Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera from Ocean Drilling Program (ODP) Sites 752, 1168 and 1139, southern Indian Ocean

Abstract

Deep-sea benthic foraminifera provide important markers of
environmental conditions in the deep-ocean basins where their assemblage
composition and test chemistry are influenced by ambient physical and
chemical conditions in bottom-water masses. However, all foraminiferal
studies must be underpinned by robust taxonomic approaches. Although many
parts of the world's oceans have been examined, over a range of geological
timescales, the Neogene benthic foraminifera from the southern Indian Ocean
have only been recorded from a few isolated sites. In this study, we have
examined 97 samples from Neogene sediments recovered from three ODP sites in
the southern Indian Ocean (Sites 752, Broken Ridge; 1139, Kerguelan Plateau; 1168,
west Tasmania). These data cover a range of palaeolatitudes and water depths
during the Miocene. More than 200 species of benthic foraminifera were
recorded at each site and, despite their geographic and bathymetric
separation, the most abundant taxa were similar at all three sites. Many of
these species range from late Oligocene to early Pliocene
demonstrating relatively little faunal turnover of the most abundant taxa
during the key palaeoclimatic shifts of the Miocene. We illustrate and
document the occurrence of the 52 most abundant species (i.e. those with
>1 % abundance) encountered across the three study sites.

Full text

J. Micropalaeontology, 38, 189–229, 2019
https://doi.org/10.5194/jm-38-189-2019
© Author(s) 2019. This work is distributed under
the Creative Commons Attribution 4.0 License.
Latest Oligocene to earliest Pliocene deep-sea
benthic foraminifera from Ocean Drilling Program (ODP)
Sites 752, 1168 and 1139, southern Indian Ocean
Dana Ridha, Ian Boomer, and Kirsty M. Edgar
Earth Sciences, School of Geography, Earth and Environmental Sciences,
University of Birmingham, Edgbaston, B15 2TT, UK
Correspondence: Ian Boomer (i.boomer@bham.ac.uk)
Received: 26 July 2019 – Revised: 1 November 2019 – Accepted: 11 November 2019 – Published: 16 December 2019
Abstract. Deep-sea benthic foraminifera provide important markers of environmental conditions in the deep-
ocean basins where their assemblage composition and test chemistry are influenced by ambient physical and
chemical conditions in bottom-water masses. However, all foraminiferal studies must be underpinned by robust
taxonomic approaches. Although many parts of the world’s oceans have been examined, over a range of geolog-
ical timescales, the Neogene benthic foraminifera from the southern Indian Ocean have only been recorded from
a few isolated sites. In this study, we have examined 97 samples from Neogene sediments recovered from three
ODP sites in the southern Indian Ocean (Sites 752, Broken Ridge; 1139, Kerguelan Plateau; 1168, west Tasma-
nia). These data cover a range of palaeolatitudes and water depths during the Miocene. More than 200 species
of benthic foraminifera were recorded at each site and, despite their geographic and bathymetric separation, the
most abundant taxa were similar at all three sites. Many of these species range from late Oligocene to early
Pliocene demonstrating relatively little faunal turnover of the most abundant taxa during the key palaeoclimatic
shifts of the Miocene. We illustrate and document the occurrence of the 52 most abundant species (i.e. those with
>1 % abundance) encountered across the three study sites.
1 Introduction
The Neogene can be informally subdivided into an early
warm interval and late cool interval (Flower and Kennett,
1993). Superimposed on this long-term cooling is the Mid-
Miocene climatic optimum (MMCO) between ∼15 and
17 Ma, marking the warmest interval of the Neogene (Zachos
et al., 2001). Cooling following the MMCO culminated at
∼13–14 Ma in major growth of the East Antarctic Ice Sheet
(EAIS). Further cooling coincided with increasing intensifi-
cation of Antarctic deep-water formation and growth of the
AIS and Greenland ice sheet (Flower and Kennett, 1993,
1994, 1995).
Profound palaeoclimatic and palaeoceanographic changes
in the Neogene drove progressive and significant turnover
of deep-sea benthic foraminiferal communities from the so-
called transitional fauna characterising the late Oligocene–
early Miocene to the late Neogene fauna of the late Miocene
and finally the establishment of recognisably modern assem-
blages in the mid-Pleistocene (Jones, 1994; Kaiho, 1994).
The first major transitional interval occurred in the early-to-
middle Miocene, between ∼13–17 Ma (Kaiho, 1994), span-
ning the onset of the MMCO, subsequent East Antarctic
Ice Sheet growth, and associated changes in water mass
character and upwelling intensity (Woodruff, 1985). Further
changes in assemblages that occur in the late Miocene, ∼8–
10 Ma, relate to global cooling and expansion of the Antarc-
tic ice sheet (Gupta et al., 2004). Finally, the last global ex-
tinction of benthic foraminifera (∼20 % of deep-sea gen-
era) occurred in the late Pliocene–mid-Pleistocene (Hayward
et al., 2012) and is variously attributed to increasing ocean
productivity and intensification of low-oxygen zones and
changes in the food type and supply, due to the expansion
of the Antarctic and later Northern Hemisphere ice sheets
and global cooling (Hermoyian and Owen, 2001; Gupta et
al., 2004; Smart et al., 2007).
Published by Copernicus Publications on behalf of The Micropalaeontological Society.

190 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Superimposed on this long-term global pattern of benthic
foraminifer assemblage change are distinct basinal and lati-
tudinal differences in the timing of certain biotic events. For
instance, the global decline in deep-sea benthic foraminifer
species abundance and richness occurs much earlier (late
Oligocene–Miocene) in the Southern Ocean than in the In-
dian Ocean (late Miocene; Hayward et al., 2012). However,
the response of deep-sea benthic fauna in the Indian Ocean to
these global changes remains relatively poorly constrained.
Previous records of Miocene benthic foraminifera from
the central and southern Indian Ocean are largely single-
site reports with a focus on palaeoceanographical interpre-
tations, though some provide useful illustration for compar-
ison with the present study. Boersma (1990) reported ben-
thic foraminifera from 12 ODP sites from leg 115 across the
central Indian Ocean, where quantitative studies of the ben-
thic foraminifera indicated faunal separation at intermediate,
deep and abyssal water depths from the late Oligocene on-
wards, and hence the development of a more strongly ver-
tically stratified ocean. Nomura (1991) systematically de-
scribed the most abundant Oligocene-to-Pleistocene ben-
thic foraminifera species and assemblage changes at ODP
Sites 754 (Broken Ridge) and 756 (Ninetyeast Ridge) in the
eastern Indian Ocean, linking them with palaeoceanographic
changes.
There have been two studies focussing on the Kergue-
len Plateau. Schroder-Adams (1991) investigated middle-
Eocene-to-Holocene benthic foraminifera from Sites 736,
738 and 744 on the Kerguelen Plateau (Fig. 1), de-
scribing and illustrating many species. Significant faunal
changes (and species replacements) were noted in the mid-
dle Eocene, late-Eocene–early-Oligocene, middle Miocene
and late Miocene. Mackensen (1992) quantitatively studied
benthic foraminiferal assemblages from the lower Miocene
through to upper Pleistocene interval from ODP Site 747
(Central Kerguelen Plateau) and Sites 748 and 751 (South-
ern Kerguelen Plateau).
Gupta and Satapathy (2000) studied abyssal benthic
foraminifera from 31 samples of the latest Miocene to Pleis-
tocene from Deep Sea Drilling Project (DSDP) Site 236,
north of the Mascarene Plateau. This study investigated the
link between benthic foraminifera distribution, productivity
and deep-ocean thermohaline circulation in the west-central
Indian Ocean during that period.
The main aim of this work is to provide a taxonomic
and stratigraphic reference for the most common Miocene
benthic foraminifera encountered in the southern Indian
Ocean, supported by scanning electron microscopy (SEM)
images. The study deals with foraminifera encountered in
ODP Holes 752A, 1139A and 1168A, all in the southern part
of the Indian Ocean (Fig. 1). Of these, the latter two have
not previously been investigated for benthic foraminiferal
assemblages; whilst Neogene benthic foraminifera assem-
blages have been used for palaeoceanographic reconstruc-
tions in Hole 752A (Singh et al., 2012), no systematic species
description or reference images were provided. The present
study expands our understanding of the spatial and temporal
distribution of the most abundant benthic foraminifera from
the deep-sea Miocene sediments of this region.
2 Materials and methods
ODP Site 752 is situated on the Broken Ridge in the south-
eastern Indian Ocean (30◦53.47500 S, 93◦34.65200 E; Fig. 1)
at a current water depth of 1086 m (Shipboard Scientific
Party, 1989). The palaeodepth reconstruction for the site
is upper bathyal in the late Oligocene shallowing to mid-
lower bathyal in the Miocene (Shipboard Scientific Party,
1989). The current study covered a 79.7m interval of sedi-
ments from Hole 752A, which includes the latest Oligocene
through to early Pliocene ages; the average core recovery
is 70.6 %, composed mainly of foraminifer and nannofos-
sil ooze. An age model for Hole 752A was developed us-
ing the calcareous nannofossil datums from the initial report
(Shipboard Scientific Party, 1989), with revised ages from
the Astronomically Tuned Neogene Time Scale (ATNT2012)
in GTS2012 (Hilgen et al., 2012).
ODP Site 1139 is located west of Kerguelen Island
(50◦1102000 S, 63◦5504000 E; Fig. 1) with a modern water
depth of 1415 m consistent with a bathyal palaeodepth
throughout the late Oligocene to earliest late Miocene (Ship-
board Scientific Party, 2000). The late Oligocene to earli-
est late Miocene interval was sampled from Hole 1139A,
comprising 167.3 m with average core recovery of 62 %.
The sediments are foraminifera-bearing nannofossil ooze,
nannofossil-bearing clay, and claystone and nannofossil-
bearing ooze and chalk. An age model has been estab-
lished using planktic foraminifera (Shipboard Scientific
Party, 2000) and calcareous nannofossil datums (Persico et
al., 2003) with ages revised using ATNT2012 in GTS2012
(Hilgen et al., 2012).
ODP Site 1168 is situated at a middle bathyal water depth
(2463 m) on the western margin of Tasmania (42◦3804000 S,
144◦2503000 E; Fig. 1) (Shipboard Scientific Party, 2001). The
sedimentary interval studied in Hole 1168A covers 422.8m
with an average recovery of 98%. This interval covers the
latest Oligocene through to early Pliocene, which is mainly
composed of nannofossil ooze, silty clay and claystone. The
age model for Hole 1168A uses planktic foraminifera, cal-
careous nannofossils and geomagnetic polarity datums cal-
culated by Stickley et al. (2004) with revised ages from
ATNT2012 in GTS2012 (Hilgen et al., 2012).
In total, 97 samples were selected from the study sites (30,
25 and 42 samples at ODP Sites 752, 1139 and 1168, re-
spectively). Sediment samples were processed by first dis-
aggregating them in water and then wet sieving at 63µm.
Samples were then dried overnight in an oven at 40◦C,
the dry >63 µm sieved size fraction was then split until an
aliquot of >300 benthic foraminifer specimens was achieved
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 191
Figure 1. Location map of the study area. Studied ODP sites are marked as stars (752, 1168 and 1139), additional sites referred to in the
text are located by dots (after Weatherall et al., 2015). Sites 752, 754 and 756 (leg 121), Site 1168 (leg 189), Site 1139 (leg 183), Sites 705–
716 (leg 115) and Deep Sea Drilling Project (DSDP) Site 236 (leg 24). The isobath adjacent to the Kerguelen Plateau, Broken Ridge and
Ninetyeast Ridge is 3000 m, while the isobath near the Maldives, Chagos, Saya de Malha, Nazareth and Seychelles banks is 2000 m.
and subsequently picked clean of all benthic foraminifer.
The 52 most abundant species encountered (of more than
500 species provisionally determined across all three sites)
were selected for this work as they represent ∼79 % of the
total benthic foraminifera specimens found across all sites.
Scanning electron microscope images of representative (un-
coated) specimens of each selected species were taken on a
Phenom Desktop SEM at the University of Birmingham. The
total size range of specimens observed within the study was
also recorded by measuring the longest axis of the specimens
using a reticle on a stereo microscope.
3 Foraminiferal assemblages
In the current study the generic and suprageneric classifica-
tion of the foraminifera follows Loeblich and Tappan (1988,
1994). The classification of Loeblich and Tappan (1994) is
broadly the same as that of Loeblich and Tappan (1988) with
some modification of high-level categories as given in Loe-
blich and Tappan (1992). Specifically, the elevation of the
foraminifera to a higher taxonomic rank by Lee (1990) to
the class Foraminiferea was accepted by Loeblich and Tap-
pan (1992) and Kaminski (2004). The key sources for species
identification are Wright (1978), Hayward, (2002), Hayward
et al. (2012) and Holbourn et al. (2013). Descriptions are
based on existing literature with additional observations from
this study.
Benthic foraminifer assemblages are relatively well-
preserved with specimens largely complete and surficial de-
tails preserved. Assemblages are diverse with more than 200
species identified in total in each site (500 within the entire
study) and are dominated by calcareous benthic foraminifera.
This diversity is higher than previously documented in the In-
dian Ocean (e.g. Boersma, 1990; Nomura, 1991; Mackensen,
1992; Schroder-Adams, 1991). The 52 selected species de-
scribed here are assigned to 33 different genera and are sub-
divided into four orders, 15 superfamilies, 20 families and 16
subfamilies. Four recorded species are left in open nomen-
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192 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
clature, two each from Bolivina and Gyroidinoides. All taxa
described are calcareous hyaline forms with the exception of
one agglutinated species. Representatives of the Buliminida
(31 species) constitute the majority of those recorded, espe-
cially in ODP Hole 752A, but they are also both abundant
and diverse in Holes 1139A and 1168A. Species of the Ro-
taliida are the next most important components of the benthic
foraminifer assemblages.
4 Taxonomy
Class Foraminifera J. J. Lee, 1990
Order Buliminida Fursenko, 1958
Superfamily Bolivinoidea Glaessner, 1937
Family Bolivinidae Glaessner, 1937
Genus Bolivina d’Orbigny, 1839b
Bolivina dilatata Reuss 1850
Figure 3: 1a–b
1850 Bolivina dilatata Reuss: p. 381, pl. 48, fig. 15a–c
1949 Bolivina subspathulata Boomgaart, p. 112, pl. 12,
fig. 4
2016 Bolivina dilatata Reuss; Lei and Li, p. 202, pl. 2,
figs. a–h
Description
The test is biserial elongate, has perforate hyaline wall, the
shape of the chambers is triangular to trapezoidal. The su-
tures are depressed. The aperture is a slit-like to arcuate
opening with a thin tooth plate in the middle. The aperture
location is terminal.
Dimensions
The species length ranges from 125 to 250 µm in the current
study.
Stratigraphic range
This species is recorded in the Aquitanian of 1139 (ODP Site
1139) and late Oligocene (Chattian) to early Pliocene (Zan-
clean) at 1168. This species is not recorded at 752, consistent
with earlier studies (Nomura, 1995).
Remarks
This is considered a cosmopolitan species (Lei and Li, 2016)
but to the best of our knowledge this is the first time it has
been recorded in the Indian Ocean.
Bolivina huneri Howe, 1939
Figure 3: 2a–b
1939 Bolivina huneri Howe: p. 66, pl. 9, figs. 3, 4
1983 Bolivina huneri Howe; Tjalsma and Lohmann,
p. 23, pl. 11, fig. 5
1987 Bolivina huneri Howe; Miller and Katz, p. 124,
pl. 1, figs. 7a–b
2013 Bolivina huneri Howe; Holbourn et al., p. 68,
figs. 1–3
Description
Very densely ornamented biserial species with tapered test.
The test surface has a reticulate ornamentation largely ob-
scuring sutures with the exception of the final chamber. The
inflated chambers are elongated. The aperture is situated at
the base of the last chamber and is arcuate with wide aper-
ture view and tooth plate.
Dimensions
The species length ranges from 175 to 250 µm in the current
study.
Stratigraphic range
This species is recorded from the Chattian through to early
Burdigalian of 752. At 1168 the species is confined to
the Miocene and ranges from Burdigalian to Tortonian; the
species is not recorded at 1139. These ranges fall within the
middle-Eocene–late-Miocene (Tortonian) range determined
by Holbourn et al. (2013).
Remarks
The distribution is cosmopolitan (Holbourn et al., 2013)
and has previously been recorded on the Kerguelen Plateau,
southern Indian Ocean, at ODP Sites 748 and 751 (Mack-
ensen, 1992) and 736, 738 and 744 (Schroder-Adams, 1991).
Bolivina reticulata Hantken, 1875
Figure 3: 3a–b
1875 Bolivina reticulata Hantken: p. 65, pl. 15, fig. 6
1978 Bolivina reticulata Hantken; Wright, p. 711, pl. 2,
figs. 8–10
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 193
Description
Broad biserial test with wide, slightly inflated, elongate
chambers, characteristically covered by narrow anastomos-
ing ribs. The test is slightly perforated and has a narrow keel
around the periphery. Depressed and limbate sutures are ob-
served. The aperture is sub-rounded and located at the base
of the last chamber with an internal tooth plate. The species
is characterised by its sub-rounded shape in side view and
slightly compressed in cross section.
Dimensions
The species length ranged from 150 to 375 µm in the current
study.
Stratigraphic range
The species is recorded at 752 from the Chattian to Serraval-
lian, from the Miocene and early Pliocene (Zanclean) at 1168
and Langhian–Tortonian at 1139.
Remarks
This species is recorded in the Mediterranean (Wright, 1978),
South Atlantic Ocean (Tjalsma, 1983). This species has a
highly variable morphology both in terms of test shape and
ornamentation (Wright, 1978). This species is included by
some authors as a junior synonym of Latibolivina subretic-
ulata (Parr, 1932) by Loeblich and Tappan (1994) and Hol-
bourn et al. (2013), and it is primarily recorded from the trop-
ical Indian Ocean and Pacific Ocean (Holbourn et al., 2013).
Thus, this species likely has a global distribution.
Bolivina cf. reticulata Hantken, 1875
Figure 3: 5a–b
cf. 1875 Bolivina reticulata Hantken: p. 65, pl. 15, fig. 6
cf. 1978 Bolivina reticulata Hantken; Wright, p. 711,
pl. 2, figs. 8–10
cf. 2013 Latibolivina subreticulata (Parr); Holbourn et
al., p. 327, figs. 1–4
Description
The test is biserial, with hyaline wall composition. The
chamber shape is elongate to trapezoid. The suture is de-
pressed and limbate. The ornamentation is of anastomosing
ribs. The aperture is basal and rounded with a tooth plate.
Dimensions
The species length ranges from 125 to 375 µm in the current
study.
Stratigraphic range
The species is only observed at 752 through the Miocene and
early Pliocene (Zanclean). This species is highly abundant in
the mid-to-late Miocene.
Remarks
This species is similar to Bolivina reticulata (Hantken, 1875)
but B. reticulata has a broader (particularly towards the
base) and relatively more elongate test, with a more com-
pressed periphery in apertural view. According to Wright
(1978), Bolivina reticulata varies in shape from an elongate
form with limbate sutures to specimens with little ornamen-
tation. This species is also similar to Bolivina viennensis
(Marks, 1951) reported in the Polish Carpathian Foredeep
Basin (Central Paratethys) by Peryt (2013). This form has so
far only recorded in the Indian Ocean.
Bolivina sp. 3
Figure 3: 4a–c
Description
The test is biserial and elongate with smooth surface. Some
specimens are highly perforated. The chambers are elongate
to trapezoid in shape. The sutures are slightly depressed. The
aperture is sub-rounded and located at the base of the last
chamber.
Dimensions
The species length ranged from 150 to 250 µm in the current
study.
Stratigraphic range
The species is only observed in the late Oligocene (Chattian)
of 752. At 1139 it occurs between the Chattian and Serraval-
lian. At 1168 the species ranges throughout the Miocene and
into the early Pliocene (Zanclean).
Remarks
This species differs from Bolivina dilatata (Reuss, 1850), by
having a longer test relative to its width (2.5 times long as
wide rather than 2 :1); the sutures are extremely shallow. The
current species is similar to the Bolivina cf. dilatata maxima
(p. 148, fig. 7, pl. k) recorded in the Central Paratethys (Peryt,
2013) but the latter has more depressed sutures. The cur-
rent species differs from Bolivina spathulata (Williamson,
1858), which has sharp edge in the apertural view, whereas
specimens here have a sub-rounded edge in apertural view.
Also, specimens from the China seas (Lei and Li, 2016) have
an anastomosing imperforate costae ornamentation, and they
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194 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
are much larger ∼550 µm (2 times longer than those ob-
served here). This species is so far only been recorded in the
southern Indian Ocean (current study).
Bolivina sp. 9
Figure 3: 6a–b
Description
The test is finely perforated, smooth and elongate, with a
hyaline wall composition. Chambers are broad, trapezoid and
biserially arranged, with a depressed suture between cham-
bers. Aperture is loop-shaped with basal position. The test
length is nearly 3 times longer than the width. Test is tapered
at both ends.
Dimensions
The species length ranges from 125 to 250 µm in the current
study.
Stratigraphic range
The species occurs through the Miocene and into the early
Pliocene (Zanclean) at 752. The species is very abundant in
the early Burdigalian at 752. Only very few specimens have
been recorded at 1168, mainly in the Chattian and between
the Burdigalian to Serravallian with a very few records in
the early Pliocene (Zanclean). This species is not recorded at
1139.
Remarks
This species is similar to Bolivinellina pseudopunctata (Alve
and Murray, 2001) recorded in intertidal environments from
southern England (Ibid; pl. 2, figs. 7–9). The current species
has a wider loop-shaped aperture with the last chamber
slightly irregular, while B. pseudopunctata has a narrower,
slightly compressed aperture and the shape of the chambers
are consistently trapezoidal. This species differs from Bo-
livina spathulata (Williamson, 1858) by having a big and
rounded (loop-shaped) aperture on the apertural face rather
than a small and narrow loop-shaped aperture. Also, Boliv-
ina spathulata specimens from the China seas (Lei and Li,
2016) have an anastomosing imperforate costae ornamenta-
tion, not been observed in the current study.
Bolivina viennensis Marks, 1951
Figure 3: 7a–b
1951 Bolivina viennensis Marks: p. 60, pl. 7, fig. 1
1991 Brizalina albatrossi (Cushman); Nomura, p. 53,
pl. 1, fig. 3
2005 Bolivina viennensis Marks; Popescu and Crihan,
p. 380, pl. 1, figs. 14–16
2008 Bolivina viennensis Marks; Baldi and Hoheneg-
ger, p. 2, fig. 3.a
Description
The test is biserial, elongated. Chambers elongated to trape-
zoid, sutures depressed. Surface ornamentation reticulate
showing an anastomosing pattern. Aperture rounded, termi-
nal. Normal pores are relatively few and concentrated near
the margin of the chambers.
Dimensions
The species length ranges from 175 to 575 µm in the current
study.
Stratigraphic range
This species is only present at 752 and ranges from the Burdi-
galian to Messinian, reappearing in the early Pliocene (Zan-
clean).
Remarks
This species is similar to Bolivina cf. reticulata but the length
of Bolivina viennensis is nearly 3 times longer than the width,
and the test has a sharp end near the proloculus, while Boliv-
ina cf. reticulata is shorter and is sub-rounded near the pro-
loculus. This species is recorded as Bolivina viennensis in
the southern Vienna Basin (Austria) (Baldi and Hohenegger,
2008) and in Romania (Popescu and Crihan, 2005). Similar
specimens are recorded as Brizalina albatrossi in the east-
ern Indian Ocean at Site 754 in the Broken Ridge (Nomura,
1991).
Genus Bolivinellina Saidova, 1975
Bolivinellina cf. B. translucens (Phleger & Parker,
1951)
Figure 3: 8a–c
cf. 1951 Bolivina translucens Phleger & Parker: p. 15,
pl. 7, figs. 13, 14
cf. 1988 Bolivinellina translucens (Phleger & Parker);
Loeblich and Tappan: pl. 547, figs. 6, 7
cf. 1999 Bolivina translucens (Phleger & Parker);
Ohkushi et al., p. 139, pl. 2, figs. 11, 12
Description
Test is elongate, triserial to biserial, narrow and oval in
section. Chambers are narrow and moderately inflated; su-
tures are depressed and oblique. Wall calcareous, hyaline and
smooth; a few pores apparent near the sutures. Aperture is
basal and slit-to-loop-shaped with a tooth plate at the base of
the aperture. Test nearly 3–4 times longer than wide.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 195
Dimensions
The species length ranges from 175 to 475 µm in this study.
Stratigraphic range
The species is only found at Site 752 where it is distributed
throughout the succession, with the greatest abundance in the
late Miocene (Messinian) and early Pliocene (Zanclean).
Remarks
The chambers appear triserially arranged in the initial part of
the test, with inflated chambers which become less inflated
later towards the aperture. Bolivinellina translucens is widely
distributed in Gulf of Mexico (Phleger and Parker, 1951) and,
more locally, has been recorded along the central West Coast
of India (Mazumder et al., 2003) and throughout the Indian
Ocean (current study).
Superfamily Bolivinitoidea Cushman, 1927
Family Bolivinitidae Cushman, 1927
Genus Abditodentrix Patterson, 1985
Abditodentrix pseudothalmanni (Boltovskoy & de
Kahn, 1981)
Figure 3: 9a–b
1981 Bolivinita pseudothalmanni Boltovskoy & de
Kahn: p. 44–46, pl. 1, figs. 1–5
1985 Abditodentrix asketocomptella Patterson, p. 139,
pl. 1, figs. 1–9
1988 Abditodentrix pseudothalmanni (Boltovskoy & de
Kahn); Loeblich and Tappan, p. 503, pl. 554, figs. 1–5
cf. 1991 Brizalina thalmanni (Renz, 1948), Nomura,
p. 53, pl. 1, fig. 4
Description
Test elongate, biserial and sagittate. The test is flattened in
side view with a distinctive truncate margin giving it a rect-
angular appearance in section. Chambers are elongated to
trapezoid. The sutures are depressed. Aperture is in a basal
position and elliptical in shape with tooth plate infolded on
one side. The surface is densely covered with elevated retic-
ulate ornamentation.
Dimensions
The species length ranges from 100 to 425 µm in this study.
Stratigraphic range
Occurs in the late Oligocene (Chattian) in all study sites,
ranging into the early late Miocene (early Tortonian) in 1139
and the early Pliocene (Zanclean) at 752 and 1168.
Remarks
Most of the observed specimens are highly compressed in
cross section and densely ornamented in the initial portion,
near the proloculus, with simple reticulation or no ornamen-
tation near the aperture. This species is recorded in the south-
west Pacific Ocean (Hayward et al., 2002) and the south-west
Atlantic Ocean (Loeblich and Tappan, 1988). It is recorded
as Brizalina cf. thalmanni in the eastern Indian Ocean (No-
mura, 1991).
Superfamily Buliminoidea Jones in Griffith and Hen-
frey, 1875
Family Buliminidae Jones, 1875
Genus Bulimina d’Orbigny, 1826
Bulimina truncana Gümbel, 1868
Figure 3: 10a–b
1868 Bulimina truncana Gümbel: p. 727, pl. 2, figs. 77a,
b
1884 Bulimina rostrata Brady: p. 408, pl. 5, figs. 14, 15
1927 Bulimina alazanensis Cushman: p. 161, pl. 25,
fig. 4
1940 Bulimina bremneri Finlay: p. 455, pl. 64, figs. 84–
86
1991 Bulimina alazanensis Cushman: Schroder-Adams,
p. 624, pl. 1, fig. 5
1991 Bulimina truncana Gümbel: Hermelin, p. 65, pl. 1,
figs. 10, 16
2002 Bulimina truncana Gümbel: Hayward et al., pl. 1,
figs. 9, 10
Description
Test is triserial, elongate and sub-triangular in cross section,
tapered in outline. Rounded to sub-rounded in the apertural
end and pointed in the apical end. The chambers are inflated
with obscured sutures. The longitudinal costae occur on all
chambers (although the upper portion of the final three cham-
bers is smooth) and join together to form a spinose protru-
sion from the earliest chambers. The aperture is loop-shaped
with internal tooth plate and surrounded by apertural lip. The
greatest width of the test is towards the apertural end.
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196 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Dimensions
The species length ranges from 125 to 625 µm in the current
study.
Stratigraphic range
The species is recorded from the late Oligocene to Pliocene
at 752 and 1168, whereas it is confined to the Miocene (Aqui-
tanian to Serravallian) at 1139.
Remarks
Boersma (1986) noted that Bulimina truncana has a highly
variable morphology and thus multiple synonyms are based
on differences in the number and extent of costae, their height
and orientation. Of these, small forms with fewer costae that
almost completely cover the final chamber have been of-
ten been attributed to Bulimina rostrata, whereas those with
more costae, that also tend to be wavy, are assigned to Bu-
limina alazanensis. Furthermore, B. rostrata also achieves
its maximum width towards the middle of the test and is
overall sub-circular in cross section compared to B. alaza-
nensis where the maximum test width occurs closer to the
apertural end and the test has a more sub-triangular cross
section. B. truncana is very similar to B. alazanaensis but
was separated out based on having fewer, straighter costae
that begin on the first or second chambers rather than the
third (Boersma, 1986). Ultimately overlapping species oc-
currences at the same sites, difficulties in consistently dif-
ferentiating taxa and missing holotype specimens (B. alaza-
nensis) have led to them being synonymised (Hayward et al.,
2019a) in the World Register of Marine Species (WoRMS).
Here, tests are usually sub-triangular in cross section, hav-
ing parallel, regularly formed costae with the largest width
of the test towards that apertural end rather than the middle,
and hence they are closest to the forms included in B. trun-
cana. This species is considered to be cosmopolitan and has
been recorded in Southwest Pacific Ocean (Hayward et al.,
2002), Tasman sea, Coral Sea (Boersma, 1986) and Indian
Ocean (Nomura, 1995).
Bulimina cf. B. elongata d’Orbigny, 1846
Figure 3: 11a–b
cf. 1846 Bulimina elongata d’Orbigny: p. 187, pl. 11,
figs. 19, 20
cf. 1826 Bulimina elongata d’Orbigny: (Nomen
nudum), p. 269
cf. 2004 Bulimina elongata d’Orbigny: De Man et al.,
p. 187 , pl. 2, fig. 9
cf. 2008 Bulimina elongata d’Orbigny: Kender et al.,
p. 513, pl. 17, figs. 4, 5
Description
Elongate, narrow (3 times as long as wide), triserial test be-
coming nearly biserial in final whorl. Little tapering of test.
Chambers are flattened and broad. Sutures are depressed. The
aperture is basal and rounded. The surface is mainly smooth
with very few perforations.
Dimensions
The species length ranges from 150 to 400 µm in this study.
Stratigraphic range
This species ranges from the late Oligocene (Chattian) to
mid-Burdigalian at 1168, extending into the early Tortonian
at 1168. This supports the range (mid-Eocene to Pleistocene)
identified by Thomas (1990). This species is not present at
752.
Remarks
This species differs from Bulimina elongata sensu stricto
(see Holbourn et al., 2013) in that the chambers are less in-
flated (have a nearly polygonal outline) and a much slower
rate of chamber enlargement. Holbourn et al. (2013) consider
Bulimina elongata a cosmopolitan species.
Bulimina gibba Fornasini, 1902
Figure 3: 12
1902 Bulimina gibba Fornasini: p. 378, pl. O, figs. 32–
34
2005 Bulimina gibba Fornasini; Rasmussen, p. 86,
pl. 11, fig. 5
2013 Bulimina gibba Fornasini; Holbourn et al., p. 97,
figs. 1–3
Description
Triserial, elongate, fusiform shape. Nearly circular in cross
section with narrow apical tip. Chambers are inflated with
clearly depressed sutures and strongly overlapping. The aper-
ture is loop-shaped, located at the base of the last chamber
with apertural lip merging with an internal tooth plate. The
test surface is relatively smooth with perforation distributed
across the chambers.
Dimensions
The species length ranges from 125 to 300 µm in the current
study.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 197
Stratigraphic range
This species occurs from the late Oligocene (Chattian) to
early Tortonian at 1139. This species is not recorded at ei-
ther 752 or 1168.
Remarks
This species is commonly found and described (e.g. Hol-
bourn et al., 2013) as having an apical protrusion from the
proloculus; however, in this study, no spinose protrusions are
observed in any specimen. This absence cannot be explained
by preservation. This species has previously only been re-
ported from the Atlantic (Holbourn et al., 2013), but the au-
thors noted that it may well range further, which is confirmed
in this study. Holbourn et al. (2013) report this species from
the Miocene to Holocene but this work suggests it may occur
earlier in the Indian Ocean.
Bulimina striata d’Orbigny in Guérin-Méneville, 1832
Figure 3: 13
1832 Bulimina striata d’Orbigny in Guérin-Méneville:
p. 18, pl. 3, fig. 16
1852 Bulimina costata d’Orbigny: p. 194
1922 Bulimina inflata var. mexicana Cushman: p. 95,
pl. 21, fig. 2
1953 Bulimina mexicana Cushman; Phleger and others,
p. 33, pl. 6, fig. 27
1954 Bulimina striata mexicana Cushman; Parker,
p. 511, pl. 6, fig. 24
1957 Bulimina australis Vella: p. 32, pl. 8, figs. 168–
169
1992 Bulimina striata d’Orbigny in Guérin-Méneville:
Mackensen, p. 668, pl. 1, fig. 5
2003 Bulimina striata d’Orbigny in Guérin-Méneville:
Hayward et al., p. 517
2011 Bulimina striata d’Orbigny in Guérin-Méneville:
Kuppusamy et al., p. 46, pl. 2, figs. 7, 8
2013 Bulimina striata d’Orbigny in Guérin-Méneville:
Mancin et al., p. 64, pl. 2, fig. 3
Description
Test is conical, triserial, and semicircular in cross section;
the initial part of the test (near proloculus) is acute. Cham-
bers are inflated overhanging previous chambers, becoming
broader and less ornamented near the aperture. The sutures
are depressed but largely obscured by ornamentation in the
earlier portion of the test. Longitudinal costae extend from
the lower half of the chambers to overhang preceding cham-
bers ending in acute points. The aperture is loop-shaped and
bordered by a lip with an internal tooth plate.
Dimensions
The species length ranges from 100 to 500 µm in this study.
Stratigraphic range
The species is recorded throughout the whole succession
(late Oligocene–Pliocene) at 752. At 1139 and 1168 this
species occurs from the late Oligocene (Chattian) disappear-
ing from 1139 in the Langhian and early Tortonian at 1168.
Remarks
This species is cosmopolitan. This species is identified
around New Zealand (Debenay, 2012; Hayward et al., 2003)
and by Kuppusamy et al. (2011) in north-west Atlantic
Ocean. Many older Indian Ocean publications report this
species as B. mexicana (e.g. Nomura, 1995), which does have
a global distribution (Holbourn et al., 2013). Holbourn et
al. (2013) consider the range of this species to be Miocene–
Holocene; here it is found in the late Oligocene.
Superfamily Turrilinoidea Cushman, 1927
Family Turrilinidae Cushman, 1927
Subfamily Turrilininae Cushman, 1927
Genus Eubuliminella Revets, 1993
Eubuliminella exilis (Brady, 1884)
Figure 4: 1
1884 Bulimina elegans var. exilis Brady: p. 399, pl. 50,
figs. 5, 6
1927 Buliminella subfusiformis var. tenuata Cushman:
p. 149, pl. 2, fig. 9
1930 Buliminella subfusiformis (Cushman); Cushman
and Moyer, p. 56, pl. 7, fig. 20
1947 Bulimina exilis Brady; Cushman and Parker,
p. 123, pl. 28, figs. 27, 28
1947 Bulimina subfusiformis var. tenuata (Cushman);
Cushman and Parker, p. 124, pl. 28, fig. 29
1993 Eubuliminella exilis (Brady); Revets, p. 141, pl. 1,
figs. 1–7
2015 Eubuliminella exilis (Brady); Setoyama & Kamin-
ski, p. 8, figs. 4.2, 8.7
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198 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Description
Elongate, narrow test that is triserial in the latter part. The
test length is about 4 times as long as it is wide. Ovate and
little inflated chambers separated by curved and depressed
sutures. The aperture is basal and loop-shaped, but from the
apertural opening the tooth plate is not clear.
Dimensions
The species length ranges from 150 to 550 µm in the current
study.
Stratigraphic range
The stratigraphic range of the species at 1168 is early
Miocene (Burdigalian) to the early Pliocene (Zanclean); Hol-
bourn et al. (2013) suggested a range from the late Miocene;
the present study therefore extends the known stratigraphic
range.
Remarks
The specimens in the current study lack the spine in the initial
whorl and have more weakly inflated chambers. According
to Revets (1993) and Holbourn et al. (2013), this species can
be found worldwide.
Superfamily Cassidulinoidea d’Orbigny, 1839a
Family Cassidulinidae D’Orbigny, 1839a
Subfamily Cassidulininae d’Orbigny, 1839a
Genus Globocassidulina Voloshinova, 1960
Globocassidulina subglobosa (Brady, 1881)
Figure 4: 2
1881 Cassidulina subglobosa Brady: p. 60, illustrated
in Brady, 1884, pl. 54, fig. 17
1884 Cassidulina subglobosa Brady; Brady, p. 430,
pl. 54, fig. 17
1983 Globocassidulina subglobosa (Brady); Tjalsma
and Lohmann, p. 31, pl. 16, fig. 9
1991 Globocassidulina subglobosa (Brady); Nomura,
p. 55, pl. 5, fig. 8
2013 Globocassidulina subglobosa (Brady); Holbourn
et al., p. 264, figs. 1, 2
Description
Test is biserial and sub-globular, with a sub-circular shape
in cross section. Chambers are inflated and separated by de-
pressed, curved sutures. The test wall is smooth and only
finely perforate. The aperture is slit-like, straight or curved.
Dimensions
The species length ranges from 100 to 925 µm in this study.
Stratigraphic range
This is a long-ranging species, recorded from late Oligocene
to early Pliocene at both 752 and 1168 and late Oligocene to
the top of the studied interval (early Tortonian) at 1139.
Remarks
This species is very well-preserved in all studied sites.
This is a cosmopolitan species (Jones, 1994) that is very
long-ranging (Paleocene–Holocene; Tjalsma and Lohmann,
1983).
Genus Paracassidulina Nomura, 1983b
Paracassidulina minuta (Cushman, 1933b)
Figure 4: 3a–b
1933b Cassidulina minuta Cushman: p. 92, pl. 10,
figs. 3a–c
1951 Cassidulina crassa d’Orbigny; Phleger and
Parker, p. 26, pl. 14, figs. 4a–b
1954 Cassidulina aff. crassa d’Orbigny; Parker, p. 535,
pl. 11, fig. 1
1983b Paracassidulina minuta Cushman; Nomura,
p. 66, pl. 5, figs. 16a–c
1991 Paracassidulina minuta Cushman; Nomura, p. 55,
pl. 5, figs. 2a–b
Description
Biserial test with slightly depressed sutures. Surface is
smooth and finely perforated. Chambers are strongly curved.
Aperture is long, curved and has an apertural ridge parallel
to the chamber periphery.
Dimensions
The species length ranges from 125 to 300 µm in the current
study.
Stratigraphic range
The species ranges from Chattian to early Tortonian at 752,
Chattian to mid-Serravallian at 1139. The stratigraphic distri-
bution of the species at 1168 is early Aquitanian to Zanclean.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 199
Remarks
The chambers of the specimens found here are only slightly
inflated, this differs from Globocassidulina minuta (Cush-
man, 1933b) described from the Paumotu Islands (also called
Tuamotu Islands) in the Southern Pacific Ocean in that the
latter has more globular chambers. This species is recorded
by Nomura (1991) at Broken Ridge (Site 754) in eastern In-
dian Ocean and in the Gulf of Mexico (Poag, 2015).
Subfamily Ehrenbergininae Cushman, 1927
Genus Ehrenbergina Reuss, 1850
Ehrenbergina carinata Eade, 1967
Figure 4: 4a–b
1967 Ehrenbergina carinata Eade: p. 448, pl. 9, figs. 1–
4
1991 Ehrenbergina carinata Eade; Nomura, p. 55, pl. 5,
fig. 13
2013 Ehrenbergina carinata Eade; Hayward et al.,
p. 442, pl. 6, figs. 12, 13
Description
Test triangular in both dorsal view and cross section. Early
chambers are not inflated, while the later chambers are
slightly inflated. Well-developed peripheral keel and a spine
extending from each chamber. Dorsal view is compressed.
The sutures are flush but towards the periphery become
raised. The ventral view is characterised by central keel.
Aperture is slit-like and curved, with a narrow lip in the ven-
tral side.
Dimensions
The species length ranges from 150 to 725 µm in the current
study.
Stratigraphic range
The species range at 752 is late Miocene (Messinian) to early
Zanclean. At 1168 the species is very rare and only recorded
in the early and late Miocene (Tortonian). This species is not
recorded at 1139.
Remarks
This species is well-preserved at 752 and reaches very large
size (up to 725 µm) in some intervals, especially in the late
Miocene (Messinian). According to Eade (1967) the current
species is recorded in the Kermadec Islands, Fiji, Norfolk Is-
land, Tonga, and northern New Zealand. This species is also
recorded by Nomura (1991) at Broken Ridge (Site 754) and
New Zealand (Hayward et al., 2013).
Genus Burseolina Seguenza, 1880
Burseolina pacifica (Cushman, 1925)
Figure 4: 5a–b
1925 Cassidulina pacifica Cushman: p. 53, figs. 14–16
1925 Globocassidulina pacifica (Cushman), pl. 2,
fig. 16
1983a Burseolina pacifica (Cushman); Nomura, pl. 6,
fig. 2, pl. 21, figs. 6–10
1983b Burseolina pacifica (Cushman); Nomura, pp. 57–
60, pl. 5, figs. 1–4
1991 Burseolina cf. pacifica (Cushman); Nomura, p. 53,
pl. 5, figs. 14a–b
Description
Enrolled biserial test with rounded periphery. Sutures are
flush to the surface and thus almost indistinguishable. The
apertural face is flattened and the aperture is elongate,
curved, and with apertural flap in the lower apertural margin.
Dimensions
The species length ranges from 175 to 600 µm in this study.
Stratigraphic range
The distribution of the species at 752 ranges from the late
Oligocene (Chattian), where it is rare, with later Miocene
records from the Burdigalian to mid-Messinian. The species
is rare at 1168 in the Chattian and only appears in the late
Burdigalian in the Miocene. This species is not recorded at
1139.
Remarks
According to Cushman (1925) this species is widely dis-
tributed in deep or cool waters and it has been recorded in
the Pacific Ocean. It has also been reported from subtropical
regions, along the Pacific coast of Japan (Nomura, 1983a)
and in the Indian Ocean (Nomura, 1991). Notably the spec-
imens here are more spherical in outline than B. pacifica re-
ported by Nomura (1991). Burseolina is most abundant from
the Miocene to the present.
Burseolina cf. pacifica (Cushman, 1925)
Figure 4: 6
cf. 1925 Cassidulina pacifica Cushman: p. 53, figs. 14–
16
cf. 1925 Globocassidulina pacifica (Cushman), pl. 2,
fig. 16
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200 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
cf. 1983a Burseolina pacifica (Cushman); Nomura,
pl. 6, fig. 2, pl. 21, figs. 6–10
cf. 1983b Burseolina pacifica (Cushman); Nomura,
pp. 57–60, pl. 5, figs. 1–4
cf. 1991 Burseolina pacifica (Cushman); Nomura, p. 53,
pl. 5, figs. 4a–b
Description
Test is biserial and sub-spherical. The surface is smooth.
Flush to slightly depressed sutures. The aperture is elongated,
curved and with apertural flap in the lower apertural margin.
Dimensions
The species length ranges from 100 to 425 µm in this study.
Stratigraphic range
The range of the species at 752 is early Miocene (Aquitanian)
to early Pliocene (Zanclean). At 1139 it is very rare in the late
Oligocene (Chattian) and also in the Miocene, except for Ser-
ravallian where it has its greatest abundance. The distribution
at 1168 is mid-Burdigalian to Zanclean.
Remarks
This species differs from Burseolina pacifica by having a
relatively larger and wider aperture and broader flap. The
last chambers in this species are also usually separated from
the preceding chambers by clearly depressed sutures. This
species has a limited geographic distribution and is also ob-
served in tropical to subtropical regions, along the Pacific
coast of Japan (Nomura, 1983a) and in the Indian Ocean on
the Ninetyeast Ridge (Site 756) (Nomura, 1991).
Superfamily Pleurostomelloidea Reuss, 1860
Family Pleurostomellidae Reuss, 1860
Subfamily Pleurostomellinae Reuss, 1860
Genus Pleurostomella Reuss, 1860
Pleurostomella acuminata Cushman, 1922
Figure 4: 7a–b
1922 Pleurostomella acuminata Cushman: p. 50, pl. 19,
fig. 6
1866 Pleurostomella alternans Schwager: p. 238, pl. 6,
fig. 80
1934 Pleurostomella alternans var. telostoma Schubert;
Cushman, pl. 16, fig. 6
1978 Pleurostomella acuta Hantken; Boltovskoy, pl. 5,
fig. 42
1978 Pleurostomella dominicana Bermúdez;
Boltovskoy, pl. 6, figs. 2, 3
2011 Pleurostomella acuminata (Cushman); Kup-
pusamy et al., p. 60, pl. 9, figs. 7, 8
Description
Test is smooth, elongate and biserial, with slightly depressed
sutures. Chambers are elongate and oval. Wide terminal aper-
ture, with projecting hood on one side and two triangular or
bifid teeth on opposite side. Prolocular spine is present.
Dimensions
The species length ranges from 150 to 725 µm in the current
study.
Stratigraphic range
At 752 and 1168 it ranges from Chattian to early Zanclean
and at 1139 from Chattian to early Tortonian.
Remarks
A prolocular spine is present in most specimens but in others
it is either broken or missing. This species is cosmopolitan
and recorded throughout the Indian Ocean (Hayward, 2002).
Superfamily Stilostomelloidea Finlay, 1947
Family Stilostomellidae Finlay, 1947
Genus Orthomorphina Stainforth, 1952
Orthomorphina perversa (Schwager, 1866)
Figure 4: 8a–c
1866 Nodosaria perversa Schwager: p. 212, pl. 5,
fig. 29
1872 Nodosaria pupoides Silvestri: p. 65, pl. 6,
figs. 148–158
1937 Nodogenerina challengeriana Thalmann: p. 341,
pl. 64, figs. 25–27
1949 Nodogenerina aminaensis Bermúdez: p. 178,
pl. 11, fig. 57
1978 Orthomorphina challengeriana (Thalmann);
Boltovskoy, pl. 5, figs. 16–17
1978 Orthomorphina perversa (Schwager); Boltovskoy,
pl. 5, figs. 23–24
1992 Stilostomella sp. G. Kaiho: p. 307, pl. 5, fig. 23
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 201
2002 Orthomorphina perversa (Schwager); Hayward,
pl. 1, figs. 50–52
Description
Test uniserial, rectilinear. Chambers are ovoid in outline and
increase in height towards the last chamber. The surface is
striate with ribs perpendicularly arranged. The sutures are
straight and depressed. The aperture is rounded and termi-
nal with rim.
Dimensions
The species length ranges from 200 to 975 µm in the current
study.
Stratigraphic range
The species ranges from Chattian to early Zanclean at 752
and 1168 but only extends into the mid-Serravallian (mid-
Miocene) at 1139.
Remarks
The largest chamber is usually in the middle part of the test
and the last chamber is smaller than the penultimate one.
Sometimes this difference is very pronounced. This species
is cosmopolitan (Hayward, 2002).
Genus Siphonodosaria Silvestri, 1924
Siphonodosaria lepidula Schwager, 1866
Figure 4: 9a–b
1866 Nodosaria lepidula Schwager: p. 210, pl. 5,
figs. 27, 28
1978 Stilostomella ex.gr. lepidula (Schwager).
Boltovskoy, pl. 7, figs. 33–36
1980 Stilostomella lepidula (Schwager); Srinivasan and
Sharma, p. 46, pl. 7, figs. 1–6
1993 Stilostomella lepidula (Schwager); Gupta, 1993,
figs. 2, 1–2
1994 Nodogenerina lepidula (Schwager); Loeblich and
Tappan, p. 133, pl. 261, figs. 18–20
2002 Siphonodosaria lepidula f. lepidula (Schwager);
Hayward, p. 305, pl. 3, figs. 25–32
2012 Siphonodosaria lepidula (Schwager); Hayward et
al., p. 174, pl. 18, figs. 6–20
Description
Elongate, uniserial test, slightly arcuate shape, with cham-
bers gradually increasing in size. Chambers are spheri-
cal (campanulate or pyriform) and typically become more
widely spaced later with pronounced depressed sutures. A
ring of small spines, annularly arranged, surrounds the base
of each otherwise relatively smooth-walled chamber. Aper-
ture is terminal with a broad neck and phialine lip; there is
small tooth in the middle of the aperture giving a v-shaped
appearance to aperture.
Dimensions
The species length ranges from 200 to 1350 µm in this study.
Stratigraphic range
The distribution at 752 and 1168 is Chattian to early Zan-
clean and at 1139 is Chattian to early Tortonian.
Remarks
This species is abundant, with a cosmopolitan distribution
(Hayward et al., 2012; Holbourn et al., 2013). It is also highly
morphologically variable showing considerable variation in
ornamentation, chamber shape and the presence or absence
of a prolocular spine, and hence has been frequently recorded
under a number of different names (Hayward et al., 2012).
After detailed population level investigations, Hayward et
al. (2012) group the different morphotypes under S. lepidula
is known from the Late Cretaceous to Pleistocene, reports of
recent occurrences are considered reworked (Hayward et al.,
2012).
Siphonodosaria subspinosa (Cushman, 1943)
Figure 4: 10a–d
1934 Ellipsonodosaria sp. Cushman and Jarvis: pl. 10,
figs. 4, 5
1943 Ellipsonodosaria subspinosa Cushman: p. 92,
pl. 16, figs. 6, 7b
1983 Stilostomella subspinosa (Cushman); Tjalsma and
Lohmann, p. 36, pl. 14, figs. 16, 17
1998 Siphonodosaria subspinosa (Cushman); Robert-
son, p. 180, pl. 67, fig. 3
2010 Siphonodosaria subspinosa (Cushman); Hayward
et al., p. 129, pl. 14, figs. 20, 21
Description
Test elongate,uniserial, straight to slightly arcuate shape, cir-
cular in cross section. Chambers are spherical, separated by
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202 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
strongly depressed sutures. Short spines cover the surface.
Chambers show a relatively rapid increase in size towards
the aperture creating a tapered test outline. The aperture is
terminal with broad neck and phialine lip.
Dimensions
The species length ranges from 200 to 1125 µm in the current
study.
Stratigraphic range
The distribution at 752 and 1168 is Chattian to early Zan-
clean, and at 1139 Chattian to early Tortonian.
Remarks
Specimens are very well-preserved and abundant at Site 752.
Holbourn et al. (2013) comment that this species shows large
variability in the size and ornamentation of the test, with
short spines commonly found on the test sometimes converg-
ing to form fine costae in the lower portion of the test. This
is consistent with observations in this study. Distribution is
worldwide (Holbourn et al., 2013).
Genus Stilostomella Guppy, 1894
Stilostomella cf. S. annulifera Cushman and Bermúdez,
1936
Figure 4: 11a–b
cf. 1936 Ellipsonodosaria annulifera Cushman and
Bermúdez: p. 28, pl. 5, figs. 8, 9
1978 Stilostomella cf. S. annulifera Wright: p. 717,
pl. 8, fig. 2
cf. 1990 Stilostomella annulifera (Cushman and
Bermúdez); Thomas, 1990, p. 590, pl. 1, fig. 4
cf. 1998 Siphonodosaria annulifera (Cushman and
Bermúdez); Robertson, p. 178, pl. 66, figs. 1, 2
Description
Test elongate, rectilinear, arcuate in overall shape with a
smooth wall texture. Chambers are inflated to sub-spherical.
Sutures are limbate and shallow. Chambers become bigger
and more elongated towards apertural end. Aperture is ter-
minal with phialine lip.
Dimensions
The species length ranges from 225 to 1250 µm in this study.
Stratigraphic range
The distribution of the species at 1139 is Chattian to early
Tortonian and at 1168 and 752 from Chattian to early Zan-
clean.
Remarks
Specimens here largely lack the prolocular spine because
of preservation (see Fig. 4: 11a). This species is similar to
Strictocostella scharbergana Neugeboren, 1856, but the last
chamber of S. annulifera is slightly bigger and more inflated
with a shorter neck. Some publications (e.g. Hayward et al.,
2012) synonymise these two species and use S. scharber-
gana.S. consobrina differs from this species by having su-
tures which are flusher to the test, as shown in Holbourn et
al. (2013). It has also been observed that the chambers of this
species are particularly large in those specimens recorded in
Mediterranean Sea (Wright, 1978). S. cf. S. annulifera has
a cosmopolitan distribution (Holbourn et al., 2013) includ-
ing records in the Mediterranean Sea (Wright, 1978) and
Antarctica (Thomas, 1990). S. annulifera ranges from the
early Eocene to middle Miocene (Holbourn et al., 2013), but
S. scharbergana is considered to range from the early Pale-
ocene to mid-Pleistocene (Hayward et al., 2012), more con-
sistent with this extended younger occurrence.
Genus Strictocostella Patterson, 1987
Strictocostella matanzana (Palmer & Bermúdez, 1936)
Figure 4: 12a–c
1936 Ellipsonodosaria?matanzana Palmer &
Bermúdez: p. 298, pl. 18, fig. 12
1940 Ellipsonodosaria matanzana Palmer and
Bermúdez; Palmer, pl. 15, fig. 1
1957 Siphonodosaria cf. matanzana (Palmer and
Bermúdez); Todd, pl. 67, fig. 7
1975 Stilostomella lepidula Saidova: p. 315, pl. 87,
fig. 6
1998 Siphonodosaria matanzana (Palmer and
Bermúdez); Robertson, pl. 67, fig. 2
2007 Myllostomella matanzana (Palmer and
Bermúdez); O’Neill et al., p. 1085, pl. 1, figs. 22–
23
2010 Strictocostella matanzana (Palmer and
Bermúdez); Hayward et al., p. 128, figs. 14, 27–
28
2012 Strictocostella matanzana (Palmer and
Bermúdez); Hayward et al., p. 190, pl. 22, figs. 10–20
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 203
Figure 2. Range chart of the 52 most abundant benthic foraminifera species encountered at Ocean Drilling Program (ODP) Holes 752A,
1139A and 1168A.
Description
Test elongate, uniserial, rectilinear. The base is composed
of few small, spherical chambers which become larger and
more elongate towards the aperture. Sutures are straight and
depressed. There are small spines forming a ring towards the
base of each chamber. The aperture is terminal with a phia-
line lip and short, broad neck ornamented by pustular spines.
Dimensions
The species length ranges from 200 to 750 µm in the current
study.
Stratigraphic range
The species occurs rarely in the Chattian at Site 752 and it
is not recorded in Aquitanian–Langhian, but appears again
from early Serravallian to Zanclean. At 1139 the distribution
of the species is from Chattian to early Tortonian; it is not
recorded at 1168.
Remarks
The wall of the neck is folded, giving an undulation to the
phialine lip. S. matanzana has a cosmopolitan distribution;
sister species tend to be more prevalent in the Indian Ocean
(Hayward et al., 2012).
Strictocostella scharbergana (Neugeboren, 1856)
Figure 5: 1a–b
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204 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Figure 3. Scale bar is 100 µm. (1a–b) Bolivina dilatata, ODP Hole 1168A, 37X-6, 75–79 cm: (1a) side view; (1b) apertural view. (2a–
b) Bolivina huneri, ODP Hole 752A, 10H-5, 77–81 cm. (2a) side view; (2b) apertural view. (3a–b) Bolivina reticulata, ODP Hole 1168A,
15X-2, 76–80 cm: (3a) side view; (3b) apertural view. (4a–b) Bolivina sp. 3, ODP Hole 1168A, (4a–b) 26X-2, 73–77 cm: (4a) side view;
(4b) apertural view. (4c) 37X-6, 75–79 cm: side view. (5a–b) Bolivina cf. reticulata, ODP Hole 752A, (5a) 5H-1, 65–70cm: apertural view.
(5b) 3H-5, 62–67 cm: side view. (6a–b) Bolivina sp. 9, ODP Hole 752A, 9H-2, 104–109cm: (6a) side view; (6b) apertural view. (7a–
b) Bolivina viennensis, ODP Hole 752A, 5H-1, 65–70 cm: (7a) side view; (7b) apertural view. (8a–c) Bolivinellina cf. B. translucens, ODP
Hole 752A, 3H-1, 50–55 cm: (8a–b) side view; (8c) apertural view. (9a–b) Abditodentrix pseudothalmanni, ODP Hole 1139A, 19R-2, 60.5–
64.5 cm: (9a) side view; (9b) apertural view. (10a–b) Bulimina truncana,(10a) ODP Hole 752A, 9H-5, 65–70 cm: lateral view. (10b) ODP
Hole 1168A, 37X-6, 75–79 cm: apertural side view. (11a–b) Bulimina cf. B. elongata,(11a) ODP Hole 1168A, 23X-3, 74–78cm: apertural
side view. (11b) ODP Hole 1139A, 18R-1, 96–100 cm: apertural side view. (12) Bulimina gibba, ODP Hole 1139A, 19R-2, 60.5–64.5 cm:
lateral view. (13) Bulimina striata, ODP Hole 752A, 5H-1, 65–70 cm: apertural side view.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 205
Figure 4. Scale bar is 100 µm. (1) Eubuliminella exilis, ODP Hole 1168A, 42X-3, 70–74 cm: apertural side view. (2) Globocassidulina sub-
globosa, ODP Hole 752A, 4H-1, 70–75 cm: apertural side view. (3a–b) Paracassidulina minuta, ODP Hole 1168A, (3a) 42X-3, 70–74 cm:
apertural side view. (3b) 37X-6, 75–79, 0–1 cm: side view. (4a–b) Ehrenbergina carinata, ODP Hole 752A, 3H-1, 50–55cm: (4a) apertural
side view; (4b) side view. (5a–b) Burseolina pacifica, ODP Hole 752A, 6H-1, 70–75 cm: (5a) apertural side view; (5b) side view. (6) Burse-
olina cf. pacifica, ODP Hole 752A, 6H-1, 70–75 cm: apertural side view. (7a–b) Pleurostomella acuminata,(7a) ODP Hole 752A, 7H-1, 63–
68 cm: apertural side view. (7b) ODP Hole 1168A, 22X-2, 45–49cm: lateral view. (8a–c) Orthomorphina perversa,(8a) ODP Hole 1139A,
14R-4, 65–69 cm: side view. (8b, c) ODP Hole 752A, 3H-1, 50–55cm: (8b) side view; (8c) apertural view. (9a–b) Siphonodosaria lepidula,
ODP Hole 752A, (9a) 7H-2, 73–78 cm: side view. (9b) ODP Hole 8H-1, 100–105cm: apertural view. (10a–d) Siphonodosaria subspinosa,
(10a–b) ODP Hole 752A, 8H-6, 70–75 cm: (10a) side view; (10b) apertural view. (10c) ODP Hole 1168A, 18X-5, 80–84cm: side view.
(10d) ODP Hole 1168A, 15X-2, 76–80 cm: apertural view. (11a–b) Stilostomella cf. S. annulifera, ODP Hole 1139A, 15R-4, 41–45cm:
(11a) side view; (11b) apertural view. (12a–c) Strictocostella matanzana,(12a–b) ODP Hole 752A, 3H-5, 62–67 cm: (12a) side view;
(12b) apertural view. (12c) ODP Hole 1139A, 18R-1, 96–100 cm: side view.
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206 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Figure 5. Scale bar is 100 µm. (1a–b) Strictocostella scharbergana, ODP Hole 752A, 4H-1, 70–75 cm: (1a) side view; (1b) apertural
view. (2a–c) Trifarina angulosa,(2a) ODP Hole 752A, 6H-1, 70–75 cm: side view. (2b, c) ODP Hole 1168A, 23X-3, 74–78cm: (2b) side
view; (2c) apertural view. (3a–c) Trifarina bradyi,(3a) ODP Hole 752A, 8H-6, 70–75cm: side view. (3b) ODP Hole 1168A, 15X-2, 76–
80 cm: side view. (3c) ODP Hole 1139A, 5R-1, 121–125cm: side view. (4a–c) Siphouvigerina hispida,(4a–b) ODP Hole 1168A, 20X-3,
100–104 cm: (4a) side view; (4b) apertural view. (4c) ODP Hole 1139A, 7R-1, 111–115cm: side view. (5a–d) Uvigerina auberiana, ODP
Hole 752A, (5a) 10H-2, 60–65 cm: side view. (5b) 3H-5, 62–67cm: side view. (5c, d) 4H-2, 81–85 cm: (5c) side view; (5d) apertural
view. (6a–b) Uvigerina peregrina,(6a) ODP Hole 752A, 9H-5, 65–70cm: side view. (6b) ODP Hole 1168A, 22X-2, 45–49cm: side view.
(7a–b) Lenticulina cultrata, ODP Hole 752A, 10H-5, 77–81 cm; (7a) side view; (7b) edge view.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 207
1856 Dentalina scharbergana Neugeboren: p. 87, pl. 4,
figs. 1–4
1856 Strictocostella scharbergana Neugeboren, p. 87,
pl. 4, figs. 1–4
1866 Dentalina costai Schwager: p. 229, pl. 6, fig. 62
1884 Nodosaria consobrina (d’Orbigny); Brady, pl. 62,
figs. 23, 24
1936 Ellipsonodosaria annulifera Cushman and
Bermúdez: p. 28, pl. 5, figs. 8, 9
1960 Stilostomella consobrina (d’Orbigny); Barker,
pl. 62, figs. 23, 24
1977 Siphonodosaria consobrina (d’Orbigny);
Nishimura et al., pl. 4, figs. 16, 17
1994 Stilostomella consobrina (d’Orbigny); Jones,
1994, pl. 62, figs. 23, 24
2000 Stilostomella scharbergana (Neugeboren);
Sztràkos, pl. 5, fig. 19
2005 Myllostomella costai (Schwager); Hayward and
Kawagata, pl. 1, figs. 17, 18
2007 Strictocostella costai (Schwager); Hayward et al.,
fig. 2
2010 Strictocostella scharbergana (Neugeboren); Hay-
ward et al., p. 128, figs. 14, 28–29
2012 Strictocostella scharbergana (Neugeboren); Hay-
ward et al., p. 192, pl. 23, figs. 8–19
Description
Test arcuate, uniserial, slender. Proloculus spherical but
chambers become more ovate towards the aperture. Surface
is smooth. Sutures flush or slightly incised. Aperture is ter-
minal with a short neck and phialine lip, internal apertural
rim consists of a long, simple tooth and multiple long, fine
denticles.
Dimensions
The species length ranges from 200 to 1400 µm in the current
study.
Stratigraphic range
The species range at 752A is Chattian to early Zanclean. This
species is not recorded at 1168 and 1139.
Remarks
Wall smooth throughout, occasionally with one pustular
spine near the base of the proloculus. Hayward et al. (2012)
consider S. scharbergana conspecific with Stilostomella con-
sobrina, and in many works it is identified as S. scharber-
gana (e.g. Sztràkos, 2000; Hayward et al., 2010, 2012). Cos-
mopolitan distribution (Hayward et al., 2012).
Family Uvigerinidae Haeckel, 1894
Subfamily Angulogerininae Galloway, 1933
Genus Trifarina Cushman, 1923
Trifarina angulosa (Williamson, 1858)
Figure 5: 2a–c
1858 Uvigerina angulosa Williamson: p. 67, pl. 5,
fig. 140
1960 Angulogerina angulosa (Williamson); Barker,
pl. 74, figs. 15, 16
1994 Trifarina angulosa (Williamson); Jones, p. 86,
pl. 74, figs. 15, 16
2013 Trifarina angulosa (Williamson); Holbourn et al.,
p. 559, figs. 1–4
Description
The test is triserial in the early portion, becoming unise-
rial later. Test cross section is triangular with acute periph-
ery. The chambers are moderately inflated and increase in
size towards the apertural end. The sutures are curved and
depressed. Wall ornamented by discontinuous longitudinal
costae. The aperture is terminal, with a rounded opening atop
a short neck, with lip and internal tooth plate.
Dimensions
The species length ranges from 150 to 775 µm in the current
study.
Stratigraphic range
This species shows a staggered appearance; first appearing
sporadically at Site 752 in the Chattian then in the Aquitanian
at 1168, followed by the mid-Langhian at 1139. The last ap-
pearance is in the Tortonian at 1139 and in the Zanclean at
1168 and 752.
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208 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Remarks
Specimens from 1168 usually have the typical triangular
cross section with acute periphery but those from 1139 and
particularly 752 often appear nearly rounded in cross sec-
tion and lack the acute margin. This species is typically
recorded from the mid-Miocene to Holocene (Jones, 1994).
Thus, this study extends the species range back into the latest
Oligocene. The species is found worldwide (Holbourn et al.,
2013).
Trifarina bradyi Cushman, 1923
Figure 5: 3a–c
1923 Trifarina bradyi Cushman: p. 99, pl. 22, figs. 3, 9
1884 Rhabdogonium tricarinatum d’Orbigny; Brady,
p. 525, pl. 67, figs. 1, 3
1960 Trifarina bradyi Cushman; Barker, pl. 67, figs. 1,
3
1988 Trifarina bradyi Cushman; Loeblich and Tappan,
p. 526, pl. 574, figs. 10–13
2013 Trifarina bradyi Cushman; Holbourn et al., p. 560,
figs. 1, 2
Description
Test initially triserial becoming rectilinear and uniserial later.
Test tapered at both ends and relatively smooth, later cham-
bers. Distinctive triangular cross section with acute periph-
eral keel. Chambers are moderately inflated and increase
in size towards the aperture, separated by thick, strongly
curved, depressed sutures. The aperture is terminal with short
neck, lip and tooth plate.
Dimensions
The species length ranges from 125 to 575 µm in this study.
Stratigraphic range
At 1139 and 1168 the species is rare in the Chattian, but is
more consistently present from the Aquitanian disappearing
in the mid-Serravallian at 1139 and early Zanclean at 1169.
The species at 752 is restricted to the Burdigalian through to
Serravallian (mid-Miocene).
Remarks
Inter-site differences are observed within this species in the
Indian Ocean. Specimens from 752 and 1139 are quite sim-
ilar, with the lower half of the test having longitudinal stria-
tions, while at 1168 there are denticles or small nodes (see
Fig. 5: 3a and b). Distribution is worldwide according to
Holbourn et al. (2013) and it has previously been recorded
from the eastern Indian Ocean by Nomura (1991). As with
T. angulosa, a much earlier late Oligocene occurrence for this
species is found in the eastern Indian Ocean than previously
reported (Jones, 1994).
Subfamily Uvigerininae Haeckel, 1894
Genus Siphouvigerina Parr, 1950
Siphouvigerina hispida (Schwager, 1866)
Figure 5: 4a–c
1866 Uvigerina hispida Schwager: p. 249, pl. 7, fig. 95
1938 Uvigerina rustica Cushman and Edwards: p. 83,
pl. 14, fig. 6
1980 Euuvigerina hispida (Schwager); Srinivasan and
Sharma, pl. 7, figs. 14, 15
2002 Neouvigerina hispida (Schwager); Hayward,
p. 301, pl. 1, fig. 14
2013 Siphouvigerina hispida (Schwager); Patarroyo and
Martínez, p. 41
2013 Uvigerina hispida Schwager; Holbourn et al.
p. 592, fig. 1
Description
Triserial, elongate, nearly fusiform-shaped test with a basal
spine. Nearly circular in cross section. Chambers are in-
flated to sub-globular, gradually increasing in size towards
the aperture and separated by depressed sutures. The sur-
face is densely ornamented by acicular to coarse papillae or
spines. The aperture is terminal and rounded, with a short
neck, a phialine lip and internal tooth plate.
Dimensions
The species length ranges from 175 to 875 µm in the current
study.
Stratigraphic range
The species distribution at 752 is from early Serravallian to
late Tortonian. At 1139 the species distribution is from Chat-
tian to early Tortonian. At 1168, this species is very rare in
the Chattian but it is consistently present from the Aquitanian
to early Zanclean.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 209
Remarks
This species is generally larger at Site 1139 than at the other
study sites. This species is cosmopolitan (Hayward, 2002;
Holbourn et al., 2013) and known from the early Miocene
(Aquitanian) to recent. However, it occurs earlier (latest
Oligocene) here in the eastern sector of the Indian Ocean.
Genus Uvigerina d’Orbigny, 1826
Uvigerina auberiana d’Orbigny, 1839a
Figure 5: 5a–d
1839 Uvigerina auberiana d’Orbigny: p. 106, pl. 2,
figs. 23, 24
1884 Uvigerina asperula Czjzek; Brady, p. 578, pl. 75,
figs. 6, 8
1884 Uvigerina asperula var. auberiana Brady, p. 579,
pl. 75, fig. 9
1978 Uvigerina auberiana (d’Orbigny): Wright, p. 717,
pl. 8, fig. 10
1984 Uvigerina auberiana (d’Orbigny): Lamb and
Miller, p. 26, pl. 4, figs. 1–6
2011 Uvigerina auberiana (d’Orbigny): Kuppusamy,
p. 69, pl. 13, fig. 9
2013 Uvigerina auberiana (d’Orbigny): Holbourn et al.,
p. 584, figs. 1–3
Description
Test elongate, triserial, fusiform shape. Circular in cross sec-
tion. Depressed sutures and inflated chambers that gradually
increase in size. Small spines usually cover the whole test.
Aperture is terminal and rounded, with tubular neck, phia-
line lip and internal tooth plate.
Dimensions
The species length ranges from 200 to 525 µm in the current
study.
Stratigraphic range
The species is only recorded at 752, from Aquitanian to early
Zanclean.
Remarks
Some specimens are very densely covered in short spines,
ranging all over the test and extending to cover the neck
as well. The specimens in this study are relatively elon-
gate (∼2.5–3 times long as broad) similar to the type ma-
terial rather than the 2 :1 ratio described by Holbourn et
al. (2013). Cosmopolitan distribution according to Holbourn
et al. (2013).
Uvigerina peregrina Cushman, 1923
Figure 5: 6a–b
1923 Uvigerina peregrina Cushman: p. 166, pl. 42,
figs. 7–10
1926 Uvigerina gardnerae Cushman and Applin:
p. 175, pl. 8, figs. 16, 17
1984 Uvigerina peregrina Cushman; Lamb and Miller,
p. 36-37, pl. 8, figs. 1–3; pl. 9, figs. 1–5
1991 Uvigerina peregrina Cushman; Nomura, p. 55,
pl. 1, fig. 10
2013 Uvigerina peregrina Cushman; Hayward et al.,
p. 442, pl. 6, fig. 20
Description
Test elongate, triserial. Chambers inflated, separated by de-
pressed sutures. The test is ornamented by parallel, longitu-
dinal costae. Aperture terminal, with phialine lip and short
narrow neck.
Dimensions
The species length ranges from 175 to 700 µm in the current
study.
Stratigraphic range
The range at 752 and 1139 is Chattian to Tortonian, while at
1168 it ranges from Chattian to early Zanclean.
Remarks
The surface costae are usually non-continuous and paral-
lel. The lower portion of the test is covered in isolated
spines, which are randomly oriented; there are also in spec-
imens shown in Lamb and Miller (1984) (pl. 34–37, 8, 9).
This species is cosmopolitan (Lamb and Miller, 1984) with
records including the Indian Ocean (Nomura, 1991) and New
Zealand (Hayward et al., 2013).
Order Lagenida Lankester, 1885
Superfamily Nodosarioidea Ehrenberg, 1838
Family Vaginulinidae Reuss, 1862
Subfamily Lenticulininae Chapman, Parr, and Collins,
1934
Genus Lenticulina Lamarck, 1804
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210 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Lenticulina cultrata (de Montfort, 1808)
Figure 5: 7a–b
1808 Robulus cultratus de Montfort: p. 214
1839 Robulina canariensis d’Orbigny: p. 127, pl. 3,
figs. 3, 4
1846 Robulina similis d’Orbigny: pl. 4, figs. 14, 15
2013 Lenticulina cultrata de Montfort; Hewaidy et al.,
p. 132, pl. 1, figs. 3a, b
2016 Lenticulina cultrata de Montfort; Amakrane et al.,
p. 103, pl. 6, fig. 27
Description
Test planispiral, involute, with peripheral keel. Lenticular-
shaped smooth test, circular in outline, biconvex in cross sec-
tion. Chambers are slightly inflated. Sutures are curved and
flush to the test. The primary aperture is radiate and terminal.
Dimensions
The species length ranges from 150 to 1000 µm in the current
study.
Stratigraphic range
The species distribution at 752 is Chattian to late Messinian.
The species is only present in the early-to-mid Burdigalian at
1139 and it is not recorded at all at 1168.
Remarks
This species is well-preserved and abundant at 752. It is
known from the Indian Ocean (current study) and was noted
from the west Pacific (Hanagata and Nobuhara, 2015) so
could be considered to have a limited distribution.
Subfamily Marginulininae Wedekind, 1937
Genus Amphicoryna Schlumberger, in Milne-Edwards,
1881
Amphicoryna scalaris (Batsch, 1791)
Figure 6: 1a–b
1791 Nautilus (Orthoceras) scalaris Batsch: pp. 1, 4,
pl. 2, fig. 4a, b
1826 Nodosaria longicauda d’Orbigny: p. 254
1860 Marginulina falx Jones and Parker: p. 302
1884 Nodosaria scalaris (Batsch); Brady, p. 510, pl. 63,
figs. 28–31
1960 Amphicoryna scalaris (Batsch); Barker, p. 134,
pl. 63, figs. 28–31
2013 Amphicoryna scalaris (Batsch); Holbourn et al.,
p. 42, fig. 1
Description
Test elongate, rectilinear, with pronounced basal spine.
Chambers are inflated, gradually increasing in size and sepa-
rated by straight, depressed sutures. The wall is ornamented
by long striations or fine ribs many of which are parallel and
continuous in appearance. Aperture is terminal and radiate
with ring-like ridges around the neck.
Dimensions
The species length ranges from 200 to 750 µm in this study.
Stratigraphic range
At 752 the species ranges from Chattian to Messinian. At
1139 and 1168 this species is very rare, occurring in the mid-
Burdigalian at 1139, extending to the early Zanclean at 1168.
Remarks
This species is very well-preserved and recorded in highest
abundance at 752. This species occurs worldwide according
to Holbourn et al. (2013) and is commonly found in the In-
dian Ocean.
Order Rotaliida Lankester, 1885
Superfamily Asterigerinoidea d’Orbigny, 1839a
Family Epistomariidae Hofker, 1954
Subfamily Nuttallidinae Saidova, 1981
Genus Nuttallides Finlay, 1939
Nuttallides umbonifera (Cushman, 1933a)
Figure 6: 2a–e
1933a Pulvinulinella umbonifera Cushman: p. 90, pl. 9,
fig. 9
1884 Truncatulina pygmaea Hantken; Brady, p. 666,
pl. 95, figs. 9, 10
1933a Pulvinulinella umbonifera Cushman: p. 90, pl. 9,
fig. 9
1960 Eponides bradyi Earland; Barker, pl. 95, figs. 9,
10
1987 Nuttallides umbonifera (Cushman); Miller and
Katz, p. 136, pl. 5, fig. 5
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 211
Figure 6. Scale bar is 100 µm. (1a–b) Amphicoryna scalaris, ODP Hole 752A, (1a) 10H-5, 77–81 cm: side view. (1b) 8H-1, 100–105 cm:
apertural view. (2a–e) Nuttallides umbonifera,(2a, e) ODP Hole 752A, 4H-5, 64–69 cm: (2a) umbilical view; (2e) edge view. (2b) ODP
Hole 752A, 9H-5, 65–70 cm: spiral view. (2c) ODP Hole 1168A, 13X-1, 40–44cm: spiral view. (2d) ODP Hole 1168A, 14X-4, 45–49cm:
umbilical view. (3a–c) Gyroidinoides soldanii, ODP Hole 1139A, 3R-4, 32–36 cm: (3a) spiral view; (3b) umbilical view; (3c) edge view.
(4a–c) Gyroidinoides sp. 1, ODP Hole 1139A, (4a, c) 19R-2, 60.5–64.5 cm: (4a) spiral view; (4c) edge view. (4b) 17R-3, 91–95 cm: um-
bilical view. (5a–b) Gyroidinoides sp. 2, (5a) ODP Hole 1168A, 32X-4, 80–84 cm: umbilical view. (5b) ODP Hole 1139A, 15R-4, 41–
45 cm: spiral view. (6a–d) Oridorsalis umbonatus,(6a–b) ODP Hole 1168A, 8H-1, 100–104 cm: (6a) umbilical view; (6b) spiral view.
(6c) ODP Hole 752A, 5H-1, 65–70 cm: spiral view. (6d) ODP Hole 1168A, 8H-1, 100–104cm: edge view. (7a–c) Osangularia culter, ODP
Hole 1168A, 28X-3, 75–79 cm: (7a) umbilical view; (7b) spiral view; (7c) edge view.
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212 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
1994 Osangulariella umbonifera (Cushman); Jones,
p. 99, pl. 95, figs. 9, 10
1995 Nuttallides umboniferus (Cushman); Nomura,
p. 276, pl. 3, fig. 2
Description
Lenticular, trochospiral form. Unequally biconvex test with
a sub-acute peripheral keel. About 12 crescentic chambers in
the final whorl, separated by slightly depressed, arcuate to
sinuous sutures on the umbilical side and oblique sutures on
the spiral side. Aperture is an interiomarginal slit extending
from the umbilical boss towards periphery.
Dimensions
The species length ranges from 100 to 750 µm in the current
study.
Stratigraphic range
The range of the species is Chattian to Tortonian at 1139 and
752. At 1168 it ranges from Chattian to early Zanclean.
Remarks
This species is characterised by an imperforate umbilical
boss. Nuttallides umbonifera (Cushman) is often employed
as an indicator of Antarctic Bottom Water (AABW) in the
Pacific Ocean and Indian Ocean (Corliss, 1979). This species
is long-ranging and distributed worldwide. Mackensen et
al. (1990) believed that the species is associated with car-
bonate corrosive bottom water masses in the Weddell Sea.
Distribution worldwide according to Holbourn et al. (2013)
and found throughout the Indian Ocean and Southern Ocean
(Mackensen, 1992; Nomura, 1995).
Superfamily Chilostomelloidea Brady, 1881
Family Gavelinellidae Hofker, 1956
Subfamily Gavelinellinae Hofker, 1956
Genus Gyroidinoides Brotzen, 1942
Gyroidinoides soldanii (d’Orbigny, 1826)
Figure 6: 3a–c
1826 Gyroidina soldanii d’Orbigny: p. 278
1846 Gyroidina soldanii d’Orbigny; d’Orbigny, pl. 8,
figs. 10–12
1978 Gyroidina soldanii d’Orbigny; Wright, p. 715,
pl. 5, figs. 7–9
1992 Gyroidinoides soldanii d’Orbigny; Mackensen,
p. 668, pl. 5, figs. 9, 10
1994 Gyroidinoides soldanii d’Orbigny; Jones, p. 106,
pl. 107, figs. 6, 7
Description
Test is trochospiral and planoconvex. Umbilical side is in-
volute and convex, and the spiral side is flat and evolute.
Rounded periphery. Chambers are inflated and separated by
slightly depressed sutures that are straight on umbilical side
but curved on spiral side. The surface is smooth. The aperture
is a narrow interiomarginal slit, extending from umbilicus to
the periphery. The umbilicus is open.
Dimensions
The species length ranges from 100 to 750 µm in this study.
Stratigraphic range
This species occurs in the Chattian at all three sites, ranging
to the early Zanclean at 752 and 1168, and the early Torto-
nian at 1139.
Remarks
Most of the specimens observed in this study have a planar
or slightly convex spiral side, and relatively small chambers
in the last whorl on the spiral side. Distribution worldwide
according to Holbourn et al. (2013). This species is now
considered a junior synonym of Hansenisca soldanii (Hay-
ward et al., 2019b) in the World Register of Marine Species
(WoRMS).
Gyroidinoides sp. 1
Figure 6: 4a–c
Description
Test is trochospiral, plano-convex. The spiral side is flat and
evolute, while the umbilical side is convex and involute.
Chambers are generally inflated separated by slightly de-
pressed sutures on both umbilical and spiral sides. The aper-
ture is an interiomarginal slit extending from umbilicus to
periphery. The surface is smooth.
Dimensions
The species length ranges from 100 to 550 µm in this study.
Stratigraphic range
The species is rare at 752 only occurring in the mid-
Serravallian and early Zanclean. At 1139 the species ranges
from Chattian to early Tortonian. At 1168 it is very rare in the
Chattian, with a more common range from early Burdigalian
to early Zanclean.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 213
Remarks
The chambers are more elongate and larger on the spiral side
than in Gyroidinoides soldanii and it lacks the open, narrow
umbilicus of G. soldanii and G. sp. 2. The distribution of
the species may be restricted to the current study (southern
Indian Ocean). The stratigraphic range overlaps with that of
G. soldanii.
Gyroidinoides sp. 2
Figure 6: 5a–b
Description
The test is trochospiral and planoconvex. Sutures flush to
slightly depressed on spiral side, while slightly depressed on
umbilical side. Aperture is slit-like extending from umbilicus
to periphery. The aperture has a long crescentic flap covering
the umbilicus.
Dimensions
The species length ranges from 100 to 400 µm in the current
study.
Stratigraphic range
Species ranges from Chattian to mid-Serravallian at 1139
and Chattian to early Zanclean at 1168. This species is not
recorded at 752.
Remarks
This species differs from Gyroidinoides soldanii by having
larger chambers on the spiral side, which is also more con-
vex. Gyroidinoides sp. 1 has slightly more inflated chambers
on the spiral side and a shorter, more curved apertural slit
than this species. This species may be restricted to the cur-
rent study area (southern Indian Ocean).
Family Oridorsalidae Loeblich and Tappan, 1984
Genus Oridorsalis Andersen, 1961
Oridorsalis umbonatus (Reuss, 1851)
Figure 6: 6a–d
1851 Rotalina umbonata Reuss: p. 75, pl. 5, fig. 35
1884 Pulvinulina umbonata (Reuss); Brady, p. 695,
pl. 105, fig. 2
1884 Truncatulina tenera Brady: p. 665, pl. 95, fig. 11
1932 Eponides umbonata (Reuss); Nuttall, p. 26, pl. 6,
figs. 4, 5
1983 Oridorsalis umbonatus (Reuss); Tjalsma and
Lohmann, p. 18, pl. 6, fig. 8
1988 Oridorsalis umbonata (Reuss); Whittaker, p. 137,
pl. 19, figs. 1–3
2011 Oridorsalis umbonatus (Reuss); Kuppusamy et
al., p. 59, pl. 8, figs. 23, 24
Description
Test trochospiral, lenticular, unequally biconvex, and nearly
circular in outline. The convexity of spiral side is greater than
the umbilical side. Involute in umbilical and evolute in spiral
side. Six nearly inflated chambers in the last whorl. Sutures
usually slightly depressed, curved on umbilical side and ra-
dial on spiral side. The test is smooth. The aperture is an
interiomarginal slit, extending from periphery to umbilicus.
Dimensions
The species length ranges from 100 to 750 µm in this study.
Stratigraphic range
The stratigraphic distribution of the species is Chattian to
early Zanclean at 752 and 1168, Chattian to early Tortonian
at 1139.
Remarks
This species generally has six chambers, some specimens
possess seven chambers, in the last whorl. This species is
easily identifiable and long-ranging from the Paleocene to
Holocene, and is common worldwide Holbourn et al. (2013).
Genus Osangularia Brotzen, 1940
Osangularia culter (Parker & Jones, 1865)
Figure 6: 7a–c
1865 Planorbulina farcta var. ungeriana subvar. culter
Parker and Jones: pp. 382, 421, pl. 19, fig. 1
1866 Anomalina bengalensis Schwager, p. 259, pl. 7,
fig. 111
1884 Truncatulina culter (Parker and Jones); Brady,
p. 668, pl. 96, fig. 3
1953 Osangularia culter (Parker and Jones); Phleger et
al., p. 42, pl. 9, figs. 11–16
1960 Osangularia bengalensis (Schwager); Barker,
pl. 96, fig. 3
1994 Osangularia bengalensis (Schwager); Jones,
p. 100, pl. 96, fig. 3
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214 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Description
Test is trochspiral, lenticular and unequally biconvex; the
umbilical side is more convex than spiral side. The umbil-
ical side is involute, while the spiral side is evolute. Acute
periphery with a serrated keel. The slightly inflated nearly 10
chambers, separated by slightly elevated, curved sutures on
the spiral side, and straight with slightly depressed sutures
on umbilical side. The test is smooth with some perforation
in the chambers on spiral side. Aperture is slit-like interi-
omarginal, extending from umbilicus to periphery.
Dimensions
The species length ranges from 125 to 1000 µm in the current
study.
Stratigraphic range
The range of the species is Chattian to early Zanclean at 752,
Chattian to early Tortonian at 1139. At 1168 it ranges from
Aquitanian to Langhian, and then mid-Tortonian to early
Zanclean.
Remarks
Osangularia culter is one of several species dominating in-
termediate water assemblages in the Miocene (Woodruff,
1985). Distribution cosmopolitan (Holbourn et al., 2013).
Superfamily Discorboidea Ehrenberg, 1838
Family Rosalinidae Reiss, 1963
Genus Gavelinopsis Hofker, 1951
Gavelinopsis lobatulus (Parr, 1950)
Figure 7: 1
1913 Discorbina praegeri (Heron-Allen and Earland):
p. 122, pl. 10, figs. 8–10
1950 Discorbis lobatulus Parr: p. 354, pl. 13, figs. 23–
25
1960 Gavelinopsis lobatulus (Parr); Barker, p. 182,
pl. 88, figs. 1a–c
2011 Gavelinopsis lobatulus (Parr); Kuppusamy et al.,
p. 52, pl. 5, figs. 2, 3
Description
Test trochoid. The periphery is thickened and keeled. Five
slightly inflated chambers in the last whorl. Nearly flush su-
tures on spiral side, with slightly depressed sutures on um-
bilical side. The aperture is an interiomarginal slit extending
from umbilicus to periphery, having a narrow lip.
Dimensions
The species length ranges from 125 to 375 µm in the current
study.
Stratigraphic range
The stratigraphic distribution is early Langhian to early Zan-
clean at 752 and mid-Burdigalian to Mid-Serravallian at
1139. This species is not recorded at 1168.
Remarks
Most of the specimens of this species are characterised by
an umbilical plug and sharp periphery. Recorded from the
Indian Ocean and Southern ocean including the Kerguelen
Plateau (Mackensen, 1992) and eastern Indian Ocean (No-
mura, 1991). This species is considered a subjective junior
synonymy of Gavelinopsis praegeri (Hayward et al., (2019c)
in WoRMS).
Family Sphaeroidinidae Cushman, 1927
Genus Sphaeroidina d’Orbigny, 1826
Sphaeroidina bulloides d’Orbigny, 1828 in Deshayes,
1828
Figure 7: 2a–c
1828 Sphaeroidina bulloides d’Orbigny in Deshayes:
p. 267
1846 Sphaeroidina austriaca d’Orbigny: p. 284, pl. 20,
figs. 19–21
1848 Sexloculina haueri Czjzek: p. 149, pl. 13, figs. 35–
38
1884 Sphaeroidina bulloides d’Orbigny; Brady, p. 620,
pl. 84, figs. 1, 2
1949 Sphaeroidina ciperana Cushman and Todd: p. 67,
pl. 12, fig. 14
1949 Sphaeroidina compacta Cushman and Todd: p. 19,
pl. 4, fig. 14
1949 Sphaeroidina compressa Cushman and Todd:
p. 20, pl. 4, figs. 15, 16
1949 Sphaeroidina nitida Cushman and Todd: p. 20,
pl. 4, fig. 13
1953 Sphaeroidina japonica Asano: p. 19, pl. 2,
figs. 43–44
1985 Sphaeroidina bulloides d’Orbigny; Papp and
Schmid, p. 96, pl. 90, figs. 7–12
2013 Sphaeroidina bulloides d’Orbigny; Holbourn et
al., p. 520, figs. 1–3
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 215
Figure 7. Scale bar is 100 µm. (1) Gavelinopsis lobatulus, ODP Hole 752A, 3H-5, 62–67cm: umbilical view. (2a–c) Sphaeroidina bulloides,
(2a) ODP Hole 752A, 8H-1, 100–105 cm: side view. (2b) ODP Hole 1139A, 8R-1, 45–49cm: apertural side view. (2c) ODP Hole 1168A,
26X-2, 73–77 cm: apertural side view. (3a–b) Laticarinina pauperata,(3a) ODP Hole 752A, 7H-2, 73–78cm: umbilical view. (3b) ODP
Hole 1139A, 18R-1, 96–100 cm: umbilical view. (4a–b) Epistominella exigua, ODP Hole 1168A, 36X-6, 62.5–66.5cm: (4a) spiral view;
(4b) umbilical view. (5a–b) Astrononion umbilicatulum, ODP Hole 1139A, (5a) 19R-2, 60.5–64.5 cm: umbilical view. (5b) 17R-3, 91–
95 cm: edge view. (6a–b) Melonis barleeanus, ODP Hole 1139A, (6a) 4R-2, 105–109 cm: umbilical? view. (6b) 19R-2, 60.5–64.5 cm: edge
view. (7a–b) Pullenia bulloides,(7a) ODP Hole 752A, 9H-5, 65–70 cm: edge view. (7b) ODP Hole 1139A, 16R-2, 55–59 cm: edge view.
(8a–b) Pullenia quinqueloba, ODP Hole 1168A, 35X-1, 65–69 cm: (8a) edge view; (8b) umbilical view.
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216 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Description
Test is sub-globular, usually three chambers in the final
whorl. The test is variably coiled. Chambers are globular to
hemispherical, separated by slightly depressed sutures. The
wall is calcareous, imperforate and smooth. The aperture is
a crescentic opening, bordered by a lip, occurring where the
three last chambers meet.
Dimensions
The species length ranges from 125 to 725 µm in the current
study.
Stratigraphic range
The stratigraphic distribution is Chattian to Zanclean at 752
and 1168, and Chattian to Serravallian at 1139.
Remarks
The larger specimens in the current study have a thick and
opaque wall; smaller specimens are thinner-walled and ap-
pear translucent. Worldwide distribution according to Hol-
bourn et al. (2013) including previous reports from the east-
ern Indian Ocean (Nomura, 1991).
Superfamily Discorbinelloidea Sigal in Piveteau, 1952
Family Discorbinellidae Sigal in Piveteau, 1952
Subfamily Discorbinellinae Sigal in Piveteau, 1952
Genus Laticarinina Galloway and Wissler, 1927
Laticarinina pauperata (Parker & Jones, 1865)
Figure 7: 3a–b
1865 Pulvinulina repanda Fichtell and Moll
var. menardii d’Orbigny, subvar. pauperata Parker
and Jones: p. 395, pl. 16, figs. 50–51
1884 Pulvinulina pauperata (Parker and Jones); Brady,
p. 696, pl. 104, figs. 3–11
1942 Laticarinina bulbrooki Cushman and Todd: p. 19,
pl. 4, figs. 8, 9
1942 Laticarinina crassicarinata Cushman and Todd:
p. 18, pl. 4, figs. 11, 12
1949 Laticarinina pauperata (Parker and Jones);
Bermúdez, p. 309, pl. 23, figs. 43–45
1953 Laticarinina pauperata (Parker and Jones);
Phleger et al., p. 49, pl. 11, figs. 5, 6
1991 Laticarinina pauperata (Parker and Jones); No-
mura, p. 55, pl. 1, fig. 6
1992 Laticarinina pauperata (Parker and Jones); Mack-
ensen, p. 668, pl. 2, figs. 1, 2
2011 Laticarinina pauperata (Parker and Jones); Kup-
pusamy et al., p. 57, pl. 7, fig. 8
Description
Test planoconvex, slightly trochoid. The spiral side is flat;
the umbilical side is slightly convex. Chambers are inflated
but becoming progressively more loosely coiled with modest
rate of enlargement. Chambers are cuneiform on the umbil-
ical side and reniform on the spiral side, and they are sepa-
rated by radial and slightly depressed sutures. Chambers on
the ventral side are radial, while in the dorsal side is more
separated. Large, transparent peripheral keel around whole
test, projecting in front of final chamber. Aperture is on the
inner margin of last formed chamber.
Dimensions
The species length ranges from 175 to 1300 µm in the current
study.
Stratigraphic range
The range of the species at 752 is Chattian to early Burdi-
galian and then again late Langhian to early Zanclean. At
1139 the species appears in the Chattian and later from the
mid-Aquitanian to mid-Serravallian. At 1168, it is recorded
from the mid-Aquitanian to mid-Messinian.
Remarks
Most specimens are easily distinguished by the presence of a
very broad, transparent keel on the periphery.
Biogeography
This species has previously been reported from the eastern
Indian Ocean (Nomura, 1991) and Kerguelen Plateau (Mack-
ensen, 1992) but can be found worldwide (Holbourn et al.,
2013).
Family Pseudoparrellidae Voloshinova, 1952
Subfamily Pseudoparrellinae Voloshinova, 1952
Genus Epistominella Husezima and Maruhasi, 1944
Epistominella exigua (Brady, 1884)
Figure 7: 4a–b
1884 Pulvinulina exigua Brady: p. 696, pl. 103,
figs. 13–14
1951 Pseudoparrella exigua (Brady); Phleger and
Parker, p. 28, pl. 15, fig. 6
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 217
1994 Pseudoparrella exigua (Brady); Loeblich and Tap-
pan, p. 146, pl. 307, figs. 1–7
1951 Pulvinulinella exigua (Brady); Hofker, p. 322,
figs. 219–221
1960 Epistominella exigua (Brady); Barker, pl. 103,
figs. 13, 14
1992 Epistominella exigua (Brady); Mackensen, p. 668,
pl. 4, figs. 5, 6
1994 Alabaminoides exiguus (Brady); Jones, p. 103,
pl. 103, figs. 13, 14
2011 Epistominella exigua (Brady); Kuppusamy et al.,
p. 50, pl. 4, figs. 11, 12
2013 Epistominella exigua (Brady); Holbourn et al.,
p. 240, figs. 1–6
Description
Test trochoid, unequally biconvex. Slightly convex spiral
side, convex umbilical side. The periphery is acute. Five to
six moderately inflated chambers in the last whorl, separated
by curved, flush sutures on spiral side and curved, depressed
sutures on umbilical side. Aperture is an interiomarginal slit
on umbilical side extending to the face of the final and largest
chamber.
Dimensions
The species length ranges from 100 to 275 µm in the current
study.
Stratigraphic range
The species is rare at 752 ranging from early Langhian to
early Zanclean. The range is Chattian to early Tortonian at
1139 and to early Zanclean at 1168.
Remarks
The abundance of this species is employed as a proxy for
constraining the flux of pulsed organic matter (Holbourn et
al., 2013). This species is found worldwide including lo-
cal to this study in the Timor Sea (Loeblich and Tappan,
1994), Kerguelen Plateau (Mackensen, 1992) and Weddell
Sea, Antarctic (Thomas, 1990).
Superfamily Nonionoidea Schultze, 1854
Family Nonionidae Schultze, 1854
Subfamily Astrononioninae Saidova, 1981
Genus Astrononion Cushman & Edwards, 1937
Astrononion umbilicatulum Uchio, 1952
Figure 7: 5a–b
1952 Astrononion umbilicatulum Uchio: p. 36, pl. 32,
fig. 1
1978 Astrononion umbilicatulum Uchio; Wright, p. 710,
pl. 1, fig. 12
2011 Astrononion umbilicatulum Uchio; Kuppusamy et
al., p. 44, pl. 1, figs. 7–9
Description
Planispiral involute test. Periphery rounded, elliptical in lat-
eral view. Chambers large and increasing gradually in size.
Approximately seven to eight chambers in the last whorl,
separated by curved, thickened radial sutures. Aperture is a
narrow interiomarginal, crescentic slit, extending across the
periphery to the umbilicus.
Dimensions
The species length ranges from 100 to 350 µm in the current
study.
Stratigraphic range
The distribution of the species at 752 is early Aquitanian to
early Zanclean and Chattian to early Zanclean at 1168. The
range is Chattian to early Tortonian at 1139.
Remarks
The flaps that partially cover the sutures in the last whorl
leave a small cavity beneath, which look like secondary
chamberlets. This species was originally described from the
lower Pleistocene of Japan (Uchio, 1952) but has not been
widely reported from elsewhere. It is known from the In-
dian Ocean (current study), ODP Site 757 (Singh and Gupta,
2004), north-west Atlantic Ocean (Kuppusamy et al., 2011)
and the Mediterranean (Wright, 1978).
Subfamily Pulleniinae Schwager, 1877
Genus Melonis de Montfort, 1808
Melonis barleeanus (Williamson, 1858)
Figure 7: 6a–b
1858 Nonionina barleeana Williamson: p. 32, pl. 3,
figs. 68, 69
1930 Nonion barleeanum (Williamson); Cushman,
p. 11, pl. 4, fig. 5
1953 Nonion barleeanum (Williamson); Phleger et al.,
p. 30, pl. 6, fig. 4
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218 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
1979 Melonis barleeanum (Williamson); Corliss, p. 10,
pl. 5, fig. 78
1988 Melonis barleeanum (Williamson); Loeblich and
Tappan, pl. 696, figs. 5, 6
1992 Melonis barleeanus (Williamson); Mackensen,
p. 668, pl. 5, fig. 2
1997 Melonis barleeanus Williamson, 1858; Bergamin
et al., p. 38, pl. 2, figs. 1–9
Description
Test planispiral, bi-umbilicate, involute, compressed test.
The umbilicus is open. The test is elliptical, elongated in lat-
eral view, rounded periphery. Ten to 11 flattened chambers,
separated by broad, flush sutures. The test is smooth and per-
forated. The aperture is an interiomarginal slit extending to
the umbilicus, possessing an apertural lip.
Dimensions
The species length ranges from 150 to 600 µm in this study.
Stratigraphic range
The stratigraphic distribution of the species at 752 is split
into two phases, early Langhian to mid-Serravallian and then
again late Messinian to Zanclean. At 1139 it ranges from
Chattian to mid-Tortonian, while at 1168 it ranges from Chat-
tian to Zanclean.
Remarks
Forms identified here have a prominent apertural lip distin-
guishing them from morphologically similar Melonis affinis
separated out by some authors (see details in Holbourn et
al., 2013). However, we consider the morphologically simi-
lar Melonis affinis synonymous with M. barleeanus follow-
ing molecular analyses by Schweizer (2006).
Biogeography
The species is found worldwide, but Holbourn et al. (2013)
suggest that it is not present at very high latitudes. The high-
est southern-recorded occurrence is on the Kerguelen Plateau
by Mackensen (1992), which is supported by the current
study observations.
Genus Pullenia Parker & Jones in Carpenter et al., 1862
Pullenia bulloides (d’Orbigny, 1846)
Figure 7: 7a–b
1826 Nonionina bulloides d’Orbigny: p. 293
1884 Pullenia sphaeroides (d’Orbigny); Brady, p. 615,
pl. 84, figs. 12, 13
1945 Pullenia bulloides (d’Orbigny); Cushman and
Todd, p. 64, pl. 11, fig. 5
1949 Pullenia bulloides (d’Orbigny); Bermúdez, p. 276,
pl. 21, figs. 28–29
1978 Pullenia bulloides (d’Orbigny); Wright, p. 716,
pl. 7, fig. 7
2011 Pullenia bulloides (d’Orbigny); Kuppusamy et al.,
p. 62, pl. 9, figs. 18, 19
Description
Test planispiral, involute, globular, sub-circular in outline.
Four to five inflated chambers in the last whorl, separated by
flush to slightly depressed sutures. Very compact test. Aper-
ture is a narrow interiomarginal, crescentic slit, extending
from the periphery to umbilicus.
Dimensions
The species length ranges from 125 to 425 µm in this study.
Stratigraphic range
The stratigraphic distribution of the species at 752 and 1168
is Chattian to Zanclean. At 1139 the species ranges from
Chattian to mid-Tortonian.
Remarks
This species usually has a broad periphery and a smooth sur-
face. Up to five chambers are present in the final whorl com-
pared to four or five in the holotype and topotype specimens.
Distribution is worldwide (Holbourn et al., 2013) including
previous reports from the Indian Ocean by Nomura (1991)
and (Singh and Gupta, 2004).
Pullenia quinqueloba (Reuss, 1851)
Figure 7: 8a–b
1851 Nonionina quinqueloba Reuss: p. 71, pl. 5,
fig. 31a, b
1960 Pullenia subcarinata (d’Orbigny); Barker, pl. 84,
figs. 14, 15
1949 Pullenia quinqueloba (Reuss); Bermúdez, p. 276,
pl. 21, figs. 32, 33
1983 Pullenia quinqueloba (Reuss); Tjalsma and
Lohmann, p. 36, pl. 16, fig. 2
1987 Pullenia quinqueloba (Reuss); Miller and Katz,
p. 138, pl. 4, figs. 2a–b
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 219
Description
Test planispiral, involute. Sub-globular, slightly compressed
test, rounded periphery. Chambers are triangular and inflated.
Five chambers present in the final whorl, separated by de-
pressed and radial sutures. Aperture is a narrow, crescentic,
interiomarginal slit, extending from periphery to umbilicus.
Dimensions
The species length ranges from 125 to 500 µm in the current
study.
Stratigraphic range
The range of the species at 752 is Chattian to Zanclean at
1139 Chattian to mid-Serravallian and at 1168 early Aqui-
tanian to Zanclean.
Remarks
This species is distinguished from five-chambered forms of
P. bulloides by the more open coiling and less circular out-
line. This species is recorded worldwide (Holbourn et al.,
2013) but is common in the continental shelf sediments of
the Yellow Sea and the East China Sea (Lei and Li, 2016)
and in the Indian Ocean (Nomura, 1991; Singh and Gupta,
2004).
Superfamily Planorbulinoidea Schwager, 1877
Family Cibicididae Cushman, 1927
Subfamily Cibicidinae Cushman, 1927
Genus Cibicidoides Saidova, 1975
Cibicidoides bradyi (Trauth, 1918)
Figure 8: 1a–b
1918 Truncatulina bradyi Trauth, p. 235
1884 Truncatulina dutemplei (d’Orbigny); Brady,
p. 665, pl. 95, fig. 5
1976 Cibicidoides bradyi (Trauth); Pflum and Frerichs,
pl. 3, figs. 6, 7
1994 Gyroidina bradyi (Trauth); Jones, p. 99, pl. 95,
fig. 5
2013 Cibicidoides bradyi Trauth); Holbourn et al.,
p. 162, figs. 1–6
Description
Test trochospiral, unequally biconvex. The umbilical side is
involute, and the spiral side is evolute, more convex. Nine in-
flated chambers in the last whorl, separated by radial curved
sutures in umbilical side, and oblique sutures in the spiral
side. The aperture is a slit-like opening with lip extending
from periphery to spiral side.
Dimensions
The species length ranges from 100 to 450 µm in this study.
Stratigraphic range
The species first appears in the Chattian and ranges to the
Zanclean at 752 and 1168; at 1139 it ranges from Chattian to
early Tortonian.
Remarks
Distinguished from Cibicidoides robertsonianus (Brady,
1881), with which it can be confused (Holbourn et al., 2013),
by having fewer chambers (typically <10) in the last whorl,
smaller test and a more rounded periphery. Recorded world-
wide (Holbourn et al., 2013), including from the central In-
dian Ocean by Boersma (1990). Whilst this species is per-
haps most commonly referred to as C. bradyi in the literature
(e.g. Holbourn et al., 2013), accepted alternate names include
Parrelloides bradyi and Heterolepa bradyi (Hayward et al.,
2019d, in WoRMS database).
Cibicidoides mundulus (Brady, Parker & Jones, 1888)
Figure 8: 2a–c
1888 Truncatulina mundula Brady, Parker and Jones:
p. 228, pl. 45, fig. 25a–c
1953 Cibicides kullenbergi Parker in Phleger et al.:
p. 49, pl. 11, figs. 7, 8
1955 Cibicidoides mundulus (Brady, Parker and Jones);
Loeblich and Tappan, p. 25, fig. 4a–c
1960 Cibicidoides mundulus (Brady, Parker and Jones);
Barker, pl. 95, fig. 6
1992 Cibicidoides mundulus (Brady, Parker and Jones);
Mackensen, p. 668, pl. 3, figs. 5–7
Description
Unequally biconvex test, trochospiral. The umbilical side
is more convex than the spiral side. Chambers are low, in-
flated, 10 to 12 chambers in the last whorl, separated by
curved sutures. The primary aperture is a long, narrow inte-
riomarginal slit, extending from periphery to spiral side. The
test is smooth on the umbilical side and highly perforated on
the spiral side.
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220 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Figure 8. Scale bar is 100 µm. (1a–b) Cibicidoides bradyi, ODP Hole 1139A, (1a) 9R-4, 105–110 cm: umbilical view. (1b) 8R-1, 45–
49 cm: spiral view. (2a–c) Cibicidoides mundulus, ODP Hole 752A, 3H-1, 50–55 cm: (2a) umbilical view; (2b) edge view; (2c) spiral
view. (3a–c) Cibicidoides wuellerstorfi, (normal) ODP Hole 752A, 7H-2, 73–78 cm: (3a) umbilical view; (3b) edge view; (3c) spiral view.
(4a–c) Cibicidoides wuellerstorfi, (smooth) ODP Hole 752A, 5H-1, 65–70 cm: (4a) umbilical view; (4b) edge view; (4c) spiral view. (5a–
c) Planulina costata, ODP Hole 752A, 7H-2, 73–78 cm: (5a) spiral view; (5b) edge view; (5c) umbilical view. (6a–b) Eggerella bradyi, ODP
Hole 1168A; (6a) 42X-3, 70–74 cm: side view. (6b) 10H-2, 80–84cm: apertural view.
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 221
Dimensions
The species length ranges from 125 to 800 µm in this study.
Stratigraphic range
The stratigraphic distribution of the species at 752 and 1168
is Chattian to Zanclean. At 1139A it ranges from the Chattian
to early Tortonian.
Remarks
The specimens here all show a pronounced umbilical boss,
as also illustrated by Holbourn et al. (2013, p. 196, pl. 1–8)
and (Setoyama and Kaminski, 2015, p. 14, fig. 5, pl. 4a–c).
Common species found worldwide (Holbourn et al., 2013),
including in the eastern Indian Ocean (Nomura, 1991) and
Kerguelen Plateau (Mackensen, 1992).
Cibicidoides wuellerstorfi (Schwager, 1866)
Figure 8: 3a–c
1866 Anomalina wuellerstorfi Schwager: p. 258, pl. 7,
figs. 105, 107
1884 Truncatulina wuellerstorfi (Schwager); Brady,
pp. 627, 662, pl. 93, figs. 8, 9
1960 Planulina wuellerstorfi (Schwager); Barker,
pl. 93, fig. 9
1976 Cibicides wuellerstorfi (Schwager); Pflum and
Frerichs, p. 116, pl. 4, figs. 2–4
1994 Cibicidoides wuellerstorfi (Schwager); Jones,
p. 98, pl. 93, figs. 8, 9
2015 Cibicidoides wuellerstorfi (Schwager); Setoyama
and Kaminski, p. 14, fig. 5, pl. 10a–c
Description
Planispiral to low trochospiral and plano-convex test. Up to
12 elongated and curved chambers in the last whorl; increas-
ing rapidly in size. Peripheral keel so acute periphery. The su-
tures are depressed and strongly curved. The wall is coarsely
perforated on the spiral side. The aperture is a narrow slit,
having a narrow lip extending from the periphery to the spi-
ral side.
Dimensions
The species length ranges from 125 to 1000 µm in the current
study.
Stratigraphic range
The stratigraphic distribution of the coarse type at 752 is
early Aquitanian to Tortonian and mid-Aquitanian to Zan-
clean at 1168. At 1139 it ranges from mid-Burdigalian to
mid-Serravallian. The smooth type ranges from early Torto-
nian to Zanclean at 752. At 1139 it ranges from Burdigalian
to Serravallian, while at 1168 it ranges from Aquitanian to
Zanclean.
Remarks
This species has two different styles of test ornamentation
as noted by Nomura (1991) in the eastern Indian Ocean
(ODP Hole 754A) – a “smooth” and a “coarse” type. The
smooth type (pl. 6, fig. 4a–c) has a smooth umbilical side
and is relatively inflated in side view compared to the coarse
type, which is highly adorned with nodes on the umbili-
cal side and is more compressed in side view. The smooth
type is recorded at all three localities in the current study.
The coarse type Cibicidoides wuellerstorfi has previously
been assigned to three different genera; Fontobia (Schwa-
ger), Planulina Revets (1996) and Cibicidoides. Following
the molecular phylogenetic studies of Schweizer (2006) and
Schweizer et al. (2009, 2011) the latter genus has now been
accepted and applied in the literature (Setoyama and Kamin-
ski, 2015). Worldwide distribution (Holbourn et al., 2013),
widely reported in the Indian Ocean (e.g. Boersma, 1990;
Nomura, 1991).
Family Planulinidae Bermúdez, 1952
Genus Planulina d’Orbigny, 1826
Planulina costata (Hantken, 1875)
Figure 8: 5a–c
1875 Truncatulina costata Hantken: p. 73, pl. 9, fig. 2a–
c
1932 Planulina wuellerstorfi Schwager; Nuttall, p. 31,
pl. 4, figs. 14, 15
1979 Planulina wuellerstorfi Schwager; Sztràkos, p. 86,
pl. 26, fig. 4
1934 Planulina marialana Hadley: p. 27, pl. 4, figs. 4–7
1941 Planulina palmerae van Bellen: in van Bellen et
al., 1941, p. 1144, figs. 7–9
1986 Planulina costata (Hantken); van Morkhoven et
al., p. 212, pl. 72
1991 Planulina costata (Hantken); Nomura, p. 55, pl. 4,
fig. 5a–c
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222 D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera
Description
Low trochoid test. Spiral side is evolute and flattened, while
the umbilical side is involute and slightly convex. Chambers
number 10 to 12, with a low inflated shape, separated by
slightly to highly curved sutures. The test is smooth on the
umbilical side and highly perforated with small pores on the
spiral side. Aperture is slit-like, peripheral, with a small lip
extending towards the umbilicus.
Dimensions
The species length ranges from 150 to 1150 µm in the current
study.
Stratigraphic range
This species only occurs at 752 where it ranges from early
Aquitanian to early Serravallian.
Remarks
Van Morkhoven et al. (1986) considered that this species may
have evolved from Planulina cocoaensis and that P. costata
and Planulina renzi may have evolved separately through the
Oligocene and early Miocene. The current species is sim-
ilar to the smooth form of C. wuellerstorfi in overall test
shape and by having a smooth umbilical surface, but dif-
fers by having a smaller apertural opening, more compressed
test and more irregular periphery. It is similar to Planulina
karsteni (Petters and Sarmiento, 1956), but the spiral side of
P. karsteni is slightly convex, while in Planulina costata it is
plano-convex. Species is widely distributed (van Morkhoven
et al., 1986) and has previously been recorded from the east-
ern Indian Ocean (Nomura, 1991).
Order Textulariida Lankester, 1885
Superfamily Eggerelloidea Ehrenberg, 1838
Family Eggerellidae Cushman, 1937
Subfamily Eggerellinae Cushman, 1937
Genus Eggerella Cushman, 1933a
Eggerella bradyi (Cushman, 1911)
Figure 8: 6a–b
1911 Verneuilina bradyi Cushman: p. 54, fig. 87
1937 Eggerella bradyi (Cushman); Cushman, p. 52,
pl. 5, fig. 19
1951 Eggerella bradyi (Cushman); Phleger and Parker,
p. 6, pl. 3, figs. 1, 2
1960 Eggerella bradyi (Cushman); Barker, pl. 47,
figs. 4–7
1988 Eggerella bradyi (Cushman); Loeblich and Tap-
pan, p. 170, pl. 189, figs. 1, 2
1991 Eggerella bradyi (Cushman); Nomura, p. 55, pl. 1,
fig. 12a, b
Description
The test is trochoid in the initial portion, subsequently chang-
ing to a triserial arrangement. The test is nearly conical.
Chambers are highly inflated, separated by depressed su-
tures. The wall is agglutinated, with a calcareous cement.
The aperture is a semicircular slit, with narrow lip, located
at the base of the last biggest chamber.
Dimensions
The species length ranges from 125 to 550 µm in the current
study.
Stratigraphic range
The species ranges from the Chattian to Zanclean at 752 and
1168. At 1139 it ranges from Chattian to early Tortonian.
Remarks
This species is considered cosmopolitan (Kender et al., 2008)
and has previously been recorded in the Miocene of the
east Indian Ocean by Nomura (1991) and on the Kerguelen
Plateau (Mackensen, 1992).
5 Discussion and summary
Benthic foraminiferal assemblages from ODP Sites 752,
1139 and 1168 were identified from the >63 µm fraction
for the late Oligocene through to Pliocene. Assemblages are
highly diverse comprising more than 200 species at each in-
dividual site with between 41 % and 57 % of taxa at any
one site, not recorded at the other two, but these taxa rep-
resent very low proportions of the overall assemblages. Al-
though there is some faunal similarity between the three sites,
in terms of species occurrence, the faunal composition at
each site differs through the studied interval and each site
responds differently to climatic events through the Miocene.
Hole 752A yielded 268 species, of which 143 do not oc-
cur at the other two sites; at Hole 1139A 224 species were
recorded, 92 of which are only recorded at this site and in
Hole 1168A, 331 species are recorded with 189 taxa not
present at the two other sites. However, the most abundant
taxa in this study occur at all sites, with just a few exceptions.
Notably, there appear to be environmentally-cued distribu-
tions. For example, Bolivina huneri only occurs at Hole 752
and 1168 but does not occur at 1139. Bolivina dilatata is
recorded at 1139 and 1168 but not at 752. Bolivina cf. reticu-
lata and Bolivina viennensis are recorded at 752 but not at the
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D. Ridha et al.: Latest Oligocene to earliest Pliocene deep-sea benthic foraminifera 223
other two sites. Bulimina gibba is recorded at 1139 but not at
either 1168 or 752. A full discussion of the assemblage data
and palaeoceanographic implications are deferred to a later
contribution. Here we selected the 52 most abundant species,
each representing at least 1 % of the total fauna encountered
in this study, to describe and illustrate providing an impor-
tant reference work for foraminiferal studies in the southern
Indian Ocean.
Data availability. The data used to compile this paper are avail-
able at the University of Birmingham eData repository and can be
accessed at https://doi.org/10.25500/edata.bham.00000411 (Ridha
et al., 2019).
Author contributions. DR has undertaken all sample processing,
data collection and analyses under the supervision of IB and KME.
All authors have contributed to the writing of the paper.
Competing interests. The authors declare that they have no con-
flict of interest.
Acknowledgements. This research used samples and data pro-
vided by the International Ocean Discovery Program (IODP). The
authors wish to thank Chris Smart (University of Plymouth) for his
useful discussions during this project. We would also like to thank
Flavia Fiorini (Khalifa University) and a second, anonymous re-
viewer for their thoughtful comments on the original submission.
Financial support. Dana Ridha has been supported by the HCED
(The Higher Committee for Education Development in Iraq).
Review statement. This paper was edited by Laia Alegret and re-
viewed by Flavia Fiorini and one anonymous referee.
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