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Potentiality of Pteropods in reconstruction of the Quaternary climatic and

oceanographic history of the Arabian Sea

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 ISSN 0970-261X

Gond. Geol. Magz., V. 25(1), June, 2010. pp.

Special Issue on "Applied Micropaleontology"
P. Kundal and S. K. Humane (Guest Editors)

Potentiality of Pteropods in reconstruction of the Quaternary

climatic and oceanographic history of the Arabian Sea
A.D. Singh* and O. P. Singh
Centre of Advanced Study in Geology
Banaras Hindu University, Varanasi-221005 (UP)
*E-mail: arundeosingh@yahoo.com

ABSTRACT and paleoceanographers to investigate various aspects of

The utility of pteropods in the Quaternary
paleoceanographic studies has been established through
extensive studies in marginal seas like Red Sea, Gulf of Aden
and Mediterranean Sea. In the sediments of marginal areas
and topographic highs of the Arabian Sea, pteropods
constitute one of the main carbonate components, but they
have not received much attention as they deserve, until
recently. Since late nineties, there has been growing interest
among micropaleontologists and paleoceanographers to
investigate this important microfaunal group in the Arabian
Sea. Studies carried out both on recent pteropods and fossil
assemblages enabled to establish several valuable pteropod
indicators useful in deciphering history of past changes in
oceanographic and climatic conditions of the Arabian Sea.
The Quaternary pteropod records generated in recent years
provide important source of information to better
understand the nature and processes of past oceanographic
and climatic changes in this region. This paper reviews
pteropod studies carried out in Arabian Sea and evaluates
efficacy of this microfaunal group in Quaternary climatic
and oceangographic reconstructions.
Keywords: Pteropods, Quaternary Climate and Oceanography,
Arabian Sea
both modern and fossil pteropods from the Arabian Sea.
Valuable information about ecology of recent pteropods
were derived through studies on pteropod assemblages
from plankton net, sediment trap and sea bed samples of
different areas of the Arabian Sea (Sakthivel, 1973 a,b;
Auras-Schudnagies et al., 1989; Singh and Rajarama,
1997; Singh et al., 1998, 2001, 2005; Singh and Conan,
2008). These studies enabled to understand various
oceanographic factors controlling distribution of
pteropods in the Arabian Sea. Data on distribution of
recent pteropods and its relationship with various
oceanographic parameters are important, as it is used as an
analogue to infer past changes based on the faunal
variation in sedimentary records. Pteropods are often
found to be more useful than other commonly used
microfossil group like planktic foraminifera in
paleoceanographic reconstruction particularly from the
marginal Quaternary sedimentary records, because of
their abundant occurrence and more sensitiveness to
physico-chemical conditions in shallow regions.

OCEANOGRAPHY OF THE ARABIAN SEA

INTRODUCTION
Pteropods, holoplanktic microgastropods with
aragonitic shells are common members of the calcareous
zooplankton community in open ocean environment.
They are preserved in oceanic sediments lying above the
Aragonite Compensation Depth (ACD). The Arabian Sea
is regarded as one of the most bio-productive areas of the
world (Fig. 1). In the Arabian Sea, modern ACD varies
from about 600 m (in western region) to 400-500 m (in
northern and eastern regions) (Millero et al., 1998; Sabine
et al., 2002). Thus, well-preserved pteropods shells in the
Arabian Sea sediments are limited to topographic highs
and marginal areas, where they constitute one of the major
biotic components. Pteropods are considered to be very
sensitive to the marine environments and have been
extensively used for Quaternary paleoceanographic and
paleoclimatic reconstructions in the Red Sea, the
Mediterranean Sea and the Gulf of Aden. In recent years,
there is growing interest among the micropaleontologists
The Arabian Sea is characterized by seasonal reversal
of monsoon driven winds during summer and winter,
resulting spatial changes in surface circulation,
hydrography and biological productivity (Wyrtki, 1971).
In summer monsoon (June to September), southwesterly
winds blow along the Somali Coast generating large scale
coastal upwelling in the western and northern Arabian
Sea. During this period, the South Equatorial Current
intensifies, northern branch of which forms the Somali
Current as part of anticyclonic southwest monsoon
circulation. During winter monsoon (December to
March), moderate northeasterly winds blow over the
northern Arabian Sea and a cyclonic circulation develops.
Northeast monsoon winds are too weak to generate strong
coastal upwelling and offshore Ekman pumping. During
this season, sea-surface temperature off Pakistan and
northern Indian coasts decreases due to winter cooling
causing convective mixing. Winter sea-surface cooling
effect is also observed in the northwestern part of the
 A.D. Singh and O. P. Singh

Fig. 1. Important pteropod studies in the Arabian Sea. [1. Auras-Schudnagies et al., 1989; 2. Bharti, 2008; 3. Herman and Rosenberg,
1969; 4. Ivanova, 1999; 5. Klocker et al., 2006; 6. Mohan et al., 2006; 7. Reichart et al., 1998; 8. Reichart et al., 2002; 9. Sakthivel, 1969;
10. Sakthivel, 1973; 11. Singh, 1998; 12. Singh, 2007; 13. Singh and Rajarama, 1997; 14. Singh and Conan, 2008; 15. Singh et al., 1998;
16. Singh et al., 2005; 17. Singh et al., 2007; 18. Singh et al., 2006; 19. Singh et al., 2001; 20. Singh et al., 2000; 21. von Rad et al., 1999].

Arabian Sea. Biologial productivity in western and
northern Arabian Sea is very high in summer monsoon,
whereas during winter, a moderate increase in
productivity is observed in northeastern Arabian Sea.
During the inter-monsoon periods, November to
December and April to May, the water column in entire
Arabian Sea is stratified and characterized by nutrient-
poor condition.
The water masses in the Arabian Sea are a
combination of locally and externally generated water
masses (Schott and McCreary, 2001). The surface water
called Arabian Sea Water (ASW) is generated during
winter. Below the surface waters, North Indian Central
Water (NICW), and an old Indian Central Water (ICW)
ventilates the thermocline of the Arabian Sea. ICW is
formed in the subtropics of the southern hemisphere and
spread westwards with South Equatorial Current (SEC).
Antarctic Intermediate Water (AIW) and Red Sea Water
(RSW) with similar density, flow at intermediate depth.
Deep waters of the Arabian Sea are a mixture of
Circumpolar Deep Water (CDW) and the overlying
intermediate waters (Schott and McCreary, 2001).
PTEROPOD ASSEMBLAGE OF THE ARABIAN SEA
The pteropod assemblages recovered from plankton
tows, sediment trap and sea bottom sediments of the
Arabian Sea are characterized by low diversity and
composed of both epipelagic and mesopelagic forms.
Limacina inflata (d’Orbigny), Limacina trochiformis
(d’Orbigny), Creseis acicula (Rang), Creseis virgula
(Rang) forma virgula (Rang), Creseis virgula (Rang)
forma conica Eschscholtz, Creseis chierchiae (Boas),
Clio convexa (Boas), Clio pyramidata Linnaeus,
Cavolinia longirostris (De Blainville), and Diacria
quadridentata (de Blainville), Styliola subula (Quoy and
Gaimard) and Hyalocylix striata (Rang) are common
pteropod species in the Arabian Sea. Other rare species
are: Dicaria trispinosa (de Blainville), Cavolinia gibbosa
(Rang) and Limacina bulimoides (d’Orbigny).
The classification of the pteropods has gone through
several stages of development since Pelseneer’s (1888)
creation of two suborders: Thecosomata and
Gymnosomata. Meisenheimer (1905) divided the
Thecosomata into Euthecosomata and Pseudothe-
 Pteropods and Quaternary climatic of the Arabian
cosomata. Further refinements at the family and
subfamily levels have been introduced by Tesch (1904,
1913, 1948), Van der Spoel (1967), Bé and Gilmer (1977)
and Van der Spoel and Boltovskoy (1981). Most
commonly used and widely accepted systematic
classification is by Bé and Gilmer (1977) and Van der
Spoel and Boltovskoy (1981).
Knowledge of the regional distribution pattern of
recent pteropods and its correlation with the physico-
chemical conditions of the water column is essential in
order to investigate their potentiality as indicators of
environmental changes. Several studies have been carried
out on living as well as dead pteropod assemblages in
marginal seas to understand the relationships between
their distribution pattern and environmental conditions
(Rosenberg and Herman, 1965; Reiss et al., 1980;
Weikert, 1987; Almogi-Labin, 1984; Auras-Schudnagies
et al., 1989). Less effort has been directed towards the
investigation of Arabian Sea pteropod populations.
Sakthivel (1969, 1973a, b) made attempts to study living
pteropods of the Arabian Sea, the Bay of Bengal and the
southwestern parts of the Indian Ocean. Important
contributions on the Arabian Sea Recent pteropods made
in the last few years are by Auras-Schudnagies et al.
(1989), Singh and Rajarama (1997), Singh et al. (1998,
2005) and Singh and Conan (2008).
RECORD OF PTEROPOD FLUX IN ARABIAN SEA
Evaluation of production, accumulation and
dissolution of calcium carbonate in ocean basins is
essential to better understand the marine carbonate
system, which is prerequisite for modeling related to
climate and atmospheric CO2. Aragonite, a metastable
polymorph of CaCO3, is an important phase of marine
carbonate. As shelled pteropods (Euthecosomatous
pteropods) are known to be the major source to marine
carbonates, there is increasing interests among
paleoceanographers to estimate pteropod flux in ocean
basins and its seasonal variation particularly in high
productive areas e.g. the Arabian Sea. With the
development of sediment traps, particle flux studies got a
great impetus in recent years. The available carbonate
particle flux data from moored sediment traps deployed in
the Arabian Sea are primarily limited to calcitic biogenic
components, thus neglecting aragonitic component
(Conan et al., 2002). Only a few studies were made on
pteropod fluxes in the western side of the Arabian Sea
(Mohan et al., 2006; Singh and Conan, 2008). These
studies recorded seasonal and inter-annual variations in
pteropod species in response to hydrographic changes
linked to the seasonal monsoonal variations. Singh and
Conan (2008) estimated that about 22.5 % of the total
CaCO3 produced in western Arabian Sea was contributed
by aragonitic pteropods. Therefore, aragonitic
component of CaCO3 plays a major role in regulating
carbon sequestration by the Arabian Sea. As the aragonite
is more soluble than calcite, calcification of pteropods is
expected to be severely affected by rising atmospheric
CO2 concentration in recent time that would result
acidification of seawater and decline of the saturation state
with respect to aragonite. To have better understanding of
biogeochemical processes and carbon cycle in the Arabian
Sea basin wide, pteropod flux studies need to be extended
to other regions.
DISTRIBUTION OF PTEROPODS IN RECENT
SEDIMENTS
Auras-Schudnagies et al. (1989) recorded the
distribution of pteropods in surface sediments of the
northwestern Arabian Sea. This study was limited to
deeper regions. As pteropods are preserved in sea bottom
sediments lying above the ACD, they abundantly occur in
marginal sediments of the Arabian Sea. Therefore,
Arabian Sea margins are ideal locations to study
distribution pattern of pteropods. Among the few studies
made on recent pteropods from marginal areas, the most
important was by Singh and co-workers (Singh et al.,
1998, 2005) who mapped distribution of pteropod species
over entire western continental shelf of India and
correlated them with modern bathymetric and
hydrographic conditions.
Species of pteropods in surficial sediment samples
over the western Indian shelf fall into the two major
groups: mesopelagic (deeper water dwellers) and
epipelagic (shallow water dwellers). The quantitative data
reflect that the relative abundances of epipelagic and
mesopelagic components in assemblage vary with the
water depths across the shelf. All those species recorded
from <50 m depths are exclusively epipelagic, whilst
assemblages from >50 m depths comprise both
mesopelagic and epipelagic species. The abundance of
mesopelagic pteropods increases with the depths, which
suggests that the vertical depth habit of pteropods is an
important factor controlling their abundance pattern
across the western continental shelf of India.
The majority of pteropod species encountered in the
Arabian Sea have a wide range of salinity tolerance, as
they recorded frequently in high salinity waters of the Red
Sea (Weikert, 1987) and also in the south China Sea where
salinity is as low as ~30 ‰ (Rottman, 1980). Thus, it is
unlikely that the differences in absolute salinity of surface
waters are responsible for pteropod distribution pattern in
surficial sediments. Instead, a prominent variation in the
vertical (surface-subsurface water) salinity gradient
across the basin might have profoundly influenced the
abundance of pteropods (Singh et al., 2005). A high
vertical salinity gradient as observed along the southwest
coast of India could be an obstacle for migratory
pteropods. Pteropod studies by Singh et al. (1998, 2005)
also demonstrated sensitiveness of various pteropod
species to variations in bathymetry and hydrographic
parameters such as temperature, salinity and oxygen
concentration. It was found that distribution patterns of
species like L. inflata, C. acicula and C. chierchaiae in
marginal areas are controlled by bathymetry and oxygen
concentration of seawater has strong influence upon the
 A.D. Singh and O. P. Singh
Fig. 2. Latitudinal variations of relationship between L. inflata / Creseis spp. ratios and water depths. [A : southern sector, B : Central
sector, C : Northern sector of western continental margin of India] (Data Source, Singh et al., 2005).
distribution patterns of L. inflata, C. convexa and S.
subula. Thus, distribution pattern studies of recent
pteropods in the Arabian Sea enabled to identify various
pteropod indicators useful in reconstruction of
paleoceanographic history from fossil assemblages. The
data on modern pteropods of the Arabian Sea generated in
recent years (Singh et. al., 1998, 2005) are being
effectively used as analogue to deduce past changes based
on the faunal variation in sedimentary records of the
western continental margin (Bharti, 2008; Singh, 2007;
Singh et al., 2001, 2006 , 2007)
PTEROPOD AS SEA LEVEL INDICATORS
The data on distribution of pteropods in shelf
sediments off west coast of India clearly indicate certain
species (L. inflata, Creseis acicula, C.chierchiae) to be
depth sensitive (Herman and Rosenberg ,1969; Singh et
al., 1998). In general, an opposite trend in relative
abundance of L. inflata (mesopelagic) and Creseis spp.
(epipelagic) across the shelf was noticed, indicating
bathymetric control on their distributions. Therefore,
these species are considered as potential bathymetric
indicators. Singh et al. (1998) prepared a model on
relationship between variations in Limacina inflata /
Creseis spp. ratio and water depths across the western
continental shelf of India and applied this model to fossil
records to decipher changes in sea level in the region
during the late Quaternary (Singh et al., 2001).
Furthermore, it was suggested that sensitivity of these
pteropod species to water depth changes limited to open
shelf regimes. In their recent study, Singh et al. (2005)
documented regional variation in response of depth
sensitive pteropod species to the bathymetric conditions.
The range of variation in L. inflata/Creseis spp. ratio in
response to bathymetric change conspicuously differs
along the shelf with its maximum 1.5 in northern, 4 in
central and 7 in southern sectors of the western continental
shelf of India (Fig. 2). This latitudinal variation was
suggested to be attributed to regional difference in
hydrographic parameters of sea surface waters (mainly
salinity) along the west coast of India. Thus, a model on
L.inflata/ Creseis spp. ratio and water depth relationship is
suggested to be applicable in deciphering paleo-
bathymetry from the sediment cores of the same regions.
LATE QUATERNARY SEA – LEVEL HISTORY
ALONG THE CENTRAL WEST COAST OF INDIA
BASED ON PTEROPODS
Using model developed by Singh et al. (1998) for the
relations between modern bathymetric conditions and the
L. inflata/ Creseis spp. ratio to the pteropod assemblages
of late Quaternary sedimentary records, Singh et al.
(2001) reconstructed history of sea level changes along
the central west coast of India for the last 30 kyr BP
(Fig. 3). The study revealed that sea level was around 25 m
below the present mean sea level (PMSL) at ~ 36 kyr BP.
Sea level lowered at slower rate between 25 kyr BP. At ~
30 kyr BP, the sea level stood at around 40 m below
PMSL. During the last glacial maximum, sea level was at
~ 100 m below PMSL. Between 10 and 15 kyr BP, sea
level rose rapidly from ~ 80 m to ~ 20 m. Since 7 kyr BP,
rise of sea level has been at slow pace.
USEFULNESS OF PTEROPODS IN QUATERNARY
PALEOCENOGRAPHY WITH SPECIAL REFERENCE
TO THE ARABIAN SEA
Variations in pteropod assemblages (species
abundance and preservation state of shells) through time
 Pteropods and Quaternary climatic of the Arabian
Fig. 3. Inferred sea level variation along the northern Kerala
coast during the last 36 kyr, based on the pteropod record (Data
Source, Singh et al., 1998, 2001).
have been widely used in reconstruction of the Quaternary
oceanographic history of the marginal seas surrounding
the Arabian Sea (e.g. Almogi- Labin, 1984; Almogi-Labin
et al., 1986, 1988, 1991, 1998). In recent years, serious
efforts have been made to investigate fossil pteropods of
the Arabian Sea in understanding various aspects of
oceanographic changes during the Quaternary (Reichart
et al., 1998, 2002; von Rad et al., 1999; Klocker et al.,
2006; Singh, 1998, 2007; Singh et al., 2006, 2007;
Ivanova, 1999; Bharti, 2008). A summary of important
findings made through theses studies are discussed
below:
PRESERVATION STATE OF PTEROPOD SHELLS
Several studies have revealed variation in pteropod
shell preservation in sediments relating to past changes in
carbonate chemistry of intermediate deep water masses
associated with fluctuations in climatic condition and
circulation pattern (e.g. Almogi-Labin et al., 1986, 1998;
Reichart et al., 1998, Singh et al., 2006; Singh, 1998,
2007). The established micropaleontological parameters
indicating preservation condition of pteropod shells
which are commonly used are: (1) abundance ratio of
pteropods (Pt) and planktic foraminifera (Pf), and (2)
transparency of pteropod shells. As compared to calcitic
planktic foraminifera, aragonitic pteropods are more
susceptible to dissolution. Thus, abundance ratio of
pteropods to planktic foraminifera [(Pt/Pt+Pf)%] indicate
degree of dissolution/preservation of pteropod shells
(Fig. 4). High values of pteropod to planktic foraminifera
abundance ratio are indicative of better preservation
condition, whereas low values suggest a greater loss of
pteropod shells due to dissolution.
Pteropod shells in sediments are characterised by
various states of preservation ranging from well preserved
transparent shells to very poorly preserved broken white
shells. Among pteropod population, L. inflata and C.
acicula are known to be dissolution prone species (Singh
et al., 2005). Thus, variation pattern of the percentage
abundance of transparent shells in total population of
these species indicate preservation condition of aragonite
in sediments (Fig. 4). Preservation of pteropod shells in
sediments depends upon the saturation state of the
overlying bottom water mass with respect to aragonite.
Arabian Sea is characterized by strong seasonal variation
in monsoon induced primary productivity. High primary
productivity results oxygen minimum condition in middle
of the water column between 150 and 1200 m depths
called Oxygen Minimum Zone (OMZ) (Altabet et al.,
1995). The seawater in the OMZ is undersaturated with
respect to aragonite due to consumption of dissolved
oxygen, lowering the ph and carbonate saturation because
of addition of CO2 and oxidation of ammonia to nitrate
(Canfield and Raiswell, 1991). The Arabian Sea because
of its unique geographical setting is poorly ventilated.
Thus, the dissolved oxygen concentration or in other
words the strength of the OMZ controls the depth of
aragonite compensation and consequently the
preservation of aragonite (pteropod shells) in sediments
(Singh, 2007). Therefore, variation in the intensity of the
OMZ directly results into changes in the ACD. As
preservation of pteropod (aragonite) in sediments is
related to the ACD, record of pteropod abundance spikes
in sediment cores can be valuable imprints of changes in
the OMZ intensity and the ACD.
LATE QUATERNARY VARIABILITY PATTERN OF
PTEROPOD ABUNDANCE AND PRESERVATION
RECORDS IN RELATION TO CHANGES IN OMZ
INTENSITY , ACD AND MONSOON STRENGTH
In recent years, several workers have investigated
abundance and preservation records of pteropods in late
Quaternary sediments of western (Ivanova, 1999; Klocker
et al., 2006), northern (Reichart et. al., 1998; von Rad et
al., 1999) and eastern Arabian Sea (Singh et al., 2006;
Singh, 1998,2007; Bharti, 2008) in order to understand
variability patterns of the OMZ intensity, ACD and its
relation to the changes in monsoonal strength, deep water
circulation and ventilation. Thus, pteropod record holds a
key to infer oxygenation history.
Based on their detailed studies of pteropod
assemblages from late Quaternary sediment records of
western continental margins of India, Singh et al. (2006)
and Singh (2007) documented pteropod rich intervals at
certain times (Fig. 4). These pteropod abundance spikes
correspond to aragonite peaks suggesting pteropod shells
as a prime source of aragonite to the sediments. However,
there could be other minor contributors of aragonite like
molluscans other than pteropods or/and corals, as
reflected by the aragonite profile of the examined core
 A.D. Singh and O. P. Singh

Fig. 4. Records of absolute abundance of pteropods, relative abundance of transparent shells of a dissolution susceptible species L.
inflata, abundance ratio of pteropod to planktic foraminifera and ratio of aragonite/calcite wt. percentages in a gravity core retrieved
from the Arabian Sea Indian margin off Goa (Data source, Singh, 2007).

showing presence of aragonite in the late Holocene
sediment, even though pteropod shells are absent. The
predominance of transparent shells in pteropod
population during intervals of aragonite maxima indicates
that the aragonite spikes in core are the results of
enhanced preservation (Fig. 4). The radiocarbon
chronology of these aragonite layers in sediment cores
suggest that these events are coeval with the Heinrich
events of north Atlantic associated with cold phases of
northern Hemisphere (Singh et al., 2006; Singh, 2007;
Bharti, 2008). A comparison of these millennial events
with the records from the western (Ivanova, 1999;
Klocker et al., 2006) and northern Arabian Sea (Sirocko et
al., 1993, von Rad et al., 1999) indicates that these events
are isochronous within the resolution of the radiocarbon
dates in the Arabian Sea. Interestingly, carbonate rich and
organic carbon (OC) poor sediments characterize the
intervals of pteropod abundance maxima (Singh, 2007;
Singh et al., 2006). The OC poor, bioturbated sediments
are deposited under depositional conditions of increased
oxygen level of water column and reduced productivity.
The Arabian Sea productivity is mainly governed by the
summer monsoon winds. During northern Hemisphere
cold periods, the strength of summer monsoon and
associated wind induced productivity was weakened
(Reichart et al., 1998; Singh et al., 2006). The low surface
productivity and increased oxygen concentrations during
these periods might have resulted a weak OMZ inducing
lowering of the ACD in the Arabian Sea and thus better
preservation of pteropod shells. On the other hand,
intervals of poor preservations of pteropods indicate
periods of strong OMZ and shallow ACD. High primary
productivity due to strong monsoonal winds, results
pronounced oxygen depletion in the water column,
consequently an intensified OMZ. A strong OMZ leads
shallow aragonite lysocline causing poor preservation of
pteropod shells in sediments. Singh et al. (2006) also
suggested that incoming oxygenated waters originated
outside of the basin could also influence water column
oxygen condition in the Arabian Sea.
 Pteropods and Quaternary climatic of the Arabian
RECORD OF PTEROPODS IN DEEP – SEA
SEDIMENTS: NEOTECTONIC IMPLICATIONS
The occurrence of pteropod shells in deep-sea
sediments below aragonite compensation depth is a rare
and unusual event. Singh et al. (2000) in their recent
study, reported pteropod rich layers from a deep-sea
sediment core (Lat. 15° 35. 52’ N, Long. 68° 35.09 ‘E;
3820 m water depth) of the eastern Arabian Sea spanning
the last 135 kyr. The pteropod assemblage comprises of
both the epipelagic and mesopelagic forms. It was
intriguing to record well-preserved aragonite shells from
3.8 km below the present day ACD estimated as at about
500 m. There could be two possibilities for deep
occurrence of pteropod shells: 1) considerable deepening
of the ACD during certain periods, 2) transportation of the
pteropod shells to deeper region from the shallow area.
Singh et al. (2000) ruled out first possibility because
available data on the ACD variations during the
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ACKNOWLEDGEMENTS
Pteropod studies carried out by ADS and his co-
workers were funded by the Department of Ocean
Development, the Department of Science and Technology,
New Delhi and the Indian Space Research Organization.
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