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Journal of Foraminiferal Research, v. 33, no. 2, p. 144–154, April 2003
WATER-DEPTH DISTRIBUTION OF MIOCENE PLANKTONIC FORAMINIFERA
FROM ODP SITE 744, SOUTHERN INDIAN OCEAN
WOJCIECH MAJEWSKI1
Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warszawa, Poland
ABSTRACT
Planktonic foraminifera were sampled from the strati-
graphic interval between 21 and 10 Ma at ODP Site 744
(southern Indian Ocean). The water-depth hierarchy of
18 taxa was established using d18O measurements and
morphological indicators. The latter were derived from
modern foraminifera, and included the presence of
spines, test size, wall texture, chamber shape, and type
of coiling.
The shallow-water fauna includes predominantly
small, microperforate species belonging to Globigerinita,
Tenuitellinata, Tenuitella, as well as small Globigerina
and Neogloborotalia. The intermediate group comprises
heavily ornamented microperforates (Tenuitella and
Turborotalita) as well as large Globigerina and possibly
Neogloboquadrina. The deep-water taxa include large
forms such as Globorotalia, Catapsydrax and possibly
Turborotalita.
There is general agreement between isotopic and mor-
phological proxies of water-depth habitats among the
analyzed Miocene planktonic foraminifera from south-
ern high latitudes. Large and deep-water dwelling fo-
raminifera have significantly shorter stratigraphic rang-
es compared to small foraminifera inhabiting shallow-
waters. This relationship is probably due to common
cryptic speciation among planktonic foraminifera, which
may be more pronounced among small taxa of simple
test morphology.
INTRODUCTION
SITE LOCATION AND MIOCENE CLIMATE
ODP Site 744 (latitude 61834.669S and longitude
80835.469E) is located near the southern edge of the Ker-
guelen Plateau, southern Indian Ocean, at a present water-
depth of 2307m, and is ;1500 km NNE from Prydz Bay,
East Antarctica. The scope of this study spans a large por-
tion of the Miocene Epoch between 21.0 and 10.0 Ma. Dur-
ing this interval the region around and including ODP Site
744 was separated from the Antarctic continental shelf by
an abyssal deep sea basin which prevented significant input
of terrigenous material. For this reason the Miocene sedi-
ments in the ODP Site 744 region are purely biogenic and
consist of nannofossil and diatom ooze.
The Miocene was an eventful epoch in Earth history es-
pecially in the Southern Ocean. The opening of the Drake
Passage near the Oligocene-Miocene boundary (Barker and
Burrell, 1976) was the principal factor driving long-term
Neogene global cooling. Also, the main growth phase of the
Antarctic ice sheet took place during the middle Miocene
1 E-mail: wmaj@twarda.pan.pl
144
pg 144 # 1
at ;14Ma (Shackleton and Kennett, 1975). It coincided
with significant intensification of the Circum-Antarctic Cur-
rent, which in turn led to the widespread development of
stratigraphic hiatuses at southern high latitudes (Keller and
Barron, 1987; Flower and Kennett, 1994). This major stage
of East Antarctic Ice Sheet growth was directly preceded by
the mid-Miocene climate optimum (;16 Ma), apparently
the warmest period of the entire Neogene (Savin, 1977).
Nevertheless, the ultimate reasons for the progressive mid-
Miocene warming and cooling are still a topic of debate
(Pagani and others, 1999).
d18O WATER-DEPTH RANKING VERSUS MORPHOLOGY
Reconstructing foraminiferal paleoecology is an essential
prerequisite for the correct interpretation of their extensive
fossil record, which in turn contributes a great deal to our
understanding of the Miocene climatic processes and re-
cords. One of the critical elements of planktonic foraminif-
eral ecology is their distribution within the water column.
Keller (1985) and Gasperi and Kennett (1993) reconstructed
depth habitats of some Miocene planktonic foraminifera, but
they did not include high latitude assemblages, which were
widespread throughout the Southern Ocean. This study is
intended to address this deficiency.
Urey (1947, 1948) and subsequent researchers established
that the oxygen isotopic composition of calcite is dependent
on temperature. Following this discovery, Emiliani (1954)
conducted the first investigation of depth preferences among
planktonic foraminifera, based on oxygen isotopic ratios of
tests from core samples. Since that time, numerous studies
of water-depth distribution of planktonic foraminifera have
been carried out, and assumed d18O equilibrium between
foraminiferal calcite and seawater (Schackleton, 1968;
Douglas and Savin, 1978; Deuser and others, 1981; Keller,
1985). According to this assumption, the higher the d18O
ratio of foraminiferal calcite the deeper the position of the
taxon analyzed within the water column. However, a number
of studies revealed that not all planktonic foraminifera se-
crete their shells in isotopic equilibrium with seawater
(Shackleton and others, 1973; Vergnaud Grazzini, 1976;
Williams and others, 1979; Fairbanks and others, 1980;
1982; Spero, 1998). These results suggest that a degree of
caution is necessary when d18O results are converted into
water-depth hierarchies for planktonic foraminifera. These
interpretations also highlighted the need for using alterna-
tive water depth indicators, which are independent of d18O.
Early researchers pointed out that there is a strong rela-
tionship between the position of modern foraminifera in the
water column and their test morphology (Berger, 1969; Bé,
1977; Douglas and Savin, 1978). This dependence was soon
adapted to fossil assemblages as well (Cifelli, 1969; Hart,
1980; Caron and Homewood, 1983; Jarvis and others, 1988;
Leckie, 1989). However, it is not clear whether all com-
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WATER-DEPTH DISTRIBUTION, ODP 744
pressed or keeled tests are characteristic of deep-water hab-
itats, as argued based on relationships of modern planktonic
foraminifera to bathymetry (Hart, 1980). According to iso-
topic data, some fossil morphologically complex (globoro-
taliid-like) genera could inhabit shallow-water habitats
(Shackleton and others, 1985; Corfield and Cartlidge, 1991;
Keller and others, 1992; van Eijden, 1995). A prime ex-
ample may be the morozovellids, which morphologically
resemble modern globorotaliids, but according to stable iso-
tope data could be surface dwellers. This disagreement be-
tween water-depth foraminiferal habitats derived from the
morphologic and oxygen-isotopic indicators is especially
evident among Paleogene and Cretaceous taxa (Keller and
others, 1992; van Eijden and Ganssen, 1995; Pearson, 1998;
Norris and Wilson, 1998). Fortunately, the Miocene faunas
discussed below are phylogenetically close to the extant spe-
cies, for which the general rule that morphologically com-
plex taxa inhabit deeper waters has not been questioned.
MORPHOLOGICAL INDICATORS OF WATER-DEPTH
HABITATS AND SETTLING RATES OF FORAMINIFERAL
TESTS
Living adult of foraminifera of various species inhabit
different depth-zones within the water column (Bé, 1977;
Ravelo and Fairbanks, 1992; Chaisson and Ravelo, 1997).
The bathymetric position of particular individuals usually
varies with ontogeny; numerous living taxa descend pro-
gressively through the water column with increasing matu-
rity (Fairbanks and others, 1982; Deuser, 1987).
The mechanisms by which foraminifera maintain their
position within the water column are not fully understood,
even in the case of Recent species (Lipps, 1979; Spero,
1998). As planktonic organisms, foraminifera are incapable
of active swimming but regulate buoyancy as the means of
movement within the water column (Hemleben and others,
1989). The major factors that regulate the overall buoyancy
of a single foraminifer are density of protoplasmic fluid and
test weight. Because the density of protoplasmic fluid can-
not be reconstructed in the case of fossil groups, the follow-
ing discussion will consider only test morphology and struc-
ture.
There are only brief and generalized statements in the
literature that directly address morphologic changes with
depth habitat among modern foraminiferal species. One
commonly stressed strategy for reduced sinking, and thus
maintenance of a stable position within the upper water col-
umn, is the presence of spines, as discussed in detail by
Furbish and Arnold (1997). In terms of non-spinose species,
the general opinion is that shallow-water species are glob-
ular and thin-walled in contrast to deep water forms, which
are large, flattened, thick-walled, and bear calcitic crusts
(Bé, 1977; Douglas and Savin, 1978). These morphological
trends were derived from depth habitat rankings constructed
using net tow results (Berger, 1969; Bé, 1977), oxygen iso-
topic investigations (Emiliani, 1971; Savin and Douglas,
1973; Shackleton and Vincent, 1978), or both (Fairbanks
and others, 1982).
On the other hand, measurements of settling rates of emp-
ty tests have attracted more detailed studies, and may be
treated as an approximation of the ability of various forms
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145
TABLE 1. Compilation of morphological and structural characteristics
of shallow-water (slowly settling) species as compared to deep-water
(rapidly settling) species, combined after Adelseck and Berger, 1975;
Bé, 1977; Douglas and Savin, 1978; Lipps, 1979. The properties are
arranged from top to bottom in order of decreasing significance.
Shallow-water dwellers
(slow settling)
Deep-water dwellers
(fast settling)
spinose lack of spines
small test big test
thin wall thick wall
smooth wall strongly ornamented wall
evolute coiling involute coiling
globular chambers flattened chambers
to achieve particular positions within the water column. The
morphologies of foraminiferal tests that settle relatively
slowly might correspond to shallow-water dwellers. Berger
and Piper (1972) compared results from settling experiments
with analyses of graded beds, and deduced a ranking of
modern foraminifera based on settling characteristics. Their
results were in general agreement with a similar ranking by
Parker and Berger (1971; Table 1). Other investigators con-
cluded that the most important factor governing settling
rates is the size of specimens (Adelseck and Berger, 1975),
as well as the ratio of test mass (composite of chamber
arrangement and wall thickness) to its volume, and shape
(for a review see Lipps, 1979). Moreover, the wall thickness
may be broadly approximated by its degree of ornamenta-
tion, as smooth walls tend to be thinner than those that are
heavily ornamented (Hemleben and others, 1989). Finally,
evolute shells with globular chambers are expected to pos-
sess greater volume than involute shells with flattened
chambers of similar size and wall thickness (see review by
Lipps, 1979).
EXPERIMENTAL STRATEGY
As noted above, the Miocene faunas discussed in this
study are phylogenetically close to the extant species, for
which the rule that morphologically complex taxa inhabit
deeper waters appears to be valid. Therefore, the morphol-
ogy-based water-depth ranking is considered as having
equal significance to its d18O-induced equivalent. The plank-
tonic foraminiferal taxa are ranked on the basis of morpho-
categories (Table 1), keeping in mind differences in their
significance. The results are then compared to ascertain how
consistent the various morphologic criteria are. Finally, the
composite results of morphologic analysis are compared to
the d18O ranking.
METHODS
Morphological analysis was conducted separately for
each morpho-category, which was arranged according to de-
creasing significance. These are the possession of long
spines, test size, wall ornamentation, chamber shape, and
coiling type (Table 1). Among them, the test-size analysis
was conducted quantitatively. Measurements were conduct-
ed on randomly selected populations of 18 taxa from frac-
tions greater than 63 mm. For each taxon, diameters of at
least 200 specimens were measured under a light stereo-
microscope.
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146
FIGURE 1. Location of samples analyzed for isotopic composition of oxygen and carbon in foraminifera. Sample position is related to depth in
meters below sea floor as well as the plankton-to-benthos (p/b) ratio curve. Note that the position of the samples is in areas of relatively high p/b
ratio, which indicates relatively low dissolution. Dates of the samples are according to the age model of Ramsay and Baldauf (1999).
Tests from six sediment samples were selected for stable
isotope investigation from the total of 124 available Mio-
cene samples spanning the time interval between 21.0 and
10.0 Ma. The samples were chosen according to the pres-
ervation of foraminifera, relatively high plankton-to-benthos
ratio (Fig. 1), and faunal composition. Each sediment sam-
ple provided 4 to 10 monospecific (monogeneric) samples
of foraminifera. At least one sample of Cibicidoides sp. was
analyzed from the six sets of measurements.
Stable isotope analyses were conducted on a total of 45
foraminiferal samples. The isotope analyses were performed
at the University of Florida, and run on a Finnigan-MAT
252 mass spectrometer coupled to a Kiel II carbonate prep-
aration device. The data were related to the isotopic standard
NBS-19, which was taken to have a d18O value of 22.19‰
PDB, and a d13C value of 1.92‰ PDB, through 12 standard
replicates (Appendix 1). Standard deviations, calculated
from these additional measurements for oxygen and carbon
isotopes, are 0.04‰ and 0.02‰, respectively. This indicates
high instrument precision of both d18O and d13C measure-
ments. All isotopic ratios are expressed in standard delta
notation relative to VPDB.
RESULTS
WATER-DEPTH RANKING ACCORDING TO DIFFERENT
MORPHOLOGIC CHARACTERISTICS
In the following four sections, the water-depth hierarchy
of high-latitude planktonic foraminifera is analyzed in terms
of the presence of spines, test size, wall texture, chamber
shape, and coiling type. These four morpho-categories are
arranged according to their decreasing impact on
foraminiferal position within water-column (Table 1). The
outcome of the four morphologic analyses is summarized in
Table 2.
PRESENCE OF SPINES
Presence of spine bases among the fossil species dis-
cussed is limited to Globigerina bulloides and Turborotal-
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MAJEWSKI
ita quinqueloba. They are the only taxa among all the an-
alyzed species that are expected to be able to sustain a sig-
nificantly shallower position within the water column than
would be expected from other morpho-indicators.
TEST SIZE
Size distribution diagrams of the analyzed species are pre-
sented in Fig. 2. The species may be divided into three
groups, each based on increasing size and thus on expected
progressively deeper habitat. The first group includes the
smallest taxa, whose maximum abundance is at 100 mm in
diameter. It includes G. glutinata, G. boweni, T. cf. T. cle-
menciae, Tenuitella sp., T. cf. jamesi and T. quinqueloba.
Their size distribution diagrams are artificially asymmetrical
(except G. boweni) due to the use of the 63 mm sieve for
processing sediment, which overestimated their average
size. All species within this first size-group belong to the
microperforate foraminifera.
The second group includes species with size distribution
histograms that are predominantly symmetrical and are thus
unaffected by sample processing. The diameter of highest
frequency ranges from 120 to 160 mm. This intermediate
size-group includes N. nympha, N. continuosa, T. uvula,
and the small variant of Globigerina brazieri. It is more
phylogenetically diverse than the finest-size group and in-
cludes only one species of ‘‘microperforates,’’ as well as
Globigerina and Neogloboquadrina. When looking at the
size histograms, little variation in size occurs among the
species belonging to this size-group.
The largest size-group includes the eight remaining taxa
of Globigerina, Globorotalia, and Catapsydrax, which are
all potentially deep-water dwellers. They are considerably
larger in diameter than the species in the two previous
groups. Moreover, they exhibit a significantly broader size
distribution and less distinctive peaks for the most abundant
diameter intervals.
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WATER-DEPTH DISTRIBUTION, ODP 744 147

TABLE 2. Comparison of vertical hierarchy rankings according to various morphological criteria discussed in this paper. The morphological criteria
are arranged with a decrease in importance to the right.

Presence of Wall Chamber

Species spines Test size texture shape Composite

G. glutina shallow shallow shallow/int. shallow

G. boweni shallow shallow shallow/int. shallow
T. uvula int. int. shallow/int. int.
T. cf. clemenciae shallow shallow shallow/int. shallow
T. cf. jamesi shallow int. shallow/int. shallow/int.
Tenuitella sp. shallow shallow shallow/int. shallow
T. quinqueloba shallow shallow int. ? shallow/int. shallow
G. bulloides shallow deep int. shallow/int. int.
G. brazieri (small) int. deep shallow/int. int.
G. brazieri (large) deep deep shallow/int. deep
N. continuosa int. deep shallow/int. int.
N. nympha int. deep shallow/int. int.
G. scitula deep int. deep deep
G. praescitula plexus deep deep deep deep
G. zealandica plexus deep deep deep deep
C. unicavus deep deep shallow/int. deep
C. dissimlis deep deep shallow/int. deep
Catapsydrax sp. deep deep shallow/int. deep

WALL TEXTURE classified as follows: Neogloboquadrina, Globigerina bra-

Different types of wall texture may be used to characterize
various phylogenetic groups within the analyzed Miocene fo-
raminifera. These groups are discussed below according to
progressive increase in complexity of their wall texture.
In general, microperforate species have smooth and finely
perforate walls. Pustules commonly occur, but their size and
concentration vary inter- and intra-specifically. Tenuitelli-
nata uvula and T. cf. T. jamesi are the two coarsely pus-
tulose species. A majority of specimens belonging to these
two taxa possesses abundant and well-developed pustules,
which may suggest an intermediate-water depth habitat. The
other species are rather delicate although many may also
carry pustules, but they are generally less pronounced and
less common than in the two species named above. There-
zieri, and the most robust Catapsydrax, which exhibits par-
ticularly massive wall ornamentation. All species, having the
cancellate type of wall, are interpreted here as deep-water
taxa.
CHAMBER SHAPE AND COILING TYPE
The genus Globorotalia alone differs significantly from
the others by its flattened, sometimes acute-edged profile.
This shape points to a generally deep-water habitat. On the
other hand, all microperforate, globigerinid, and neoglobo-
quadrinid taxa could inhabit shallower waters, as judged
from their widely understood globigerinid type of coiling.
OXYGEN ISOTOPIC RATIOS AND DEPTH HABITATS OF THE

fore, they are classified as shallow-water dwellers. On the PLANKTONIC FORAMINIFERA

other hand, Turborotalita quinqueloba possesses widely
spaced spines or a thick calcite crust, which makes it dif-
ficult to classify bathymetrically.
Globorotalia usually has a smooth test with pustules of
different sizes that are concentrated predominantly in the
umbilical and apertural area or consist of a solid, continuous
crust. Globorotalia scitula, classified here as intermediate-
water dwelling, appears to have a significantly less-devel-
oped pustulose exterior and thinner wall than the two large
and massive mid-Miocene plexi of G. praescitula and G.
zealandica. These last two robust taxa are interpreted as
being confined to deep-water habitats.
Globigerina bulloides has a typical spinose texture. It is
therefore difficult to compare it with the non-spinose taxa.
Nevertheless, if the spines are ignored, walls of Globigerina
bulloides are punctuated with irregular spine bases, which
may completely surround pores, producing delicate, rough
ridges between them. Therefore, Globigerina bulloides is
classified as occupying an intermediate-water depth.
The last group constitutes species having a cancellate wall
texture. They may be ordered by genera in respect to pro-
gressive deepening of their pore pits as well as by the regu-
larity and thickness of inter-pore polygonal ridges. They are
Stable isotopic values are listed in Appendix 1. They were
the basis for establishment of foraminiferal depth rankings
for the six sets of measurements (Fig. 3). The rankings for
each set of samples are not always consistent; therefore, a
composite or prevailing depth ranking is also presented in
Table 3. This classification into shallow, intermediate, and
deep-water habitats is relative as compared to different taxa
within a population, and it does not carry an ecological im-
plication (e.g., particular depth or position with respect to
the thermocline).
According to stable isotopic data presented in Figure 3,
the shallow-water fauna include all analyzed minute micro-
perforate species (except Turborotalita quinqueloba and
Tenuitella cf. T. jamesi), Globigerina brazieri (minute var-
iant), Neogloborotalia continuosa and possibly Neoglobo-
quadrina nympha. The intermediate group comprises of T.
cf. T. jamesi and possibly T. quinqueloba, as well as Glo-
bigerina bulloides, Globigerina brazieri (large variant) and
possibly Neogloboquadrina nympha. The deep-water taxa
include Catapsydrax sp., Catapsydrax dissimilis, Catapsy-
drax unicavus, Globorotalia zealandica plexus, Globoro-
talia praescitula plexus, Globorotalia scitula, and possibly
Turborotalita quinqueloba.
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148 MAJEWSKI

FIGURE 2. Expected depth habitats of Miocene planktonic foraminifera from ODP Site 744, according to test diameters of these taxa. The scale
in the lower right corner indicates the percentage of foraminiferal specimens within each size interval, which correspond to a 20 micron range of
test diameter. The number of counts used for construction of each histogram is indicated after a taxonomic name. Criteria, on which the Miocene
planktonic foraminiferal taxa are placed within one of the three depth-habitat groups, are discussed in text.

DISCUSSION paper. A composite water-depth ranking integrating four mor-

COMPARISON OF THE WATER-DEPTH RANKINGS DERIVED
FROM ISOTOPE AND MORPHOLOGICAL STUDIES
Table 2 shows a compilation of proposed depth habitats as
indicated by various morphological features discussed in this
pho-categories analyzed is also presented. A comparison of
results from morphologic and d18O approaches is presented
in Table 4. Most interpretations are consistent among these
two indicators. However, the d18O depth estimation of Ten-
uitellinata uvula appears to be significantly shallower than is
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WATER-DEPTH DISTRIBUTION, ODP 744 149

FIGURE 3. d18O values of different Miocene foraminifera plotted versus ages according to age model by Ramsay and Baldauf (1999). The names
of taxa are listed in the same order as their isotopic values. The benthic d18O values were corrected by 1 0.5‰ after Barrera and Huber (1991),
and are marked by black squares as well as by bold generic names. Note that the d18O scale is reversed, with the highest values at the bottom, as
they refer to deepest (coolest) water habitats. The arrows pointing at the time scale show exact age of samples analyzed.

suggested by its morphological criteria. Unfortunately, only On the other hand, Turborotalita quinqueloba and Ten-
a single stable isotopic measurement was obtained from this uitella cf. T. jamesi show slightly deeper-water habitats as
species (Table 3). Therefore, the d18O indicator of shallow- indicated by more positive d18O values than suggested by
water habitat of T. uvula is not well documented. the morphological proxy (Table 4). It appears, then, that
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150 MAJEWSKI

TABLE 3. Comparison of vertical hierarchy rankings determined from the six sets of stable isotopic measurements. The interpretations are made
separately for each set of measurements according to d18O signatures of planktonic foraminifera (Fig. 3). The last column presents composite
interpretations.

Age (Ma)
Composite
Taxon 19.4 17.8 16.5 14.4 11.4 11.0 depth ranking

G. glutinata int. shallow shallow shallow shallow

T. uvula shallow shallow
T. cf. clemenciae shallow shallow
T. cf. jamesi int. int. int.
Tenuitella sp. shallow shallow shallow shallow
T. quinqueloba int. deep deep/int.
G. bulloides int. int. deep int. int. int.
G. brazieri (small) shallow shallow shallow
G. brazieri (large) int. int. int.
N. continuosa shallow shallow shallow
N. nympha int. shallow shallow/int.
G. scitula deep deep
G. praescitula plexus deep deep int. deep
G. zealandica plexus deep deep deep
C. unicavus deep deep
C. dissimilis deep deep
Catapsydrax sp. deep deep deep

encrustation and heavily pustulose wall texture may be re- for shallow-to-intermediate depth habitats for Miocene Neo-
sponsible for this inconsistency. globoquadrina, as compared to the Recent N. pachyderma.
The assignment of Neogloboquadrina nympha and N.
continuosa as shallow-to-intermediate and shallow-water WATER-DEPTH RANKING AND RATES OF EVOLUTION
dwellers, respectively, on the basis of the d18O proxy ap-
pears to be the most controversial. These interpretations re- Figure 4 shows stratigraphic occurrences of planktonic
main uncertain, and they are even more problematic in the taxa compared to their water-depth hierarchy and d18O val-
light of the morphological proxies, especially considering ues. It appears that the shallow-water dwelling small micro-
that the various morphological indicators are also inconsis- perforates (Globigerinita, Tenuitellinata, Tenuitella) have
tent (Table 2). Bé (1977) classified their close relative (Re- the longest stratigraphic ranges, whereas large and deep-
cent Neogloboquadrina pachyderma) as deep-water dwell- water Globorotalia and Catapsydrax have the shortest. On
ers; however, others suggested a broader bathymetric distri- the other hand, the long-ranging microperforates are ex-
bution. On the other hand, the modern high latitude N. pach- pected to have the shortest life cycle and the greatest gen-
yderma (Huber and others, 2000) is significantly larger than erational turnover, according to their size, shallow position
Neogloboquadrina nympha and N. continuosa analyzed in within the water-column, and analogy to their modern rel-
this study. This size difference might support an argument atives. This suggests two possibilities: (1) the evolutionary
(mutation) rates among large deep-water dwelling forami-
nifera are greater than among the microperforates, or (2)
TABLE 4. Comparison of vertical hierarchy rankings according to cryptospeciation is more common among the shallow-water
morphology of tests and oxygen stable isotopic results. dwelling small planktonic foraminifera than among large
deep-water dwellers.
Taxon
Morphology
of test d18O values
Parker and others (1999) suggested that cosmopolitan fo-
raminiferal species were relatively small and morphologi-
Globigerinita glutinata shallow shallow cally conservative. The long and continuous ranges of the
Globigerinita boweni shallow
Tenuitellinata uvula int. shallow shallow-water dwelling microperforates provide support for
Tenuitella cf. clemenciae shallow shallow this statement; however, it does not point specifically to ei-
Tenuitella cf. jamesii shallow/int. int. ther the low evolutionary rates (1) or common cryptospe-
Tenuitella sp. shallow shallow ciation among this group of foraminifera (2). The first pos-
Turborotalita quinqueloba shallow/int. deep/int.
Globigerina bulloides int. int. sibility (1) was supported by Stanley and others (1988), who
Globigerina brazieri (small) int. shallow noted that the duration of Neogene deep-water globorotaliid
Globigerina brazieri (large) deep int. lineages is much shorter than that of the shallow-water glo-
Neogloboquadrina continuosa int. shallow bigerinid lineages. The same scenario was also favored by
Neogloboquadrina nympha int. shallow/int. Hart (1980), who also suggested greater evolutionary rates
Globorotalia scitula deep deep
Globorotalia praescitula plexus deep deep for deep-water dwellers, which were believed to be more
Globorotalia zealandica plexus deep deep advanced in an evolutionary sense, more specialized, and
Catapsydrax unicavus deep deep more susceptible to extinction. On the other hand, de Vargas
Catapsydrax dissimilis deep deep and Pawlowski (1998), who analyzed SSU rDNA partial
Catapsydrax sp. deep deep
sequences of modern planktonic foraminifera, argued for
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WATER-DEPTH DISTRIBUTION, ODP 744 151

FIGURE 4. Comparison of depth hierarchy of high-latitude planktonic Miocene foraminifera, according to their d18O values, with their strati-
graphic distribution at ODP Site 744. Note that the deep-water dwellers exhibit significantly more restricted stratigraphic ranges than taxa occupying
intermediate- and shallow-water habitats.

slower molecular evolutionary rates among extinct globoro- cryptospeciation among these groups, which leads to un-
taliids than globigerinids. Therefore, it appears that the mor- derestimation of their evolutionary turnover rates in the fos-
phological simplicity among microperforate foraminifera sil record. The last scenario appears to be likely in the light
and larger globigerinids may be the reason for common of apparently common presence of cryptospecies among
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152
modern planktonic foraminifera (Huber and others, 1997;
de Vargas and others, 1999; 2001; Darling and others, 2000)
SUMMARY
Southern Indian Ocean planktonic foraminifera from the
chronostratigraphic interval between 21 and 10 Ma (Mio-
cene) were analyzed. Water-depth ranking of 18 taxa was
established, using d18O measurements and morphological
characteristics, which include presence of spines, test size,
wall texture, chamber shape, and type of coiling. Composite
morphological indicators generally agree with isotopically
inferred water-depth habitats of the analyzed planktonic fo-
raminifera.
The shallow-water fauna includes the small microperfor-
ate species Globigerinita glutinata, Tenuitellinata uvula,
Tenuitella cf. T. clemenciae, Tenuitella sp., as well as Glo-
bigerina brazieri (small variant), Neogloborotalia contin-
uosa and possibly Neogloboquadrina nympha. The inter-
mediate group constitutes Tenuitella cf. T. jamesi and pos-
sibly Turborotalita guinqueloba, as well as Globigerina
bulloides, Globigerina brazieri (large variant) and possibly
Neogloboquadrina nympha. The deep-water taxa include
Catapsydrax sp., Catapsydrax dissimilis, C. unicavus, the
Globorotalia zealandica plexus, the Globorotalia praesci-
tula plexus, Globorotalia scitula, and possibly Turborotal-
ita quinqueloba.
Comparison of occurrences of taxa present at ODP Site
744 with their depth habitats showed that large and deep-
water dwelling foraminifera have significantly shorter strati-
graphic ranges, than small foraminifera inhabiting shallow
waters. This relationship agrees with previous suggestions,
and is probably due to common cryptospeciation among
planktonic foraminifera, which obscures true evolutionary
turnover rates due to simplicity of foraminiferal test mor-
phology.
ACKNOWLEDGMENTS
This paper is part of my dissertation completed at The
Ohio State University, Columbus, United States of America.
I would like to thank my supervisors Dr. Peter-Noel Webb
and Dr. Brian T. Huber for various support and guidance
throughout all stages of my studies. Thanks also to these
workers for proofing the manuscript’s English. The manu-
script also benefitted significantly from a review by R. C.
Thuvall. I am also very thankful to the Ocean Drilling Pro-
gram (ODP) for providing the numerous samples for this
study. Thanks are also expressed to Dr. Jason Curtis from
the University of Florida for conducting the stable isotope
measurements.
This study was financially supported by National Science
Foundation grant to Dr. Peter N. Webb (NSF/OPP 9420475,
NSF/OPP 9317979) and in part from the Cushman Foun-
dation for Foraminiferal Research (Cushman Foundation
Student Research Grant-in-aid, 1998).
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Received 4 September 2001
Accepted 2 May 2002
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154 MAJEWSKI

APPENDIX 1.

Sample Species Sediment Age Number of d18O d13C

number name sample (cm) (Ma) specimens (‰) (‰) Comments

NBS-19 22.16 1.93

NBS-19 22.19 1.92
NBS-19 22.14 1.93
NBS-19 22.21 1.93
WM-1 G. bulloides 6H-3, 60-62 11.0 10 3.09 1.28
WM-2 N. nympha 6H-3, 60-62 11.0 23 2.88 1.40
WM-3 N. continuosa 6H-3, 60-62 11.0 22 2.87 1.41
WM-4 Cibicidoides sp. 6H-3, 60-62 11.0 3 3.06 20.01
WM-5 G. scitula 7H-2, 10-12 11.4 18 3.10 1.68
WM-6 G. bulloides 7H-2, 10-12 11.4 27 2.75 1.11
WM-7 N. continuosa 7H-2, 10-12 11.4 21 2.61 1.71
WM-8 N. nympha 7H-2, 10-12 11.4 27 2.72 1.61
WM-9 G. glutinata 7H-2, 10-12 11.4 60 2.54 1.58 pustulose
WM-10 G. glutinata 7H-2, 10-12 11.4 60 2.62 1.60
WM-11 T. uvula 7H-2, 10-12 11.4 21 2.53 1.97
WM-12 T. quinqueloba 7H-2, 10-12 11.4 60 3.01 0.24
WM-13 Cibicidoides sp. 7H-2, 10-12 11.4 2 2.73 0.81
WM-14 G. bulloides 7H-5, 60-62 14.4 5 2.15 1.67
WM-15 G. zealandica pl. 7H-5, 60-62 14.4 9 2.17 1.68
WM-16 G. praescitula pl. 7H-5, 60-62 14.4 6 1.73 1.78
NBS-19 22.25 1.90
NBS-19 22.24 1.88
WM-17 G. glutinata 7H-5, 60-62 14.4 60 1.51 1.52
WM-18 Tenuitella sp. 7H-5, 60-62 14.4 60 1.61 1.49
WM-19 T. cf. jamesi 7H-5, 60-62 14.4 70 1.73 1.31
WM-20 T. cf. clemenciae 7H-5, 60-62 14.4 65 1.53 1.47
WM-21 Cibicidoides sp. 7H-5, 60-62 14.4 4 2.15 1.40
WM-22 G. zealandica pl. 8H-5, 110-112 16.5 5 1.95 1.74
WM-23 G. praescitula pl. 8H-5, 110-112 16.5 12 1.98 1.99
WM-24 T. cf. jamesi 8H-5, 110-112 16.5 52 1.87 1.44
WM-25 Tenuitella sp. 8H-5, 110-112 16.5 50 1.78 1.51
WM-26 Cibicidoides sp. 8H-5, 110-112 16.5 3 1.60 1.33
WM-27 G. bulloides 9H-5, 80-82 17.8 8 2.03 1.60
WM-28 G. praescitula pl. 9H-5, 80-82 17.8 15 2.16 1.56
WM-29 G. brazieri 9H-5, 80-82 17.8 26 1.79 1.60 small
WM-30 Catapsydrax sp. 9H-5, 80-82 17.8 19 2.30 1.17
WM-31 P. brazieri 9H-5, 80-82 17.8 15 1.91 1.76 large
WM-32 Tenuitella sp. 9H-5, 80-82 17.8 68 1.81 1.41
WM-33 G. glutinata 9H-5, 80-82 17.8 65 1.71 1.52
WM-34 T. quinqueloba 9H-5, 80-82 17.8 54 1.87 1.35
NBS-19 22.18 1.91
NBS-19 22.12 1.93
NBS-19 22.22 1.94
NBS-19 22.17 1.92
WM-35 Cibicidoides sp. 9H-5, 80-82 17.8 3 1.91 1.17
WM-36 G. brazieri 10H-6, 61-63 19.4 14 2.32 1.76 large
WM-37 G. bulloides 10H-6, 61-63 19.4 14 2.26 1.70
WM-38 C. unicavus 10H-6, 61-63 19.4 4.5 2.59 1.39
WM-39 C. dissimilis 10H-6, 61-63 19.4 5 2.66 1.54
WM-40 Catapsydrax sp. 10H-6, 61-63 19.4 5 2.55 1.59
WM-41 C. dissimilis 10H-6, 61-63 19.4 5 2.42 1.05 gametogenetic
WM-42 G. brazieri 10H-6, 61-63 19.4 18 2.14 1.58 small
WM-43 G. glutinata 10H-6, 61-63 19.4 73 2.36 1.40
WM-44 Cibicidoides sp. 10H-6, 61-63 19.4 5 2.33 0.81
WM-45 Cibicidoides sp. 7H-2, 10-12 11.4 6 2.80 0.88
NBS-19 22.22 1.90
NBS-19 22.14 1.91