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Marine Micropaleontology 81 (2011) 121

Contents lists available at ScienceDirect

Marine Micropaleontology
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / m a r m i c r o

Dissolution susceptibility of PaleoceneEocene planktic foraminifera: Implications for palaeoceanographic reconstructions


Thi Minh Phuong Nguyen a,, Maria Rose Petrizzo b, Peter Stassen a, Robert P. Speijer a
a b

Department of Earth and Environmental Sciences, K.U.Leuven, Celestijnenlaan 200E, 3001 Leuven, Belgium Universit degli Studi di Milano, Dipartimento di Scienze della Terra, Ardito Desio, via Mangiagalli 34, 20133 Milano, Italy

a r t i c l e

i n f o

a b s t r a c t
We investigated shell characteristics and differential dissolution susceptibility of planktic foraminiferal species derived from upper Paleocene and lower Eocene deep-sea sequences, Ocean Drilling Program (ODP) Site 865 (Allison Guyot) and Sites 1209B, 1210B and 1212A (Shatsky Rise) in the North Pacic Ocean. The purposes of this study are: 1) assessing the effects of differential dissolution on upper Paleocenelower Eocene planktic foraminiferal assemblages, at species level and within different biozones, to quantify dissolution susceptibility of genera and species; 2) investigating the differences in shell characteristics; 3) revealing the relationship between shell parameters and dissolution robustness of taxa, and 4) identifying the key shell parameter(s) inuencing the dissolution susceptibility of foraminiferal taxa. Two independent experiments were carried out, one focusing on gradual qualitative deterioration of taxa by dissolution and the other documenting the weight loss of taxa. Shell parameters such as wall thickness, porosity and pore size were determined through Scanning Electron Microscopy (SEM) and image analysis (JMicroVision). We found that the large muricate Acarinina and Morozovella are most resistant, followed by the cancellate Subbotina and the small muricate Igorina, conrming results of previous work. At species level, the thickwalled 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 formosagracilis, 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.

Article history: Received 11 January 2011 Received in revised form 24 June 2011 Accepted 8 July 2011 Available online xxxx Keywords: dissolution susceptibility planktic foraminifera North Pacic Ocean Drilling Program PaleoceneEocene paleoceanography paleoclimatology taphonomy

1. Introduction 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 oors. 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). 0377-8398/$ see front matter 2011 Elsevier B.V. All rights reserved. doi:10.1016/j.marmicro.2011.07.001

1980; ari et al., 2005), making them excellent paleoenvironmental recorders in Quaternary and deep-time studies (e.g. Berger, 1968; Mix, 1989). More specically, planktic foraminifera provide information 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. B, 1968; Berger, 1968; Herman and Bogyo, 1980). However, only a small proportion of all shells sinking to the sea oor are ultimately preserved in the sedimentary record, because of carbonate dissolution in the water column, at the sea oor and within the sediment. Dissolution of planktic foraminifera in the deep sea is a

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2 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

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 difcult 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 reect the biocoenosis and underlying environmental signals (e.g. van der Zwaan et al., 1990). Since differential susceptibility of planktic foraminifera to dissolution will inuence the composition of fossil assemblages, the reliability of paleoenvironmental reconstructions, based on planktic foraminiferal assemblages may be signicantly 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 specic 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 lithied 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-

blages from stratigraphic intervals that record remarkable environmental 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 signicantly, both these muricate, symbiontbearing, 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 Pacic 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, Rhl et al., 2004; Petrizzo, 2005) and the Eocene Thermal Maximum 2 (ETM 2 Shipboard Scientic Party, 2003). The selected sedimentary sequences contain various indications suggesting that partial dissolution prevailed in the Pacic Ocean during the PaleoceneEocene interval (Sager et al., 1993; Colosimo et al., 2006). This study is aimed at: 1) assessing the effects of differential dissolution on upper Paleocenelower 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) inuencing 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.

1212

Fig. 1. Late Paleocene paleogeography (Hay et al., 1999) and locations of ODP Sites 865 (Allison Guyot, equatorial Pacic Ocean), 1209, 1210 and 1212 (Shatsky Rise, northwestern Pacic Ocean).

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 Table 1 List of samples, taxa and size fractions used in the qualitative shell deterioration experiment (experiment 1). Sample Zone/ Taxa subzone name Size fraction (m) 125 250 330 400 490 N550 250 330 400 490 550 x x 6364 198-1209B23H-1, 6364 198-1209B23H-2, 101102 198-1209B23H-2, 101102 198-1209B23H-2, 101102 198-1210B19H-CC 198-1210B19H-CC 198-1210B19H-CC 198-1212A9H-CC 198-1212A9H-CC Table 1 (continued) Sample Zone/ Taxa subzone name Size fraction (m) 125 250 330 400 490 N550 250 330 400 490 550 x 3

P4c

143-865B11H-4, 4850 143-865B11H-4, 4850 143-865B11H-4, 4850 143-865B12H-1, 130132 143-865B12H-1, 130132 143-865B12H-1, 130132 143-865B12H-1, 130132 143-865B13H-1, 124125 143-865B13H-1, 124125 143-865B13H-1, 124125 143-865B13H-1, 124125 143-865B13H-2, 5052 143-865B13H-2, 5052 143-865B13H-2, 5052 143-865B13H-3, 5052 143-865B13H-3, 5052 143-865B13H-4, 5052 143-865B13H-4, 5052 143-865B13H-4, 5052 143-865B13H-4, 5052 143-865B14H4, 4749 143-865B14H-4, 4749 198-1209B23H-1, 6364 198-1209B23H-1,

P6a (E3) P6a (E3) P6a (E3) P5 (E2)

Acarinina subsphaerica Morozovella subbotinae Subbotina velascoensis Acarinina soldadoensis Morozovella occlusa Morozovella aequa Subbotina velascoensis Acarinina soldadoensis Igorina pusilla x x

Subbotina velascoensis Acarinina subsphaerica Morozovella occlusa Subbotina velascoensis Acarinina soldadoensis Morozovella subbotinae Subbotina velascoensis Acarinina soldadoensis Subbotina velascoensis x x

P4b

P4b

P4b

P5 (E2)

P5 (E2)

P5 (E2)

P6a (E3) P6a (E3) P6a (E3) P6b (E4) P6b (E4)

x x x x x x x

x x x x x

x x

P5

P5

Table 2 List of samples, taxa used in the weight loss experiment (experiment 2). x x Samples Zone/subzone Taxa name Size fraction (m) 125 330 x x x x x x x x x x x x 330 490 x x x x x x x

P5

Morozovella acuta Subbotina velascoensis Acarinina nitida Morozovella velascoensis Subbotina velascoensis Igorina pusilla x x x

P5

x 143-865B-11H-4, 4850 143-865B-11H-4, 4850 143-865B-11H-4, 4850 143-865B-12H-1, 130132 143-865B-12H-1, 130132 143-865B-12H-1, 130132 143-865B-12H-1, 130132 143-865B-13H-1, 124125 143-865B-13H-1, 124125 143-865B-13H-1, 124125 143-865B-13H-1, 124125 143-865B-13H-1, 124125 143-13H-2, 5052 143-13H-2, 5052 143-865B-13H-3, 5052 143-865B-13H-3, 5052 143-865B-13H-3, 5052 143-865B-13H-3, 5052 143-865B-13H-4, 5052 143-865B-13H-4, 5052 143-865B-13H-4, 5052 143-865B-13H-4, 5052 143-865B-13H-4, 5052 143-865B-13H-4, 5052 143-865B-14H4, 4749 143-865B-14H4, 4749 143-865B-14H4, 4749 143-865B-14H4, 4749 198-1209B-23H1, 6364 198-1212A-9H-CC 198-1212A-9H-CC 198-1212A-9H-CC 198-1212A-9H-CC 198-1212A-9H-CC 198-1212A-9H-CC 198-1210B-19H-CC 198-1210B-19H-CC 198-1210B-19H-CC 198-1210B-19H-CC 198-1210B-19H-CC P6a (E3) P6a (E3) P6a (E3) P5 (E2) P5 (E2) P5 (E2) P5 (E2) P5 P5 P5 P5 P5 P4c P4c P4c P4c P4c P4c P4c P4c P4c P4c P4c P4c P4a P4a P4a P4a P4c P6b (E4) P6b (E4) P6b (E4) P6b (E4) P6b (E4) P6b (E4) P6a (E3) P6a (E3) P6a (E3) P6a (E3) P6a (E3) Acarinina soldadoensis Morozovella subbotinae Subbotina velascoensis Acarinina soldadoensis Morozovella occlusa Morozovella aequa Subbotina velascoensis Acarinina nitida Igorina pusilla Igorina tadjikistanensis Morozovella acuta Subbotina velascoensis Acarinina nitida Morozovella velascoensis Acarinina nitida Igorina albeari Igorina tadjikistanensis Subbotina velascoensis Acarinina nitida Igorina pusilla Igorina albeari Igorina tadjikistanensis Morozovella acuta Subbotina velascoensis Igorina pusilla Igorina albeari Igorina tadjikistanensis Morozovella acuta Morozovella pasionensis Acarinina soldadoensis Acarinina subsphaerica Igorina broedermanni Morozovella subbotinae Morozovella formosagracilis Subbotina velascoensis Acarinina soldadoensis Acarinina subsphaerica Igorina broedermanni Morozovella subbotinae Subbotina velascoensis

P4c

P4c

P4c

P4c

P4c

Subbotina velascoensis Acarinina nitida Igorina pusilla x

P4c

P4c

P4c

Morozovella acuta Subbotina velascoensis x x

P4c

x x x x x x x x x x x x x x x x x x

x x x x x

x x

x x x x x x x x x x x x x x

P4a

Igorina x tadjikistanensis Igorina pusilla x

P4a

P4c

Acarinina subsphaerica Morozovella pasionensis

P4c

x x x x x

(continued on next part of this page)

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4 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

2. Material and methods 2.1. Material The planktic foraminiferal taxa used in the experiments were derived from ODP Leg 143, Hole 865B, Allison Guyot in the equatorial Pacic Ocean and from ODP Leg 198, Holes 1209B, 1210 and 1212A, Shatsky Rise in the North Pacic 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 identications (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 inlling. However, the sedimentary successions from which they were retrieved also show changes in fragmentation 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 13002000 m (Bralower et al., 1995; Sager et al., 1999; Shipboard Scientic Party, 2002). 2.2. Dissolution experiments To construct a dissolution susceptibility ranking scheme for planktic taxa, two independent experiments were carried out. The rst 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 rst soaked in water and washed through a 38 m 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). Wellpreserved empty specimens were picked from six different size fractions of 125250 m, 250330 m, 330400 m, 400490 m, 490550 m and N550 m. Where possible, 1020 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 disintegrated. Every 2 h, specimens were cleaned through a 38 m sieve using distilled water, dried, and their state of preservation was reassessed by binocular microscope. A foraminiferal preservation scale modied 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 = 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 = CRPS (size m)/nr. of size fractions, m = size fraction of 125250 m, 250330 m, 330400 m, 400490 m, 490 550 m and N550 m.

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 rst 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 125330 m and 330490 m. To assess the general fate of the species, all specimens encountered in the sub-samples (about 1000 specimens) were used in the experiment, without distinguishing 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 t0 is the time before taxa are exposed to acid treatment. CRRW = 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 = (CRRWy Ny)/ Ny, in which CRRWy indicates the CRRW of taxa in sample y and Ny is the number of specimens of taxa in sample y.

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Plate 1. SEM images illustrating the species concepts applied in this study. 1) Igorina pusilla, sample 865B-13H-1, 124125 cm. 2) Morozovella subbotinae, sample 1210B-19H-CC. 3) Igorina albeari, sample 1209B-22H-3, 111112 cm. 4) Igorina tadjikistanensis, sample 1209B-23H-2, 148149 cm. 5) Igorina broedermanni, sample 865B-8H-2, 5052 cm. 6) Morozovella velascoensis, sample 865B-13H-2, 5052 cm. 7) Morozovella aequa, sample 1209B-23H-2, 101102 cm. 8) Morozovella pasionensis, sample 1209B-23H-1, 6364 cm. a, umbilical view; b, lateral view; c, spiral view. Scale bars: 100 m.

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6 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

Plate 2. SEM images illustrating the species concepts applied in this study. 1) Morozovella occlusa, sample 1209B-23H-2, 101102 cm. 2) Acarinina nitida, sample 1209B-23H-2, 91 92 cm. 3) Acarinina subsphaerica, sample 1209B-23H-2, 101102 cm. 4) Acarinina soldadoensis, sample 865B-11H-4, 4850 cm. 5) Subbotina velascoensis, 1209B-23H-1, 6364 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 m.

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 7

Data viewer (Object extraction) Class Whole image (image surface %)

Class 1 54.45 17.17 758

Poligon 1(image surface %) Poligon 1 count

1 D measurement Parameter Value (m)

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). Measurements 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 signicantly 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 nonparametric 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. 3AD). 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. 3AC, 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, respectively, 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

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8 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

1 0.8

1 0.8

RPS

0.4 0.2 0

RPS

0.6

0.6 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

12

16

20

24

28

32

36

40

Exposure time (hours)


A. soldadoensis 400-490 m M. subbotinae 400-490 m S. velascoensis 400-490 m

Exposure time (hours)


A. soldadoensis 330-400 m M. occlusa 330-400 m M. aequa 330-400 m S. velascoensis 330-400 m

1 0.8

1 0.8

RPS
0 2 4 6 8 10 12 14 16 18 20

RPS

0.6 0.4 0.2 0

0.6 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)


A. nitida 125-250 m S. velascoensis 125-250 m I. pusilla 125-250 m

Exposure time (hours)


A.soldadoensis M.subbotinae S.velascoensis

E
1 0.8

RPS

0.6 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 Morozovella Igorina Subbotina

Fig. 3. AC: 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. Aspecies from sample 1210B-19H-CC (Shatsky Rise), size fraction of 400490 m; Bspecies from sample 865B-12H-1, 130132 cm (Allison Guyot), size fraction of 330400 m; Cspecies from sample 865B-13H-4, 5052 cm (Allison Guyot), size fraction of 125250 m; Ddifferential reduction in Relative Preservation Score between species from sample 1210B-19H-CC when all size fractions are combined together and Eidem 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). 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

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 3A3B). 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 125250 m and 250330 m as well as at the combined

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 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 (m) 125250 Acarinina species Taxa name A. nitida CRPS 4.0 Igorina species Taxa name I. pusilla I. tadjikistanensis I. pusilla I. tadjikistanensis CRPS 3.5 3.4 5.1 5.1 M. acuta M. pasionensis M. occlusa M. aequa M. subbotinae M. velascoensis M. acuta M. occlusa M. pasionensis M. aequa M. subbotinae M. velascoensis M. subbotinae M. velascoensis 4.6 5.7 5.7 8.0 8.9 6.6 7.1 7.4 8.0 8.7 10.3 7.7 15.8 9.0 Morozovella species Taxa name CRPS Subbotina species Taxa name S. velascoensis CRPS 3.4

250330

330400

A. nitida A. subsphaerica A. soldadoensis A. nitida A. subsphaerica A. soldadoensis

6.0 8.0 9.8 8.6 10.6 11.8

S. velascoensis

4.9

S. velascoensis

6.8

400490

A. nitida A. subsphaerica A. soldadoensis

11.5 15.4 16.1

S. velascoensis

8.7

490550

A. soldadoensis

20.6

N 550

size fraction (Table 3A3B) 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 signicant 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 3A3B). 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. Intraspecic differential dissolution between samples and locations. Our results also highlight intraspecic dissolution susceptibility 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, 130132 cm and 865B-13H-1, 124125 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 of genera and species ranked in decreasing order of dissolution resistance, based on the average CRPS from the start until the end of experiment 1. Note that the data presented here are a combination of different size fractions from different samples. Taxa name Acarinina Morozovella Subbotina Igorina Number of specimens Time (hours) taxa persist Average CRPS 303 279 284 90 84 78 36 22 84 56 56 42 32 34 30 34 36 26 22 22 11.5 8.8 6.0 4.3 14.6 11.4 11.7 7.5 8.4 7.8 6.8 6.6 6.0 5.9 4.3 4.2

A. soldadoensis 172 A. subsphaerica 61 M. subbotinae 97 A. nitida 70 M. aequa 30 M. velascoensis 52 M. pasionensis 30 M. occlusa 40 S. velascoensis 284 M. acuta 30 I. pusilla 60 I. tadjikistanensis 30

3.2.1. Average genus and species weights Our measurements show a considerable difference in average initial weight between genera and species. In the size fractions N125 m Acarinina is the heaviest genus, with an average weight of 13.6 g/specimen, followed by Morozovella (12.2 g), Subbotina (7.9 g) and Igorina (6.2 g) (Table 4). Among Acarinina, A. soldadoensis is the heaviest species (average weight 16.5 g), followed by A. subsphaerica (13.4 g) and A. nitida (11.3 g). Igorina albeari is the heaviest Igorina (7.1 g), followed by I. pusilla (6.3 g), I. tadjikistanensis (5.7 g) and I. broedermanni (5.5 g). The morozovellids can be divided into two groups: a heavy group, composed of the large species M. formosa-gracilis, M. subbotinae, M. aequa, M. velascoensis and M. pasionensis (average weights 11.417.1 g) and a second lighter group consisting of M. occlusa and M. acuta (7.68.6 g/specimen; Table 4 and Appendix A). 3.2.2. Weight loss by dissolution The weight loss experiments show that dissolution causes differential decreases in weight among genera (Fig. 6). The weight of Acarinina reached zero after 58 h of exposure while for Morozovella,

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10 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

A
1 0,8 0,6 0,4 0,2 0

susceptible (Table 4). In contrast, Morozovella species show large differences in weight loss. M. subbotinae was the most robust of the Morozovella species investigated, taking longest to dissolve and with highest RRW throughout the experiment. M. occlusa and M. acuta dissolve most rapidly and have the lowest RRW. Consequently,

RPS

A
0 10 20 30 40 50 60 70 80
A. nitida

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

90

Exposure time (hours)


A. soldadoensis A. subsphaerica

B
1 0,8 0,6 0,4 0,2 0 0 0 10 20 30 40 50

RPS

Exposure time (hours)


A. soldadoensis330-400 m, 198-1210B-19H-CC A. soldadoensis 330-400 m, 198-1212A-9H-CC A. soldadoensis 330-400 m, 143-865B-12H-1, 130-132 A. soldadoensis 330-400 m, 143-865B-13H-1, 124-125

RPS

8
I. pusilla

10

12

14

16

18

20

22

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

Exposure time (hours)


I. tadjkistanensis

1 0,8

RPS

0,6 0 0,4 0,2 0 0 4 8 12 16 20 24 28 32 36 40 44 48 52 56

RPS

Exposure time (hours)


M. subbotinae 490-550 m, 198-1210B-19H-CC M. subbotinae 490-550 m, 143-865B-11H-4, 48-50

10

20

30

40

50

60

M. subbotinae 400-490 m, 198-1210B-19H-CC M. subbotinae 400-490 m, 143-865B-11H-4, 48-50

Exposure time (hours)


M. acuta M. pasionensis M. aequa M. subbotinae M. occlusa M. velascoensis

C
1 0,8 0,6 0,4 0,2 0

Fig. 4. Differential reduction in the mean Relative Preservation Score with exposure 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.

Subbotina and Igorina it took only 36, 34 and 28 h, respectively. During the experimental process, the relative remaining weight (RRW) and the mean CRRW of Acarinina are highest, followed by those of Morozovella, Subbotina and Igorina, respectively (Fig. 6 and Table 4). As Acarinina has the highest original starting weight this nding is expected. Among Acarinina, the weight of A. nitida dropped to zero after just 36 h of the experiment (Fig. 6). The weight of A. subsphaerica and A. soldadoensis decreased more slowly and they fully fragmented after 44 h and 58 h, respectively. This results in the low mean CRRW of A. nitida in comparison with the other Acarinina species (Table 4). The Igorina species dissolved at similar rates (Fig. 6), resulting in nearly identical CRRW values (5.35.6), with I. albeari being slightly less

RPS

12

16

20

24

28

32

Exposure time (hours)


S. velascoensis 330-400 m, 198-1209B-23H-1, 63-64 S. velascoensis 330-400 m, 143-865B-13H-2, 50-52

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 susceptibility than the same taxa from Sites 1209, 1210 and 1212. AA. soldadoensis in size fraction of 330400 m from Zones E2-E3. BM. subbotinae in size fractions of 400 490 m and 490550 m from Zone E3 (subzone P6a). CS. velascoensis in size fraction of 330400 m from Subzone P4c.

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 Table 4 List of genera and species ranked in decreasing order of dissolution resistance, based on the mean CRRW during experiment 2. Taxon Acarinina Morozovella Subbotina Igorina A. soldadoensis A. subsphaerica M. subbotinae M. aequa A. nitida M. formosa-gracilis M. velascoensis M. pasionensis S. velascoensis I. albeari I. pusilla I. tadjikistanensis I. broedermanni M. acuta M. occlusa Number of specimens 2969 2696 2912 3263 884 1024 1097 68 1061 59 269 45 2912 1209 661 710 683 1000 158 Average original weight (g) 13.6 12.2 7.9 6.2 16.5 13.4 15.2 14.7 11.3 17.1 14.6 11.4 7.9 7.1 6.3 5.7 5.5 8.6 7.6 Time (hours) taxa persist 58 36 34 28 58 44 36 32 36 34 30 28 34 28 24 22 20 20 18 CRRW 9.4 7.0 6.0 5.5 11.3 9.3 9.0 7.9 7.7 7.1 6.9 6.7 6.0 5.6 5.4 5.4 5.3 5.2 4.7 11

genus Morozovella of which M. formosa-gracilis is the largest (average size 480 m), followed by M. velascoensis (440 m) and M. subbotinae (430 m). The mid-size group of taxa (300400 m) consists of the remaining Morozovella species, the Acarinina species and S. velascoensis. Igorina yields the smallest taxa, with average size ranges from 200 to 250 m. 3.3.2. Wall thickness Our measurements on wall thickness of the last normal chamber of each specimen reveal signicant differences between the taxa studied, both at genus and species level (Fig. 7B, Appendix C). Based on the average thickness, these taxa can be divided into two main groups: 1) the thick-walled group that exclusively consists of Acarinina species, of which A. soldadoensis has wall thickness of 31 m (as well as the larger variation); A. subsphaerica and A. nitida showing thickness of 24 m; 2) and a thin-walled group, of which large Morozovella species such as M. aequa, M. subbotinae, and M. formosa-gracilis yield an average wall thicknesses of 1419 m. The test walls of Igorina spp., S. velascoensis, and of the other morozovellids are slightly thinner (913 m). 3.3.3. Shell porosity and pore size Results indicate considerable dissimilarity in the proportion of porosity on the external test surface (% pores) and pore sizes between species (Fig. 7CD). S. velascoensis has the highest porosity (23%) and pore size (N7 m 2), but it also shows the largest variation. Acarinina species exhibit quite low and equable porosities, from 11% in A. subsphaerica to 12% in A. nitida. The pore sizes of this group are intermediate, which range around 5 m 2. The difference in porosity 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 m 2. Morozovellids yield porosities ranging from 9% (M. occlusa) to 16% (M. velascoensis) and have the smallest pores (34 m 2) (Fig. 7C and Appendices D and E).

M. occlusa and M. acuta have the lowest mean CRRW (4.7 and 5.2, respectively), followed by M. pasionensis, M. velascoensis, M. formosagracilis and M. aequa (6.7 to 7.9). M. subbotinae has the highest mean CRRW (9.0), being twice as robust as the most susceptible M. occlusa and M. acuta (Table 4 and Fig. 6). 3.3. Shell characteristics 3.3.1. Size Results on the variation in size (largest diameter) between species (Fig. 7A, Appendix B) show that most of the large species belong to the

ac uta I. b roe de rm an I. t ni ad jiki sta ne I. p ns us is illa I. a lbe ari S. ve las co en M. sis pa sio ne ns M. is ve las co en M. sis for mo sa -gr A. ac nit ilis ida

sp ec im oc en clu s sa

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50 100 0

aC

na

bb oti

tot

SU

M.

M.

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M.

Time (hours) taxa exposure in to acid treatment

0 4 8 12 16 20 24 28 32 36 40 44 48 52 56 60
0 50 100 0 750015000 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

A.

su bs p

Fig. 6. Differential weight loss among species by dissolution. Horizontal axis of the rst 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.

so lda d
50% lost of total CaCO3 and total number of specimens of each species

ae qu a

al C

of

su

50 100

oe

ns

O3

is

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12 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

A 600
500

40

Size of Species (m)

30 400 Shell wall thickness (m)

300

20

200

10

100

0
os agr M ac .v ilis el as co M . s ens is ub bo A. tin so ae ld ad oe ns is M .a M eq .p ua as io ne S. ns ve is la sc oe A. ns su is bs ph ae ric a M .a cu M . o ta cc lu sa A. ni tid a I. al I. be br ar oe i de rm an ni I. I. pu ta si dj lla ik is ta ne ns is

0
ae si s I. al be .p ar as i io I. ne ta ns dj ik is is ta ne ns is I. I. pu br si oe lla de rm an M ni .o cc lu sa M .a cu ta ilis ac ic a si s iti d ua tin si s en co el as S. v el as a en eq er A. n bo gr ha co en

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.a

sp

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A. s

A. s

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.f

Taxa name
Mean size of species Standard deviation

.f

Taxa name
Mean shell wall thickness Standard deviation

C
30 25

D 10
8

20 Porosity (% pore) 2 Pore area (m ) 6

15

10

0
ve la sc M oe .v ns el is as co en si s I. al be ar i M .a cu M ta .s ub I. bo ta dj tin ik ae is ta ne I. ns br oe is de rm M an .p ni as io ne ns is M I. .f pu or si m ll os a agr ac ilis A. ni tid A. a so ld ad A. oe su ns bs is ph ae ric a M .a eq ua M .o cc lu sa

0
ns is I. pu ad si jik lla is ta ne ns is I. al I. be br ar oe i de rm A. an su ni bs ph ae ric a A. ni M tid .s a ub bo A. tin so ae ld ad oe ns is M .a eq ua M .f M or .a m cu os ta agr M ac .v ilis el as co M en .p si as s io ne ns is M .o cc lu sa I. t as co e

S.

S.

ve l

Taxa name
Mean % pore Standard deviation Mean pore area Standard deviation

Taxa name

6 Number of chambers

0
is pu br si oe lla de rm an ni M .o cc lu sa I. al M be .v ar el i as I. co ta en dj si ik s is M ta .f ne or ns m is os agr A. ac su ilis bs ph ae ric a M .a cu ta A. ni tid M .s a ub bo A. tin so ae ld ad oe ns is M .a S. eq ua ve la sc oe ns is on en s

.p

as i

I.

I.

Taxa name
Mean number of chambers Standard deviation

Fig. 7. Shell characteristics among species: Athe size of species is measured as the largest diameter of individual specimens; Bthe wall thickness is measured on the last preserved chamber of specimens; Cshell porosity is the percentage of total pore area on the external surface of the specimens (% pore); Dthe 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 Ethe number of chambers in the last whorl.

3.3.4. Number of chambers in the last whorl M. pasionensis has the highest number (on average 7.7 chambers) and S. velascoensis the least (3.5 chambers) (Fig. 7E, Appendix F). Igorina species and M. occlusa, M. velascoensis, M. formosa-gracilis have more or less

the same number of chambers in the last whorl, ranging from 5.9 to 6.4. Acarinina species and M. acuta, M. subbotinae and M. aequa have 4 to 5.1 chambers in the last whorl. The largest variation in number of chambers is observed in M. velascoensis and M. occlusa, ranging from 5 to 8 chambers.

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 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 CRRW CRPS Wall thickness Original weight Size Porosity Pore area Number of chambers 0.90 0.95 0.85 0.60 0.21 0.13 0.46 CRPS 0.00 0.83 0.94 0.76 0.31 0.24 0.41 Wall thickness 0.00 0.00 0.74 0.47 0.28 0.19 0.52 Original weight 0.00 0.00 0.01 0.87 0.12 0.31 0.43 Size 0.04 0.00 0.12 0.00 0.20 0.38 0.33 Porosity 0.51 0.33 0.38 0.71 0.53 0.24 0.10 Pore area 0.68 0.46 0.56 0.33 0.22 0.44 0.35 Number of chambers 0.13 0.19 0.08 0.16 0.30 0.76 0.27

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. 8AB). A regression analysis for 15 species was carried out (Table 6) in which the original weight of species was considered as a

dependent variable, while the shell wall thickness and the mean size were considered as independent variables. The results indicate a coefcient 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. Independent variables: wall thickness, size. N: 15 Multiple R: 0.966 Multiple R2: 0.933 Multiple R2 adjusted: 0.923 ANOVA: F: 83.986 p: 8.788E-08 Coeff. Std. err. t p 0.003 0.000 0.000

A 20
15

Original weight (g)

10

y = 0.0419x - 3.0491 R2 = 0.8075

Constant 4.449 1.226 3.630 Wall thickness 0.234 0.049 4.786 Size 0.036 0.004 9.659 Weight = 4.449 + (0.234 wall thickness) + (0.036 size) 2. Dependent variable: CRPS. Independent variables: wall thickness, size N: 12 Multiple R: 0.927 Multiple R2: 0.860 Multiple R2 adjusted: 0.828 ANOVA F: 27.436 p: 1.480E-04 Coeff. Std.err. t 1.497 5.088 3.324

0 100

200

300

400

500

Average size of species (m)


Original weight Linear (Original weight)

B 20
15

p 0.169 0.001 0.009

Original weight (g)

Constant 2.609 1.742 Wall thickness 0.294 0.058 Size 0.018 0.005 CRPS = 2.609 + (0.294 wall thickness) + (0.018 size) 3. Dependent variable: CRRW.

10

y = 0.3991x + 4.8851 R2 = 0.4146

0
0 5 10 15 20 25 30 35

Shell wall thickness (m)


Original weight Linear (Original weight)

Independent variables: wall thickness, size N: 15 Multiple R: 0.954 Multiple R2: 0.910 Multiple R2 adjusted: 0.895 ANOVA F: 60.259 p: 5.514E-07 Coeff. Std. err. t 2.348 8.807 3.120 p 0.037 0.000 0.009

Fig. 8. The relationship between the original weight of species and their average size (A), and wall thickness (B). R2 value is very high in A and much lower in B, demonstrating that a larger part of the variation in the weight of species is explained by the linear relation with their size and a smaller part of this variation is explained by the linear relation with wall thickness.

Constant Wall thickness Size CRRW = 1.513 + (0.227

1.513 0.644 0.227 0.026 0.006 0.002 wall thickness) + (0.006 size).

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14 T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121

A
Shell wall thickness (m)

40

B
Shell wall thickness(m)

40

30

30

20

20

10

y = 1.891x + 0.8423 R 2 = 0.6717

10

y = 3.2854x - 7.7508 R 2 = 0.8361


12

0 2 4 6 8 10 12 14 16 18

0 4 6 8 10

CRPS
Mean wall thickness Standard deviation Linear (mean wall thickness) Mean wall thickness Standard deviation Linear (mean wall thickness)

CRRW

C
Original weight (g)

20

D
Original weight (g)

20

15

15

10

10

y = 1.0393x + 2.8589 2 R = 0.7532

y = 1.7945 - 1.5467 x 2 R = 0.6493

0 2 4 6 8 10 12 14 16 18

0
4 6 8 10 12

CRPS

CRRW

E
Size of species (m)

600

F 600
500

500

Size of species (m)

400

400

300

300

200

200

100

y = 16.939x + 200.83 2 R = 0.4561

100

y = 27,269x + 143,1 2 R = 0,3262


4 6 8 10 12

0
2 4 6 8 10 12 14 16

CRPS
Mean size Standard deviation Linear (mean size) Mean size Standard deviation Linear (mean size)

CRRW

Fig. 9. Cumulative Relative Preservation Scores (CRPS) and Cumulative Relative Remaining Weight (CRRW) plotted vs. wall thickness (gures A and B), initial weight (gures C and D) and size (gures E and F). R2 values are very high in gures A, B, C, D and lower in gures 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 this linear regression.

Weight of taxa = 4:449 + 0:234shell wall thickness + 0:036size 3.5. Relationships between dissolution susceptibility and shell parameters In order to examine the relationship between dissolution susceptibility and shell parameters, we analyzed the correlation between CRPS, CRRW and the shell parameters (Table 5). Results

show that CRPS and CRRW are closely correlated (0.90). Wall 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 both preservation scores is shown in Fig. 9AB. 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. 9CD. In comparison with wall thickness and original weight, the average size of taxa expresses a slightly weaker relationship with CRRW and CRPS (0.60 and 0.76 respectively). Even though, smaller species in most cases are more susceptible to dissolution than the larger species (Fig. 9EF).

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T.M.P. Nguyen et al. / Marine Micropaleontology 81 (2011) 121 15

A 30
25

B
y = -0.2959x + 15.551
Porosity (% pore)

30

R = 0.0691
Porosity (% pore)
20

25

y = -0.3797x + 15.816 R = 0.0493


2

20

15

15

10

10

0 2 4 6 8 10 12 14 16

0 4 6 8 10 12

CRPS
Mean porosity Standard deviation Linear (mean porosity) Mean porosity Standard deviation Linear (mean porosity)

CRRW

C
10

D
y = -0.0826x + 4.8789
8

10

y = -0.0684x + 4.7184
Pores size (m2)
8

R = 0.0337

R = 0.0088

Pores size (m )

0 2 4 6 8 10 12 14 16

0 4 6 8 10 12

CRPS
Mean pores size Standard deviation Linear (mean pores size Mean pores size Standard deviation Linear (mean pores size)

CRRW

E
Number of chambers

F
Number of chambers

y = -0,2744x + 7,272
6 6

2 R = 0.2113

y = -0.1614x + 6.5061 2 R = 0.1793


2 4 6 8 10 12 14 16

0
4 6 8 10 12

CRPS
Mean number of chambers Standard deviation Linear (mean number of chambers) Mean number of chambers Standard deviation

CRRW

Linear (mean number of chambers)

Fig. 10. Dissolution indicators (CRPS and CRRW) vs. porosity (gures A and B), pore size (gures C and D) and number of chambers in the nal whorl (gures 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 nal 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 signicant differences in these shell parameters between resistant species and vulnerable species, as indicated in Fig. 10AF. 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 coefcient 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

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y = 0.5958x + 2.4045 R2 = 0.8879

4 2 4 6 8 10 12 14 16 18

CRPS

CRRW Linear (CRRW)


Fig. 11. Cumulative Relative Remaining Weight (CRRW) plotted vs. Cumulative Relative Preservation Score (CRPS). R2 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

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 and their weight is in good agreement with in-situ studies on 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 around the Tethys (e.g. Canudo et al., 1995; Arenillas and Molina, 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 oor (e.g., Ruddy, 1997). Because lighter shells (or taxa) generally descend more slowly to the seaoor, 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 lter and leads to a relative increase of heavy specimens and taxa in sediments. 4.3. Wall thickness and size dependency

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 R 2 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 rst 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 intermediate 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. 9CD and Table 4). As heavier species consist

CRRW

Our experiments demonstrate that the dissolution susceptibility of planktic foraminiferal species strongly depends on their wall thickness (Table 5 and Fig. 9AB), 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 ndings are also in line with quantitative distribution data from ancient sediments (e.g. Sliter, 1995) and modern deposits (e.g., Sliter, 1975; Conan et al., 2002). In those records, thinwalled 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, 2 such as water depth, chemical composition of water column [CO3 ], temperature and light, (B and Hemleben, 1970; Berger, 1971; Anderson, 1975; Schiebel et al., 2007). This variation leads to differences in the weight of species between different stratigraphic

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intervals and locations. The variable original weights of taxa used in our experiments conrmed 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 intraspecic dissolution susceptibility, 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 reect 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 B, 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 inated 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 (Rhl 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 worldwide. Examples are the Early Late Paleocene Event (ELPE; Rhl et al., 2004; Petrizzo, 2005), PaleoceneEocene Thermal Maximum (PETM; e. g. Bralower et al., 1995; Zachos et al., 2001, 2003, 2005), Eocene Thermal Maximum 2 (ETM-2; Shipboard Scientic 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 lowmid 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 deepdwelling 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 ndings. 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 Scientic 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 Wall thickness (m) A. nitida A. soldadoensis A. subsphaerica I. albeari I. broedermanni I. pusilla I. tadjikistanensis M. acuta M. aequa M. formosa-gracilis M. occlusa M. pasionensis M. subbotinae M. velascoensis S. velascoensis 24.0 30.9 24.2 11.7 9.8 9.9 10.2 9.2 19.1 13.8 9.4 10.5 16.5 12.6 12.2 Size (m) 285 405 338 253 217 202 199 323 380 481 298 373 428 440 353 Weight (g) 11.3 16.5 13.4 7.1 5.5 6.3 5.7 8.6 14.7 17.1 7.6 11.4 15.2 14.6 7.9 CRPS 7.5 14.6 11.4 CRRW 7.7 11.3 9.3 5.6 5.3 5.4 5.4 5.2 7.9 7.1 4.7 6.7 9.0 6.9 6.0 Calculated Weight (g) 11.3 17.2 13.3 7.3 5.6 5.0 5.0 9.2 13.6 15.9 8.4 11.3 14.7 14.2 11.0 CRPS 9.6 13.8 10.6 5.4 4.2 3.9 4.0 5.9 9.8 10.1 5.5 7.2 9.9 9.0 7.3 CRRW 8.7 11.0 9.1 5.7 5.0 5.0 5.0 5.6 8.1 7.6 5.4 6.2 7.8 7.0 6.4

4.3 4.2 5.9 8.4 6.6 6.8 11.7 7.8 6.0

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2002). A complex interaction between these factors results in 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 experimental results have a strong bearing on the interpretation of foraminiferal dissolution in natural environments.

Weight (g)

0
ad A. oe su ns bs is ph ae ric A. a ni ti d a M .a eq M ua .s M ub .f bo or m t os ina e agr M ac .v ilis el as co S. en ve si s la sc oe ns is I. al be M .p ar i as io I. ne ta dj ns ik is is ta ne ns is I. pu I. br si lla oe de rm an ni M .o cc lu sa M .a cu ta

Taxa name

Measured weight Calculated weight

B 12

0
a ub bo m t os ina e agr M ac .v ilis el as co S. en ve si s la sc oe ns is I. al be M .p ar i as io I. ne ta dj ns ik is is ta ne ns is I. pu I. br si lla oe de rm an ni M .o cc lu sa M .a cu ta si s ic a tid a en ph a ni da do A. .a eq u er

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 reect 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 Scientic 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,

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. We apply the formulas presented in Sections 3.4 and 3.5 to the measured shell wall thickness and size of early Paleogene species, to see how the weight, CRPS and CRRW of species calculated from these formulas t 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 species were used, therefore it is more representative for the 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 m, only slightly bigger than the average size of the smallest group (Igorina species) but it has a thick wall (~24 m, (Appendices B and C). These values in M. subbotinae are 428 m and 16.5 m, 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 m, we nd that our regression formulas are valid when the size of a species is 125 m. This is also the size fraction that we used in our experiments.

CRRW

A.

so ld

su bs

so l

.s

A.

A.

M .f

or

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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 lithied sediments is limited (Malmgren, 1987; Petrizzo et al., 2008). Our ndings 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 boundary 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 ndings 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 fulll 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 quantication 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 PaleoceneEocene 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 experiments well reect 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

alteration by dissolution should be part of every paleoenvironmental reconstruction based on quantitative foraminiferal records. Acknowledgments The paper greatly beneted 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 nancial support to R.P. Speijer. Financial support of MIUR-PRIN 2007 to M.R. Petrizzo is acknowledged. Appendix A. Supplementary data Supplementary data to this article can be found online at doi:10. 1016/j.marmicro.2011.07.001. References
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