Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113

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Palaeogeography, Palaeoclimatology, Palaeoecology

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 / p a l a e o

The significance of molluscs as paleoecological indicators of freshwater systems in

central-western Argentina
Claudio G. De Francesco ⁎, Gabriela S. Hassan
CONICET-Centro de Geología de Costas y del Cuaternario, Universidad Nacional de Mar del Plata, CC 722, 7600 Mar del Plata, Argentina

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

Article history: The objectives of this study were to (a) analyze the distribution pattern of molluscs in freshwater systems
Received 4 September 2008 from central-western Argentina, (b) investigate the key environmental factors affecting the species
Received in revised form 15 January 2009 distribution, and (c) compare Quaternary and modern mollusc assemblages in order to evaluate the extent
Accepted 19 January 2009
and limitations of freshwater molluscs as paleobioindicators. A total of 45 freshwater habitats were sampled
for living molluscs. At each selected site, physical and chemical parameters were quantified, and living
Keywords:
Freshwater molluscs
molluscs collected. Principal Component Analysis (PCA) was used for the ordination of sampling sites based
Distribution patterns on environmental variables. To explore the relationships between environmental variables and mollusc
Paleoecology assemblages, a Canonical Correspondence Analysis (CCA) was performed. Data on fossil mollusc assemblages
Quaternary from three Quaternary alluvial successions outcropping in the area were combined with the modern data
Mendoza set by means of a Detrended Correspondence Analysis (DCA). Eight taxa were identified: Lymnaea viator,
Argentina Heleobia hatcheri, Heleobia parchappii, Heleobia aff. parchappii, Chilina mendozana, Biomphalaria peregrina,
Physa acuta, and a bivalve attributed to the family Sphaeriidae. Except for H. hatcheri and P. acuta, the
species were also recorded as fossils in the area. Overall, the occurrence of molluscs in central-western
Argentina appeared to be related to calm vegetated shallow water bodies. With the exception of H. parchappii
that only occurred in saline waters (8–16 mS cm− 1), the mollusc assemblages were restricted to low salinities
(below 1.9 mS cm− 1). Two main groups, indicative of different energetic environmental conditions related to
the stability of the flow regime, were recognized: (1) L. viator, P. acuta, and B. peregrina occurred at lentic
habitats (shallow lakes) or calm backwaters (pools) within lotic systems such as streams and rivers, (2) H.
hatcheri, H. aff. parchappii, C. mendozana, and Sphaeriidae dominated in lotic habitats where a directional
water flow exists. These same groups were recognized in the fossil record allowing the reconstruction of
habitats that differed in the magnitude of the water flow.
© 2009 Elsevier B.V. All rights reserved.

1. Introduction information significantly restricts the quality and extent of paleoen-

Molluscs are among the most ubiquitous fossils present in
Quaternary non-marine sediments, being found in a wide variety of
deposits including fluvial, lacustrine, glaciolacustrine and paludal
sediments (Miller and Bajc, 1990). They constitute an invaluable
source of data for reconstructing past environments, in particular the
hydrodynamics of past water bodies. However, the absence of
ecological data for many species as well as a general inadequacy in
our understanding of the factors controlling their distribution has
limited their broad use as paleoecological indicators. This situation is
particularly worrying in Argentina because the ecology of freshwater
molluscs has been poorly studied and the ecological requirements of
species remain in many cases nearly unknown. This lack of modern
⁎ Corresponding author. Tel.: +54 223 4754060; fax: +54 223 4753150.
E-mail addresses: cgdefra@mdp.edu.ar (C.G. De Francesco), ghassan@mdp.edu.ar
(G.S. Hassan).
vironmental information that can be gathered from the mollusc
assemblages preserved in Quaternary alluvial successions.
Recently, paleoecological analysis of Pleistocene and Holocene
alluvial successions cropping out in the semiarid region of Mendoza
(33–35° S, 69° W) revealed the occurrence of at least six mollusc
species. As most of them are common in freshwater habitats from
other geographic regions of Argentina (Pampa or Patagonia) the scarce
ecological information available was used to reconstruct the past
hydrodynamics of these successions. As a result, the Quaternary
mollusc assemblages preserved in the basin of Río Tunuyán (northern
Mendoza), which were dominated by semi-aquatic snails (Lymnaea
viator), suggested the development of damp habitats that were
occasionally submerged (De Francesco and Zárate, 2006; De Francesco
et al., 2007). On the other hand, the mollusc assemblages preserved in
the basin of Río Atuel (southern Mendoza) that showed a higher
diversity, with the presence of species not represented in northern
Mendoza (e. g., Chilina mendozana), suggested the development of

0031-0182/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2009.01.003
106 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
shallow vegetated habitats subject to episodic flooding events
although deeper than those represented in the north (Dieguez et al.,
2004; De Francesco and Dieguez, 2006).
Because of the lack of information on modern species distribu-
tions in the area, we cannot predict either the magnitude of the water
bodies developed in the past or the mean values of the main
environmental parameters that may have characterized them. Some
of the species represented as fossils were recently recorded living in
Bañado Carilauquen, a tributary of Llancanelo Lake in southern
Mendoza (Ciocco and Scheibler, 2008). In this exploratory work, the
distribution of molluscs appeared to be related to a gradient of
conductivity and hardness. Yet, as Bañado Carilauquen constitutes
only an isolated case within the total range of freshwater habitats in
Mendoza Province, this study does not provide information on the
factors that affect the distribution of molluscs at a regional scale. At
present, there is no available information on the distribution of
freshwater molluscs in other types of freshwater systems (streams,
rivers, shallow lakes, dams, canals) in Mendoza Province. Therefore,
an understanding of the factors influencing local distribution of
modern species in Mendoza becomes a key issue in assessing the
value of freshwater molluscs as paleoecological indicators in the
area.
The aim of the present study is to assess the paleoecological
significance of freshwater molluscs in the semiarid region of Mendoza
in order to support paleoenvironmental reconstructions in the area. In
particular, the present contribution aims to: (a) analyze in detail the
distribution pattern of molluscs in freshwater systems of the area,
(b) investigate the key environmental factors affecting the species
distributions, and (c) compare Quaternary and modern mollusc
assemblages in order to infer the nature of past habitats and, thereby,
evaluate the extent and limitations of freshwater molluscs as
paleobioindicators.
Fig. 1. Location map of modern (circles) and Quaternary (squares) study sites in central-western Argentina.
2. Study area
The province of Mendoza lies within the dominion of the South
American Arid Diagonal, a region considered to have been climatically
sensitive to the latitudinal shift of the Pacific and Atlantic anticyclonic
centres during the late Pleistocene and the Holocene (Abraham de
Vázquez et al., 2000). The climate of the region is semiarid with a main
annual rainfall of 250 mm in the eastern piedmont (Capitanelli, 2005).
Regionally, the study area is cut across by several fluvial systems
(Tunuyán, Diamante, and Atuel rivers) with their headwaters located
in the high Andes Cordillera (Fig. 1). The river water discharges depend
on glacial melting and the mainly winter precipitation of Pacific origin.
When reaching the piedmont, these rivers form very extensive and
complex alluvial fans. On the margins of these rivers several
Quaternary alluvial deposits composed of mollusc shells are con-
tinuously exposed along several kilometers (Zárate, 2002).
3. Materials and methods
3.1. Modern dataset
3.1.1. Field sampling
In total, 45 sites were selected to represent the maximum
heterogeneity of aquatic environments in the area (Fig. 1), that is,
the whole range of variation in morphology, substrate, and flow (e.g.,
streams, rivers, shallow lakes, ponds, dams, canals). As the study
aimed to understand the value of molluscs as paleoecological
indicators in the area, all the selected sites were located within the
same area in which Quaternary molluscs crop out (see De Francesco
and Dieguez, 2006; De Francesco et al., 2007). The basins of Río
Tunuyán, Río Diamante, Río Atuel, and Río Grande were included. The
sites selected represented a wide range of disturbance levels
 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
(anthropogenic impact) from almost pristine environments (Andes
Cordillera) to streams running across cities or through areas of low to
moderate cattle-raising and agriculture (basin of Río Tunuyán).
At each site, we quantified current velocity, water temperature, pH,
and conductivity. Current velocity was measured using a neutrally
buoyant sphere and calculating the time it took the sphere to move
5 m. Temperature, pH, and conductivity were measured with field
instruments. Aquatic vegetation cover at each sampling site was
estimated visually, and a nominal variable erected for statistical
purposes (0 = no vegetation, 1 = low vegetation cover, and 2 = high
vegetation cover).
One subsurface water sample (1 l) was taken at each site to assess
chemical parameters. Water samples were collected in polyethylene
bottles and stored in ice until they were transported to the laboratory.
A sediment sample (ca. 0.5 kg) was taken for grain size analysis and
organic content.
We carefully inspected up to 20 m of the shores of every water
body investigated, searching for molluscs. Living molluscs were
searched for among the submerged vegetation, under stones, or on
the substratum. Molluscs were collected both by hand and with the
aid of sieves (0.5 mm). In addition, surrounding land was searched for
empty shells. Those sites where neither live specimens nor dead shells
were present were considered as uninhabited by molluscs. Because of
the different substrata sampled (surface, submerged and emergent
vegetation), relative abundance of molluscs was estimated by
reference to search effort (number of snails caught per hour) following
Martín et al. (2001). The collected molluscs were identified back at the
laboratory.
3.1.2. Laboratory analyses
Water samples were analyzed within fifteen days of collection for
concentrations of nitrate (NO−3), sulfate (SO2− −
4 ), chloride (Cl ), fluoride
(F−), phosphate (PO3− −2 −
4 ), carbonate (CO3 ), bicarbonate (HCO3), magne-
2+ 2+
sium (Mg ), calcium (Ca ), and silica (SiO2). Chemical analyses were
Fig. 2. Shells of the Quaternary and extant freshwater mollusc taxa identified in central-western Argentina. A: Lymnaea viator; B: Biomphalaria peregrina; C: Chilina mendozana;
D: Heleobia parchappii; E: Heleobia aff. parchappii; F: Sphaeriidae; G: Heleobia hatcheri; H: Physa acuta. Scale bars: 1 mm.
107
performed using standard methods: chloride following the Mhor
method, sulfate by turbidimetry, calcium and magnesium by com-
plexometric titrations with EDTA, potassium by flame spectrometry,
silica by the silicomolibdate method, fluoride by the zirconyl chloride
method and nitrate by the brucine method (APHA, 1992).
Sediment grain size was analyzed with the dry-sieving technique
of Folk (1968). It was quantified as the proportion of gravel (N2 mm),
coarse sand (N500 μm), medium sand (250–499 μm), fine sand (125–
249 μm), very fine sand (62–124 μm), and mud (silt and clay, b62 μm).
Additionally, the organic content of each sample was estimated using
the loss-on-ignition method (LOI) for 4 h at 550 °C and water content
(humidity) was calculated by drying the sediment for 24 h at ca. 105 °C
(Heiri et al., 2001).
Molluscs were identified to species (when possible) and their
relative abundance calculated. Gastropod identification was based on
de Castellanos and Gaillard (1981), de Castellanos and Landoni (1981),
Fernández (1981), Gaillard and de Castellanos (1976), and Rumi
(1991).
3.2. Application to the fossil record
Fossil data were obtained from previous published works. Three
different alluvial successions were included: (1) La Bomba section (LB)
(De Francesco et al., 2007), (2) Puesto Moya (PM), and (3) Puesto
Vicencio (PV) (De Francesco and Dieguez, 2006). The LB succession,
which is Pleistocene, is located in the proximity of Río Tunuyán,
whereas PM and PV, with Holocene shells, are located in southern
Mendoza (Fig. 1). All discrete levels (n = 17) containing mollusc shells
were considered.
3.2.1. La Bomba (Pleistocene)
This alluvial succession (33°28′ S, 69°03′ W) crops out on the right
margin of Arroyo La Estacada, a tributary of Río Tunuyán (in the area of
modern sites 1 and 2; Fig. 1). The succession was analyzed in detail by
108 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113

De Francesco et al. (2007). The stratigraphic section, mainly composed 4. Results

of fine sands, interbedded with levels of silty clays and organic matter,
records the interval between circa 35 14C ka B.P. and 31 14C ka B.P. and 4.1. Modern mollusc fauna
has a total height of 3 m. It consists of 14 discrete levels containing
freshwater mollusc shells, all of which were included in the present Only 25 of the 45 sites had live molluscs. In total, 3278 specimens
analysis (LB1–LB14). Shells showed an excellent preservation (Fig. 2). belonging to 8 species (7 gastropods and 1 bivalve) were recorded in
Most specimens were complete, exhibiting neither abrasion nor the studied sites. The gastropods were Lymnaea viator (Lymnaeidae),
dissolution, which suggests that they were deposited in the same Heleobia hatcheri, Heleobia parchappii, Heleobia aff. parchappii
environment where they lived. For further information on stratigra- (Cochliopidae), Chilina mendozana (Chilinidae), Biomphalaria pere-
phy and paleoecology see De Francesco et al. (2007). grina (Planorbidae), and Physa acuta (Physidae). The bivalve was
attributed to the family Sphaeriidae (Fig. 2). Recently, Ciocco (2008)
3.2.2. Puesto Moya and Puesto Vicencio (Holocene) pointed out the presence of the bivalve Pisidium chiquitanum in
These two alluvial successions crop out on the right margin of Río Mendoza, which may correspond to the same species recorded here as
Atuel in southern Mendoza, in the proximity of Laguna Llancanelo, in Sphaeriidae. However, we need to perform additional comparative
an area of broad meandering sections. The mollusc assemblages have studies in order to reliably assign these specimens to P. chiquitanum.
been analyzed in detail by De Francesco and Dieguez (2006). The PM The three commonest species were H. hatcheri, L. viator and P. acuta.
succession is late Holocene whereas the PV succession is early The species richness, Shannon diversity, Simpson's diversity, species
Holocene. Only one level from PM and two levels from PV (PV4 and equitability, and evenness indices were not significantly different
PV7) contained mollusc shells, and were included in the present between northern and southern sites (Table 1). Therefore, it can be
analysis. The PM succession (35°15′ S, 69°14′ W) has a total height of concluded that the assemblages represented in the different areas are
4.67 m and is composed of 8 sedimentary levels, of which only one not significantly different, although in many cases the taxonomic
(81–112 cm depth) contained freshwater mollusc shells. This level composition differed among sites even within the same stream.
corresponds to a hydromorphic paleosol. The PV succession (35°13′ S,
69°06′ W) has a total height of 7.84 m and is composed of 9 4.2. Environmental variables
sedimentary levels, of which two (PV4 and PV7) contained mollusc
shells. PV4 (97–207 cm) is a sedimentary bank composed of clayey The standard product-moment correlation analysis of environ-
sediments, with sand interbedded. PV7 (located at 310–338 cm) is a mental data permitted reducing the 25 variables (Table 2) to 13.
hydromorphic paleosol, similar to that in PM. Shells from PM and PV Hardness, Ca2+, Mg2+, Cl− and SO2− 4 showed a significant correlation
−
exhibited significant shell abrasion (Fig. 2). In addition, most speci- (r N 0.64, p b 0.001) with conductivity. CO2−
3 , and HCO3 were correlated
mens were fragmented, suggesting some degree of post-mortem with pH (r N 0.51, p b 0.001). Coarse sand was correlated with gravel
transportation. Therefore, assemblages from PM and PV probably and very fine sand (r N 0.62, p b 0.001). Medium sand was correlated
represent deposits transported by currents that were dropped on the with fine sand (r = 0.58, p b 0.001). Mud was correlated with gravel,
flood plain. For further information on stratigraphy and paleoecology fine sand, very fine sand, humidity, and LOI (r N 0.61, p b 0.001).
see De Francesco and Dieguez (2006). Therefore, the reduced matrix included conductivity, pH, temperature,
−
depth, current velocity, vegetation cover, NO2− 3−
3 , PO4 , F , SiO2, coarse
3.3. Data analysis sand, medium sand, and mud.
The first two components of the PCA ordination (Fig. 3A) accounted
Standard product-moment correlation analyses were conducted to for 63.8% of the variation in the data. The first axis explained 44.1% of
identify strongly intercorrelated environmental variables, permitting the total variation and exhibited a very high positive correlation
some of them to be omitted from subsequent statistical analyses. (r = 0.99) with vegetation cover (Fig. 3B) and, to a lesser extent, with
Principal Component Analysis (PCA) was used for ordination of SiO2, medium sand, and mud (r ~ 0.37). In addition, this axis was
sampling sites based on the reduced set of environmental variables negatively correlated with conductivity (r = −0.44), current velocity
measured. Environmental data were log transformed, as log (x + 1). (r = −0.32) and depth (r = −0.20). The second axis, which explained
For each site inhabited by molluscs, the species richness (S), 19.7% of total variation, was positively correlated with mud (r = 0.81),
Shannon diversity index (H′), Simpson's diversity index (D), species depth (r = 0.44), and NO2− 3 (r = 0.36), and negatively with coarse sand
evenness (E), and equitability (J) were calculated. In order to test for (r = 0.87) and medium sand (r = 0.50). The first axis allowed differ-
differences between northern and southern sites (Río Tunuyán versus entiating sites inhabited by molluscs from uninhabited ones. In fact,
Río Atuel basins), Mann–Whitney tests were performed. All these the sites inhabited by molluscs were located on the right side of the
analyses were carried out with the computer program PAST v 1.81 plot, and characterized as shallow (b1 m) highly vegetated water
(Hammer et al., 2008). bodies with low conductivity (b1.9 mS cm− 1) and nil or low current
To explore the relationships between environmental variables and velocity (0–0.7 m s− 1). In addition, these sites displayed finer
the mollusc assemblages, a Canonical Correspondence Analysis (CCA) sediments and were more enriched in SiO2. On the other hand,
was performed. A series of partial CCAs, run with one explanatory those sites characterized by nil or low vegetation cover, deeper
variable at a time, was used to separate the total variation in mollusc
abundance into components that represent the unique contributions
Table 1
of individual environmental variables, the contribution of covariance
Mean values, standard deviations (SD) and range of the species richness (S), Shannon
between variables and the unexplained variance (Bocard et al., 1992). diversity index (H′), Simpson diversity index (D), evenness (E), and equitability (J) in
Statistical significance was assessed by unrestricted Monte Carlo tests northern (n = 12) versus southern (n = 13) sites (n. s.: not significant)
(full model) involving 999 permutations at p ≤ 0.001.
Site Northern sites Southern sites Mann– p
Fossil mollusc assemblages were combined with the modern data Whitney
Mean ± SD Range Mean ± SD Range
set by means of Detrended Correspondence Analysis (DCA). In order to (U)
define groups of major similarities, a Cluster Analysis (CA) based on Richness (S) 2.42 ± 1.08 1–4 2.23 ± 1.09 1–4 70 0.68 (n.s.)
the Bray–Curtis similarity measure was performed with the program Shannon (H′) 0.40 ± 0.31 0.00–0.99 0.37 ± 0.39 0.00–0.95 60 0.34 (n.s.)
PAST ver. 1.81 (Hammer et al., 2008). All ordinations (except CA) were Simpson (D) 0.23 ± 0.20 0.00–0.58 0.22 ± 0.24 0.00–0.58 76 0.93 (n.s.)
Evenness (E) 0.73 ± 0.23 0.42–1.00 0.76 ± 0.19 0.54–1.00 68 0.60 (n.s.)
performed using the program CANOCO version 4.5 (ter Braak and
Equitability (J) 0.40 ± 0.32 0.00–0.99 0.35 ± 0.33 0.00–0.71 70 0.68 (n.s.)
Šmilauer, 1998).
Table 2
Values obtained for environmental variables in the 45 sampling sites. Cond: conductivity (mS cm− 1), T: water temperature (°C), Depth (cm), Cu: current velocity (m s− 1), Veg: vegetation cover, Hard: hardness (mg l− 1 of CaCO3), Hum: humidity
(%), LOI: organic content (%). Categories of grain size (gravel, CS: coarse sand, MS: medium sand, FS: fine sand, VFS: very fine sand, and mud) are expressed in %. Concentrations of ions are expressed in mg l− 1. Altitude is expressed in meters
above sea level

Site Altitude Cond pH T Depth Cu Veg Cl− SO2−

4 NO−3 PO3−
4 F− CO2−
3 HCO−3 SiO2 Ca2+ Mg2+ Hard Hum LOI Gravel CS MS FS VFS Mud
1—La Estacada 860.1 1.15 8.16 21.2 33 0.30 0 66.0 610.0 14.80 0.21 0.54 0.0 216.3 46.4 44.0 64.3 378.0 19.98 1.90 76.24 2.77 1.19 2.97 6.23 10.59
2—Puente El Zampal 923.9 1.41 8.24 16.7 34 0.30 0 112.0 795.0 7.93 0.10 1.69 0.0 370.0 63.3 49.7 125.7 648.0 21.67 2.02 76.24 2.77 1.19 2.97 6.23 10.59
3—Puente Roto 936.8 1.40 8.31 25.8 43 0.60 0 112.0 650.0 10.00 0.12 1.70 0.0 216.3 34.0 38.3 114.0 571.0 20.01 1.65 76.24 2.77 1.19 2.97 6.23 10.59

C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
4—Arroyo Guajardino 927.3 1.01 8.05 20.7 27 0.55 1 66.0 455.0 11.34 0.04 1.93 0.0 340.0 18.5 35.6 48.7 292.0 20.80 1.19 41.51 5.31 11.42 18.51 11.76 11.47
5—Guajardino (nac.) 1080.3 0.33 8.25 13.5 6 0.00 2 13.1 123.0 6.80 0.01 1.45 0.0 123.6 10.7 10.0 38.1 184.0 57.87 7.87 0.00 0.00 25.55 25.57 24.02 24.85
6—Río Las Tunas 1143.9 0.30 8.25 22.8 25 1.50 0 13.1 127.0 0.00 0.00 0.67 0.0 69.5 10.8 16.8 34.0 184.0 8.35 1.32 74.56 15.08 8.88 1.33 0.13 0.02
7—Torrecillas 1 1178.5 0.17 8.10 16.7 32 1.00 1 19.8 19.6 0.50 0.00 1.12 0.0 108.1 12.6 12.0 17.7 104.1 20.00 1.70 63.35 9.51 7.72 9.88 5.12 4.42
8—Río Las Tunas 2 927.3 0.10 7.89 19.9 125 0.00 2 59.3 376.0 0.50 0.00 2.89 0.0 340.0 62.8 38.9 1.6 606.0 42.65 4.70 0.00 0.76 1.60 12.75 37.85 47.02
9—Río Las Tunas 3 908.4 0.64 8.06 18.2 40 1.00 2 26.3 280.0 0.50 0.00 1.76 0.0 231.7 48.0 34.9 68.5 373.0 40.00 5.00 0.00 0.76 1.60 12.75 37.85 47.02
10—Arroyo medio 1 932.9 1.69 7.91 18.8 15 0.00 2 85.7 20.2 0.50 0.94 2.15 0.0 494.4 86.5 103.5 200.0 1095.0 40.97 7.99 17.95 8.23 18.93 17.84 16.19 20.85
11—Arroyo medio 2 932.9 0.75 8.05 17.1 150 0.27 2 39.5 286.0 5.35 0.22 1.24 0.0 239.4 56.3 22.0 63.8 321.0 68.61 5.31 0.00 0.93 13.48 24.31 23.09 38.19
12—Torrecillas 2 943.6 0.51 8.16 15.6 80 0.27 1 29.6 176.0 5.35 0.00 1.48 0.0 216.3 42.5 22.2 79.0 385.0 27.01 2.37 0.00 0.00 0.05 2.16 26.40 71.39
13—Vista Flores 965.4 1.21 7.87 16.0 6 0.10 1 145.0 525.0 0.50 0.21 0.70 0.0 409.4 61.4 33.8 88.0 451.0 41.78 7.81 3.79 1.84 6.88 19.93 19.30 48.25
14—Río Tunuyán 893.0 0.82 8.22 18.2 200 1.50 0 102.2 393.0 5.50 1.18 1.03 0.0 154.5 16.5 21.2 68.0 336.0 20.81 0.85 52.15 0.29 2.13 17.04 18.34 10.05
15—Arroyo Guiñazu 868.2 0.74 8.10 16.5 49 0.43 2 26.3 324.0 0.00 2.54 1.54 0.0 301.2 48.7 23.2 43.6 240.0 37.89 4.27 7.80 10.23 20.14 26.36 16.54 18.92
16—Arroyo Caroca 871.5 0.81 7.88 17.0 75 1.00 2 59.3 204.5 0.50 0.00 2.38 0.0 224.0 54.6 28.2 83.8 420.0 46.22 3.53 8.05 5.21 14.61 19.73 18.43 33.96
17—Acequia Claro 874.4 0.76 8.10 16.7 15 0.67 2 52.7 202.7 3.75 1.56 1.51 0.0 193.1 53.0 23.2 27.6 173.0 35.80 2.55 8.80 6.91 13.27 25.46 19.72 25.83
18—Arroyo Claro 874.4 1.18 7.93 18.1 50 0.67 1 115.4 481.0 2.46 1.89 1.91 0.0 278.1 52.8 28.5 50.4 281.4 35.80 2.55 8.80 6.91 13.27 25.46 19.72 25.83
19—Arroyo Yaucha 973.9 0.21 7.90 10.1 43 1.35 0 9.9 46.7 3.65 0.40 0.00 0.0 231.7 20.0 17.2 28.1 160.0 20.74 0.88 0.00 0.57 15.70 50.96 24.47 8.29
20—Presa del Tigre 914.5 0.71 7.71 19.0 20 0.38 2 85.7 315.0 0.00 0.17 1.62 0.0 131.3 12.7 20.5 63.1 314.0 9.92 1.00 74.56 15.08 8.88 1.33 0.13 0.02
21—Arroyo Gaby 863.4 0.76 8.17 19.3 85 0.16 2 98.9 117.0 11.70 0.00 1.17 0.0 168.0 11.6 35.1 58.8 333.0 24.34 3.95 0.45 3.74 7.56 35.08 30.52 22.64
22—Presa Nihuil 1300.5 1.16 8.04 18.3 20 0.00 2 165.0 67.8 3.75 0.00 1.17 0.0 168.0 11.0 76.3 105.6 630.0 10.55 1.15 74.56 15.08 8.88 1.33 0.13 0.02
23—Los Molles 1938.1 0.79 7.81 21.6 26 0.00 2 115.4 68.8 4.97 0.00 1.55 0.0 594.0 34.5 31.8 58.4 323.0 49.89 4.32 23.58 19.35 15.14 13.62 10.15 18.16
24—Río Salado 1932.8 2.05 7.85 14.9 150 1.00 0 646.4 17.8 3.23 0.00 1.45 0.0 92.7 9.7 62.4 52.5 375.0 22.80 1.74 0.00 0.59 16.21 57.20 14.21 11.78
25—Vertiente Molles 1956.4 0.62 7.47 14.5 3 1.00 0 214.0 11.2 0.96 0.00 0.64 0.0 610.0 16.5 62.4 43.6 338.0 4.67 1.21 94.81 3.18 1.05 0.69 0.18 0.08
26—Niña Encantada 2092.9 0.76 7.72 11.7 55 0.67 2 26.3 392.0 12.8 0.00 0.70 0.0 231.0 28.5 133.0 1.08 337.0 47.58 3.33 23.58 19.35 15.14 13.62 10.15 18.16
27—Llancanelo 1336.1 8.68 7.87 19.9 25 0.00 0 427.4 1670.0 1.41 0.00 2.42 0.0 139.0 5.6 90.0 89.7 598.0 34.19 2.95 0.27 5.05 8.45 21.52 23.08 41.63
28—Agua Botada 1969.4 0.59 7.92 19.9 10 0.75 0 13.2 174.0 0.00 0.00 2.35 0.0 471.0 41.3 16.0 53.2 262.0 9.92 0.65 61.60 26.34 8.14 3.30 0.41 0.21
29—Bañados Grande 1593.4 1.05 7.34 14.3 15 0.00 2 58.2 253.2 0.00 0.00 0.76 0.0 410.0 35.5 25.2 97.0 467.3 67.52 8.45 11.94 13.47 26.31 26.58 12.49 9.20
30—Río Malargüe 1538.1 0.55 8.00 16.5 150 0.35 0 49.4 156.0 0.00 0.17 0.75 0.0 285.0 42.6 27.2 42.2 244.0 21.33 2.23 63.34 9.51 7.73 9.88 5.12 4.42
31—Vertiente Malargüe 1537.6 0.72 7.93 16.0 5 0.10 2 27.0 124.0 0.00 0.00 1.30 0.0 278.0 38.0 13.3 51.5 248.0 26.66 2.23 0.00 1.61 16.60 26.89 26.12 28.78
32—El Chacay 1425.4 0.34 8.10 11.2 28 1.10 2 10.0 114.0 0.00 0.00 0.40 0.0 162.2 17.0 5.97 44.0 198.0 7.61 0.96 64.81 8.08 6.85 10.13 4.29 5.83
33—Laguna Blanca 1637.8 3.97 8.04 15.6 60 0.00 0 465.0 90.0 0.00 0.00 4.88 0.0 84.0 21.2 43.0 356.0 1546.6 19.46 0.94 22.20 8.31 34.65 34.39 0.22 0.21
34—Arroyo Sosneado 1612.8 0.79 8.11 14.3 14 0.17 2 39.5 256.0 0.00 0.00 0.69 0.0 160.0 34.4 27.2 83.2 415.0 24.49 2.38 26.36 13.68 19.22 24.04 9.41 7.29
35—Vertiente Sosneado 1790.3 0.80 8.24 19.3 7 0.17 2 26.3 342.0 0.00 0.08 1.45 0.0 226.0 50.7 36.5 60.5 343.6 17.09 1.69 26.69 11.64 12.69 26.69 13.14 7.14
36—Laguna Sosneado 1 2112.6 0.22 9.40 22.3 20 0.10 2 46.1 79.5 0.00 0.13 0.50 50.8 58.6 43.7 17.2 24.3 144.4 56.09 5.58 0.00 1.04 10.99 28.30 34.54 25.13
37—Laguna Sosneado 2 2137.2 0.22 9.00 15.9 50 0.00 2 49.4 58.1 0.00 0.00 0.00 50.2 67.0 32.6 18.0 26.2 154.4 64.74 5.84 3.24 7.12 14.88 22.90 25.12 26.74
38—Vertiente Cañon 934.3 2.11 8.19 18.9 10 0.17 0 178.1 690.0 11.86 0.60 1.61 0.0 595.0 17.8 21.3 69.7 344.2 17.16 1.18 67.28 27.56 3.68 0.71 0.31 0.46
39—Río Atuel 1068.2 1.47 7.55 19.8 50 0.67 0 185.0 692.0 0.00 0.00 0.61 0.0 243.0 22.2 33.6 172.0 800.3 10.62 1.30 44.37 40.82 11.74 2.21 0.39 0.48
40—Salinas Diamante 1290.5 30.00 7.55 16.1 15 0.00 0 30000 51530 0.00 0.00 1.88 0.0 150.8 13.2 260.0 1858.5 8391.6 20.38 1.37 5.27 7.28 6.86 18.78 41.04 20.77
41—Las Aguaditas 552.9 1.86 7.81 15.5 50 0.10 2 335.7 680.0 1.37 0.00 0.52 0.0 360.0 37.8 22.6 200.6 892.6 21.12 3.52 38.26 17.07 18.17 14.77 5.67 6.06
42—Monte Comán 523.9 1.40 8.11 18.4 150 0.67 0 234.1 4775.0 2.51 0.05 0.84 0.0 234.6 36.6 34.0 72.6 387.6 27.31 1.78 0.00 0.00 1.82 28.27 49.10 20.81
43—Salina La Horqueta 411.1 9.83 8.26 20.8 35 0.00 0 2440.0 4405.0 0.00 0.27 0.05 0.0 142.0 10.0 51.8 324.0 1481.5 15.94 1.04 9.24 16.30 7.54 34.95 22.54 9.42
44—Río Desaguadero 410.1 1.70 8.17 19.9 150 0.40 0 442.0 118.0 0.00 0.30 0.00 0.0 217.8 16.4 29.8 158.4 734.5 27.06 1.95 0.00 0.00 0.05 2.36 29.79 67.80
45—Arroyo Bebedero 402.7 16.00 7.66 20.6 20 0.00 0 7717.0 2140.0 0.00 0.02 1.39 0.0 150.8 72.7 345.0 695.0 3764.0 32.95 5.44 0.00 1.68 32.62 21.54 23.04 21.12

109
110 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113

Fig. 3. (A) Principal components analysis (PCA) biplot for sites and environmental variables. Sites inhabited by molluscs are indicated by white circles, uninhabited sites by black
circles (B) Correlations of environmental variables with the first component of the PCA.

(N1.5 m), or displaying higher conductivity (N4 mS cm− 1) and current sediments. In addition, they appeared to be related to higher tempera-
velocity (N1 m s− 1) were uninhabited by molluscs and located on the tures (13.5–22.3 °C) and higher depths (6–125 cm). On the other hand,
left side of the PCA plot. The unique sites with high conductivity values Heleobia hatcheri, Chilina mendozana, Heleobia aff. parchappii and the
(8–16 mS cm− 1) that contained live snails were represented by Laguna sphaeriids dominated in less vegetated lotic water bodies with coarser
Llancanelo (27) and Arroyo Bebedero (45). Both sites contained only sediments and lower pH (7.3–8.2). In addition, these species appeared to
specimens of the mud snail Heleobia parchappii, not represented at be more tolerant to slightly higher conductivity (0.5–1.2 mS cm− 1), and
any other site. As these two sites represent outliers from the modern lower temperatures (14.3–19 °C).
data set (because they represent brackish water conditions) they were
discarded from subsequent statistical analyses. Although site 25
displayed low conductivity, it was located on the left side of the plot
mainly because of the absence of vegetation and the exclusively
dominance of hard substrata (boulders).

4.3. Relationships between mollusc assemblages and environmental

variables

The variables of the reduced matrix obtained from standard

product-moment correlation analyses were used as representatives of
the environmental data to relate to the abundance of molluscs in the
CCA (Fig. 4). The 13 variables explained 78% of variation in the data.
The unrestricted Monte Carlo test for all canonical axes was
significant (F = 2.46; p b 0.001). Partial CCAs showed that the total
explained variance was mainly composed of pH (18.2%), NO−3 2 (12.3%),
conductivity (11.8%), temperature (10.7%), F− (9.8%), mud (6.9%), coarse
sand (6.9%), depth (6.7%), vegetation cover (6.6%), SiO2 (6.4%), and
current speed (4.8%). The first two axes of the CCA accounted for 43.7%
of variation in the abundance of molluscs and 55.9% of the species–
environment relationship (Table 3). The species–environment corre-
lations of CCA axis 1 (r = 0.96) and axis 2 (r = 0.95) were high and
indicated a strong relationship of mollusc abundance to environ-
mental variables.
The CC1× CC2 plot showed that the species Lymnaea viator, Physa
Fig. 4. Canonical correspondence analysis (CCA) ordination plot showing the relationship
acuta, and Biomphalaria peregrina dominated in highly vegetated sites between sampling sites (white circles), species (black squares) and environmental
−1
characterized by high pH (7.7–9.4) and NO2− 3 (0.5–12.8 mg l ), on fine variables (arrows). For variable abbreviations see Table 1.
 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113 111

Table 3 These sites represent streams dominated by coarse sediments (mainly

Summary statistics for the first two axes of CCA, with the 13 selected environmental boulders), moderate conductivity (0.87 ± 0.26 mS cm− 1), and mean
factors
depths of 45 cm. They were dominated by Heleobia aff. parchappii and
CCA axes Axis 1 Axis 2 H. hatcheri. On the other hand, fossil level PV4 (located above PV7) was
Eigenvalues (λ) 0.766 0.697 similar to modern sites 26, 34 and 20. These sites represent shallower
Species–environment correlations 0.959 0.954 vegetated streams (mean depth= 29 cm) characterized by moderate
Cumulative % variance
water velocity (0.41 ± 0.25 m s− 1) and lower conductivity (0.75 ± 0.04 mS
– Of species abundance 22.9 43.7
– Of species–environment relationship 29.3 55.9 cm− 1). They were dominated by H. hatcheri, C. mendozana and
Total inertia 3.35 Sphaeriidae.
Sum of all canonical eigenvalues 2.61 Most levels from the LB section (LB1–LB3, LB7–LB9; LB10, LB12,
LB13) and PM were similar to modern site 36 (Laguna El Sosneado). This
freshwater vegetated shallow lake is located in the proximities of the
High Cordillera (Fig. 1) and, therefore, is an environment with very low
4.4. Application to the fossil record anthropogenic impact. Snails (mostly L. viator) were found exclusively
on the submerged vegetation present in a very shallow area (20 cm
In total, 4831 fossil shells were combined with the modern data set. deep) where a small stream flows into the lake. The remaining levels
According to the cluster analysis and DCA, 7 groups were recognized, of from LB (LB4–LB6, LB11, LB14) did not have a modern analogue in the
which only 3 included both living and fossil samples (Fig. 5). Results present model. These levels were dominated by B. peregrina, a species
showed that fossil level PV7 was similar to modern sites 9, 16 and 22. that did not dominate in any modern site.

Fig. 5. (A) Detrended correspondence analysis (DCA) ordination plot of modern (white circles) and Quaternary (black circles) mollusc samples. (B) Cluster analysis (CA) of modern and
Quaternary mollusc samples based on Bray–Curtis similarity measures. Groups defined by CA at similarities N 0.5 are encircled in DCA. PV: Puesto Vicencio (Holocene), PM: Puesto
Moya (Holocene), LB: La Bomba (Pleistocene).
112 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
5. Discussion and conclusions
The PCA clearly discriminated between sites inhabited by molluscs
and uninhabited sites. The former was characterized by high
vegetation cover, low conductivity and current speed as well as
predominance of fine sediments (mainly mud). Depth, substrate, flow,
and vegetation have been considered important in explaining snail
distributions (see Dillon, 2000 and references therein). Pulmonate
snails are primarily adapted to calmer water, with few species
inhabiting lotic environments, their occurrence in rivers restricted to
calm backwaters and eddies (Dillon, 2000). Therefore, molluscs in the
semiarid region of central-western Argentina occur primarily in calm,
vegetated, shallow water bodies. With the exception of a few species
that can tolerate high conductivity such as Heleobia parchappii (see
De Francesco and Isla, 2004), assemblages are also confined to low
salinities (below 1.9 mS cm− 1).
Heleobia parchappii was the only mollusc species recorded in saline
waters (8–16 mS cm− 1). In similar saline habitats in central-western
Argentina, Doering (1884) reported Heleobia occidentalis. Based on
their similar shell morphology, Gaillard and de Castellanos (1976)
suggested that H. occidentalis could be a geographical variety of
H. parchappii adapted to saline waters. De Francesco (2007) suggested
that they could be synonyms. Ciocco and Scheibler (2008) found no
conchological differences between the two species. We did not find
any significant morphological or anatomical (penial structure)
difference between them concluding that they may be synonyms. In
the Pampean region, H. parchappii occurs both in fresh and saline
water. However, when the species occurs in saline waters it is the only
mollusc species present (De Francesco and Isla, 2004; Peretti, 2005)
while in fresh water it is usually accompanied by other mollusc
species. Ciocco and Scheibler (2008) found that along a gradient of
conductivity in the Bañado Carilauquen, H. parchappii coexisted with
other species only in the headwaters and middle reaches at low
salinities (0.9–1.6 mS cm− 1) but was the only species found in the
extremely saline waters (8.0–19.5 mS cm− 1) of the outlet into the lake.
Therefore, although we cannot morphologically distinguish the
populations of H. parchappii in fresh water from those inhabiting
saline waters, the composition of the assemblage may be indicative of
these two sorts of habitats, which has important implications for the
value of freshwater mollusc assemblages as indicators of past
continental salinities.
Although the shell morphology of Heleobia aff. parchappii is different
from typical H. parchappii, the anatomy of the penial complex is similar.
We suggest that this species may be a variety of H. parchappii. It is
known that Heleobia displays a significant phenotypic plasticity in shell
morphology (De Francesco, 2007). Another possibility is that this taxon
is an undescribed species, ecologically similar to H. parchappii. A third
possibility is that this taxon corresponds to Heleobia kuesteri as recently
suggested by Masi and Ciocco (2008). Heleobia kuesteri is a problematic
species that was originally described for Mendoza, of which there is no
updated taxonomic or ecological information. In fact, it has been
considered as species inquirenda (Cazzaniga, 1981) because of the
difficulties for assessing its taxonomic identity. Although we cannot be
sure of the taxonomic identity of this taxon with the scattered
observations presented here, the presence of this same morphology
as fossil in the area (it was published as Heleobia sp. or Heleobia cf. H.
australis by De Francesco and Dieguez, 2006) allows their utility as a
reliable bioindicator with important consequences for the reconstruc-
tion of past environments. In other words, despite its taxonomic
identity the ecological characteristics of this taxon are still an
informative indicator of the original depositional environment.
The CCA clearly indicated a strong relationship between mollusc
taxa and environmental variables. Two main groups can be recognized:
(Group 1) the species L. viator, P. acuta, and B. peregrina dominated in
highly vegetated sites characterized by high pH and NO2− 3 , and lower
current velocity, on fine sediments, and (Group 2) the two Heleobia
species (H. hatcheri and H. aff. parchappii) together with C. mendozana,
and the sphaeriids dominated in less vegetated lotic water bodies with
higher current velocity and coarser sediments, as well as lower pH. In
addition, these species appeared to be more tolerant to slightly higher
conductivity. Even though the waters of the Bañado Carilauquen are
clear and variations in pH scarcely significant, Ciocco and Scheibler
(2008) also observed a predominance of L. viator and B. peregrina at
higher pH, and in turbid waters, whereas C. mendozana and H. hatcheri
dominated in more clear and oxygenated waters, with high HCO−3. These
results are in agreement with previous studies that suggested the
occurrence of L. viator and B. peregrina in lentic bodies (shallow
lakes, ponds) of areas containing submerged aquatic vegetation
(de Castellanos and Landoni, 1981; Rumi, 1991) as well as the presence
of H. hatcheri and C. mendozana in rivers and streams (Gaillard and de
Castellanos, 1976; de Castellanos and Gaillard, 1981). Thus, from a
paleoecological standpoint, we conclude that these two assemblages
are indicative of different energetic environments related to the
stability of the flow regime. The assemblage dominated by L. viator, P.
acuta and B. peregrina represents lentic habitats (shallow lakes) or calm
backwaters (pools) within lotic systems such as streams and rivers; the
assemblage dominated by H. hatcheri, H. aff. parchappii, C. mendozana,
and Sphaeriidae indicates lotic habitats where a directional water flow
(of variable velocity) exists.
The two main groups observed in the modern dataset were
recognized in the fossil record. According to the DCA, the mollusc
assemblages preserved in Puesto Vicencio (PV7 and PV4) are included in
the group 2 mentioned above. Consequently, they represent shallow
vegetated streams characterized by moderate current velocity and low
conductivity. According to the clusters observed, a subtle difference
between both levels can be inferred. This is due to the occurrence of
some L. viator specimens in PV4, which may be consequence of
taphonomic alteration. According to the poor preservation of these
shells (in most cases, the weathered shells simply disintegrate when
touched), it is probable that they represent a mixed assemblage,
including specimens that would have been deposited after some
transport by currents. Kotzian and Simões (2006) found that small
scale lateral transport of shells does occur in fluvial systems. The
occurrence of L. viator here suggests shallow water bodies probably
connected to the main lotic system. This mixed assemblage probably
represents a taphocoenosis deposited during flooding events. Moreover,
the whole PV succession may reflect the variations in water flow that
occurred in the floodplain during the Holocene in Río Atuel. An
important conclusion is that the molluscs preserved in this sort of
alluvial succession represent mixed assemblages that suffered some
degree of taphonomic alteration. Because of the natural dynamics of the
fluvial environment, it is expected that molluscs usually be subject to
transport and redeposition.
The LB and PM successions represent lentic habitats with nil or very
low water velocity (Group 1). These habitats would have been vegetated
shallow lakes within the floodplain and not connected to the fluvial
system. The excellent preservation of the shells from LB supports this
interpretation (see De Francesco et al., 2007). As L. viator inhabits the
submerged vegetation in very shallow areas where small streams flow
into the lake, shells may be deposited in situ and, thus, represent
parautochthonous assemblages. Some LB levels (LB4–LB6, LB11, LB14),
dominated by B. peregrina, did not have a modern analogue in the
present model. However, B. peregrina is widely distributed in shallow
lakes from La Pampa and Buenos Aires Province (Rumi, 1991; Peretti,
2005), regions characterized by higher humidity than the region
studied here. Thus, these LB levels probably indicate relatively wetter
conditions, which agrees with the previous interpretation, based on the
whole assemblage together with the isotopic evidence, that the studied
section records a mild interval corresponding to an interstadial (Marine
isotope stage 3) of the last glacial cycle (De Francesco et al., 2007).
Although the results obtained here indicated that mollusc species
are useful for inferring past habitats in the semiarid region of
 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
Mendoza, the presence of species not recorded as fossils in the area (H.
hatcheri and P. acuta) introduces a complication in using the modern
assemblage to infer paleoenvironments. P. acuta (= P. cubensis) is an
introduced species widely distributed in Argentina (Núñez and
Pelichotti, 2003). This species has a superior reproductive capacity
compared to other pulmonate snails, as well as ability to migrate
upstream and to quickly recolonize a water-body (see de Cock and
Wolmarans, 2007 and references therein). Consequently, it may have a
negative impact on indigenous freshwater molluscs in particular and
on the biodiversity of freshwater habitats in general. One important
caveat regarding paleoenvironmental interpretations is the lack of
knowledge of the ecological role played in the past by other modern
coexisting species such as L. viator and B. peregrina. It is likely that, in
the absence of P. acuta, these species would have occupied a different
range of habitats or, eventually, experienced an increase in their
distribution. Similarly, the absence of H. hatcheri in the past suggests
that the species colonized Mendoza in recent times (in fact, the
species was not recorded as fossil in Mendoza even in younger alluvial
deposits of up to ca. 500 years B.P.; De Francesco and Zárate, 2006), so
the habitats they occupy today would have been absent in the past or
occupied by a different species, which cannot be ascertained. Even
though the total biodiversity has suffered changes in recent times,
many of them probably as a consequence of human activities, the good
news is that mollusc assemblages still constitute reliable bioindicators
of past environmental conditions and can be used with confidence.
Acknowledgements
Financial support for this study was provided by Agencia Nacional
de Promoción Científica y Tecnológica (PICT 16-20706). We thank
Adolfo Gil, Gustavo Neme, Miguel Giardina, and Alejandra Guerci
(Museo de Historia Natural de San Rafael) for support during field
work. We acknowledge Gustavo Bernava for chemical analyses, and
Mauricio Quiroz for providing data on altitude. Robert Cowie and
Néstor Ciocco improved the manuscript with constructive reviews.
The authors are members of Consejo Nacional de Investigaciones
Científicas y Técnicas (CONICET).
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