Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil

Quaternary International xxx (2016) 1e13

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Palaeoenvironmental interpretations based on molluscs from midHolocene lacustrine limestones, Mato Grosso do Sul, Brazil Giselle Utida a, *, Emiliano Castro Oliveira b, Maurice Tucker c, Setembrino Petri d, Paulo Cesar Boggiani d ~o em Geoquímica e Geotecto
ria, Sa ~o Paulo, SP CEP 05508
s-graduaça ^nica, Instituto de Geoci^ Programa de Po encias, Rua do Lago, 562, Cidade Universita ~o Paulo, Brazil 080, Universidade de Sa b ~o Paulo, Brazil Departamento de Ci^ encias do Mar, Instituto do Mar, Universidade Federal de Sa c School of Earth Sciences, University of Bristol, BS8 3RW, England, UK d
ria, Sa ~o Paulo, SP CEP 05508-080, Departamento de Geologia Sedimentar e Ambiental, Instituto de Geoci^ encias, Rua do Lago, 562, Cidade Universita ~o Paulo, Brazil Universidade de Sa a

a r t i c l e i n f o

a b s t r a c t

Article history: Received 30 November 2015 Received in revised form 1 November 2016 Accepted 6 November 2016 Available online xxx

In central Brazil there are continental carbonate rocks in three different geological contexts e the
city (Xaraie
s Formation), the mid-Holocene limestones lenses in Pleistocene wackestones in the Corumba the Pantanal Plain (Pantanal do Miranda) and old and modern tufas in the Serra da Bodoquena (Serra da Bodoquena Formation). We present here a systematic and taphonomic study of the molluscs found in the carbonate deposits in the Serra da Bodoquena and the limestones lenses of the Pantanal do Miranda, which provide information concerning the carbonate palaeoenvironments of these two distinct areas. Seven molluscan species were found: the gastropods Marisa sp., Pomacea canaliculata, an unidentified hydrobiid, Idiopyrgus sp., Biomphalaria sp. and Megalobulimus sp., and the bivalve Pisidium sp. The gastropods include species still living in the area today, which supports a Holocene age for the limestones, as indicated by published 14C dates. Overall, the data obtained from the Serra da Bodoquena carbonates suggest a freshwater environment with minimal current-wave action, in an area probably fed by springs, close to a terrestrial environment, in view of the assemblage composed of Pomacea canaliculata, the unidentified hydrobiid, Idiopyrgus sp., Biomphalaria sp. and Megalobulimus sp. The lacustrine palaeoenvironment in the Pantanal do Miranda area was different, as indicated by the presence of Marisa sp. and absence of shallow-water species such as Biomphalaria sp. The gastropods from Serra da Bodoquena and Pantanal do Miranda are predominantly aquatic and amphibious; they are probably younger than
area. the terrestrial gastropods reported from fine-grained limestones in the Corumba © 2016 Elsevier Ltd and INQUA. All rights reserved.

Keywords: Gastropods Taphonomy Tufa Lake Serra da Bodoquena Pantanal

1. Introduction Continental Quaternary carbonate deposits occur widely in state of Mato Grosso do Sul (Brazil) and were first described by Almeida
as the Xaraie
s Formation. Almeida (1945) (1945) from Corumba distinguished four categories of rocks: calcareous tufa with fossil plants, soft porous tufa, conglomerate with calcareous cement, and a micritic limestone with fossil gastropods. In the micritic limestone, Mendes (in Almeida, 1945) identified the land gastropods

* Corresponding author. E-mail addresses: [email protected] (G. Utida), [email protected] (E.C. Oliveira), [email protected] (M. Tucker), [email protected] (S. Petri), [email protected] (P.C. Boggiani).

Stenogyra (Opeas) misera, Zonitoides sp. and Bulimulus sp. (Binney and Bland, 1869; Oliveira and Almeida, 1999). Almeida (1945) suggested a Pleistocene or Pliocene age for these limestones, based on the gastropods, because they are living taxa. To the south in the Serra da Bodoquena, Almeida (1965) also
s Formation assigned Quaternary carbonate deposits to the Xaraie and later the Holocene age was confirmed (Turcq et al., 1987; Boggiani et al., 1998). Sallun Filho et al. (2009a) proposed a sepa
s Formation to a new one, ration of these deposits from the Xaraie the Serra da Bodoquena Formation, due to differences in lithological characteristics, the existence of clear and abrupt contacts, and the relatively easy mapping of the unit. Boggiani et al. (2002) discovered some gastropods in these deposits, Biomphalaria sp., Physa sp. and Aquidauania sp. These species are typically from

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Please cite this article in press as: Utida, G., et al., Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.11.007

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freshwater but they have an amphibious habit (Davis, 1979; Malek, 1983; Utzinger et al., 1997; Oliveira and Almeida, 1999; Utzinger and Tanner, 2000). Gastropods also are preserved in wetlands of Pantanal, in Pantanal do Miranda within lenticular carbonate deposits and in calcium carbonate layers in margins and bottoms of recent ponds of alkaline waters (Turcq et al., 1987; Assine and Soares, 2004). Turcq et al. (1987) described the lenticular carbonates of Pantanal do
s Formation, and bearing the Miranda as belonging to the Xaraie gastropods Pomacea sp. and Biomphalaria sp. (Boggiani and Coimbra, 1995; Boggiani et al., 1998). Utida et al. (2009, 2012)
s described freshwater micro- and macro-fossils from the Xaraie Formation in the Pantanal, and suggested a lacustrine palaeoenvironment for carbonate deposition. These fossiliferous micritic carbonates are widespread between the Serra da Bodoquena and Pantanal do Miranda. They contain a diversity of fossils but molluscs are found only in the upper part of the deposits where microfossils were not found (Utida et al., 2012). The presence of molluscs may indicate a different palaeoenvironment from that where the microfossils occur. We present here a taphonomic study of the molluscs found in
s Formation collected in Serra the carbonate deposits of the Xaraie da Bodoquena and the carbonate lenses of the Pantanal do Miranda, which provide information concerning the carbonate palaeoenvironments of these two distinct areas, Serra da Bodoquena and Pantanal do Miranda.

in these carbonates, and suggested that they were deposited on an alluvial plain. According to the recent calibration curve of Hogg et al. (2013), this age would recalculated to 5931 (±473 year cal BP). Sallun Filho et al. (2009b) confirmed the Holocene 14C ages for ~o Calc

s, with a range of 2850 to shells from the Mineraça ario Xarae 2720 cal yr BP (in the upper part of the deposit), to 6530 to 6310 cal yr BP (at a depth of 3 m into the deposit), but they interpreted the facies as lacustrine (Sallun Filho et al., 2009b; Utida, 2009). The Pantanal do Miranda is located just to the south of the extensive alluvial fan of the Taquari River, and limestones are exposed on the Park Road of Pantanal Sul, and elsewhere in the area. These rocks are mostly lenticular deposits, and they are generally quartz and fossil-rich wackestones, from 0.3 to 1.0 m thick and a lateral extension of 10e15 m, rarely up to 50 m (Fig. 3 c, d). These Pantanal do Miranda limestones were probably deposited in a lacustrine palaeoenvironment, when fluvial influence was minimal (Almeida, 1965; Boggiani et al., 1998; Ribeiro et al., 2001), during the mid-Holocene according to a14C date of 3910 (±100 years BP) (Boggiani et al., 1998), or 4255 (±279 years cal BP), according to the calibration curve of Hogg et al. (2013). They were deposited in localized areas upon a laterally-extensive unconsolidated fine quartz sand surface, probably in the lower reaches of an alluvial fan. The Pantanal limestones were probably deposited under a more arid climate, which existed during the period ~5600 to 2600 years BP (Mcglue et al., 2012).

2. Regional setting

3. Material and methods

The Serra da Bodoquena is on the southern margin of the Pantanal and is a range of hills 200 km in length and 400e800 m in height. The Quaternary carbonates overlie Neoproterozoic/Edia
Group (Fig. 1); the latter have caran carbonates of the Corumba been extensively dissolved by groundwater to create major cave systems. Emerging groundwater from the carbonates provides bicarbonate (and calcium) to the rivers and lakes of the area, where tufa and micrite are widely precipitated (Almeida, 1965; Boggiani and Coimbra, 1995; Sallun Filho, 2005; Sallun Filho and Karmann, 2007). The sedimentary characteristics of the studied deposits are different in terms of their geology and geomorphology. The Serra da Bodoquena is much affected by normal faults with a NNW-SSE direction in the northern part and NW-SE, NE-SW and NS directions in the south-central portion. This range is made up of Neoproterozoic carbonate basement rocks, occurring up to 600 m above sea level, with overlying occurrences of Quaternary continental carbonates. The Pantanal do Miranda is located in the Pantanal basin, with altitudes of 90 m approximately, directly on the oldest lobe (Pleistocene) of a megafan of the Taquari River (Assine and Soares, 2004); in this region there are many occurrences of restricted lenses of continental carbonates (Fig. 2). The studied fossils of the Serra da Bodoquena occur within the ~o Calca
rio Xarae
s. The carbonate deposit deposit of the Mineraça here is composed of unconsolidated fossiliferous wackestone in the form of tabular and lenticular beds, 0.5e3.0 m thick. This facies crops out over Middle Pleistocene wackestone and a thin bed of oncolitic rudstone, and is commonly covered by boundstone (soft tufa) younger than 5500 years (Fig. 2) (Ribeiro et al., 2015; Oliveira et al., 2016). These limestones are beige to grey in colour, with 60e95% calcium carbonate and some terrigenous sediment, mostly quartz silt, but also some clay (Fig. 3 a, b). They mostly occur close to rivers where they have been exposed by erosion through meandering, but they are also seen in quarries where they are being exploited for agriculture (Utida et al., 2012). Turcq et al. (1987) obtained an age of 5200 (±230 years BP) from 14C dating of shells

Fossiliferous micritic limestones were collected from 14 sites in the Serra da Bodoquena and Pantanal do Miranda (Table 1, Figs. 1
rio Xarae
s (site 1) and and 3. The deposits of the Mineraç~ ao Calca Pantanal do Miranda (site 13) were the same outcrops where Sallun Filho et al. (2009b) and Boggiani and Coimbra (1995) collected shells and obtained the carbon ages described above. The fossils studied from the Serra da Bodoquena belong to the lacustrine facies association of wackestones, oncolitic rudstones and fossil-rich wackestones Oliveira et al., 2016 (Fig. 2). The thickness of this association ranges between 15 cm and 6 m and it covers an area of between 50 m2 and 20 km2. The fact that this facies association is dominated by micritic carbonate, with ostracods, gastropods and bivalves, and is located in low, flat areas near the base of hills, in the form of discontinuous and elongated lenses, is compatible with a lacustrine environment (Utida et al., 2012). The fossils studied from the Pantanal do Miranda belong to a palustrine facies association, which is formed of quartz sand-rich wackestones, with a thickness ranging between 1.5 and 0.5 m, and covering an average area of around 200 m2. The textures include rhizoliths (and preserved roots) and mud cracks, and wellrounded and well-sorted quartz sandy sediments of the Pantanal carbonates (Oliveira et al., 2016). The geographical location of these occurrences, in the oldest lobe of the Taquari River megafan, suggests the introduction of carbonate-rich groundwater from the Neoproterozoic carbonate basement. 3.1. Fossil sampling Samples from fourteen sites (Table 1) were collected manually
rio where carbonates are cropping out. At the Mineraç~ ao Calca
s (site 1, Fig. 3 a), samples were collected by hand from the Xarae first 2 m at the top of the outcrop, and by auger drilling where there is no exposure, resulting in a composite section of about 5 m depth, with carbonates collected at each 0.5 m. In the laboratory shells were extracted manually or with tools such as tweezers, brushes and sieves, from carbonate hand samples of around 100 g. Molluscs

Please cite this article in press as: Utida, G., et al., Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.11.007

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Fig. 1. Study area. a) Serra da Bodoquena and Pantanal do Miranda localities. b) Serra da Bodoquena, circles indicate areas of micritic limestone, numbers indicate study sites. Based on Boggiani et al. (2002) and Sallun Filho (2005).


/Lad
ario plateau. Fig. 2. - Stratigraphy of the continental carbonates in the Serra da Bodoquena, Pantanal do Miranda and Corumba

Please cite this article in press as: Utida, G., et al., Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.11.007

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rio Xarae
s, at Fazenda Sa ~o Geraldo, Bonito-MS. b) Detail of the carbonate with shells in Bonito. c) Outcrop of Pantanal do Miranda Fig. 3. Field view of exposures. a) Mineraç~ao Calca limestone. d) Detail of the limestone with shells in Pantanal do Miranda.

Table 1 List of examined sites with geographical coordinates (longitude and latitude) and repository number of carbonate samples at the Litoteca of Geosciences Institute of S~ ao Paulo University. Examined sites Site

UTM WGS84 zone 21k

1 2 3 4 5 6 7 8 9 10 11 12 13 14

544652 545726 550753 551813 552247 551846 526992 564035 536210 536885 536396 529466 496138 495429

Repository 7650534 7649713 7656577 7677369 7665262 7677392 7670727 7665028 7718678 7721952 7723881 7728054 7834854 7847272

07TUF01 07TUF02 07TUF19 07TUF04A 07TUF16B 07TUF06A 07TUF07A 07TUF17A 07TUF15 07TUF12 07TUF13 07TUF14A Ponto32 Ponto38

were counted when at least the shell apexes were preserved and identified based on the work of Knight et al. (1960); Cox et al. (1969); Cowie and Thiengo (2003) and Simone (2006). They were classified according to the nomenclature of Bouchet and Rocroi (2005) for gastropods and Bouchet and Rocroi (2010) and Bieler et al. (2010) for bivalves. This analysis was used only for qualitative data as the number of specimens found was not statistically significant. Shells are kept in the Scientific Collection of the Geosciences Institute of S~ ao Paulo University (Appendix). All carbonate samples were deposited in the scientific collection of the Litoteca of Geo~o Paulo University (USP), Brazil (Table 1). sciences Institute of Sa 3.2. Taphonomic analysis ~o Calca
rio An oriented sample was collected from the Mineraça
s (site 1, Fig. 3a) near the top of the deposit for taphonomic Xarae

analysis and palaeoenvironmental interpretation. It was a block 32
16
25 cm in size, and the first 7 cm were analysed in layers ~1.5 cm thick. All shells in the layers were counted and analysed for their fragmentation, alteration (bioerosion, abrasion, corrosion), packing (dispersed to dense) and size. Fragmentation and alteration were analysed by categories of percentage of area affected by estimation. Fragmented shells were identified taxonomically when at least the apexes were presented and were classified with more than 90% of area fragmented; when the apexes were not preserved, shells were just counted as fragments. Only the right valves of bivalves were counted and analysed. These taphonomic signatures were adapted from Kidwell and Holland (1991); Kowalewski et al. ~ es (2006). (1994); Henderson et al. (2002) and Kotzian and Simo Spearman rank correlation coefficients were used to measure the correlation between species and taphonomic attributes. Only elongated shelled species (Idiopyrgus sp. and unidentified hydrobiid) were analysed for their orientation, taking note of the apex direction, the pointed end of the shell, as discussed by Tucker (2011). The Rayleigh z test was applied to evaluate a random distribution of shells, considering a 0.05 significance level for a critical value of z. Rose diagrams and Rayleigh tests were performed using the ORIANA 4 Statistical Program. The oriented block of carbonate sample and shells recovery from it were deposited in the scientific collection of the Litoteca of Geosciences Institute of S~ ao Paulo University (USP), Brazil (repository name “bloco”). 4. Results 4.1. Mollusc identification Shells recovered from the Serra da Bodoquena and Pantanal limestones are represented by 6 gastropod taxa and one bivalve taxon (Table 2). Gastropods identified are Pomacea canaliculata, Biomphalaria sp., Idiopyrgus sp., Marisa sp., Megalobulimus sp. and an unidentified hydrobiid, and the bivalve is Pisidium sp. (Figs. 4 and 5). Shells are kept in the Scientific Collection of the

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Table 2 Taxa occurrence at each site studied. *occurrence of microfossils: ostracods, characean algae and microgastropods (Utida et al., 2012). Taxa occurrence Site

Depth

Taxa P. canaliculata

Biomphalaria sp.

Unidentified hydrobiid

Idiopyrgus sp.

1

5.0 4.5 4.0 3.5 3.0* 2.5* 2.0* 1.5 1.0 0.5 0.0

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

2 3 4 5 6 7 8 9 10 11 12 13 14

* * * *

x

x x x x x x x x x

Pisidium sp.

Marisa sp.

Megalobulimus sp. 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

~o Paulo University (Appendix) but not all Geosciences Institute of Sa specimens were preserved due to their fragility and because of this, some sites do not have a representative of all species collected. 4.2. Taphonomy A total of 996 molluscan shells were recovered from the ori~o Calc

s (Fig. 6c). Around 49% of ented block from Mineraça ario Xarae them belong to Idiopyrgus sp. (Fig. 4d), followed by Biomphalaria sp. (Fig. 5a). with 19%. Pomacea canaliculata (Fig. 4a) and the unidentified hydrobiid (Fig. 4c) each represent 13%. The only bivalve identified was Pisidium sp. (Fig. 5c), representing 4% of the shells. The packing of the shells in the sediment is weak to dispersed with 90% of the shells supported by the carbonate matrix (Fig. 6a). Only 9% of shells were densely packed. Bioclast size distribution is concentrated between 1 and 9.5 mm for shell height (Fig. 6b), representing 95% of the total bioclasts. This group is composed predominantly of Biomphalaria sp., Idiopyrgus sp., the unidentified hydrobiid and Pisidium sp., with size variations from 1
1.5 to 9
5 mm (shell height x width) (Fig. 6d). The largest species is P. canaliculata with a distribution of shell height x width from 2.8
2.3 mm to 40.0
33.7 mm (Fig. 6d). P. canaliculata height distribution indicates a preference for small shells (~40%) and the high standard deviation suggests a non-normal distribution of sizes (Fig. 6e), although a higher number of observations would be necessary to confirm this. The other taxa have a distribution closest to the normal distribution around the mean size, as indicated by the low standard deviation (Fig. 6e). Only 20.6% of the total specimens have some kind of alteration (absence of bioerosion, 3.5% abrasion, 2.1% carbonate encrustation and 15% corrosion). Carbonate encrustations consist of very fine tubes, quite distinct from the carbonate matrix but they do not represent alteration. Fragmentation has a low index; 50% of the shells are not fragmented and only around 10% have more than 50% of the shells fragmented. In view of the small proportion of abraded shells, abrasion and corrosion are considered as just one attribute,

x x x

x x x

termed shell alteration. Shells of Idiopyrgus sp. are the most affected by alteration with around 20% of individuals affected across 10% of their area, although 71% of shells are well-preserved. The taxon also has 45% of shells unfragmented and 28% of them fragmented in less than 10% of their area (Fig. 7). These taphonomic attributes of Idiopyrgus sp. are not correlated (Spearman ¼ 0.115; p ¼ 0.011). More than 85% of shells of Biomphalaria sp. have no alteration and the other 23% were affected in less than 10% of their area. Biomphalaria sp. is the species with fewer fragmented shells; more than 62% of them are unfragmented and 24% have less than 10% of their areas fragmented (Fig. 7). These taphonomic attributes of Biomphalaria sp. are not correlated (Spearman ¼ 0.048; p ¼ 0.506). The unidentified hydrobiids have 81% of their shells unaffected by alteration, 13% affected in less than 10% of their area and less than 5% affected in more than 50% of their area (Fig. 7). Fragmented shells of the unidentified hydrobiid affected in 10% of their area are equivalent in percentage of specimens unfragmented, around 40% of shells (Fig. 7). These taphonomic attributes of the unidentified hydrobiids are not correlated (Spearman ¼ 0.250; p ¼ 0.004). Pomacea canaliculata represent 83% of shells unaffected by alteration and 10% with alteration in less than 10% of their area. This species has only 29% of shells unfragmented and there is a gradual distribution of percentages of shell in fragmentation categories, reaching 9% of shells with more than 90% of their area fragmented (Fig. 7). These taphonomic attributes of P. canaliculata are not correlated (Spearman ¼ 0.099; p ¼ 0.255). The bivalve Pisidium sp. has 95% and 65% of shells unaffected by alteration and fragmentation, respectively. With the very small number of specimens involved, this result is not reliable (Fig. 7). Orientation analysis and Rayleigh z test (z ¼ 0.634; p ¼ 0.531; a ¼ 0.05; critical z value ¼ 2.993) indicate that there is no mean direction of shells, since the z value was lower than the critical z value (Fig. 8). Therefore, there is no preferred orientation of shells; they have a random distribution.

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Fig. 4. Molluscs identified in carbonates from Mato Grosso do Sul. a) Pomacea canaliculata. b) Marisa sp. c) Unidentified hydrobiid. d) Idiopyrgus sp.

5. Discussion 5.1. Mollusc assemblage The planorbid Biomphalaria sp. was found in almost all deposits and is a good indicator of a quiet, shallow-water lacustrine environment with little turbulence, as would occur in areas which are

temporarily flooded (Utzinger et al., 1997; Brown et al., 1998; Utzinger and Tanner, 2000). Biomphalaria sp. and Pomacea canaliculata are widely distributed in the carbonates studied from the Serra da Bodoquena, and their living representatives are easily found in the region and throughout South America (Davis, 1979; Malek, 1983). Shells of P. canaliculata occur at site 1 from 3.0 to 5.0 m depth in

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Fig. 5. Molluscs identified in carbonates from Mato Grosso do Sul. a) Biomphalaria sp. b) Megalobulimus sp. The star represents carbonate sediment that is filling the shell. c) Pisidium sp., internal and external views of left and right valves.

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rio Xarae
s (site 1). a) Shell packing. b) Bioclast size distribution. c) Mollusc taxa disFig. 6. Graphs of taphonomic attributes. Data obtained from carbonates of Mineraç~ao Calca tribution. d) Size distribution of species e) Histogram of heights of mollusc species.

the limestone (Table 2) and these may be related to more arid periods, since then these gastropods would have buried themselves in the sediment to prevent their drying out. This habit is likely to have

given rise to high mortality rates (Kretzschmar and Heckman, 1995), although there is no evidence for this. In addition, permanent submersion of the Pomacea egg masses leads to death of the

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Fig. 7. Percentage of shells affected by fragmentation and alteration distributed by taxon. Key indicates the percentage of area fragmented. Shells were classified with more than 90% of their area fragmented when only the apexes were preserved and could be identified.

Fig. 8. Histogram of orientation of shell apexes obtained from the oriented sample collected at site 1.

embryos, although individual eggshells are not affected for at least two months (Pizani et al., 2005). This indicates that these freshwater gastropods lived close to the margins of lakes and in areas of non-permanent water, so then they could easily have had access to the land to lay their eggs. These gastropods are typical of tropical to subtropical climates and environments of low to insignificant water energy. However, they are not good indicators of other environmental parameters since they are able to adapt to a range of conditions, such as areas of low humidity and river systems, and the adults can tolerate exposure to the air and salinity fluctuations (Heckman, 1998; Cowie and Thiengo, 2003; Pizani et al., 2005). The diet of Pomacea sp. is varied; they are able to eat periphyte and macrophyte debris, and organic matter, according to their phase of development (Hirai, 1988). Hydrobiid gastropods, known as spring-snails, are exclusively

aquatic and their presence indicates permanent waters, inhabiting especially the areas around springs (Hershler, 1998). They have an important palaeontological significance because during their development they lack a free-swimming dispersal phase, so that they have limited mobility and a low dispersal potential. Dispersion of hydrobiids may have occurred during floods, transporting them from one lake to another (Worthington Wilmer et al., 2008, 2011). They were found at sites 1, from 2.0 to 5.0 m depth in the limestone, and sites 4, 6, 9, 10 and 11 (Table 2). The absence of the unidentified hydrobiid at the other sites suggests that these deposits might have formed during periods of non-permanent water or more marginal lake conditions. The pomatiopsid Idiopyrgus sp. occurs only in South America. It has an amphibious habit and is usually found associated with Biomphalaria sp. (Davis, 1979; Malek, 1983). Idiopyrgus sp. and the unidentified hydrobiid are vertically distributed in the limestones of site 1 from 2.0 to 5.0 m depth. The assemblage formed of these species, Idiopyrgus sp., Biomphalaria sp., P. canaliculata and the unidentified hydrobiid, was found from 2.0 to 5.0 m depth at site 1, and at sites 4, 6, 9 and 10 (Table 2). Considering their stratigraphic position and molluscan characteristics, they might represent a late phase of carbonate deposition during an episode of shallow water. Specifically, the presence of the unidentified hydrobiid may suggest that the studied region consisted of pools and springs at the time, and that for P. canaliculata, that there was aquatic vegetation as source of food for them (Cruz et al., 2015). The base of the limestone deposit at site 1 has no shells preserved (Table 2) and also no microfossils (Utida et al., 2012). This is likely indicating an unsuitable environment for these organisms or unfavourable conditions for their shell preservation. The carbonates of the Pantanal do Miranda are characterized by the presence of ampullariid Pomacea canaliculata and Marisa sp. Marisa species lay their eggs directly on the water in a gelatinous mass, whereas Pomacea species lay eggs with a carbonate coating on twigs or other surfaces near the water (Pilsbry, 1933). Possibly the carbonates from the Pantanal, where only Marisa sp. was found (Site 14), were some distance from vegetation. In such a situation, P. canaliculata could not lay eggs and this would have allowed Marisa sp. to dominate the molluscan fauna. Except for locality 14,

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G. Utida et al. / Quaternary International xxx (2016) 1e13

the deposit at 13 in the Pantanal and recent sediments nearby have large concentrations of P. canaliculata; this is probably a consequence of being situated closer to a lake margin. The presence of P. canaliculata could indicate a flooded area, such as a lake, with small islands with vegetation, where it lived and laid its eggs. Marisa sp. would have lived in the more open, quiet water between the islands, and its eggs could have floated gently on the surface of the water. Pisidium sp. belongs to one group of very small, cosmopolitan and freshwater bivalves. It is typically a burrowing bivalve, although the shell does not appear to have scars of syphons (Kuiper, 1983; Meyrick and Preece, 2001; Meyrich and Karrow, 2007). Examples were only found at several outcrops, probably because elsewhere they were not noticed due to their small size. At site 1, they occur between 2.0 and 3.0 m depth in the limestone; this is the same interval interpreted by Utida et al. (2012) as a clear-water, shallow lacustrine palaeoenvironment with aquatic vegetation because of the presence of microfossils, including ostracods, characean algae and some microgastropods. Finally Megalobulimus sp. belongs to a South American group of land snails, very common in forested areas. It has been found in Argentina, Paraguay and subtropical regions of Brazil (Bequaert, 1948; Sawaya and Petersen, 1962; Simone and Mezzalira, 1994). Its occurrence in the carbonate deposits may indicate close proximity to vegetation. 5.2. Taphonomic attributes
rio Xarae
s has the best preserved micritic carMineraç~ ao Calca bonates in the Serra da Bodoquena with a complete succession. Shells are preserved just near the top of these carbonates. Thus, the ~o Calca
rio Xarae
s was selected oriented block collected at Mineraça as the best representative of the upper part of the carbonate sequence. Weak to dispersed packing might indicate that the deposit was not a result of a high bioclastic input, hydraulic transport or failure of sediment supply (Kidwell and Holland, 1991). Although the size distribution of bioclasts is concentrated around a height of 10 mm, which could indicate a degree of sorting, it is easily observed that this is the natural size-range (Simone, 2006) of the majority of species found, predominantly Idiopyrgus sp., Biomphalaria sp. and the unidentified hydrobiids. The size of these species is also similar to a normal distribution, indicating that there was not a preference for a particular size (Fig. 6e). The fossil assemblage is dominated by Idiopyrgus sp., ~49% of the total shells, present from 2.0 to 5.0 m depth in the carbonate de~o Calca
rio Xarae
s (Table 2, site 1). Its abundance posit of Mineraça reflects favourable environmental conditions towards the end of carbonate deposition; this same environment can be attributed to most parts of the other deposits (Table 2). The biggest bioclasts are the shells of Pomacea canaliculata, distributed from 2.3
2.8 mm to 45
30 mm (Fig. 6d). These bioclasts are very different from a normal distribution with a high standard deviation and size selection around 5 mm height. The juveniles of P. canaliculata, defined as specimens with dimensions half of the average values (Yanes et al., 2008), are 71% of the total specimens with a size lower than 11
9 mm (height x width). One possible explanation of the size selection and absence of adult snails could be shell predation by snail kites (these are observed in the region today). According to De Francesco et al. (2006), these raptor birds prefer adult snails larger than 40 mm. Considering the larger size of P. canaliculata shells relative to that of the other taxa, their scarceness in the fossil record might reflect a lower abundance of adults (Martín and De Francesco,

2006) and/or a lower preservation rate (Yanes et al., 2008). In this last case, a low preservation might be indicated by a higher percentage of fragmented shells than the other taxa and only 5% of unfragmented shells are adults (Fig. 7). Probably, the majority of the large adult shells were completely fragmented as a result of the longer time they were subjected to reworking and abrasion. Although there is a higher fragmentation percentage of alteration for these P. canaliculata shells, it is not different from the other species (Fig. 7), probably due to their thicker shells. Shells affected by fragmentation were not affected by alteration to the same degree, according to the Spearman correlation. This suggests that fragmentation and alteration may have been produced by different actions. Biomphalaria sp. had the highest percentage of unfragmented shells, very different from P. canaliculata, the largest one. This is probably due to their shape being less influenced by hydrodynamic effects, since shells would have been lying on the lake floor allowing a faster burial in the sediment. There is also the possibility of a mixed assemblage. The highest percentages of shells affected by alteration belong to Idiopyrgus sp., ~30% of total specimens, whereas the other taxa are similarly distributed around 20e25% of shells affected by alteration. Based on this, fragmentation might be caused by intrinsic factors, especially the size-range of each species, favouring a longer time of reworking for the larger shells. On the other hand, high numbers of shells unaffected by alteration might be driven by extrinsic factors, such as a low energy environment, indicated by the shell assemblage, and environmental conditions that contributed to shell preservation. This hypothesis is supported by the polymodal pattern of shells that could be attributed to a lower water velocity that was not sufficient to orientate or transport shells, which would have then led to shell damage. It is clear from the degree of fragmentation and orientation associated with a polymodal pattern of shell apexes that the bioclasts were autochthonous in origin. The low alteration and generally well-preserved nature of the shells, also demonstrates the absence of any significant transport (De Francesco et al., 2007). The fabric and arrangement of the shells in the limestone are weak to dispersed, and most are supported by the fine-grained and/or sandy matrix. These data are consistent with an environment of little or no current/wave activity, as one would expect in a shallow lake or close to a spring. 6. Conclusions Molluscs identified in mid-Holocene limestones of Mato Grosso do Sul in the Serra da Bodoquena suggest a shallow and quiet freshwater environment occupied by aquatic vegetation, probably similar to springs. The fossil assemblage is characterized by the gastropods Idiopyrgus sp., Biomphalaria sp., Pomacea canaliculata and an unidentified hydrobiid. The occurrence of these species indicates a clear shallow-water lacustrine palaeoenvironment, with higher amounts of vegetation and organic matter debris, as they are food source for molluscs. The vegetation density is probably the difference between the palaeoenvironment with molluscs from that with microfossils, previously described by Utida et al. (2012). The interpretation of quiet waters as the palaeoenvironment of deposition for the micritic carbonates from the Serra da Bodoquena is also corroborated by taphonomic data that demonstrate a nonpreferred orientation of shells and low rates of shell damage. This environment is assumed to have existed during the last 3 m of carbonate deposition, since this mollusc assemblage was not found in the lower parts of the succession, and it is widespread across the Serra da Bodoquena. In the Pantanal do Miranda the occurrence of Marisa sp. in fine-

Please cite this article in press as: Utida, G., et al., Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.11.007

G. Utida et al. / Quaternary International xxx (2016) 1e13

grained limestones indicates a generally quiet water-body that was quite remote from the lake margin, with little current-wave activity and occupied by vegetation. There were some differences in palaeoenvironment between the Pantanal do Miranda and Serra da Bodoquena, although both were lacustrine aquatic environments rich in carbonate. There are no species indicating shallow waters in the Pantanal area, and the wackestones with quartz sand suggest more agitated waters than in the Serra da Bodoquena; this is probably associated with seasonal floods, as is typical for palustrine areas.

11

with reviews and comments; Geraldo Majella Pinheiro, the owner ~o Calc

s, for allowing access to conduct the of the Mineraça ario Xarae study in the mine area and assistance of their staff; CNPq for the financial support (proc. 479500/2007-0); Conselho Nacional de
gico (CNPq) and Coordenaça ~o Desenvolvimento Científico e Tecnolo de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the Master Scholarships.

Acknowledgements Appendix The authors are grateful to Gabriella T. Fontaneta for collaboration with the locality map; William Sallun Filho for collaboration ~o Paulo University. List of identified shell samples and respective collection numbers deposited at Scientific Collection of the Geosciences Institute of Sa Class Gastropoda Cuvier, 1797 Clade Caenogastropoda Cox, 1960 Family Ampullariidae Gray, 1824 Genus Pomacea Perry, 1810 Pomacea canaliculata Lamarck, 1828 Site

Collection no

Total no

1 4 5 6 7 8 9 10

GP/1E 5700, GP/1E 5701, GP/1E 5722, GP/1E 5725, GP/1E 5726, GP/1E 5730, GP/1E 5731, GP/1E 5732 GP/1E 5704, GP/1E 5736 GP/1E 5714 GP/1E 5738 GP/1E 5706 GP/1E 5716, GP/1E 5717 GP/1E 5711 GP/1E 5707 Genus Marisa Gray, 1824 Marisa sp. GP/1E 5720. Family Hydrobiidae Stimpson, 1865 Genus and species unidentified GP/1E 5735 GP/1E 5739 GP/1E 5713 GP/1E 5709 Family Pomatiopsidae Stimpson, 1865 Genus Idiopyrgus Pylsbry, 1911 Idiopyrgus sp. GP/1E 5702, GP/1E 5724, GP/1E 5726, GP/1E 5727 GP/1E 5718 GP/1E 5713 GP/1E 5709 Order Pulmonata Cuvier In Blainville, 1814 Suborder Basommatophora Family Planorbidae Rafinesque, 1815 Genus Biomphalaria Preston, 1910 Biomphalaria sp. GP/1E 5702, GP/1E 5723, GP/1E 5726, GP/1E 5729, GP/1E 5730, GP/1E 5731, GP/1E 5732 GP/1E 5735 GP/1E 5715 GP/1E 5738 GP/1E 5719 GP/1E 5712 GP/1E 5708 Family Megalobulimidae Leme, 1973 Genus Megalobulimus K. Miller, 1878 Megalobulimus sp. GP/1E 6517 GP/1E 6518

41 6 3 3 3 9 4 3

13

4 6 9 10

1 8 9 10

1 4 5 6 8 9 10

1 12 Class Bivalvia Linnaeus, 1758 Clade Heterodonta Neumayr, 1884 Family Pisidiidae Gray, 1857 Genus Pisidium C. Pfeiffer, 1821 Pisidium sp. 4 GP/1E 5734 6 GP/1E 5737

6

14 88 4 18

40 2 5 10

110 9 10 32 12 15 6

1 1

1 9

Please cite this article in press as: Utida, G., et al., Palaeoenvironmental interpretations based on molluscs from mid-Holocene lacustrine limestones, Mato Grosso do Sul, Brazil, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.11.007

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