Globorotalia menardii complex (panel c), Gs. sacculifer δ13C (‰) (panel b) and δ18O (‰) (panel a) from Site
214. Data in panels a–d are from Gupta and Dhingra (2004). A major switch in the proxy record is visible during ~1.2–0.9 Myr ago.

have remained so during the past 2.4 Myr. Organic carbon flux
linked to changes in the monsoon strength and trade wind
intensity appears to be the primary factor controlling changes in
benthic foraminiferal assemblages in the southeastern Arabian
Sea and eastern Indian Ocean. Our study thus supports the recent
observations by Gupta et al. (2003) and Gupta and Thomas
(2003) suggesting that changes in the tropical monsoons were
closely related to changes in the Northern Hemisphere ice vol-
ume both at orbital and suborbital time scales. The high vari-
ability in the tropical deep-sea environments occurred at a time
when the Earth's climate was experiencing large scale turnovers
due to the increased intensity of glacial–interglacial cycles.
244 A.K. Gupta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 261 (2008) 234–245
Acknowledgements
Deep Sea Drilling Project is thanked for providing samples
for the present study. This study was funded by DST, New Delhi
(No. SR/S4/ES-46/2003).
Appendix A. Supplementary data
Supplementary data associated with this article can be found,
in the online version, at doi:10.1016/j.palaeo.2008.01.012.
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