Marine Micropaleontology 81 (2011) 1–21

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Marine Micropaleontology
j o ur nal ho mep ag e : www.e lsevier. com/l o cate/ m arm icro

Dissolution susceptibility of Paleocene–Eocene planktic foraminifera: Implications

for palaeoceanographic reconstructions
Thi Minh Phuong Nguyen a,⁎, Maria Rose Petrizzo b, Peter Stassen a, Robert P. Speijer a
a
Department of Earth and Environmental Sciences, K.U.Leuven, Celestijnenlaan 200E, 3001 Leuven, Belgium
b
Università degli Studi di Milano, Dipartimento di Scienze della Terra, “Ardito Desio”, via Mangiagalli 34, 20133 Milano, Italy

a r t i c l e in fo abstract

Article history: We investigated shell characteristics and differential dissolution susceptibility of planktic foraminiferal
Received 11 January 2011 species derived from upper Paleocene and lower Eocene deep-sea sequences, Ocean Drilling Program (ODP)
Received in revised form 24 June 2011 Site 865 (Allison Guyot) and Sites 1209B, 1210B and 1212A (Shatsky Rise) in the North Pacific Ocean. The
Accepted 8 July 2011
purposes of this study are: 1) assessing the effects of differential dissolution on upper Paleocene–lower
Eocene planktic foraminiferal assemblages, at species level and within different biozones, to quantify
Keywords:
dissolution susceptibility of genera and species; 2) investigating the differences in shell characteristics; 3)
dissolution susceptibility
planktic foraminifera
revealing the relationship between shell parameters and dissolution robustness of taxa, and 4) identifying the
North Pacific key shell parameter(s) influencing the dissolution susceptibility of foraminiferal taxa.
Ocean Drilling Program Two independent experiments were carried out, one focusing on gradual qualitative deterioration of taxa by
Paleocene–Eocene dissolution and the other documenting the weight loss of taxa. Shell parameters such as wall thickness,
paleoceanography porosity and pore size were determined through Scanning Electron Microscopy (SEM) and image analysis
paleoclimatology (JMicroVision).
taphonomy We found that the large muricate Acarinina and Morozovella are most resistant, followed by the cancellate
Subbotina and the small muricate Igorina, confirming results of previous work. At species level, the thick-
walled Acarinina soldadoensis, Acarinina subsphaerica and the large Morozovella subbotinae are the most
resistant species. Most of the large Morozovella species such as Morozovella aequa, Morozovella formosa-
gracilis, Morozovella velascoensis and Morozovella pasionensis, together with Acarinina nitida show
intermediate dissolution resistance, whereas the small muricate Igorina species, the cancellate Subbotina
velascoensis and the thin-walled Morozovella acuta and Morozovella occlusa are the most vulnerable species.
We propose a formula for calculating the dissolution resistance of taxa based on their wall thickness and
size — two key parameters in dissolution resistance of a species. Application of this formula reveals good
agreement between the calculated and measured dissolution resistance, indicating its robustness.
Furthermore, the agreement between our experimental results, in-situ experimental results on live
foraminifera and natural quantitative/qualitative records suggests that our experiments accurately mimic
natural dissolution processes. Consequently, these experimental results strongly bear on interpretations of
foraminiferal dissolution in natural environments, especially in studies on early Paleogene climatic events
that are often associated with dissolution phenomena. More generally, a proper assessment of taphonomic
alteration by dissolution should be part of every paleoenvironmental reconstruction based on quantitative
foraminiferal records.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction 1980; Žarić et al., 2005), making them excellent paleoenvironmental

Planktic foraminifera are among the most abundant organisms in
the open ocean and their shells often form a major component of
calcareous sediments on the ocean's floors. The sensitivity of planktic
foraminifera to the properties of ambient sea water has been
extensively demonstrated (e.g. Berger, 1970, 1975; Coulbourn et al.,
⁎ Corresponding author. Tel.: + 32 16 326452; fax: + 32 16 322980.
E-mail addresses: thiminhphuong.nguyen@ees.kuleuven.be,
hoathachthao2001@yahoo.com (T.M.P. Nguyen).
recorders in Quaternary and deep-time studies (e.g. Berger, 1968;
Mix, 1989). More specifically, planktic foraminifera provide informa-
tion on upper ocean environments in which they calcify and play an
important role in interpretations of paleoenvironmental conditions at
the time of deposition (e.g. Bé, 1968; Berger, 1968; Herman and
Bogyo, 1980).
However, only a small proportion of all shells sinking to the sea
floor are ultimately preserved in the sedimentary record, because of
carbonate dissolution in the water column, at the sea floor and within
the sediment. Dissolution of planktic foraminifera in the deep sea is a

0377-8398/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.marmicro.2011.07.001
2 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
widespread phenomenon (e.g. Berger, 1970; Peterson and Prell, 1985;
Conan et al., 2002). In ocean cores, complete dissolution is easily
recognized by the absence of calcareous foraminifera in clay beds (e.g.
Spiegler and Jansen, 1989). Less pervasive dissolution is, however,
more difficult to recognize and is often ignored, although various
subtle indications, such as increased shell fragmentation, decreased
planktic/benthic (P/B) ratios and carbonate content are frequently
documented in the geological records (Melillo, 1988; Weaver and
Raymo, 1989; Iaccarino and Proto-Decima, 1990; Premoli Silva and
Spezzaferri, 1990; Chen and Farrell, 1991; Brunner, 1992; Basov,
1995; Sliter, 1995).
In paleoenvironmental studies, the basic assumption is that fossil
assemblages truthfully reflect the biocoenosis and underlying environ-
mental signals (e.g. van der Zwaan et al., 1990). Since differential
susceptibility of planktic foraminifera to dissolution will influence the
composition of fossil assemblages, the reliability of paleoenvironmental
reconstructions, based on planktic foraminiferal assemblages may be
significantly reduced. For this reason, an objective assessment of
potential taphonomic alteration by dissolution should be part of every
paleoenvironmental reconstruction based on quantitative foraminiferal
data, from deep-sea basins and continental margin settings.
Most studies dealing with planktic foraminiferal dissolution are
restricted to recent settings and Quaternary sequences. Research carried
out on pre-Quaternary stratigraphic intervals is relatively rare and
focused on specific qualitative and quantitative aspects of dissolution
(e.g. Jenkins and Orr, 1971; Violanti et al., 1979; Malmgren, 1987;
Petrizzo et al., 2008). Most knowledge on dissolution susceptibility of
extant planktic foraminifera is derived from quantitative analysis, and in
rare cases, from experimental studies. A general outcome of these
studies is that dissolution susceptibility depends on the characteristics of
the foraminiferal shell, such as surface/volume ratio, microstructural
features, or porosity (e.g. Sliter et al., 1975; Thunell, 1976; Corliss and
Honjo, 1981). A commonly used index for foraminiferal dissolution is
the percentage of planktic foraminiferal fragments in the assemblage
(Berger, 1973; Cullen and Prell, 1984; Peterson and Prell, 1985; Le and
Thunell, 1996). Yet, this index does not unequivocally characterize
dissolution levels (Petrizzo et al., 2008). For instance, fragment ratios
can drop rapidly in samples that are severely affected by dissolution. In
addition, the fragmentation index cannot be employed in lithified
sediments that require more intense processing than soft mud,
contributing to fragmentation.
This study is part of a larger effort dealing with the effects of
differential dissolution on benthic and planktic foraminiferal assem-
1212
Fig. 1. Late Paleocene paleogeography (Hay et al., 1999) and locations of ODP Sites 865 (Allison Guyot, equatorial Pacific Ocean), 1209, 1210 and 1212 (Shatsky Rise, northwestern
Pacific Ocean).
blages from stratigraphic intervals that record remarkable environmen-
tal changes. The general aims are to: 1) reveal the effects of dissolution
on the composition of foraminiferal assemblages; 2) determine which
factors control the dissolution susceptibility of taxa, and 3) improve the
reliability of paleoenvironmental reconstructions based on foraminiferal
assemblages. Our previous experimental study (Nguyen et al., 2009)
was carried out on a lower Eocene foraminiferal assemblage (benthic
and planktic) from the Esna Formation in the Dababiya section, the
Global boundary Stratotype Section and Point for the base of the Eocene
in Egypt (Dupuis et al., 2003). The results revealed, among others, that at
a generic level, Acarinina is slightly more dissolution resistant than
Morozovella, but more significantly, both these muricate, symbiont-
bearing, surface-dwelling taxa are much more resistant to dissolution
than the cancellate, asymbiotic, deep-dwelling Subbotina. Moreover,
morphology, test size and especially shell wall micro-texture seem to
play important roles in the dissolution susceptibility of planktic
foraminiferal taxa. These results motivated more detailed experimental
studies on upper Paleocene to lower Eocene planktic foraminiferal
species.
In this study, we carried out investigations on dissolution
susceptibility and shell parameters of taxa from upper Paleocene
and lower Eocene planktic foraminiferal assemblages from the Pacific
Ocean. This time slice was chosen because dissolution phenomena are
a recurrent problem in upper Paleocene to lower Eocene foraminiferal
assemblages, especially in connection with the Paleocene/Eocene
Thermal Maximum (PETM, e.g. Ernst et al., 2006; Petrizzo, 2007;
Guasti and Speijer, 2007), the Early Late Paleocene Event (ELPE, Röhl
et al., 2004; Petrizzo, 2005) and the Eocene Thermal Maximum 2
(ETM 2 — Shipboard Scientific Party, 2003). The selected sedimentary
sequences contain various indications suggesting that partial disso-
lution prevailed in the Pacific Ocean during the Paleocene–Eocene
interval (Sager et al., 1993; Colosimo et al., 2006).
This study is aimed at: 1) assessing the effects of differential
dissolution on upper Paleocene–lower Eocene planktic foraminiferal
assemblages, at species level and within different biozones, in order to
quantify dissolution susceptibility of genera and species; 2) investigating
differences in shell characteristics of planktic species; 3) revealing the
relationship between shell parameters and dissolution susceptibility;
4) identifying the key parameter(s) influencing dissolution susceptibility
of these planktic taxa. Results from these investigations form a basis for
developing a new dissolution index, which allows an objective and
quantitative evaluation of the degree of dissolution that has occurred in
planktic foraminiferal assemblages.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 3

Table 1 Table 1 (continued)

List of samples, taxa and size fractions used in the qualitative shell deterioration Sample Zone/ Taxa Size fraction (μm)
experiment (experiment 1).
subzone name
125– 250– 330– 400– 490– N 550
Sample Zone/ Taxa Size fraction (μm) 250 330 400 490 550
subzone name
125– 250– 330– 400– 490– N 550 198-1209B- P4c Subbotina x
250 330 400 490 550 23H-1, velascoensis
63–64
143-865B- P6a Acarinina x x
198-1209B- P4b Acarinina x x
11H-4, (E3) subsphaerica
23H-2, subsphaerica
48–50
101–102
143-865B- P6a Morozovella x x
198-1209B- P4b Morozovella x x
11H-4, (E3) subbotinae
23H-2, occlusa
48–50
101–102
143-865B- P6a Subbotina x
198-1209B- P4b Subbotina x
11H-4, (E3) velascoensis
23H-2, velascoensis
48–50
101–102
143-865B- P5 (E2) Acarinina x x
198-1210B- P6a Acarinina x x x
12H-1, soldadoensis
19H-CC (E3) soldadoensis
130–132
198-1210B- P6a Morozovella x x x
143-865B- P5 (E2) Morozovella x x
19H-CC (E3) subbotinae
12H-1, occlusa
198-1210B- P6a Subbotina x x x
130–132
19H-CC (E3) velascoensis
143-865B- P5 (E2) Morozovella x x
198-1212A- P6b Acarinina x x x
12H-1, aequa
9H-CC (E4) soldadoensis
130–132
198-1212A- P6b Subbotina x x x
143-865B- P5 (E2) Subbotina x x
9H-CC (E4) velascoensis
12H-1, velascoensis
130–132
143-865B- P5 Acarinina x x x
13H-1, soldadoensis
124–125
143-865B- P5 Igorina pusilla x Table 2
13H-1, List of samples, taxa used in the weight loss experiment (experiment 2).
124–125
143-865B- P5 Morozovella x x Samples Zone/subzone Taxa name Size fraction
13H-1, acuta (μm)

124–125 125– 330–

143-865B- P5 Subbotina x x 330 490
13H-1, velascoensis 143-865B-11H-4, 48–50 P6a (E3) Acarinina soldadoensis x
124–125 143-865B-11H-4, 48–50 P6a (E3) Morozovella subbotinae x x
143-865B- P4c Acarinina x x x 143-865B-11H-4, 48–50 P6a (E3) Subbotina velascoensis x x
13H-2, nitida 143-865B-12H-1, 130–132 P5 (E2) Acarinina soldadoensis x x
50–52 143-865B-12H-1, 130–132 P5 (E2) Morozovella occlusa x x
143-865B- P4c Morozovella x x x 143-865B-12H-1, 130–132 P5 (E2) Morozovella aequa x x
13H-2, velascoensis 143-865B-12H-1, 130–132 P5 (E2) Subbotina velascoensis x x
50–52 143-865B-13H-1, 124–125 P5 Acarinina nitida x x
143-865B- P4c Subbotina x x x 143-865B-13H-1, 124–125 P5 Igorina pusilla x
13H-2, velascoensis 143-865B-13H-1, 124–125 P5 Igorina tadjikistanensis x
50–52 143-865B-13H-1, 124–125 P5 Morozovella acuta x x
143-865B- P4c Igorina pusilla x x 143-865B-13H-1, 124–125 P5 Subbotina velascoensis x x
13H-3, 143-13H-2, 50–52 P4c Acarinina nitida x
50–52 143-13H-2, 50–52 P4c Morozovella velascoensis x
143-865B- P4c Subbotina x 143-865B-13H-3, 50–52 P4c Acarinina nitida x x
13H-3, velascoensis 143-865B-13H-3, 50–52 P4c Igorina albeari x
50–52 143-865B-13H-3, 50–52 P4c Igorina tadjikistanensis x
143-865B- P4c Acarinina x x 143-865B-13H-3, 50–52 P4c Subbotina velascoensis x x
13H-4, nitida 143-865B-13H-4, 50–52 P4c Acarinina nitida x x
50–52 143-865B-13H-4, 50–52 P4c Igorina pusilla x
143-865B- P4c Igorina pusilla x 143-865B-13H-4, 50–52 P4c Igorina albeari x
13H-4, 143-865B-13H-4, 50–52 P4c Igorina tadjikistanensis x
50–52 143-865B-13H-4, 50–52 P4c Morozovella acuta x x
143-865B- P4c Morozovella x 143-865B-13H-4, 50–52 P4c Subbotina velascoensis x x
13H-4, acuta 143-865B-14H4, 47–49 P4a Igorina pusilla x
50–52 143-865B-14H4, 47–49 P4a Igorina albeari x x
143-865B- P4c Subbotina x x 143-865B-14H4, 47–49 P4a Igorina tadjikistanensis x
13H-4, velascoensis 143-865B-14H4, 47–49 P4a Morozovella acuta x
50–52 198-1209B-23H1, 63–64 P4c Morozovella pasionensis x x
143-865B- P4a Igorina x x 198-1212A-9H-CC P6b (E4) Acarinina soldadoensis x x
14H4, tadjikistanensis 198-1212A-9H-CC P6b (E4) Acarinina subsphaerica x x
47–49 198-1212A-9H-CC P6b (E4) Igorina broedermanni x
143-865B- P4a Igorina pusilla x x 198-1212A-9H-CC P6b (E4) Morozovella subbotinae x x
14H-4, 198-1212A-9H-CC P6b (E4) Morozovella formosa- x
47–49 gracilis
198-1209B- P4c Acarinina x 198-1212A-9H-CC P6b (E4) Subbotina velascoensis x x
23H-1, subsphaerica 198-1210B-19H-CC P6a (E3) Acarinina soldadoensis x x
63–64 198-1210B-19H-CC P6a (E3) Acarinina subsphaerica x x
198-1209B- P4c Morozovella x x 198-1210B-19H-CC P6a (E3) Igorina broedermanni x
23H-1, pasionensis 198-1210B-19H-CC P6a (E3) Morozovella subbotinae x
63–64 198-1210B-19H-CC P6a (E3) Subbotina velascoensis x x

(continued on next part of this page)

4 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
2. Material and Thethods
2.1. Material
The planktic foraminiferal taxa used in the experiments were
derived from ODP Leg 143, Hole 865B, Allison Guyot in the equatorial
Pacific Ocean and from ODP Leg 198, Holes 1209B, 1210 and 1212A,
Shatsky Rise in the North Pacific Ocean (Fig. 1). The present water
depth for Hole 865B is 1518 m (Sager et al., 1993); for Hole 1209B is
2387 m (Takeda and Kaiho, 2007); for Hole 1210B is 2573 m
(Bralower et al., 2006) and for Hole 1212A is 2681 m (Bralower
et al., 2006).
At Hole 865B, 7 samples were retrieved from cores 11H-4, 12H-1,
13H-1, 13H-2, 13H-3, 13H-4, and 14H-4. The biozonation applied is
based on the data of Petrizzo et al. (2005), adapted to the low-latitude
zonation scheme of Berggren and Pearson (2005). The samples
span planktic foraminiferal Subzone P4a to P6a (=E3, zonation by
Berggren and Pearson, 2005). At Hole 1209B, we used 2 samples from
cores 23H-1 and 23H-2 (Zone P4). At Hole 1210B, 1 sample from core
19H-CC (Subzone P6a= E3) and at Hole 1212A, another sample from
core 9H-CC (Subzone P6b= Zone E4) were analyzed. Species identifi-
cations (Tables 1 and 2, Plates 1 and 2) follow Olsson et al. (1999),
Petrizzo (2005), and Pearson et al. (2006).
The selected samples contain rich planktic foraminiferal
assemblages. The initial preservation state of these assemblages
is good, most specimens in these assemblages are intact and
generally free of infilling. However, the sedimentary successions
from which they were retrieved also show changes in fragmenta-
tion of planktic foraminifera and changes in carbonate content,
suggesting at least partial dissolution associated with the PETM
(Sager et al., 1993; Hancock and Dickens, 2005). Deposition at
these sites took place at lower bathyal to abyssal paleodepths of 1300–
2000 m (Bralower et al., 1995; Sager et al., 1999; Shipboard Scientific
Party, 2002).
2.2. Dissolution experiments
To construct a dissolution susceptibility ranking scheme for
planktic taxa, two independent experiments were carried out. The
first experiment focused on gradual qualitative deterioration of taxa
and the second on the weight loss of taxa.
2.2.1. Experiment 1 — qualitative deterioration
The purpose of this experiment is to investigate the differential
dissolution resistance of planktic taxa, based on the deterioration of
their tests by dissolution. Samples were first soaked in water and
washed through a 38 μm sieve, and then dried on a hot plate. In total
12 species belonging to the four most common upper Paleocene to
lower Eocene genera (Acarinina, Morozovella, Subbotina and Igorina)
were used in this experiment (Table 1 and Plates 1 and 2). Well-
preserved empty specimens were picked from six different size
fractions of 125–250 μm, 250–330 μm, 330–400 μm, 400–490
μm, 490–550 μm and N 550 μm. Where possible, 10–20 intact
specimens of each species and from every size fraction were used
in the experiment. Each group of specimens was stored in separate
glass containers. Prior to the experiment, the preservation state of
these specimens was determined by binocular microscopy. During
the experiment, specimens were exposed to 5 ml of dissolution
media (a buffered acetic acid solution with a pH of 6.6) in two hour
increments until the last specimen was fully dissolved or disinte-
grated. Every 2 h, specimens were cleaned through a 38 μm sieve
using distilled water, dried, and their state of preservation was re-
assessed by binocular microscope.
A foraminiferal preservation scale modified from Boltovskoy and
Totah (1992) was employed and scores were assigned according to
the Absolute Preservation Scores (APS), ranging from 1 (=test
destroyed) to 8 (=test intact). These values were then converted
into Relative Preservation Scores (RPS), which are calculated relative
to the initial APS (see also Nguyen et al., 2009):
RPS(tx)= APS(tx)/APS(t0), in which tx is the time (measured in
hours) of experimental exposure, APS(t0) is the APS prior to the
experiment.
We also calculated the Cumulative Relative Preservation Scores
(CRPS) for each species within a size fraction, this is to express the
progressive dissolution characteristics of the taxa:
CRPS= Σ RPS(tx), x ranges from 0 to the time (hours) when the
taxa are completely dissolved or disintegrated.
Because the CRPS of a taxon differs per size fraction, we calculated
the average CRPS for the combined size. This value enables us to
compare the differential dissolution between taxa.
Average CRPS= Σ CRPS (size m)/nr. of size fractions, m= size
fraction of 125–250 μm, 250–330 μm, 330–400 μm, 400–490 μm,
490– 550 μm and N 550 μm.
2.2.2. Experiment 2 — weight loss
In the second experiment, we focused on investigating the
differential weight loss of taxa resulting from dissolution. During
the first experiment, we observed that larger specimens of a species
possess thicker test walls than smaller specimens and thus contain
more calcite, so they are expected to take longer to dissolve. This
means that the relationship between size fraction and dissolution
susceptibility of taxa is most likely related to the original weight of
the specimens. Furthermore, calcite content in the test is probably
the key factor responsible for the differential dissolution between
specimens and species (Berger, 1967). Therefore, independently
from experiment 1, we examined the role of the original test weight
on the differential dissolution susceptibility of planktic foraminiferal
species.
Another set of about 13,000 specimens, composed of 15 species
belonging to the genera Acarinina, Igorina, Morozovella and Subbotina,
were picked from splits of the same samples used in experiment 1 and
were used to evaluate the weight loss (Table 2 and Plates 1 and 2). All
specimens used in this investigation were selected from two size
fractions of 125–330 μm and 330–490 μm. To assess the general fate
of the species, all specimens encountered in the sub-samples (about
1000 specimens) were used in the experiment, without distinguish-
ing their preservation state. The experiment was carried out following
the same procedure used in experiment 1. Before and after the
experiment, the number of specimens of every species was counted
and their total weight was measured using the Micro Sartorius
scale. Then, the total weight was divided by the number of
specimens for each species to obtain the average weight of a
species at that moment. We also calculated the Relative Remaining
Weight (RRW) after every 2 h of acid exposure in comparison with
their original weight (100%) before the experiment. These values
were then summed into the Cumulative Relative Remaining Weight
(CRRW) and used to compare the weight loss among species by
dissolution.
The calculations are as follows:
RRW(tx)= TW(tx)/TW(t0), where TW is the total weight of taxa, tx
is the time that taxa are exposed to acid treatment and t 0 is the time
before taxa are exposed to acid treatment.
CRRW= Σ RRW(tx), x ranges from t0 to the time (hours) that the
taxa are completely dissolved.
Because the CRRW of a single species somewhat varies between
samples, we calculated the mean value of the CRRW for these samples.
This is needed in order to compare the differential weight loss
between taxa.
Mean CRRW= Σ (CRRWy Ny)/Σ Ny, in which CRRWy indicates
the CRRW of taxa in sample y and Ny is the number of specimens of
taxa in sample y.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 5

Plate 1. SEM images illustrating the species concepts applied in this study. 1) Igorina pusilla, sample 865B-13H-1, 124–125 cm. 2) Morozovella subbotinae, sample 1210B-19H-CC.
3) Igorina albeari, sample 1209B-22H-3, 111–112 cm. 4) Igorina tadjikistanensis, sample 1209B-23H-2, 148–149 cm. 5) Igorina broedermanni, sample 865B-8H-2, 50–52 cm.
6) Morozovella velascoensis, sample 865B-13H-2, 50–52 cm. 7) Morozovella aequa, sample 1209B-23H-2, 101–102 cm. 8) Morozovella pasionensis, sample 1209B-23H-1, 63–64 cm.
a, umbilical view; b, lateral view; c, spiral view. Scale bars: 100 μm.
6 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21

Plate 2. SEM images illustrating the species concepts applied in this study. 1) Morozovella occlusa, sample 1209B-23H-2, 101–102 cm. 2) Acarinina nitida, sample 1209B-23H-2, 91–
92 cm. 3) Acarinina subsphaerica, sample 1209B-23H-2, 101–102 cm. 4) Acarinina soldadoensis, sample 865B-11H-4, 48–50 cm. 5) Subbotina velascoensis, 1209B-23H-1, 63–64
cm.
6) Acarinina soldadoensis, sample 1210B-19H-CC. 7) Morozovella formosa-gracilis, sample 1212A-9H-CC. 8) Morozovella formosa-gracilis, sample 1210B-19H-CC. a, umbilical view;
b, lateral view; c, spiral view. Scale bars: 100 μm.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 7

Data viewer (Object extraction)

Class Class 1

Whole image (image surface %) 54.45

Poligon 1(image surface %) 17.17

Poligon 1 count 758

1 D measurement

Parameter Value (µm)

Line length 18.92385

Fig. 2. Measurements of wall thickness, shell porosity and pore size (external) using JMicroVision software (Nicolas, 2009). To determine wall thickness we used the function “1 D
measurement” and the result is indicated here as the value of “Line length” (18.92385) in the table on the left. To determine porosity and pore size, we used the function “Object
extraction”. Results of these measurements are indicated here in the upper table on the right. In this table, the value in “Poligon 1” (17.17%) indicates the percentage of pore size on
the external surface of specimen (54.45% of the whole picture) and the value in “Poligon 1 count” (758) shows the number of pores on the surface of the specimens, as indicated in
this image. Area of these pores is listed in Appendix D.

2.3. Measurement of shell parameters
To determine data on shell parameters such as wall thickness,
porosity (% of surface area occupied by pores) and pore sizes (on the
external surface), we analyzed SEM images of taxa using JMicroVision
software (Nicolas, 2009). Measured specimens were randomly chosen
from samples in which the taxa were abundant. Wall thickness was
measured in the central part of the last chamber, or on the
penultimate chamber in specimens having a very thin wall in the
last chamber related to the reproductive cycle (gametogenic
chamber). We used the function “1 D measurement” to measure the
wall thickness and “Object extraction” to measure the porosity (%
pore area of total surface area) and pore size (Fig. 2). Measure- ments
of these parameters for every taxon are listed in Appendices A to F.
Our observations showed that some taxa (for instance Subbotina
velascoensis) show fewer chambers in the last whorl than others and
when the last chamber of this species dissolved, their size significantly
decreases. So it seems that the number of chambers in the last whorl
can also be an important factor affecting the dissolution resistance of
taxa. To examine whether this is true, we counted the number of
chambers in the last whorl of each species. This is carried out, together
with measuring the size (largest diameter) of individual species using
a binocular microscope.
2.4. Data analysis
All data on CRPS, CRRW and shell parameters were analyzed
statistically, using PAST software (Hammer et al., 2001). Correlation
analysis was used to examine the degree of correlation between these
variables. As our data are not normally distributed, we used the non-
parametric correlation analysis (Spearman and Kendall correlation
measures; Gibbons, 1985) for a more powerful analysis that highlights
which parameters play the most important role in the dissolution
susceptibility of the species.
Subsequently, multiple regression analyses were performed, in
which shell parameters, showing a strong correlation with CRPS and
CRRW, were considered as independent variables while CRPS, CRRW
were considered as dependent variables. This is done in order to
quantify the effects of these parameters on the dissolution resistance
of species as well as to describe their relationship through a linear
function.
3. Results
3.1. Differential shell deterioration by dissolution (experiment 1)
3.1.1. Differential shell deterioration between genera
We investigated the variation in RPS of species belonging to
Acarinina, Morozovella, Subbotina and Igorina within a sample. Data
from individual and combined size fractions show strong species
offsets as evidenced by RPS (Fig. 3A–D). Acarinina species are generally
most dissolution resistant, followed by Morozovella subbotinae,
Morozovella aequa, Subbotina velascoensis, the other Morozovella
species and Igorina species, respectively. The differences in RPS
between taxa are prominent in larger size fractions and much less in
small size fractions (Fig. 3A–C, Table 3A). When species within a genus
from different samples and different size fractions are merged
together, the RPS of the genera vary widely. The RPS of Igorina drops
rapidly, reaching zero after 22 h of exposure, whereas the RPS of
Subbotina, Morozovella and Acarinina took 36, 56 and 84 h, respec-
tively, to reach zero (Fig. 3E). The differences in dissolution damage
between these genera leads to dissimilar average CRPS between
genera (Table 3B), which is highest (11.5) in Acarinina, and is about 1.3
times higher than that of Morozovella and 2 times higher than the CRPS
8 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21

A 1
B 1

0.8 0.8

0.6 0.6
RPS

RPS
0.4 0.4

0.2 0.2

0 0
0 4 8 12 16 20 24 28 32 36 40 44 48 52 56 60 64 68 72 76 80 84 0 4 8 12 16 20 24 28 32 36 40
Exposure time (hours) Exposure time (hours)
A. soldadoensis 400-490 µm S. velascoensis 400-490 µm A. soldadoensis 330-400 µm M. aequa 330-400 µm
M. subbotinae 400-490 µm M. occlusa 330-400 µm S. velascoensis 330-400 µm

C D
1 1

0.8 0.8

RPS
0.6 0.6
RPS

0.4 0.4

0.2 0.2

0 0
0 2 4 6 8 10 12 14 16 18 20 0 4 8 12 16 20 24 28 32 36 40 44 48 52 56 60 64 68 72 76 80 84
Exposure time (hours) Exposure time (hours)
A. nitida 125-250 µm I. pusilla 125-250 µm A. soldadoensis M.subbotinae S.velascoensis
S. velascoensis 125-250 µm

E
1

0.8

0.6
RPS

0.4

0.2

0
0 4 8 12 16 20 24 28 32 36 40 44 48 52 56 60 64 68 72 76 80 84
Exposure time (hours)
Acarinina Igorina
Morozovella Subbotina

Fig. 3. A–C: Examples of the differential reduction in Relative Preservation Score (RPS) with exposure time for some common Paleogene species in the same size fraction, from the
same sample. A—species from sample 1210B-19H-CC (Shatsky Rise), size fraction of 400–490 μm; B—species from sample 865B-12H-1, 130–132 cm (Allison Guyot), size fraction of 330–
400 μm; C—species from sample 865B-13H-4, 50–52 cm (Allison Guyot), size fraction of 125–250 μm; D—differential reduction in Relative Preservation Score between species from
sample 1210B-19H-CC when all size fractions are combined together and E—idem for genera (all species within each genus are merged) from different samples, different size fractions
combined.

of Subbotina. The small muricate Igorina shows the lowest average
CRPS, indicating that it is most dissolution prone.
3.1.2. Differential shell deterioration among species
Dissolution susceptibility of species within genera was determined
for specimens from all samples merged together, for the different size
fractions (Table 3A) and for the combined size fraction (Fig. 4 and
Table 3B).
dissolution susceptibility. The mean RPS and average CRPS values of
A. nitida are always lower than those of A. subsphaerica and
A. soldadoensis. For A. nitida, the RPS reached zero after 42 h of exposure.
A. soldadoensis lasted twice as long (84 h) and A. subsphaerica in between
(56 h) (Fig. 4A). These patterns consistently indicate, both in the different
size fractions and in the combined size fraction, that A. nitida is most
susceptible Acarinina, followed by A. subsphaerica and then A. soldadoensis
(Table 3A–3B).

3.1.2.1. Acarinina species. Three Acarinina species were used in the
investigation of shell deterioration: A. nitida, Acarinina soldadoensis
and Acarinina subsphaerica. In examinations of individual size
fractions and the combined size fraction, changes in the RPS of
these species with exposure time show considerable differences in
3.1.2.2. Igorina species. Two Igorina species were investigated: I. pusilla
and Igorina tadjikistanensis. The RPS values of these species are nearly
identical throughout the experiment, reaching zero after 22 h of
exposure (Fig. 4B). The similarity in their average CRPS for both size
fractions of 125–250 μm and 250–330 μm as well as at the combined
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 9

Table 3A
List of species ranked in decreasing order of dissolution resistance, based on the average CRPS from the start until the end of experiment 1, in different size fractions. Note that the
data presented here are a combination of different samples.

Size Acarinina species Igorina species Morozovella species Subbotina species

(μm)
Taxa name CRPS Taxa name CRPS Taxa name CRPS Taxa name CRPS

125–250 A. nitida 4.0 I. pusilla 3.5 S. velascoensis 3.4

I. tadjikistanensis 3.4

250–330 A. nitida 6.0 I. pusilla 5.1 S. velascoensis 4.9

A. subsphaerica 8.0 I. tadjikistanensis 5.1
A. soldadoensis 9.8
330–400 A. nitida 8.6 M. acuta 4.6 S. velascoensis 6.8
A. subsphaerica 10.6 M. pasionensis 5.7
A. soldadoensis 11.8 M. occlusa 5.7
M. aequa 8.0
M. subbotinae 8.9
400–490 A. nitida 11.5 M. velascoensis 6.6 S. velascoensis 8.7
A. subsphaerica 15.4 M. acuta 7.1
A. soldadoensis 16.1 M. occlusa 7.4
M. pasionensis 8.0
M. aequa 8.7
M. subbotinae 10.3
490–550 A. soldadoensis 20.6 M. velascoensis 7.7
M. subbotinae 15.8

N 550 M. velascoensis 9.0

size fraction (Table 3A–3B) demonstrate that these taxa are equally
susceptible to dissolution.
3.1.2.3. Morozovella species. Six Morozovella species were examined:
M. acuta, M. aequa, Morozovella occlusa, Morozovella pasionensis,
M. subbotinae and Morozovella velascoensis. The results indicate a
significant distinction in the solution susceptibility between
M. subbotinae and the other Morozovella species. During the
experiment, the RPS of M. subbotinae is always higher than that of
the other species and reached zero after 56 h (Fig. 4C). The decrease in
RPS is very fast in M. acuta and M. occlusa, yet slower in
M. velascoensis, M. pasionensis and M. aequa. From 26 to 34 h, the
RPS of the following species successively dropped to zero: M. acuta,
M. pasionensis, M. aequa, M. occlusa and M. velascoensis (Fig. 4C).
The observed difference in dissolution resistance between
Morozovella species is also expressed by their average CRPS values
at the individual size fractions and combined size fraction. In both
cases, this value is highest in M. subbotinae, followed by M. aequa
(Table 3A–3B). For the other taxa, comparisons between size fractions
show small deviations in their rank, with respect to the average CRPS
values (Table 3A). The combined size fraction indicates that among
Morozovella species, M. subbotinae is the most dissolution resistant,
followed by M. aequa, M. velascoensis, M. pasionensis, M. occlusa and M.
acuta, respectively (Table 3B).
3.1.2.4. Intraspecific differential dissolution between samples and
locations. Our results also highlight intraspecific dissolution suscep-
tibility between locations (Fig. 5). For instance: A. soldadoensis
specimens from sample 1210B-19H-CC are the most resistant,
followed by specimens from sample 1212A-9H-CC, 865B-12H-1, 130–
132 cm and 865B-13H-1, 124–125 cm, respectively (Fig. 5A). The
distinct difference in dissolution resistance between locations is also
observed in M. subbotinae and in S. velascoensis (Fig. 5B and C). In all
cases, the taxa from Site 865 are less dissolution resistant than the taxa
from the Shatsky Rise sites.
3.2. Differential weight loss from dissolution (experiment 2)

Table 3B List
3.2.1. Average genus and species weights
of genera and species ranked in decreasing order of dissolution resistance, based on the Our measurements show a considerable difference in average
average CRPS from the start until the end of experiment 1. Note that the data initial weight between genera and species. In the size fractions
presented here are a combination of different size fractions from different samples. N 125 μm Acarinina is the heaviest genus, with an average weight of
Taxa name Number of specimens Time (hours) taxa persist Average CRPS 13.6 μg/specimen, followed by Morozovella (12.2 μg), Subbotina
(7.9 μg) and Igorina (6.2 μg) (Table 4).
Acarinina 303 84 11.5
Morozovella 279 78 8.8
Among Acarinina, A. soldadoensis is the heaviest species (average
Subbotina 284 36 6.0 weight 16.5 μg), followed by A. subsphaerica (13.4 μg) and A. nitida
Igorina 90 22 4.3 (11.3 μg). Igorina albeari is the heaviest Igorina (7.1 μg), followed by
I. pusilla (6.3 μg), I. tadjikistanensis (5.7 μg) and I. broedermanni (5.5 μg).
A. soldadoensis 172 84 14.6
The morozovellids can be divided into two groups: a heavy group,
A. subsphaerica 61 56 11.4
M. subbotinae 97 56 11.7 composed of the large species M. formosa-gracilis, M. subbotinae,
A. nitida 70 42 7.5 M. aequa, M. velascoensis and M. pasionensis (average weights 11.4–
M. aequa 30 32 8.4 17.1 μg) and a second lighter group consisting of M. occlusa and
M. velascoensis 52 34 7.8
M. acuta (7.6–8.6 μg/specimen; Table 4 and Appendix A).
M. pasionensis 30 30 6.8
M. occlusa 40 34 6.6
S. velascoensis 284 36 6.0 3.2.2. Weight loss by dissolution
M. acuta 30 26 5.9 The weight loss experiments show that dissolution causes
I. pusilla 60 22 4.3 differential decreases in weight among genera (Fig. 6). The weight
I. tadjikistanensis 30 22 4.2
of Acarinina reached zero after 58 h of exposure while for Morozovella,
10 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21

A susceptible (Table 4). In contrast, Morozovella species show large

1 differences in weight loss. M. subbotinae was the most robust of the
Morozovella species investigated, taking longest to dissolve and with
0,8 highest RRW throughout the experiment. M. occlusa and M. acuta
dissolve most rapidly and have the lowest RRW. Consequently,
0,6
RPS

0,4
A
0,2 1

0 0,8
0 10 20 30 40 50 60 70 80 90
Exposure time (hours)
0,6

RPS
A. soldadoensis A. subsphaerica A. nitida

0,4
B
1 0,2

0,8 0
0 10 20 30 40 50
0,6 Exposure time (hours)
RPS

A. soldadoensis330-400 µm, 198-1210B-19H-CC

0,4 A. soldadoensis 330-400 µm, 198-1212A-9H-CC
A. soldadoensis 330-400 µm, 143-865B-12H-1, 130-132
0,2 A. soldadoensis 330-400 µm, 143-865B-13H-1, 124-125

0 B
0 2 4 6 8 10 12 14 16 18 20 22 1
Exposure time (hours)
I. pusilla I. tadjkistanensis 0,8

C 0,6
RPS

1
0,4
0,8
0,2
0,6
RPS

0
0,4 0 4 8 12 16 20 24 28 32 36 40 44 48 52 56
Exposure time (hours)
0,2 M. subbotinae 490-550 µm, 198-1210B-19H-CC
M. subbotinae 490-550 µm, 143-865B-11H-4, 48-50
0
M. subbotinae 400-490 µm, 198-1210B-19H-CC
0 10 20 30 40 50 60
M. subbotinae 400-490 µm, 143-865B-11H-4, 48-50
Exposure time (hours)
M. acuta M. aequa M. occlusa
C
M. pasionensis M. subbotinae M. velascoensis 1

Fig. 4. Differential reduction in the mean Relative Preservation Score with exposure
0,8
time between species that belong to the same genus. Note that values presented here
are combined from all size fractions and all samples for each taxon.
0,6
RPS

0,4
Subbotina and Igorina it took only 36, 34 and 28 h, respectively. During
the experimental process, the relative remaining weight (RRW) and 0,2
the mean CRRW of Acarinina are highest, followed by those of
Morozovella, Subbotina and Igorina, respectively (Fig. 6 and Table 4). 0
0 4 8 12 16 20 24 28 32
As Acarinina has the highest original starting weight this finding is Exposure time (hours)
expected. S. velascoensis 330-400 µm, 198-1209B-23H-1, 63-64
Among Acarinina, the weight of A. nitida dropped to zero after just S. velascoensis 330-400 µm, 143-865B-13H-2, 50-52
36 h of the experiment (Fig. 6). The weight of A. subsphaerica and
A. soldadoensis decreased more slowly and they fully fragmented after Fig. 5. Examples of differential dissolution susceptibility of species (same size fraction)
among samples from different locations. In general, taxa from Site 865 show higher
44 h and 58 h, respectively. This results in the low mean CRRW of
susceptibility than the same taxa from Sites 1209, 1210 and 1212. A—A. soldadoensis in
A. nitida in comparison with the other Acarinina species (Table 4). The size fraction of 330–400 μm from Zones E2-E3. B—M. subbotinae in size fractions of
Igorina species dissolved at similar rates (Fig. 6), resulting in nearly 400– 490 μm and 490–550 μm from Zone E3 (subzone P6a). C—S. velascoensis in size
identical CRRW values (5.3–5.6), with I. albeari being slightly less fraction of 330–400 μm from Subzone P4c.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 11

Table 4 genus Morozovella of which M. formosa-gracilis is the largest (average

List of genera and species ranked in decreasing order of dissolution resistance, based on size 480 μm), followed by M. velascoensis (440 μm) and M. subbotinae
the mean CRRW during experiment 2.
(430 μm). The mid-size group of taxa (300–400 μm) consists of the
Taxon Number of Average original Time (hours) CRRW remaining Morozovella species, the Acarinina species and S. velascoensis.
specimens weight (μg) taxa persist Igorina yields the smallest taxa, with average size ranges from 200 to
Acarinina 2969 13.6 58 9.4 250 μm.
Morozovella 2696 12.2 36 7.0
Subbotina 2912 7.9 34 6.0
Igorina 3263 6.2 28 5.5 3.3.2. Wall thickness
A. soldadoensis 884 16.5 58 11.3 Our measurements on wall thickness of the last normal chamber of
A. subsphaerica 1024 13.4 44 9.3
each specimen reveal significant differences between the taxa
M. subbotinae 1097 15.2 36 9.0
M. aequa 68 14.7 32 7.9 studied, both at genus and species level (Fig. 7B, Appendix C). Based
A. nitida 1061 11.3 36 7.7 on the average thickness, these taxa can be divided into two main
M. formosa-gracilis 59 17.1 34 7.1 groups: 1) the thick-walled group that exclusively consists of
M. velascoensis 269 14.6 30 6.9 Acarinina species, of which A. soldadoensis has wall thickness of
M. pasionensis 45 11.4 28 6.7
S. velascoensis 2912 7.9 34 6.0 31 μm (as well as the larger variation); A. subsphaerica and A. nitida
I. albeari 1209 7.1 28 5.6 showing thickness of 24 μm; 2) and a thin-walled group, of which
I. pusilla 661 6.3 24 5.4 large Morozovella species such as M. aequa, M. subbotinae, and
I. tadjikistanensis 710 5.7 22 5.4 M. formosa-gracilis yield an average wall thicknesses of 14–19 μm.
I. broedermanni 683 5.5 20 5.3
The test walls of Igorina spp., S. velascoensis, and of the other
M. acuta 1000 8.6 20 5.2
M. occlusa 158 7.6 18 4.7 morozovellids are slightly thinner (9–13 μm).

3.3.3. Shell porosity and pore size

Results indicate considerable dissimilarity in the proportion of
M. occlusa and M. acuta have the lowest mean CRRW (4.7 and 5.2, porosity on the external test surface (% pores) and pore sizes
respectively), followed by M. pasionensis, M. velascoensis, M. formosa- between species (Fig. 7C–D). S. velascoensis has the highest porosity
gracilis and M. aequa (6.7 to 7.9). M. subbotinae has the highest mean (23%) and pore size (N 7 μm2), but it also shows the largest variation.
CRRW (9.0), being twice as robust as the most susceptible M. occlusa and Acarinina species exhibit quite low and equable porosities, from 11%
M. acuta (Table 4 and Fig. 6). in A. subsphaerica to 12% in A. nitida. The pore sizes of this group are
intermediate, which range around 5 μm2. The difference in porosity
3.3. Shell characteristics between Igorina species is also minor, varying from 13% in I. pusilla
to 15% in I. albeari and pore sizes range from 5 to 6 μm2 .
3.3.1. Size Morozovellids yield porosities ranging from 9% (M. occlusa) to 16%
Results on the variation in size (largest diameter) between species (M. velascoensis) and have the smallest pores (3–4 μm2) (Fig. 7C and
(Fig. 7A, Appendix B) show that most of the large species belong to the Appendices D and E).
Time (hours) taxa exposure in to acid treatment

0
4
8
12
16
20
50% lost of total CaCO3 and total number
24 of specimens of each species
28
32
36
40
44
48
52
56
60
0 50 100 0 7500 15000 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100 0 50 100

Fig. 6. Differential weight loss among species by dissolution. Horizontal axis of the first graph indicates the percentage of total CaCO3 of all species and the second graphs shows the
number of all specimens used in the experiment. For the other graphs, horizontal axis shows the weight percentage of species. Data presented here resulted from the combination of
all size fractions of species, from different samples. Species are arranged from left to right in increasing order of Cumulative Relative Remaining Weight (CRRW). The subhorizontal
line connects the levels at which 50% of the CaCO3 and number of specimens are lost.
12 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21

A 600 B 40

500
30
Size of Species (m)

Shell wall thickness (m)

400

300 20

200
10
100

0 0

Taxa name Taxa name

Mean size of species Mean shell wall thickness

Standard deviation Standard deviation

C D 10
30

25 8

20
Pore area (m ) 6
Porosity (% pore)

15
4
10

2
5

0 0

Taxa name Taxa name

Mean % pore Mean pore area
Standard deviation Standard deviation

E
8

6
Number of chambers

Taxa name

Mean number of chambers

Standard deviation

Fig. 7. Shell characteristics among species: A—the size of species is measured as the largest diameter of individual specimens; B—the wall thickness is measured on the last preserved
chamber of specimens; C—shell porosity is the percentage of total pore area on the external surface of the specimens (% pore); D—the average size of a pore on the outer surface is
calculated as the total surface area of pores divided by the number of the pores and E—the number of chambers in the last whorl.

3.3.4. Number of chambers in the last whorl the same number of chambers in the last whorl, ranging from 5.9 to 6.4.
M. pasionensis has the highest number (on average 7.7 chambers) and Acarinina species and M. acuta, M. subbotinae and M. aequa have 4 to 5.1
S. velascoensis the least (3.5 chambers) (Fig. 7E, Appendix F). Igorina chambers in the last whorl. The largest variation in number of chambers is
species and M. occlusa, M. velascoensis, M. formosa-gracilis have more or less observed in M. velascoensis and M. occlusa, ranging from 5 to 8 chambers.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 13

Table 5
Correlation between CRRW, CRPS and shell parameters. Correlation values are given in the lower triangle of the matrix, and the probabilities that these parameters are uncorrelated
to each other are given in the upper part.

CRRW CRPS Wall thickness Original weight Size Porosity Pore area Number of chambers

CRRW 0.00 0.00 0.00 0.04 0.51 0.68 0.13

CRPS 0.90 0.00 0.00 0.00 0.33 0.46 0.19
Wall thickness 0.95 0.83 0.01 0.12 0.38 0.56 0.08
Original weight 0.85 0.94 0.74 0.00 0.71 0.33 0.16
Size 0.60 0.76 0.47 0.87 0.53 0.22 0.30
Porosity −0.21 −0.31 −0.28 −0.12 0.20 0.44 0.76
Pore area 0.13 −0.24 0.19 −0.31 −0.38 0.24 0.27
Number of chambers −0.46 −0.41 −0.52 −0.43 −0.33 −0.10 −0.35

3.4. Relationships between weight and other shell parameters
Our correlation analysis reveals a high value (0.87) between the
weight and the size of the species (Table 5). For weight and wall
thickness this value is slightly lower (0.74). The correlation values
between the original weight and the number of chambers in the last
whorl, the average pore area and the porosity are negative and much
lower (−0.43, −0.31 and −0.12, respectively). When the original
weights of the species are plotted vs. average size and wall thickness it
is shown that the larger species are the heavier ones. In most cases,
these are also the ones with the thicker test walls of the last chambers
(Fig. 8A–B).
A regression analysis for 15 species was carried out (Table 6) in
dependent variable, while the shell wall thickness and the mean
size were considered as independent variables. The results indicate
a coefficient of 0.234 between the weight and the shell wall
thickness and of 0.036 between the weight and the size. Based on
these values, the regression for the weight of species and their shell
wall thickness and size can be formulated as below. R 2 for this
regression analysis is very high (0.923), indicating that more than
Table 6
Multiple-regression results.
1. Dependent variable: original weight.

which the original weight of species was considered as a Independent variables: wall thickness, size.
N: 15
Multiple R: 0.966
Multiple R 2: 0.933
A 20 Multiple R 2 adjusted: 0.923
ANOVA:
F: 83.986
15 p: 8.788E-08
Original weight (µg)

Coeff. Std. err. t p

10 Constant −4.449 1.226 −3.630 0.003

Wall thickness 0.234 0.049 4.786 0.000
Size 0.036 0.004 9.659 0.000
5 y = 0.0419x - 3.0491 Weight= −4.449 +(0.234 wall thickness)+(0.036 size)
R2 = 0.8075
2. Dependent variable: CRPS.
0 Independent variables: wall thickness, size
100 200 300 400 500
N: 12
Average size of species (µm)
Multiple R: 0.927
Original weight Multiple R 2: 0.860
Multiple R 2 adjusted: 0.828
Linear (Original weight)
ANOVA
F: 27.436
B p: 1.480E-04
20
Coeff. Std.err. t p

Constant −2.609 1.742 −1.497 0.169

15
Wall thickness 0.294 0.058 5.088 0.001
Original weight (µg)

Size 0.018 0.005 3.324 0.009

CRPS = −2.609 +(0.294 wall thickness) +(0.018 size)
10
3. Dependent variable: CRRW.

y = 0.3991x + 4.8851 Independent variables: wall thickness, size

5
N: 15
R2 = 0.4146
Multiple R: 0.954
Multiple R2: 0.910
0
0 5 10 15 20 25 30 35 Multiple R 2 adjusted: 0.895
Shell wall thickness (µm) ANOVA
F: 60.259
Original weight
p: 5.514E-07
Linear (Original weight)
Coeff. Std. err. t p
Fig. 8. The relationship between the original weight of species and their average size Constant 1.513 0.644 2.348 0.037
(A), and wall thickness (B). R2 value is very high in A and much lower in B, Wall thickness 0.227 0.026 8.807 0.000
demonstrating that a larger part of the variation in the weight of species is explained by Size 0.006 0.002 3.120 0.009
the linear relation with their size and a smaller part of this variation is explained by the CRRW = 1.513 +(0.227 wall thickness)+(0.006 size).
linear relation with wall thickness.
14 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21

A 40
B 40

Shell wall thickness(µm)

Shell wall thickness (µm)

30 30

20 20

10 y = 1.891x + 0.8423 10
y = 3.2854x - 7.7508
2 2
R = 0.6717 R = 0.8361

0 0
2 4 6 8 10 12 14 16 18 4 6 8 10 12
CRPS CRRW
Mean wall thickness Mean wall thickness

Standard deviation Standard deviation

Linear (mean wall thickness) Linear (mean wall thickness)

C D 20
20

15 15

Original weight (µg)

Original weight (µg)

10 10

y = 1.0393x + 2.8589 y = 1.7945x - 1.5467

2 5 2
5 R = 0.7532 R = 0.6493

0 0
2 4 6 8 10 12 14 16 18 4 6 8 10 12
CRPS CRRW

E F 600
600

500 500
Size of species (µm)

Size of species (µm)

400 400

300 300

200 200
y = 16.939x + 200.83
2 y = 27,269x + 143,1
100 R = 0.4561 100
2
R = 0,3262
0 0
2 4 6 8 10 12 14 16 4 6 8 10 12

CRPS CRRW
Mean size Mean size

Standard deviation Standard deviation

Linear (mean size) Linear (mean size)

Fig. 9. Cumulative Relative Preservation Scores (CRPS) and Cumulative Relative Remaining Weight (CRRW) plotted vs. wall thickness (figures A and B), initial weight (figures C and
D) and size (figures E and F). R2 values are very high in figures A, B, C, D and lower in figures E and F, indicating that a major part of variation in CRPS and CRRW is explained by the
linear relations with wall thickness and weight while a smaller part of the dissolution indicators is explained by the linear relations with size.

90% of the variation in the original weight of taxa is explained by show that CRPS and CRRW are closely correlated (0.90). Wall
this linear regression. thickness has a strong bearing on both CRPS and CRRW as indicated
by the high correlation values of 0.83 for CRPS and 0.95 for CRRW.
Indeed, the distinct linear relationship between wall thickness and
Weight of taxa = −4:449 + ð0:234mshell wall thicknessÞ both preservation scores is shown in Fig. 9A–B.
+ ð0:036msizeÞ The original weight of the taxa also shows a strong correlation
with both CRRW, CRPS (0.85 and 0.94, respectively). In all cases, the
heavy species are most dissolution resistant, as indicated in Fig. 9C–
3.5. Relationships between dissolution susceptibility and shell parameters D. In comparison with wall thickness and original weight, the
average size of taxa expresses a slightly weaker relationship with
In order to examine the relationship between dissolution CRRW and CRPS (0.60 and 0.76 respectively). Even though,
susceptibility and shell parameters, we analyzed the correlation smaller species in most cases are more susceptible to dissolution
between CRPS, CRRW and the shell parameters (Table 5). Results than the larger species (Fig. 9E–F).
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 15

A B 30
30

y = -0.2959x + 15.551 y = -0.3797x + 15.816

25 25
2
R = 0.0691

Porosity (% pore)
2
R= 0.0493
Porosity (% pore)

20 20

15 15

10 10

5 5

0 0
2 4 6 8 10 12 14 16 4 6 8 10 12
CRPS CRRW

Mean porosity Mean porosity

Standard deviation Standard deviation

Linear (mean porosity) Linear (mean porosity)

C D 10
10
y = -0.0826x + 4.8789 y = -0.0684x + 4.7184
2
2
R = 0.0337 8 R = 0.0088

Pores size (µm2)

8
Pores size (µm2)

6 6

4 4

2 2

0 0
2 4 6 8 10 12 14 16 4 6 8 10 12

CRPS CRRW
Mean pores size Mean pores size

Standard deviation Standard deviation

Linear (mean pores size Linear (mean pores size)

E 8
F
8
y = -0,2744x + 7,272
2
R = 0.2113
6 6
Number of chambers
Number of chambers

4 4

2 y = -0.1614x + 6.5061 2
2
R = 0.1793

0 0
2 4 6 8 10 12 14 16 4 6 8 10 12
CRPS CRRW
Mean number of chambers Mean number of chambers

Standard deviation Standard deviation

Linear (mean number of chambers) Linear (mean number of chambers)

Fig. 10. Dissolution indicators (CRPS and CRRW) vs. porosity (figures A and B), pore size (figures C and D) and number of chambers in the final whorl (figures E and F). At all graphs,
regression lines are nearly parallel to X-axis and R2 values are very low, demonstrating that there are no general relations between the dissolution susceptibility of species and their
porosity, pore size or number of chambers.

The number of chambers in the final whorl presents a weak,
negative correlation with the CRPS and the CRRW (−0.41 and −0.46,
respectively). Similarly, test porosity and pore size indicate a very
weak and negative relationship, yet with high un-correlation values
for the CRPS and the CRRW (upper part of Table 5). There are no
significant differences in these shell parameters between resistant
species and vulnerable species, as indicated in Fig. 10A–F.
To quantify the effects of shell parameters on the dissolution
susceptibility of taxa, as well as to describe the relationship between
them, we carried out multi-regression analyses, in which the CRPS
and the CRRW are considered as dependent variables. Wall thickness
and average size are the largest controls on the CRPS and CRRW and
are considered as independent variables. We do not take the original
weight into this analysis since this parameter is derived from the
combination of wall thickness and size, as mentioned above. The
results show that 83% of CRPS and 90% CRRW are explained by the
combination between these two structural parameters. Regression
coefficient values between the shell wall thickness and the CRPS,
CRRW are 0.294 and 0.227, respectively. The corresponding values
between the mean size and the solution preservation indicators are
16 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
12
10
CRRW
8 and their weight is in good agreement with in-situ studies on
y = 0.5958x + 2.4045
R2 = 0.8879
6
4 around the Tethys (e.g. Canudo et al., 1995; Arenillas and Molina,
2 4 6 8 10 12 14 16
CRPS
CRRW
Linear (CRRW)
Fig. 11. Cumulative Relative Remaining Weight (CRRW) plotted vs. Cumulative Relative
Preservation Score (CRPS). R 2 value is very high, indicating a high agreement in
dissolution ranking for species within the two experiments.
0.018 and 0.006 (Table 6). Linear regression formulas between the
CRPS, CRRW and these shell parameters can be expressed as follows:
a/ CRPS= −2.609 +(0.294 wall thickness)+(0.018 size)
b/ CRRW= 1.513 +(0.227 wall thickness)+(0.006 size)
4. Discussion
4.1. Relative ranking of dissolution susceptibility
We have constructed a dissolution ranking for some common late
Paleocene to early Eocene planktic foraminiferal species. Similarity in
the ranking schemes derived from both experiments, shown by the
high correlation and R2 values between CRPS and CRRW (0.90, Table 5
and 0.89, Fig. 11), highlights the reliability of our proposed scheme.
Accordingly, the large and muricate Acarinina is most robust, followed
by Morozovella. The cancellate Subbotina is intermediate in dissolution
susceptibility and the small muricate Igorina is the most dissolution
vulnerable genus.
At species level, in both experiments, Acarinina species show a
consistent ranking: in ascending order of dissolution susceptibility these
are: A. soldadoensis, A. subsphaerica and A. nitida. For Igorina, both results
show similar susceptibility for I. pusilla and I. tadjikistanensis. Although
I. albeari and I. broedermanni were not assessed in the first experiment,
their similar CRRW values indicate that there is a little difference in
dissolution susceptibility between the Igorina species.
Within Morozovella, we observed an overall similar dissolution
ranking in both experiments. Accordingly, M. subbotinae and M. aequa
are the most resistant species. Morozovella formosa-gracilis was not
examined in the shell destruction experiment, nevertheless the
resemblance in CRRW between this species, M. pasionensis and M.
velascoensis suggests that these three species have the same interme-
diate dissolution robustness. The two smaller taxa, M. acuta and M.
occlusa show higher dissolution susceptibility in both assessments.
Among the species combined, the thick-shelled A. soldadoensis, A.
subsphaerica and the strongly muricate M. subbotinae indicate high
dissolution robustness. All the other large Morozovella species,
together with A. nitida show intermediate dissolution susceptibility.
The cancellate S. velascoensis, the small muricate Igorina species and
the two small, thin-walled M. acuta and M. occlusa reveal high
dissolution vulnerability in both assessments.
4.2. Relationship between dissolution resistance and weight
Our results show that lighter species are dissolved more rapidly
than heavier ones (Fig. 9C–D and Table 4). As heavier species
consist
of more calcite, it could be expected that dissolution media need a
longer time to break down the shells. This demonstrates that
progressive dissolution results in the relative enrichment of heavier
species (and specimens) within the assemblage.
The observed relationship between dissolution resistance of taxa
selective dissolution of modern planktic foraminifera (Berger,
1967). It is also a reasonable explanation for the relative enrichment
of Acarinina and Morozovella in partially dissolved Paleogene
planktic foraminiferal assemblages in records from various locations
18
1996; Berggren and Ouda, 2003; Luciani et al., 2007) and ODP sites
(e.g. Basov, 1995; Kelly, 2002; Lyle et al., 2002). The direct
relationship between the weight of taxa and their dissolution
susceptibility in our controlled laboratory environment is clear. In
natural environments, in which the time that taxa are exposed to
dissolution agents is not controlled, it can be expected that this
connection could even be stronger. Generally, dissolution of
foraminiferal tests is considered to be a taphonomic process (e.g.
Martin and Liddell, 1991), but shell loss from dissolution already
starts in the water column from partial resorption of the planktic
shells and is mediated through biodegradation at the sea floor (e.g.,
Ruddy, 1997). Because lighter shells (or taxa) generally descend
more slowly to the seafloor, they are exposed longer in the water
column and thus, are more prone to early dissolution (e.g. Martin
et al., 1995; Schiebel et al., 2007). This process provides a
taphonomic filter and leads to a relative increase of heavy speci-
mens and taxa in sediments.
4.3. Wall thickness and size dependency
Our experiments demonstrate that the dissolution susceptibility of
planktic foraminiferal species strongly depends on their wall thickness
(Table 5 and Fig. 9A–B), and that planktic assemblages becoming
relatively enriched in thick-walled individuals as dissolution progresses.
Within a species, larger specimens are often heavier and possess
thicker test walls and as a result, they are the more dissolution
resistant. This is not directly applicable between species, because
species can be larger, but also thinner than others. In this case, the
overwhelming role of shell wall thickness on the dissolution
resistance of taxa will be weakened and size can become a more
dominant control on dissolution susceptibility. For example the shells
of M. subbotinae and M. aequa are thinner than A. nitida but these two
Morozovella species are more dissolution resistant. The difference in
size between these three taxa is a reasonable explanation for this
discrepancy as the Morozovella species are generally larger than
A. nitida (Fig. 7A). This resulted in less original weight of A. nitida
(Table 4), and consequently, less dissolution resistance of A. nitida in
comparison with M. subbotinae and M. aequa.
The close link between wall thickness, size fraction and solution
susceptibility of taxa in our experiment is supported by previous
laboratory studies (Nguyen et al., 2009) as well as by in-situ studies,
based on both modern planktic (Berger, 1967; Berger, 1970; Conan
et al., 2002) and benthic (Corliss and Honjo, 1981) foraminiferal
assemblages. Our findings are also in line with quantitative distribu-
tion data from ancient sediments (e.g. Sliter, 1995) and modern
deposits (e.g., Sliter, 1975; Conan et al., 2002). In those records, thin-
walled and small planktic and benthic foraminifera were documented
as more susceptible to carbonate dissolution than other thick-walled
and larger taxa.
In addition, the size and shell-thickness of species can vary in time
and space, depending on the surrounding environmental parameters,
such as water depth, chemical composition of water column [CO3−2],
temperature and light, … (Bé and Hemleben, 1970; Berger, 1971;
Anderson, 1975; Schiebel et al., 2007). This variation leads to
differences in the weight of species between different stratigraphic
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21 17

intervals and locations. The variable original weights of taxa used in
our experiments confirmed this for Paleogene taxa. In most cases,
specimens of taxa from Site 865 are lighter than specimens from the
Shatsky Rise sites. This results in intraspecific dissolution susceptibil-
ity, as observed in our experiments (Fig. 5) as well as in in-situ studies
of Berger (1967).
4.4. Independency of porosity and number of chambers
Theoretically, higher porosity, larger pore size and low number of
chambers result in lower calcite content of the shell, more surface area
subject to dissolution, and thus should reflect low dissolution
resistance. In the modern ocean, the more dissolution-prone planktic
foraminiferal species are generally small and have thin walls and large
pores, whereas the less solution-susceptible species are bigger and
have thick walls and small pores (Sliter et al., 1975). However, in our
experiments, the impact of number of chambers in the last whorl,
shell porosity and pore size on the dissolution resistance of taxa is low
(Table 5). This can be expected, as the interrelationships among shell
parameters are complex, and their roles in dissolution resistance of
taxa may vary widely, depending on the taxa and the local conditions
(e.g. Hemleben and Bé, 1979; Herrero and Canales, 2002). Based on
our observations, we believe that porosity, pore size and number of
chambers in the last whorl, however, do play a role in dissolution
susceptibility of S. velascoensis. This species is quite large and average
wall thickness is intermediate in comparison with other taxa
(Appendices B and C). In addition, the chambers are highly inflated
and increase rapidly in size leading to only 3.5 chambers in the last
whorl. This set of parameters leads to a lower calcite content in the
shells of S. velascoensis compared to the other taxa studied (Appendix
A), and a relatively low dissolution resistance.
4.5. Effects of dissolution on the composition of planktic foraminiferal
assemblages
The early Paleogene deep-sea record provides evidence of episodic
dissolution events in pelagic carbonate sequences (Röhl et al., 2004;
Colosimo et al., 2006), presumably resulting from lysocline and
Carbonate Compensation Depth (CCD) shoaling. These events are
coincident with postulated hyperthermal events documented world-
wide. Examples are the Early Late Paleocene Event (ELPE; Röhl et al.,
2004; Petrizzo, 2005), Paleocene–Eocene Thermal Maximum
(PETM;
e. g. Bralower et al., 1995; Zachos et al., 2001, 2003, 2005), Eocene
Thermal Maximum 2 (ETM-2; Shipboard Scientific Party, 2003;
Lourens et al., 2005), and X-event or Eocene Thermal Maximum 3
(ETM-3; Kroon et al., 2007; Agnini et al., 2009). The upper Paleocene–
lower Eocene foraminiferal assemblages often show evidence of
dissolution at different intensity levels and in different locations from
low–mid latitudes (see e.g. Basov, 1995; Urquhart, 2001; Huber, 1991;
Kelly, 2002; Kelly et al., 2005; Colosimo et al., 2006; Kaiho et al., 2006;
Petrizzo, 2007). In these assemblages, consistent with partial
dissolution, Acarinina species (e.g. A. soldadoensis, A. mckannai,
A. esnaensis) and robust Morozovella species (e.g. M. aragonensis,
M. aequa, M. subbotinae) are often the main components, while fragile
taxa such as Subbotina spp., Igorina spp., Globanomalina spp.,
Chiloguembelina spp. as well as some vulnerable Morozovella species
(M. acuta, M. occlusa, M. edgari) are rare or absent. Although in some
cases this faunal composition may be accounted for by a primary
ecological control on assemblages such as the anomalously high
abundance in the PETM interval of large M. velascoensis at
Shatsky Rise (Petrizzo, 2007) or of robust Acarinina in the Weddell
Sea (ODP Site 690; Kelly, 2002), our results indicate that the
interpretation of such assemblages requires careful scrutiny of the
preservation state.
Previous studies on modern foraminifera document that deep-
dwelling taxa were generally more resistant to dissolution than
surface-dwelling taxa (Berger, 1968; Coulbourn et al., 1980). By
analogy, the surface-dwelling Paleogene genera Morozovella and
Acarinina have long been assumed to be more susceptible to
dissolution, compared to the deep-dwelling Subbotina (Boersma and
Premoli-Silva, 1983; Berggren and Norris, 1997) and this is contrary to
our findings. However, observations on modern assemblages also
indicate that the more dissolution-prone planktonic foraminiferal
species are generally small and have thin walls, whereas the less
solution-susceptible species are bigger and have thick walls (Sliter et
al., 1975). This is in agreement with our results on the role of shell size
and wall thickness on dissolution susceptibility of taxa. Conversely,
the observed increase in relative abundance of Subbotina in some
intervals that had experienced dissolution in Paleogene (e.g., Bernaola
et al., 2007; Guasti and Speijer, 2007) is rather indicative of an initial
strong dominance of this taxon than an indicator of dissolution
(Nguyen et al., 2009).
Moreover, some observations of severely dissolved planktic
foraminiferal assemblages during the onset of the ELPE at Shatsky
Rise (Shipboard Scientific Party, 2002; Petrizzo, 2005) seem to
contradict our empirically derived species dissolution susceptibilities.
In these studies, I. tadjikistanensis and I. pusilla are inferred as
dissolution resistant species and conversely, I. albeari, morozovellids,
acarininids, globanomalinids and subbotinids are interpreted as the
more dissolution-susceptible taxa. However, Petrizzo (2005) also

Table 7
Comparison between the measured and the calculated weight, CRPS and CRRW using formulas in Sections 3.4 and 3.5.

Measured Calculated

Wall thickness (μm) Size (μm) Weight (μg) CRPS CRRW Weight (μg) CRPS CRRW

A. nitida 24.0 285 11.3 7.5 7.7 11.3 9.6 8.7

A. soldadoensis 30.9 405 16.5 14.6 11.3 17.2 13.8 11.0
A. subsphaerica 24.2 338 13.4 11.4 9.3 13.3 10.6 9.1
I. albeari 11.7 253 7.1 5.6 7.3 5.4 5.7
I. broedermanni 9.8 217 5.5 5.3 5.6 4.2 5.0
I. pusilla 9.9 202 6.3 4.3 5.4 5.0 3.9 5.0
I. tadjikistanensis 10.2 199 5.7 4.2 5.4 5.0 4.0 5.0
M. acuta 9.2 323 8.6 5.9 5.2 9.2 5.9 5.6
M. aequa 19.1 380 14.7 8.4 7.9 13.6 9.8 8.1
M. formosa-gracilis 13.8 481 17.1 7.1 15.9 10.1 7.6
M. occlusa 9.4 298 7.6 6.6 4.7 8.4 5.5 5.4
M. pasionensis 10.5 373 11.4 6.8 6.7 11.3 7.2 6.2
M. subbotinae 16.5 428 15.2 11.7 9.0 14.7 9.9 7.8
M. velascoensis 12.6 440 14.6 7.8 6.9 14.2 9.0 7.0
S. velascoensis 12.2 353 7.9 6.0 6.0 11.0 7.3 6.4
18 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
A 2002). A complex interaction between these factors results in
20
15
Weight (µg)
10
5 experimental results have a strong bearing on the interpretation of
Taxa name
Measured weight
Calculated weight
B 12
8 We apply the formulas presented in Sections 3.4 and 3.5 to the
CRRW
4
0 species were used, therefore it is more representative for the
Taxa name
Measured CRRW
Calculated CRRW
Fig. 12. Measured data vs. calculated data on weight (A) and Cumulative Relative
Remaining Weight (B). Species are arranged from left to right in decreasing order of
measured wall thickness. The difference between the measured data and the calculated
data is generally small, pointing to the robustness and reliability of our regression
formulas on the weight and the dissolution ranking schemes of species.
observed that the ELPE assemblages are made up mainly of heavily
overgrown igorinids whereas the other genera are almost absent in
the assemblages. Outside the ELPE, igorinids are less common and
show no shell thickening. The unusual dominance of thick-walled
igorinids within the ELPE might reflect their facility of adapting to
changes in water masses circulation (change in deep water circulation
and/or variation in surface water productivity) and in carbonate
saturation (see Shipboard Scientific Party, 2002; Petrizzo, 2005). This
record indirectly emphasizes the importance of assessing the wall
thickness in the determination of the dissolution susceptibility of taxa,
as observed in our experiments.
4.6. Implications for dissolution estimates in natural environments
4.6.1. Experimental vs. natural dissolution
In natural environments, dissolution of foraminiferal assemblages is
controlled by two groups of factors: 1) factors concerning the shell of
taxa (surface/volume ratio, wall thickness, size, porosity, morphology),
and 2) factors concerning the surrounding environment prior to, during
and after deposition (physico-chemical parameters of sea-water,
interstitial water or ground water, biological productivity and activity,
sedimentation conditions; e.g. Conan et al., 2002; Herrero and Canales,
differences between natural dissolution and dissolution in a controlled
experimental setup, in which only shell parameters and exposure time
are considered. Yet, the close agreement between our experimental
results and in-situ experimental results (e.g. Berger, 1967, 1970) as well
as natural quantitative/qualitative records indicative of dissolution
(Sliter, 1975; Petrizzo et al., 2008) suggest that our experiments
accurately mimic a combination of natural processes, and thus the
foraminiferal dissolution in natural environments.
4.6.2. Weight and dissolution ranking scheme for planktic foraminiferal
assemblages
Since it is our overall aim to develop a dissolution index that can
easily be employed or adapted to any quantitative foraminiferal study,
we need parameters that satisfy the following requirements: 1) a
strict relationship with the differential dissolution susceptibility of
taxa, and 2) applicability in a routine procedure of quantitative
foraminiferal analysis. The close connection between dissolution
resistance of species and weight, wall thickness and size indicates that
these are prime parameters for the evaluation of dissolution in
foraminiferal assemblages.
measured shell wall thickness and size of early Paleogene species, to
see how the weight, CRPS and CRRW of species calculated from these
formulas fit with measured data. Despite the very good agreement
between CRPS and CRRW, we prefer to use the formula developed for
CRRW (3.5.b) rather than that for CRPS (3.5.a), because CRRW is
gained from the experiment in which all the specimens of each
composition of the assemblage than the CRPS. In these formulas,
only wall thickness and size are included. The reason for this selection
is that these parameters satisfy our two requirements listed above.
Even though the weight of species directly controls the dissolution
resistance of taxa, we do not include this parameter in the regression
formulas of CRPS and CRRW because the weight of taxa is directly
derived from the combination of wall thickness and size (Tables 5, 6
and Fig. 8). Consequently, if weight would be included in these
formulas, the role of wall thickness and size would be doubled.
Results from the application of the regression formulas 3.4 and 3.5.b
indicate good agreement between the measured and the calculated
weight and the CRRW (Table 7 and Fig. 12). However, a discrepancy is
observed for S. velascoensis, as the calculated weight is much higher
than the measured weight (Fig. 12A). This result is expected
considering the size and the wall thickness of S. velascoensis as
discussed in Section 4.4. The measured values for CRRW are in line
with the calculated values, except for M. subbotinae and A. nitida
(Fig. 12B). The calculated CRRW in M. subbotinae is about 15% lower
than the measured CRRW and vice versa in A. nitida. The average size of
A. nitida is 285 μm, only slightly bigger than the average size of the
smallest group (Igorina species) but it has a thick wall (~24 μm,
(Appendices B and C). These values in M. subbotinae are 428 μm and
16.5 μm, respectively. It seems with those two species, that the size and
the wall thickness affect the CRRW in a way that differs from the other
species and our regression formula could not accurately take this into
account. Our regression formula seems to provide an overestimation for
the role of wall thickness and an underestimation for size in these
particular examples.
The negative constants in formulas (see Sections 3.4 and 3.5.a)
demonstrate that our regression formulas can be applied only within
certain boundary conditions. At the condition that values in weight,
CRPS and CRRW are ≥0 and that wall thickness is at least 1 μm, we
find that our regression formulas are valid when the size of a species is
≥ 125 μm. This is also the size fraction that we used in our
experiments.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
The discrepancies between the measured and the calculated data
in weight and CRRW underline the fact that our formulas for
calculating weight and the dissolution ranking of taxa should be
employed only after understanding the shell parameters of taxa.
Overall, however, the excellent agreement between the calculated
and the measured values in weight and CRRW indicates that the
dissolution susceptibility ranking of foraminifera can be determined
using the experimentally developed regression formulas.
4.6.3. Application of dissolution ranking formula
So far, most of our knowledge on the dissolution susceptibility of
planktic foraminifera is restricted to recent taxa (e.g. Berger, 1967,
1970; Sliter et al., 1975). Understanding the dissolution resistance of
extinct planktic species in older, especially lithified sediments is
limited (Malmgren, 1987; Petrizzo et al., 2008). Our findings can be
applied to elucidate the taphonomic overprint related to partial
dissolution of Paleogene planktic foraminiferal assemblages. For
instance, common observations across the Paleocene/Eocene bound-
ary at ODP sites, such as increased relative abundance of species of
Acarinina and Morozovella, increased fragmentation of Subbotina
together with low P/B ratio, low foraminiferal numbers, and low
relative abundances of Subbotina species (e.g. Kelly, 2002; Kaiho et al.,
2006; Petrizzo, 2007) likely indicate partial dissolution. These findings
may be extrapolated to different environmental settings or time
intervals characterized by different taxa with similar test
characteristics.
In most quantitative foraminiferal studies, wall thickness and size
of the common taxa are not routinely determined. Yet, as these
parameters fulfill our requirements of easy collection and robust
statistical relationships, a dissolution susceptibility ranking for
benthic and planktic foraminiferal assemblages from different
stratigraphic intervals can be achieved in relatively limited time.
This enables a dissolution quantification based on changes in relative
abundance of dissolution resistant taxa (having thick-walled and
large size) in any quantitative foraminiferal study.
5. Conclusions
We propose a dissolution susceptibility ranking for common
Paleocene–Eocene planktic foraminiferal species, based on the
differential destruction and weight loss of these taxa through
dissolution. Among these species, the thick-walled A. soldadoensis
and A. subsphaerica and the large M. subbotinae are the most resistant
species. Most of the large Morozovella species such as M. aequa,
M. formosa-gracilis, M. velascoensis and M. pasionensis, together with
A. nitida have intermediate dissolution susceptibility. Small, muricate
Igorina species, together with the cancellate S. velascoensis and the
thin-walled M. acuta and M. occlusa are the most dissolution
vulnerable species.
We found a strict dependency of dissolution resistance on weight,
wall thickness and size. The agreement between our experimental
results and in-situ experimental results on recent foraminifera as well
as natural quantitative/qualitative records suggests that our experi-
ments well reflect natural dissolution processes. Consequently, our
results provide important insight into the impact of partial dissolution
on foraminiferal assemblages in natural environments, especially
during Paleogene hyperthermal events that are often associated with
dissolution phenomena.
A formula for assessing dissolution resistance of individual taxa is
proposed. Application of this formula shows a good agreement
between the calculated and the measured dissolution resistance,
indicating its robustness and reliability. With this approach, based on
the wall thickness and size, dissolution susceptibility assessments for
foraminiferal assemblages from different areas and stratigraphic
intervals can be achieved. A proper assessment of taphonomic
19
alteration by dissolution should be part of every paleoenvironmental
reconstruction based on quantitative foraminiferal records.
AcknowledgThents
The paper greatly benefited from constructive comments by two
anonymous reviewers and useful suggestions by the Editor Frans
Jorissen. This research used samples provided by the Ocean Drilling
Program (ODP) sponsored by the U. S. National Science Foundation
(NSF) and participating countries under management of Joint
Oceanographic Institutions (JOI), Inc. We thank the Research
Foundation — Flanders (FWO — G.0422.10) and the K.U.Leuven
Research Fund for financial support to R.P. Speijer. Financial support of
MIUR-PRIN 2007 to M.R. Petrizzo is acknowledged.
Appendix A. SuppleThentary data
Supplementary data to this article can be found online at doi:10.
1016/j.marmicro.2011.07.001.
References
Agnini, C., Macri, P., Backman, J., Brinkhuis, H., Fornaciari, E., Giusberti, L., Luciani, V., Rio,
D., Sluijs, A., Speranza, F., 2009. An early Eocene carbon cycle perturbation at
52.5 Ma in the Southern Alps: chronology and biotic response. Paleoceanography
24 (PA2209). doi:10.1029/2008PA001649.
Anderson, J.B., 1975. Factors controlling CaCO 3 dissolution in the Weddell Sea from
foraminiferal distribution patterns. Marine Geology 19 (5), 315–332.
Arenillas, I., Molina, E., 1996. Biostratigrafía y evolucíon de las asociaciones de
foraminiferos planctónicos del transito Paleoceno–Eoceno en Alamedilla (Cordilleras
Bétias). Revista Espanola de Micropaleontologica 18 (1), 75–96.
Basov, I.A., 1995. Paleogene planktonic foraminifer biostratigraphy of Sites 883 and 884,
Detroit Seamount (subartic Pacific). Proceedings of the Ocean Drilling Program,
Scientific Results, 145 Available from World Wide Web:http://www-odp.tamu.
edu/publications/145_SR/VOLUME/CHAPTERS/sr145_09.pdf.
Bé, A.W.H., 1968. Shell porosity of recent planktonic foraminifera as a climatic index.
Science 161 (3844), 881–884.
Bé, A.W.H., Hemleben, C., 1970. Calcification in a living planktic foraminifera
Globigerinoides sacculifer. Neues Jahrbuch fűr Geologie und Paleontologie, Abhan-
dlungen 134, 221–234.
Berger, W.H., 1967. Foraminifera ooze, solution at depth. Science 156 (3773), 383–385.
Berger, W.H., 1968. Planktonic foraminifera — selective solution and paleoclimatic
interpretation. Deep-Sea Research 15 (1), 31–43.
Berger, W.H., 1970. Planktonic foraminifera — selective solution and lysocline. Marine
Geology 8 (2), 111–138.
Berger, W.H., 1971. Sedimentation of planktonic foraminifera. Marine Geology 11 (5),
325–358.
Berger, W.H., 1973. Deep-sea carbonates: Pleistocene dissolution cycles. Journal of
Foraminiferal Research 3 (4), 187–195.
Berger, W.H., 1975. Deep Sea Carbonates Dissolution Profile from Foraminiferal
Preservation: Cushman Foundation for Foraminiferal Research, Special Publication,
13, pp. 82–86.
Berggren, W.A., Norris, R.D., 1997. Biostratigraphy, phylogeny and systematics of
Paleocene trochospiral planktic foraminifera. Micropaleontology 43 (Suppl. 1), 1–
116.
Berggren, W.A., Ouda, K., 2003. Upper Paleocene–lower Eocene planktonic
foraminif- eral biostratigraphy of the Dababiya section, Upper Nile Valley (Egypt).
In: Ouda, K., Aubry, M.-P. (Eds.), The Upper Paleocene–Lower Eocene of the
Upper Nile Valley: Part 1. Stratigraphy: Micropaleontology, 49, pp. 61–92. Suppl.
1.
Berggren, W.A., Pearson, P.N., 2005. A revised tropical to subtropical Paleogene
planktonic foraminiferal zonation. Journal of Foraminiferal Research 35 (4), 279–
298.
Bernaola, G., Baceta, J.I., Orue-Etxebarria, X., Alegret, L., Martin-Rubio, M., Arostegui, J.,
Dinares-Turell, J., 2007. Evidence of an abrupt environmental disruption during the
mid-Paleocene biotic event (Zumaia section, western Pyrenees). Geological Society
of America Bulletin 119 (7-8), 785–795.
Boersma, A., Premoli-Silva, I., 1983. Paleocene planktonic foraminiferal biogeography
and the paleoceanography of the Atlantic-Ocean. Micropaleontology 29 (4), 355–
381.
Boltovskoy, E., Totah, V.I., 1992. Preservation index and preservation potential of some
foraminiferal species. Journal of Foraminiferal Research 22 (3), 267–273.
Bralower, T.J., Zachos, J.C., Thomas, E., Parrow, M., Paull, C.K., Kelly, D.C., Premoli Silva, I.,
Sliter, W.V., Lohmann, K.C., 1995. Late Paleocene to Eocene paleoceanography of
the equatorial Pacific Ocean: stable isotopes recorded at Ocean Drilling Program
Site 865, Allison Guyot. Paleoceanography 10 (4), 841–865.
Bralower, T.J., Premoli Silva, I., Malone, M.J., 2006. Leg 198 synthesis: a remarkable 120-
m.y. record of climate and oceanography from Shatsky Rise, northwest Pacific
Ocean. In: Bralower, T.J., Premoli Silva, I., Malone, M.J. (Eds.), Proceedings of the
Ocean Drilling Program, Scientific results, 198. Available from World Wide Web
http://www-odp.tamu.edu/publications/198_SR/synth/synth.htm.
20 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
Brunner, C.A., 1992. Paleoenvironment and biostratigraphy of foraminifers at Sites 794,
795, 796 and 797 in the Japan Sea. Proceedings of the Ocean Drilling Program,
Scientific Results, 127/128, Part 1. Available from World Wide Web http://www-
dp.tamu.edu/publications/127128SR_1/VOLUME/CHAPTERS/sr127128_12.pdf.
Canudo, J.I., Keller, G., Molina, E., Ortiz, N., 1995. Planktic foraminiferal turnover and
δ13 C isotopes across the Paleocene–Eocene transition at Caravaca and Zumaya,
Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 114 (1), 75–100.
Chen, M.-T., Farrell, J., 1991. Planktonic foraminifer faunal variations in the
Northeastern Indian Ocean: a high resolution record of the past 800,000 years
from Site 758. Proceedings of the Ocean Drilling Program, Scientific Results, 121.
Available from World Wide Web http://www-odp.tamu.edu/publications/121_SR/
VOLUME/CHAPTERS/sr121_04.pdf.
Colosimo, A.B., Bralower, T.J., Zachos, J.C., 2006. Evidence for lysocline shoaling at the
Paleocene/Eocene Thermal Maximum on Shatsky Rise, northwest Pacific. Pro-
ceedings of the Ocean Drilling Program, Scientific Results, 198. Available from
World Wide Web http://www-odp.tamu.edu/publications/198_SR/VOLUME/
CHAPTERS/112.PDF.
Conan, S.M.H., Ivanova, E.M., Brummer, G.J.A., 2002. Quantifying carbonate dissolution
and calibration of foraminiferal dissolution indices in the Somali Basin. Marine
Geology 182 (3–4), 325–349.
Corliss, B.H., Honjo, S., 1981. Dissolution of deep-sea benthonic foraminifera.
Micropaleontology 27 (4), 356–378.
Coulbourn, W.T., Parker, F.L., Berger, W.H., 1980. Fauna and solution patterns of planktic
foraminifera in surface sediments of the North Pacific. Marine Micropaleontology 5,
329–399.
Cullen, J.L., Prell, W.L., 1984. Planktonic foraminifera of the Northern Indian Ocean —
distribution and preservation in surface sediments. Marine Micropaleontology 9
(1), 1–52.
Dupuis, C., Aubry, M.-P., Steurbaut, E., Berggren, W.A., Ouda, K., Magioncalda, R.,
Cramer, B.S., Kent, D.V., Speijer, R.P., Heilmann-Clausen, C., 2003. The Dababiya
Quarry section: lithostratigraphy, clay mineralogy, geochemistry and paleontology.
In: Ouda, K., Aubry, M.-P. (Eds.), The Upper Paleocene–Lower Eocene of the Upper
Nile Valley: Part 1. Stratigraphy: Micropaleontology, 49, pp. 41–59. Suppl. 1.
Ernst, S.R., Guasti, E., Dupuis, C., Speijer, R.P., 2006. Environmental perturbation in the
southern Tethys across the Paleocene/Eocene boundary (Dababiya, Egypt):
foraminiferal and clay mineral records. Marine Micropaleontology 60 (1), 89–
111.
Gibbons, J.D., 1985. Nonparametric Statistical Inference. Second Edition, Revised and
Expanded. Marcel Dekker, New York. 488 pp.
Guasti, E., Speijer, R.P., 2007. The Paleocene–Eocene thermal maximum in Egypt and
Jordan: an overview of the planktic foraminiferal record. In: Monechi, S., Coccioni,
R., Rampino, M. (Eds.), Large ecosystem perturbations: Causes and Consequences:
The Geological Society of America, Special Paper, 424, pp. 53–67.
Hammer, Ø., Harper, D.A.T., Ryan, P.D., Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001.
PAST: Paleontological Statistics Software Package for education and data analysis.
Palaeontologia Electronica 4 (1) 9pp. Available from World Wide Web http:// palaeo-
electronica.org/2001_1/past/issue1_01.htm.
Hancock, H.J.L., Dickens, G.R., 2005. Carbonate dissolution episodes in Paleocene and
Eocene sediment, Shatsky Rise, West-Central Pacific. Proceedings of the Ocean
Drilling Program, Scientific Results, 198. Available from World Wide Web http://
www-odp.tamu.edu/publications/198_SR/VOLUME/CHAPTERS/116.PDF.
Hay, W.W., DeConto, R., Wold, C.N., Wilson, K.M., Voigt, S., Schulz, M., Wold-Rossby, A.,
Dullo, W.C., Ronov, A.B., Balukhovsky, A.N., Soeding, E., 1999. Alternative global
Cretaceous paleogeography. In: Barrera, E., Johnson, C.C. (Eds.), The Evolution of
the Cretaceous Ocean/Climate System: Geological Society of America, Special
Paper, 332, pp. 1–47.
Hemleben, C., Bé, A.W.H., 1979. “Dissolution” effects induced by shell resorption during
gametogenesis in Hastigerina pelagica (d'Orbigny). Journal of Foraminiferal
Research 9 (2), 118–124.
Herman, Y., Bogyo, T.P., 1980. Quantitative paleoclimatology: micropaleontologic data
from deep sea cores. Marine Geology 36 (1–2), 23–34.
Herrero, C., Canales, M.L., 2002. Taphonomic processes in selected Lower and Middle
Jurassic foraminifera from the Iberian range and Basque–Cantabrian Basin (Spain).
Journal of Foraminiferal Research 32 (1), 22–42.
Huber, B.T., 1991. Paleogene and early Neogene planktonic foraminifer biostratigraphy
of Sites 738 and 744, Kerguelen plateau (Southern Indian ocean). Proceedings of
the Ocean Drilling Program, Scientific Results, 119 Available from World Wide
Web http://www-odp.tamu.edu/publications/119_SR/VOLUME/CHAPTERS/
sr119_24.pdf.
Iaccarino, S., Proto-Decima, F., 1990. Distribution patterns of Neogene benthic
foraminifers in Sites 717, 718 and 719 (Leg 116). Proceedings of the Ocean Drilling
Program, Scientific Results, 116. Available from World Wide Web http://www-
odp.tamu.edu/publications/116_SR/VOLUME/CHAPTERS/sr116_18.pdf.
Jenkins, D.G., Orr, W.N., 1971. Cenozoic planktonic foraminiferal zonation and the
problem of test dissolution. Micropaleontology 3 (3), 301–304.
Kaiho, K., Takeda, K., Petrizzo, M.R., Zachos, J.C., 2006. Anomalous shifts in tropical
Pacific planktonic and benthic foraminiferal test size during the Paleocene–Eocene
Thermal Maximum. Palaeogeography, Palaeoclimatology, Palaeoecology 237 (2–
4), 456–464.
Kelly, D.C., 2002. Response of Antarctic (ODP Site 690) planktonic foraminifera to the
Paleocene–Eocene Thermal Maximum: faunal evidence for ocean/climate change.
Paleoceanography 17 (4), 1071. doi:10.1029/2002PA000761.
Kelly, D.C., Zachos, J.C., Bralower, T.J., Schellenberg, S.A., 2005. Enhanced terrestrial
weathering/runoff and surface-ocean carbonate production during the recovery
stages of the Paleocene–Eocene Thermal Maximum. Paleoceanography 20.
doi:10.1029/2005PA001163.
Kroon, D., Zachos, J.C., Leg 208 Scientific Party, 2007. Leg 208 synthesis: Cenozoic
climate cycles and excursions. In: Kroon, D., Zachos, J.C., Richter, C. (Eds.),
Proceedings of the Ocean Drilling Program, Scientific Results, 208. Available from
World Wide Web http://www-odp.tamu.edu/publications/208_SR/synth/synth.
htm:1-55.
Le, J.N., Thunell, R.C., 1996. Modelling planktic foraminiferal assemblage changes and
application to sea surface temperature estimation in the western equatorial Pacific
Ocean. Marine Micropaleontology 28 (3–4), 211–229.
Lourens, L.J., Sluijs, A., Krun, D., Zachos, J.C., Thomas, E., Röhl, U., Bowles, J., Raffi, I., 2005.
Astronomical pacing of late Palaeocene to early Eocene global warming events.
Nature 435 (7045), 1083–1087.
Luciani, V., Giusberti, L., Agnini, C., Backman, J., Fornaciari, E., Rio, D., 2007. The
Paleocene–Eocene Thermal Maximum as recorded by Tethyan planktonic forami-
nifera in the Forada section (northern Italy). Marine Micropaleontology 64 (3–4),
189–214.
Lyle, M., Wilson, P.A., Janecek, T.R., 2002. Proceedings of the Ocean Drilling Program,
Initial Report, 199. Available from World Wide Web http://www-odp.tamu.edu/
publications/199_IR/VOLUME/CHAPTERS/IR199_01.PDF.
Malmgren, B.A., 1987. Differential dissolution of Upper Cretaceous planktonic
foraminifera from a temperate region of the South Atlantic Ocean. Marine
Micropaleontology 11 (4), 251–271.
Martin, R.E., Liddell, W.D., 1991. Taphonomy of foraminifera in modern carbonate
environments: implication for the formation of foraminiferal assemblages. In:
Donovan, S.K. (Ed.), Fossilization: The Processes of Taphonomy. Belhaven Press,
London, pp. 170–194.
Martin, R.E., Harris, M.S., Liddell, W.D., 1995. Taphonomy and time-averaging of
foraminiferal assemblages in Holocene tidal flat sediments, Bahia la Choya, Sonora,
Mexico (northern Gulf of California). Marine Micropaleontology 26 (1–4), 187–
206.
Melillo, A.J., 1988. Neogene planktonic foraminifer biostratigraphy, ODP Leg. 101,
Bahamas. Proceedings of the Ocean Drilling Program, Scientific Results, 101.
Available from World Wide Webhttp://www-odp.tamu.edu/publications/101_SR/
VOLUME/CHAPTERS/sr101_01.pdf.
Mix, A.C., 1989. Pleistocene paleoproductivity: evidence from organic carbon and
foraminiferal species. In: Berger, W.H., Smetacek, V.S., Wefer, G. (Eds.), Productivity
of the Ocean: Present and Past. Dahlem Workshop Reports: Life Sciences Research
Report, 44, pp. 313–340.
Nguyen, T.M.P., Petrizzo, M.R., Speijer, R.P., 2009. Experimental dissolution of a fossil
foraminiferal assemblage (Paleocene–Eocene Thermal Maximum, Dababiya,
Egypt): implications for paleoenvironmental reconstructions. Marine Micropale-
ontology 73 (3–4), 241–258.
Nicolas, R., 2009. JMicroVision: Image Analysis Toolbox for Measuring and Quantifying
Components of High-Definition Images. Version 1.2.7. [Online] Available from
World Wide Web http://www.jmicrovision.com.
Olsson, R.K., Hemleben, C., Berggren, W.A., Huber, B.T., 1999. Atlas of Paleocene
planktonic foraminifera. Smithsonian Contributions to Paleobiology 85 252pp..
Pearson, P.N., Olsson, R.K., Huber, B.T., Hemleben, C., Berggren, W.A., 2006. Atlas of
Eocene Planktonic Foraminifera: Cushman Foundation for Foraminiferal Research,
Special Publication, 41. 514 pp.
Peterson, L.C., Prell, W.L., 1985. Carbonate dissolution in recent sediments of the eastern
equatorial Indian Ocean: preservation patterns and carbonate loss above the
lysocline. Marine Geology 64 (3–4), 259–290.
Petrizzo, M.R., 2005. An early late Paleocene event on Shatsky Rise, northwest Pacific
Ocean (ODP Leg 198): evidence from planktonic foraminiferal assemblages. In:
Bralower, T.J., Premoli Silva, I., Malone, M.J. (Eds.), Proceedings of the Ocean Drilling
Program, Scientific Results, 198. Available from World Wide Web: http://www-
odp.tamu.edu/publications/198_SR/102/102.htm.
Petrizzo, M.R., 2007. The onset of the Paleocene–Eocene Thermal Maximum (PETM) at
Sites 1209 and 1210 (Shatsky Rise, Pacific Ocean) as recorded by planktonic
foraminifera. Marine Micropaleontology 63 (3–4), 187–200.
Petrizzo, M.R., Premoli Silva, I., Ferrari, P., 2005. Data report: Paleogene planktic
foraminifera biostratigraphy ODP Leg 198 Holes 1209A, 1210A and 1211A (Shatsky
Rise, Northwest Pacific Ocean). Proceedings of the Ocean Drilling Program,
Scientific Results, 198. Available from World Wide Web http://www-odp.tamu.
edu/publications/198_SR/VOLUME/CHAPTERS/110.PDF.
Petrizzo, M.R., Leoni, G., Speijer, R.P., De Bernardi, B., Felletti, F., 2008. Dissolution
susceptibility of some Paleogene planktonic foraminifera from ODP Site 1209
(Shatsky Rise, Pacific Ocean). Journal of Foraminiferal Research 38 (4), 357–371.
Premoli Silva, I., Spezzaferri, S., 1990. Paleogene planktonic foraminifer biostratigraphy
and paleoenvironmental remarks on Paleogene sediments from Indian Ocean Sites,
Leg 115. Proceedings of the Ocean Drilling Program, Scientific Results, 115.
Available from World Wide Web http://www-odp.tamu.edu/publications/115_SR/
VOLUME/CHAPTERS/sr115_19.pdf.
Röhl, U., Westerhold, T., Bralower, T.J., Petrizzo, M.R., Zachos, J.C., 2004. An early late
Paleocene global dissolution event and new constraints for an astronomically-
tuned early Paleogene time scale. ICP VIII, 8th International Conference on
Paleoceanography, Biarritz, France, 5–10 September, 2004.
Ruddy, G., 1997. An overview of carbon and sulfur cycling in marine sediments. In:
Jickells, T.D., Rae, J.E. (Eds.), Biogeochemistry of Intertidal Sediments. Cambridge
University Press, Cambridge, pp. 98–118.
Sager, W.W., Winterer, E.L., Firth, J.V., 1993. Initial Report, Proceedings of the Ocean
Drilling Program, Initial Reports, 143. Available from World Wide Web http:// www-
odp.tamu.edu/publications/143_IR/143ir.htm.
Sager, W.W., Kim, J., Klaus, A., Nakanishi, M., Khankishieva, L.M., 1999. Bathymetry of
Shatsky Rise, northwest Pacific Ocean: implications for ocean plateau development
at a triple junction. Journal of Geophysical Research 104, 7557–7576.
 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 1–21
Schiebel, R., Barker, S., Lendt, R., Thomas, H., Bollmann, J., 2007. Planktic foraminiferal
dissolution in the twilight zone. Deep-Sea Research II. Topical studies in
Oceanography 54 (5–7), 676–686.
Shipboard Scientific Party, 2002. Leg 198 summary. Initial report. In: Bralower, T.J.,
Premoli Silva, I., Malone, M.J. (Eds.), Proceedings of the Ocean Drilling Program,
198. Available from World Wide Web http://www-odp.tamu.edu/publications/
198_IR/VOLUME/CHAPTERS/IR198_01.PDF.
Shipboard Scientific Party, 2003. Leg 208 Preliminary Report: Ocean Drilling Program.
Available from World Wide Web http://www-odp.tamu.edu/publications/prelim/
208_prel/208PREL.PDF.
Sliter, W.V., 1975. Foraminiferal life and residue assemblages from Cretaceous slope
deposits. Geological Society of America Bulletin 86, 897–906.
Sliter, W.V., 1995. Cretaceous planktonic foraminifers from Sites 865, 866 and 869: a
synthesis of Cretaceous pelagic sedimentation in the Central Pacific Ocean Basin.
Proceedings of the Ocean Drilling Program, Scientific Results, 143. Available from
World Wide Web http://www-odp.tamu.edu/publications/143_SR/VOLUME/
CHAPTERS/sr143_02.pdf.
Sliter, W.V., Bé, A.W.H., Berger, W.H., 1975. Dissolution of Deep-Sea Carbonates:
Cushman Foundation for Foraminiferal Research, Special Publication, 13. 159pp.
Spiegler, D., Jansen, E., 1989. Planktonic foraminifer biostratigraphy of Norwegian sea
sediments. ODP Leg 104. Proceedings of the Ocean Drilling Program, Scientific
Results, 104. Available from World Wide Web http://www-odp.tamu.edu/
publications/104_SR/VOLUME/CHAPTERS/sr104_34.pdf.
Takeda, K., Kaiho, K., 2007. Faunal turnovers in central Pacific benthic foraminifera
during the Paleocene–Eocene Thermal Maximum. Palaeogeography, Palaeoclima-
tology, Palaeoecology 251 (2), 175–197.
Thunell, R.C., 1976. Calcium–carbonate dissolution history in late Quaternary deep-sea
sediments, Western Gulf of Mexico. Quaternary Research 6 (2), 281–297.
21
Urquhart, E., 2001. Campanian to Miocene planktonic foraminifers from the Iberia
abyssal plain. In: Beslier, M.O., Whitmarsh, R.B., Wallace, P.J., Girardeau, J. (Eds.),
Proceedings of the Ocean Drilling Program, Scientific Results, 173. Available from
World Wide Web http://www-odp.tamu.edu/publications/173_SR/VOLUME/
CHAPTERS/SR173_09.PDF.
van der Zwaan, G.J., Jorissen, F.J., de Stigter, H.C., 1990. The depth dependency of
planktonic/benthic foraminiferal ratios: constraints and applications. Marine
Geology 95 (1), 1–16.
Violanti, D., Premoli Silva, I., Cita, M.B., Kersey, D., Hsű, K.J., 1979. Quantitative
characterization of carbonate dissolution facies of the Atlantic Tertiary sediments.
An attempt. Rivista Italiana di Paleontologia e Stratigrafia 85 (2), 517–548.
Weaver, P.P.E., Raymo, M.E., 1989. Late Miocene to Holocene planktonic foraminifers
from the equatorial Atlantic, Leg 108. Proceedings of the Ocean Drilling Program,
Scientific Results, 108. Available from World Wide Web http://www-odp.tamu.
edu/publications/108_SR/VOLUME/CHAPTERS/sr108_05.pdf.
Zachos, J.C., Pagani, M., Sloan, L., Thomas, E., Billups, K., 2001. Trends, rhythms, and
aberrations in global climate 65 Ma to present. Science 292 (5517), 686.
Zachos, J.C., Wara, M.W., Bohaty, S., Delaney, M.L., Petrizzo, M.R., Brill, A., Bralower, T.J.,
Premoli Silva, I., 2003. A transient rise in tropical sea surface temperature during
the Paleocene–Eocene Thermal Maximum. Science 302 (5650), 1551–1554.
Zachos, J.C., Röhl, U., Schellenberg, S.A., Sluijs, A., Hodell, D.A., Kelly, D.C., Thomas, E.,
Nicolo, M., Raffi, I., Lourens, L.J., McCarren, H., Kroon, D., 2005. Rapid acidification of
the ocean during the Paleocene–Eocene Thermal Maximum. Science 308 (5728),
1611–1615.
Žarić, S., Donner, B., Fischer, G., Mulitza, S., Wefer, G., 2005. Sensitivity of planktic
foraminifera to sea surface temperature and export production as derived from
sediment trap data. Marine Micropaleontology 55 (1–2), 75–105.