Quaternary International 437 (2017) 186e198

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Quaternary International
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Palaeoenvironmental interpretations based on molluscs from mid-

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

a r t i c l e i n f o a b s t r a c t

Article history: In central Brazil there are continental carbonate rocks in three different geological contexts e the
Received 30 November 2015 Pleistocene wackestones in the Corumba
city (Xaraie

s Formation), the mid-Holocene limestones lenses in
Received in revised form the Pantanal Plain (Pantanal do Miranda) and old and modern tufas in the Serra da Bodoquena (Serra da
1 November 2016
Bodoquena Formation). We present here a systematic and taphonomic study of the molluscs found in the
Accepted 6 November 2016
Available online 1 March 2017
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
Keywords:
Gastropods
hydrobiid, Idiopyrgus sp., Biomphalaria sp. and Megalobulimus sp., and the bivalve Pisidium sp. The gas-
Taphonomy tropods include species still living in the area today, which supports a Holocene age for the limestones, as
Tufa indicated by published 14C dates. Overall, the data obtained from the Serra da Bodoquena carbonates
Lake suggest a freshwater environment with minimal current-wave action, in an area probably fed by springs,
Serra da Bodoquena close to a terrestrial environment, in view of the assemblage composed of Pomacea canaliculata, the
Pantanal unidentified hydrobiid, Idiopyrgus sp., Biomphalaria sp. and Megalobulimus sp. The lacustrine palae-
oenvironment 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
the terrestrial gastropods reported from fine-grained limestones in the Corumb
a area.
© 2016 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Stenogyra (Opeas) misera, Zonitoides sp. and Bulimulus sp. (Binney
and Bland, 1869; Oliveira and Almeida, 1999). Almeida (1945)
Continental Quaternary carbonate deposits occur widely in state suggested a Pleistocene or Pliocene age for these limestones,
of Mato Grosso do Sul (Brazil) and were first described by Almeida based on the gastropods, because they are living taxa.
(1945) from Corumba
as the Xaraie

s Formation. Almeida (1945) To the south in the Serra da Bodoquena, Almeida (1965) also
distinguished four categories of rocks: calcareous tufa with fossil assigned Quaternary carbonate deposits to the Xaraie
s Formation
plants, soft porous tufa, conglomerate with calcareous cement, and and later the Holocene age was confirmed (Turcq et al., 1987;
a micritic limestone with fossil gastropods. In the micritic lime- Boggiani et al., 1998). Sallun Filho et al. (2009a) proposed a sepa-
stone, Mendes (in Almeida, 1945) identified the land gastropods ration of these deposits from the Xaraie
s Formation to a new one,
the Serra da Bodoquena Formation, due to differences in litholog-
ical characteristics, the existence of clear and abrupt contacts, and
* Corresponding author. the relatively easy mapping of the unit. Boggiani et al. (2002)
E-mail addresses: [email protected] (G. Utida), [email protected] discovered some gastropods in these deposits, Biomphalaria sp.,
(E.C. Oliveira), [email protected] (M. Tucker), [email protected] (S. Petri), Physa sp. and Aquidauania sp. These species are typically from
[email protected] (P.C. Boggiani).

http://dx.doi.org/10.1016/j.quaint.2016.11.007
1040-6182/© 2016 Elsevier Ltd and INQUA. All rights reserved.
 G. Utida et al. / Quaternary International 437 (2017) 186e198 187

freshwater but they have an amphibious habit (Davis, 1979; Malek, in these carbonates, and suggested that they were deposited on an
1983; Utzinger et al., 1997; Oliveira and Almeida, 1999; Utzinger alluvial plain. According to the recent calibration curve of Hogg
and Tanner, 2000). et al. (2013), this age would recalculated to 5931 (±473 year cal
Gastropods also are preserved in wetlands of Pantanal, in Pan- BP). Sallun Filho et al. (2009b) confirmed the Holocene 14C ages for
tanal do Miranda within lenticular carbonate deposits and in cal- shells from the Mineraça ~o Calc

s, with a range of 2850 to
ario Xarae
cium carbonate layers in margins and bottoms of recent ponds of 2720 cal yr BP (in the upper part of the deposit), to 6530 to
alkaline waters (Turcq et al., 1987; Assine and Soares, 2004). Turcq 6310 cal yr BP (at a depth of 3 m into the deposit), but they inter-
et al. (1987) described the lenticular carbonates of Pantanal do preted the facies as lacustrine (Sallun Filho et al., 2009b; Utida,
Miranda as belonging to the Xaraie
s Formation, and bearing the 2009).
gastropods Pomacea sp. and Biomphalaria sp. (Boggiani and The Pantanal do Miranda is located just to the south of the
Coimbra, 1995; Boggiani et al., 1998). Utida et al. (2009, 2012) extensive alluvial fan of the Taquari River, and limestones are
described freshwater micro- and macro-fossils from the Xaraie
s exposed on the Park Road of Pantanal Sul, and elsewhere in the
Formation in the Pantanal, and suggested a lacustrine palae- area. These rocks are mostly lenticular deposits, and they are
oenvironment for carbonate deposition. generally quartz and fossil-rich wackestones, from 0.3 to 1.0 m
These fossiliferous micritic carbonates are widespread between thick and a lateral extension of 10e15 m, rarely up to 50 m (Fig. 3 c,
the Serra da Bodoquena and Pantanal do Miranda. They contain a d). These Pantanal do Miranda limestones were probably deposited
diversity of fossils but molluscs are found only in the upper part of in a lacustrine palaeoenvironment, when fluvial influence was
the deposits where microfossils were not found (Utida et al., 2012). minimal (Almeida, 1965; Boggiani et al., 1998; Ribeiro et al., 2001),
The presence of molluscs may indicate a different palaeoenviron- during the mid-Holocene according to a14C date of 3910 (±100
ment from that where the microfossils occur. years BP) (Boggiani et al., 1998), or 4255 (±279 years cal BP), ac-
We present here a taphonomic study of the molluscs found in cording to the calibration curve of Hogg et al. (2013). They were
the carbonate deposits of the Xaraie
s Formation collected in Serra deposited in localized areas upon a laterally-extensive unconsoli-
da Bodoquena and the carbonate lenses of the Pantanal do Miranda, dated fine quartz sand surface, probably in the lower reaches of an
which provide information concerning the carbonate palae- alluvial fan. The Pantanal limestones were probably deposited
oenvironments of these two distinct areas, Serra da Bodoquena and under a more arid climate, which existed during the period ~5600
Pantanal do Miranda. 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 Pan- Fossiliferous micritic limestones were collected from 14 sites in
tanal and is a range of hills 200 km in length and 400e800 m in the Serra da Bodoquena and Pantanal do Miranda (Table 1, Figs. 1
height. The Quaternary carbonates overlie Neoproterozoic/Edia- and 3. The deposits of the Mineraç~ ao Calc

ario Xarae
s (site 1) and
caran carbonates of the Corumb
a Group (Fig. 1); the latter have Pantanal do Miranda (site 13) were the same outcrops where Sallun
been extensively dissolved by groundwater to create major cave Filho et al. (2009b) and Boggiani and Coimbra (1995) collected
systems. Emerging groundwater from the carbonates provides bi- shells and obtained the carbon ages described above.
carbonate (and calcium) to the rivers and lakes of the area, where The fossils studied from the Serra da Bodoquena belong to the
tufa and micrite are widely precipitated (Almeida, 1965; Boggiani lacustrine facies association of wackestones, oncolitic rudstones
and Coimbra, 1995; Sallun Filho, 2005; Sallun Filho and Karmann, and fossil-rich wackestones Oliveira et al., 2016 (Fig. 2). The thick-
2007). ness of this association ranges between 15 cm and 6 m and it covers
The sedimentary characteristics of the studied deposits are an area of between 50 m2 and 20 km2. The fact that this facies
different in terms of their geology and geomorphology. The Serra da association is dominated by micritic carbonate, with ostracods,
Bodoquena is much affected by normal faults with a NNW-SSE gastropods and bivalves, and is located in low, flat areas near the
direction in the northern part and NW-SE, NE-SW and NS di- base of hills, in the form of discontinuous and elongated lenses, is
rections in the south-central portion. This range is made up of compatible with a lacustrine environment (Utida et al., 2012).
Neoproterozoic carbonate basement rocks, occurring up to 600 m The fossils studied from the Pantanal do Miranda belong to a
above sea level, with overlying occurrences of Quaternary conti- palustrine facies association, which is formed of quartz sand-rich
nental carbonates. The Pantanal do Miranda is located in the Pan- wackestones, with a thickness ranging between 1.5 and 0.5 m,
tanal basin, with altitudes of 90 m approximately, directly on the and covering an average area of around 200 m2. The textures
oldest lobe (Pleistocene) of a megafan of the Taquari River (Assine include rhizoliths (and preserved roots) and mud cracks, and well-
and Soares, 2004); in this region there are many occurrences of rounded and well-sorted quartz sandy sediments of the Pantanal
restricted lenses of continental carbonates (Fig. 2). carbonates (Oliveira et al., 2016). The geographical location of these
The studied fossils of the Serra da Bodoquena occur within the occurrences, in the oldest lobe of the Taquari River megafan, sug-
deposit of the Mineraça ~o Calca
rio Xarae
s. The carbonate deposit gests the introduction of carbonate-rich groundwater from the
here is composed of unconsolidated fossiliferous wackestone in the Neoproterozoic carbonate basement.
form of tabular and lenticular beds, 0.5e3.0 m thick. This facies
crops out over Middle Pleistocene wackestone and a thin bed of 3.1. Fossil sampling
oncolitic rudstone, and is commonly covered by boundstone (soft
tufa) younger than 5500 years (Fig. 2) (Ribeiro et al., 2015; Oliveira Samples from fourteen sites (Table 1) were collected manually
et al., 2016). These limestones are beige to grey in colour, with where carbonates are cropping out. At the Mineraça ~o Calc

ario
60e95% calcium carbonate and some terrigenous sediment, mostly Xarae
s (site 1, Fig. 3 a), samples were collected by hand from the
quartz silt, but also some clay (Fig. 3 a, b). They mostly occur close to first 2 m at the top of the outcrop, and by auger drilling where there
rivers where they have been exposed by erosion through is no exposure, resulting in a composite section of about 5 m depth,
meandering, but they are also seen in quarries where they are being with carbonates collected at each 0.5 m. In the laboratory shells
exploited for agriculture (Utida et al., 2012). Turcq et al. (1987) were extracted manually or with tools such as tweezers, brushes
obtained an age of 5200 (±230 years BP) from 14C dating of shells and sieves, from carbonate hand samples of around 100 g. Molluscs
188 G. Utida et al. / Quaternary International 437 (2017) 186e198

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).

/Lada
Fig. 2. - Stratigraphy of the continental carbonates in the Serra da Bodoquena, Pantanal do Miranda and Corumba
rio plateau.
 G. Utida et al. / Quaternary International 437 (2017) 186e198 189

Fig. 3. Field view of exposures. a) Mineraç~ao Calca

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
limestone. d) Detail of the limestone with shells in Pantanal do Miranda.

Table 1 analysis and palaeoenvironmental interpretation. It was a block

List of examined sites with geographical coordinates (longitude and latitude) and 32
16
25 cm in size, and the first 7 cm were analysed in layers
repository number of carbonate samples at the Litoteca of Geosciences Institute of
~1.5 cm thick. All shells in the layers were counted and analysed for
S~
ao Paulo University.
their fragmentation, alteration (bioerosion, abrasion, corrosion),
Examined sites packing (dispersed to dense) and size. Fragmentation and alter-
Site UTM WGS84 zone 21k Repository ation were analysed by categories of percentage of area affected by
1 544652 7650534 07TUF01 estimation. Fragmented shells were identified taxonomically when
2 545726 7649713 07TUF02 at least the apexes were presented and were classified with more
3 550753 7656577 07TUF19 than 90% of area fragmented; when the apexes were not preserved,
4 551813 7677369 07TUF04A shells were just counted as fragments. Only the right valves of bi-
5 552247 7665262 07TUF16B
6 551846 7677392 07TUF06A
valves were counted and analysed. These taphonomic signatures
7 526992 7670727 07TUF07A were adapted from Kidwell and Holland (1991); Kowalewski et al.
8 564035 7665028 07TUF17A (1994); Henderson et al. (2002) and Kotzian and Simo ~ es (2006).
9 536210 7718678 07TUF15 Spearman rank correlation coefficients were used to measure the
10 536885 7721952 07TUF12
correlation between species and taphonomic attributes.
11 536396 7723881 07TUF13
12 529466 7728054 07TUF14A Only elongated shelled species (Idiopyrgus sp. and unidentified
13 496138 7834854 Ponto32 hydrobiid) were analysed for their orientation, taking note of the
14 495429 7847272 Ponto38 apex direction, the pointed end of the shell, as discussed by Tucker
(2011). The Rayleigh z test was applied to evaluate a random dis-
tribution of shells, considering a 0.05 significance level for a critical
were counted when at least the shell apexes were preserved and value of z. Rose diagrams and Rayleigh tests were performed using
identified based on the work of Knight et al. (1960); Cox et al. the ORIANA 4 Statistical Program.
(1969); Cowie and Thiengo (2003) and Simone (2006). They were The oriented block of carbonate sample and shells recovery from
classified according to the nomenclature of Bouchet and Rocroi it were deposited in the scientific collection of the Litoteca of
(2005) for gastropods and Bouchet and Rocroi (2010) and Bieler Geosciences Institute of S~ ao Paulo University (USP), Brazil (re-
et al. (2010) for bivalves. This analysis was used only for qualita- pository name “bloco”).
tive data as the number of specimens found was not statistically
significant.
4. Results
Shells are kept in the Scientific Collection of the Geosciences
Institute of S~
ao Paulo University (Appendix). All carbonate samples
4.1. Mollusc identification
were deposited in the scientific collection of the Litoteca of Geo-
~o Paulo University (USP), Brazil (Table 1).
sciences Institute of Sa
Shells recovered from the Serra da Bodoquena and Pantanal
limestones are represented by 6 gastropod taxa and one bivalve
3.2. Taphonomic analysis taxon (Table 2). Gastropods identified are Pomacea canaliculata,
Biomphalaria sp., Idiopyrgus sp., Marisa sp., Megalobulimus sp. and
An oriented sample was collected from the Mineraça ~o Calca

rio an unidentified hydrobiid, and the bivalve is Pisidium sp. (Figs. 4

s (site 1, Fig. 3a) near the top of the deposit for taphonomic
Xarae and 5). Shells are kept in the Scientific Collection of the
190 G. Utida et al. / Quaternary International 437 (2017) 186e198

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. Pisidium sp. Marisa sp. Megalobulimus sp.

1 5.0 x x x x x
4.5 x x x x
4.0 x x x x
3.5 x x x
3.0* x x x x
2.5* x x x x
2.0* x x x x
1.5 x
1.0
0.5
0.0
2 x x
3 x
4 * x x x x x x
5 * x x
6 * x x x x x x
7 * x
8 x x x
9 x x x x x
10 x x x x
11 x x x x
12 x
13 x x x
14 x

~o Paulo University (Appendix) but not all

Geosciences Institute of Sa termed shell alteration.
specimens were preserved due to their fragility and because of this, Shells of Idiopyrgus sp. are the most affected by alteration with
some sites do not have a representative of all species collected. 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%
4.2. Taphonomy of their area (Fig. 7). These taphonomic attributes of Idiopyrgus sp.
are not correlated (Spearman ¼ 0.115; p ¼ 0.011).
A total of 996 molluscan shells were recovered from the ori- More than 85% of shells of Biomphalaria sp. have no alteration
ented block from Mineraç~ ao Calc

ario Xarae
s (Fig. 6c). Around 49% of
and the other 23% were affected in less than 10% of their area.
them belong to Idiopyrgus sp. (Fig. 4d), followed by Biomphalaria sp. Biomphalaria sp. is the species with fewer fragmented shells; more
(Fig. 5a). with 19%. Pomacea canaliculata (Fig. 4a) and the uniden- than 62% of them are unfragmented and 24% have less than 10% of
tified hydrobiid (Fig. 4c) each represent 13%. The only bivalve their areas fragmented (Fig. 7). These taphonomic attributes of
identified was Pisidium sp. (Fig. 5c), representing 4% of the shells. Biomphalaria sp. are not correlated (Spearman ¼ 0.048; p ¼ 0.506).
The packing of the shells in the sediment is weak to dispersed The unidentified hydrobiids have 81% of their shells unaffected
with 90% of the shells supported by the carbonate matrix (Fig. 6a). by alteration, 13% affected in less than 10% of their area and less
Only 9% of shells were densely packed. Bioclast size distribution is than 5% affected in more than 50% of their area (Fig. 7). Fragmented
concentrated between 1 and 9.5 mm for shell height (Fig. 6b), shells of the unidentified hydrobiid affected in 10% of their area are
representing 95% of the total bioclasts. This group is composed equivalent in percentage of specimens unfragmented, around 40%
predominantly of Biomphalaria sp., Idiopyrgus sp., the unidentified of shells (Fig. 7). These taphonomic attributes of the unidentified
hydrobiid and Pisidium sp., with size variations from 1
1.5 to hydrobiids are not correlated (Spearman ¼ 0.250; p ¼ 0.004).
9
5 mm (shell height x width) (Fig. 6d). The largest species is Pomacea canaliculata represent 83% of shells unaffected by
P. canaliculata with a distribution of shell height x width from alteration and 10% with alteration in less than 10% of their area. This
2.8
2.3 mm to 40.0
33.7 mm (Fig. 6d). P. canaliculata height species has only 29% of shells unfragmented and there is a gradual
distribution indicates a preference for small shells (~40%) and the distribution of percentages of shell in fragmentation categories,
high standard deviation suggests a non-normal distribution of sizes reaching 9% of shells with more than 90% of their area fragmented
(Fig. 6e), although a higher number of observations would be (Fig. 7). These taphonomic attributes of P. canaliculata are not
necessary to confirm this. The other taxa have a distribution closest correlated (Spearman ¼ 0.099; p ¼ 0.255).
to the normal distribution around the mean size, as indicated by the The bivalve Pisidium sp. has 95% and 65% of shells unaffected by
low standard deviation (Fig. 6e). alteration and fragmentation, respectively. With the very small
Only 20.6% of the total specimens have some kind of alteration number of specimens involved, this result is not reliable (Fig. 7).
(absence of bioerosion, 3.5% abrasion, 2.1% carbonate encrustation Orientation analysis and Rayleigh z test (z ¼ 0.634; p ¼ 0.531;
and 15% corrosion). Carbonate encrustations consist of very fine a ¼ 0.05; critical z value ¼ 2.993) indicate that there is no mean
tubes, quite distinct from the carbonate matrix but they do not direction of shells, since the z value was lower than the critical z
represent alteration. Fragmentation has a low index; 50% of the value (Fig. 8). Therefore, there is no preferred orientation of shells;
shells are not fragmented and only around 10% have more than 50% they have a random distribution.
of the shells fragmented. In view of the small proportion of abraded
shells, abrasion and corrosion are considered as just one attribute,
 G. Utida et al. / Quaternary International 437 (2017) 186e198 191

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 temporarily flooded (Utzinger et al., 1997; Brown et al., 1998;

Utzinger and Tanner, 2000). Biomphalaria sp. and Pomacea canal-
5.1. Mollusc assemblage iculata are widely distributed in the carbonates studied from the
Serra da Bodoquena, and their living representatives are easily
The planorbid Biomphalaria sp. was found in almost all deposits found in the region and throughout South America (Davis, 1979;
and is a good indicator of a quiet, shallow-water lacustrine envi- Malek, 1983).
ronment with little turbulence, as would occur in areas which are Shells of P. canaliculata occur at site 1 from 3.0 to 5.0 m depth in
192 G. Utida et al. / Quaternary International 437 (2017) 186e198

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.
 G. Utida et al. / Quaternary International 437 (2017) 186e198 193

Fig. 6. Graphs of taphonomic attributes. Data obtained from carbonates of Mineraç~ao Calc
ario Xarae

s (site 1). a) Shell packing. b) Bioclast size distribution. c) Mollusc taxa dis-
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 pe- given rise to high mortality rates (Kretzschmar and Heckman,
riods, since then these gastropods would have buried themselves in 1995), although there is no evidence for this. In addition, perma-
the sediment to prevent their drying out. This habit is likely to have nent submersion of the Pomacea egg masses leads to death of the
194 G. Utida et al. / Quaternary International 437 (2017) 186e198

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.

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., Bio-
mphalaria 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
Fig. 8. Histogram of orientation of shell apexes obtained from the oriented sample characteristics, they might represent a late phase of carbonate
collected at site 1. deposition during an episode of shallow water. Specifically, the
presence of the unidentified hydrobiid may suggest that the stud-
ied region consisted of pools and springs at the time, and that for
embryos, although individual eggshells are not affected for at least P. canaliculata, that there was aquatic vegetation as source of food
two months (Pizani et al., 2005). This indicates that these fresh- for them (Cruz et al., 2015).
water gastropods lived close to the margins of lakes and in areas of The base of the limestone deposit at site 1 has no shells pre-
non-permanent water, so then they could easily have had access to served (Table 2) and also no microfossils (Utida et al., 2012). This is
the land to lay their eggs. likely indicating an unsuitable environment for these organisms or
These gastropods are typical of tropical to subtropical climates unfavourable conditions for their shell preservation.
and environments of low to insignificant water energy. However, The carbonates of the Pantanal do Miranda are characterized by
they are not good indicators of other environmental parameters the presence of ampullariid Pomacea canaliculata and Marisa sp.
since they are able to adapt to a range of conditions, such as areas of Marisa species lay their eggs directly on the water in a gelatinous
low humidity and river systems, and the adults can tolerate expo- mass, whereas Pomacea species lay eggs with a carbonate coating
sure to the air and salinity fluctuations (Heckman, 1998; Cowie and on twigs or other surfaces near the water (Pilsbry, 1933). Possibly
Thiengo, 2003; Pizani et al., 2005). The diet of Pomacea sp. is varied; the carbonates from the Pantanal, where only Marisa sp. was found
they are able to eat periphyte and macrophyte debris, and organic (Site 14), were some distance from vegetation. In such a situation,
matter, according to their phase of development (Hirai, 1988). P. canaliculata could not lay eggs and this would have allowed
Hydrobiid gastropods, known as spring-snails, are exclusively Marisa sp. to dominate the molluscan fauna. Except for locality 14,
 G. Utida et al. / Quaternary International 437 (2017) 186e198 195

the deposit at 13 in the Pantanal and recent sediments nearby have 2006) and/or a lower preservation rate (Yanes et al., 2008). In this
large concentrations of P. canaliculata; this is probably a conse- last case, a low preservation might be indicated by a higher per-
quence of being situated closer to a lake margin. centage of fragmented shells than the other taxa and only 5% of
The presence of P. canaliculata could indicate a flooded area, unfragmented shells are adults (Fig. 7). Probably, the majority of the
such as a lake, with small islands with vegetation, where it lived large adult shells were completely fragmented as a result of the
and laid its eggs. Marisa sp. would have lived in the more open, longer time they were subjected to reworking and abrasion.
quiet water between the islands, and its eggs could have floated Although there is a higher fragmentation percentage of alteration
gently on the surface of the water. for these P. canaliculata shells, it is not different from the other
Pisidium sp. belongs to one group of very small, cosmopolitan species (Fig. 7), probably due to their thicker shells.
and freshwater bivalves. It is typically a burrowing bivalve, Shells affected by fragmentation were not affected by alteration
although the shell does not appear to have scars of syphons (Kuiper, to the same degree, according to the Spearman correlation. This
1983; Meyrick and Preece, 2001; Meyrich and Karrow, 2007). Ex- suggests that fragmentation and alteration may have been pro-
amples were only found at several outcrops, probably because duced by different actions. Biomphalaria sp. had the highest per-
elsewhere they were not noticed due to their small size. At site 1, centage of unfragmented shells, very different from P. canaliculata,
they occur between 2.0 and 3.0 m depth in the limestone; this is the the largest one. This is probably due to their shape being less
same interval interpreted by Utida et al. (2012) as a clear-water, influenced by hydrodynamic effects, since shells would have been
shallow lacustrine palaeoenvironment with aquatic vegetation lying on the lake floor allowing a faster burial in the sediment.
because of the presence of microfossils, including ostracods, char- There is also the possibility of a mixed assemblage. The highest
acean algae and some microgastropods. percentages of shells affected by alteration belong to Idiopyrgus sp.,
Finally Megalobulimus sp. belongs to a South American group of ~30% of total specimens, whereas the other taxa are similarly
land snails, very common in forested areas. It has been found in distributed around 20e25% of shells affected by alteration. Based
Argentina, Paraguay and subtropical regions of Brazil (Bequaert, on this, fragmentation might be caused by intrinsic factors, espe-
1948; Sawaya and Petersen, 1962; Simone and Mezzalira, 1994). cially the size-range of each species, favouring a longer time of
Its occurrence in the carbonate deposits may indicate close prox- reworking for the larger shells. On the other hand, high numbers of
imity to vegetation. shells unaffected by alteration might be driven by extrinsic factors,
such as a low energy environment, indicated by the shell assem-
blage, and environmental conditions that contributed to shell
5.2. Taphonomic attributes
preservation. This hypothesis is supported by the polymodal
pattern of shells that could be attributed to a lower water velocity
Mineraça ~o Calca

rio Xarae
s has the best preserved micritic car-
that was not sufficient to orientate or transport shells, which would
bonates in the Serra da Bodoquena with a complete succession.
have then led to shell damage.
Shells are preserved just near the top of these carbonates. Thus, the
It is clear from the degree of fragmentation and orientation
oriented block collected at Mineraça ~o Calca
rio Xarae
s was selected
associated with a polymodal pattern of shell apexes that the bio-
as the best representative of the upper part of the carbonate
clasts were autochthonous in origin. The low alteration and
sequence.
generally well-preserved nature of the shells, also demonstrates
Weak to dispersed packing might indicate that the deposit was
the absence of any significant transport (De Francesco et al., 2007).
not a result of a high bioclastic input, hydraulic transport or failure
The fabric and arrangement of the shells in the limestone are weak
of sediment supply (Kidwell and Holland, 1991). Although the size
to dispersed, and most are supported by the fine-grained and/or
distribution of bioclasts is concentrated around a height of 10 mm,
sandy matrix. These data are consistent with an environment of
which could indicate a degree of sorting, it is easily observed that
little or no current/wave activity, as one would expect in a shallow
this is the natural size-range (Simone, 2006) of the majority of
lake or close to a spring.
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 6. Conclusions
for a particular size (Fig. 6e).
The fossil assemblage is dominated by Idiopyrgus sp., ~49% of the Molluscs identified in mid-Holocene limestones of Mato Grosso
total shells, present from 2.0 to 5.0 m depth in the carbonate de- do Sul in the Serra da Bodoquena suggest a shallow and quiet
posit of Mineraça ~o Calca
rio Xarae
s (Table 2, site 1). Its abundance freshwater environment occupied by aquatic vegetation, probably
reflects favourable environmental conditions towards the end of similar to springs. The fossil assemblage is characterized by the
carbonate deposition; this same environment can be attributed to gastropods Idiopyrgus sp., Biomphalaria sp., Pomacea canaliculata
most parts of the other deposits (Table 2). and an unidentified hydrobiid. The occurrence of these species
The biggest bioclasts are the shells of Pomacea canaliculata, indicates a clear shallow-water lacustrine palaeoenvironment, with
distributed from 2.3
2.8 mm to 45
30 mm (Fig. 6d). These higher amounts of vegetation and organic matter debris, as they are
bioclasts are very different from a normal distribution with a high food source for molluscs. The vegetation density is probably the
standard deviation and size selection around 5 mm height. The difference between the palaeoenvironment with molluscs from
juveniles of P. canaliculata, defined as specimens with dimensions that with microfossils, previously described by Utida et al. (2012).
half of the average values (Yanes et al., 2008), are 71% of the total The interpretation of quiet waters as the palaeoenvironment of
specimens with a size lower than 11
9 mm (height x width). One deposition for the micritic carbonates from the Serra da Bodoquena
possible explanation of the size selection and absence of adult is also corroborated by taphonomic data that demonstrate a non-
snails could be shell predation by snail kites (these are observed in preferred orientation of shells and low rates of shell damage. This
the region today). According to De Francesco et al. (2006), these environment is assumed to have existed during the last 3 m of
raptor birds prefer adult snails larger than 40 mm. carbonate deposition, since this mollusc assemblage was not found
Considering the larger size of P. canaliculata shells relative to in the lower parts of the succession, and it is widespread across the
that of the other taxa, their scarceness in the fossil record might Serra da Bodoquena.
reflect a lower abundance of adults (Martín and De Francesco, In the Pantanal do Miranda the occurrence of Marisa sp. in fine-
196 G. Utida et al. / Quaternary International 437 (2017) 186e198

grained limestones indicates a generally quiet water-body that was with reviews and comments; Geraldo Majella Pinheiro, the owner
quite remote from the lake margin, with little current-wave activity ~o Calc

of the Mineraça
s, for allowing access to conduct the
ario Xarae
and occupied by vegetation. There were some differences in palae- study in the mine area and assistance of their staff; CNPq for the
oenvironment between the Pantanal do Miranda and Serra da financial support (proc. 479500/2007-0); Conselho Nacional de
Bodoquena, although both were lacustrine aquatic environments Desenvolvimento Científico e Tecnolo
gico (CNPq) and Coordenaça ~o
rich in carbonate. There are no species indicating shallow waters in de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the
the Pantanal area, and the wackestones with quartz sand suggest Master Scholarships.
more agitated waters than in the Serra da Bodoquena; this is prob-
ably associated with seasonal floods, as is typical for palustrine areas.

Acknowledgements
Appendix
The authors are grateful to Gabriella T. Fontaneta for collabo-
ration 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 GP/1E 5700, GP/1E 5701, GP/1E 5722, GP/1E 5725, GP/1E 5726, GP/1E 5730, GP/1E 5731, GP/1E 5732 41
4 GP/1E 5704, GP/1E 5736 6
5 GP/1E 5714 3
6 GP/1E 5738 3
7 GP/1E 5706 3
8 GP/1E 5716, GP/1E 5717 9
9 GP/1E 5711 4
10 GP/1E 5707 3
Genus Marisa Gray, 1824
Marisa sp.
13 GP/1E 5720. 6
Family Hydrobiidae Stimpson, 1865
Genus and species unidentified
4 GP/1E 5735 14
6 GP/1E 5739 88
9 GP/1E 5713 4
10 GP/1E 5709 18
Family Pomatiopsidae Stimpson, 1865
Genus Idiopyrgus Pylsbry, 1911
Idiopyrgus sp.
1 GP/1E 5702, GP/1E 5724, GP/1E 5726, GP/1E 5727 40
8 GP/1E 5718 2
9 GP/1E 5713 5
10 GP/1E 5709 10
Order Pulmonata Cuvier In Blainville, 1814
Suborder Basommatophora
Family Planorbidae Rafinesque, 1815
Genus Biomphalaria Preston, 1910
Biomphalaria sp.
1 GP/1E 5702, GP/1E 5723, GP/1E 5726, GP/1E 5729, GP/1E 5730, GP/1E 5731, GP/1E 5732 110
4 GP/1E 5735 9
5 GP/1E 5715 10
6 GP/1E 5738 32
8 GP/1E 5719 12
9 GP/1E 5712 15
10 GP/1E 5708 6
Family Megalobulimidae Leme, 1973
Genus Megalobulimus K. Miller, 1878
Megalobulimus sp.
1 GP/1E 6517 1
12 GP/1E 6518 1
Class Bivalvia Linnaeus, 1758
Clade Heterodonta Neumayr, 1884
Family Pisidiidae Gray, 1857
Genus Pisidium C. Pfeiffer, 1821
Pisidium sp.
4 GP/1E 5734 1
6 GP/1E 5737 9
 G. Utida et al. / Quaternary International 437 (2017) 186e198 197

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