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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.
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
Fig. 1. Location map of modern (circles) and Quaternary (squares) study sites in central-western Argentina.
C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113 107
(anthropogenic impact) from almost pristine environments (Andes performed using standard methods: chloride following the Mhor
Cordillera) to streams running across cities or through areas of low to method, sulfate by turbidimetry, calcium and magnesium by com-
moderate cattle-raising and agriculture (basin of Río Tunuyán). plexometric titrations with EDTA, potassium by flame spectrometry,
At each site, we quantified current velocity, water temperature, pH, silica by the silicomolibdate method, fluoride by the zirconyl chloride
and conductivity. Current velocity was measured using a neutrally method and nitrate by the brucine method (APHA, 1992).
buoyant sphere and calculating the time it took the sphere to move Sediment grain size was analyzed with the dry-sieving technique
5 m. Temperature, pH, and conductivity were measured with field of Folk (1968). It was quantified as the proportion of gravel (N2 mm),
instruments. Aquatic vegetation cover at each sampling site was coarse sand (N500 μm), medium sand (250–499 μm), fine sand (125–
estimated visually, and a nominal variable erected for statistical 249 μm), very fine sand (62–124 μm), and mud (silt and clay, b62 μm).
purposes (0 = no vegetation, 1 = low vegetation cover, and 2 = high Additionally, the organic content of each sample was estimated using
vegetation cover). the loss-on-ignition method (LOI) for 4 h at 550 °C and water content
One subsurface water sample (1 l) was taken at each site to assess (humidity) was calculated by drying the sediment for 24 h at ca. 105 °C
chemical parameters. Water samples were collected in polyethylene (Heiri et al., 2001).
bottles and stored in ice until they were transported to the laboratory. Molluscs were identified to species (when possible) and their
A sediment sample (ca. 0.5 kg) was taken for grain size analysis and relative abundance calculated. Gastropod identification was based on
organic content. de Castellanos and Gaillard (1981), de Castellanos and Landoni (1981),
We carefully inspected up to 20 m of the shores of every water Fernández (1981), Gaillard and de Castellanos (1976), and Rumi
body investigated, searching for molluscs. Living molluscs were (1991).
searched for among the submerged vegetation, under stones, or on
the substratum. Molluscs were collected both by hand and with the 3.2. Application to the fossil record
aid of sieves (0.5 mm). In addition, surrounding land was searched for
empty shells. Those sites where neither live specimens nor dead shells Fossil data were obtained from previous published works. Three
were present were considered as uninhabited by molluscs. Because of different alluvial successions were included: (1) La Bomba section (LB)
the different substrata sampled (surface, submerged and emergent (De Francesco et al., 2007), (2) Puesto Moya (PM), and (3) Puesto
vegetation), relative abundance of molluscs was estimated by Vicencio (PV) (De Francesco and Dieguez, 2006). The LB succession,
reference to search effort (number of snails caught per hour) following which is Pleistocene, is located in the proximity of Río Tunuyán,
Martín et al. (2001). The collected molluscs were identified back at the whereas PM and PV, with Holocene shells, are located in southern
laboratory. Mendoza (Fig. 1). All discrete levels (n = 17) containing mollusc shells
were considered.
3.1.2. Laboratory analyses
Water samples were analyzed within fifteen days of collection for 3.2.1. La Bomba (Pleistocene)
concentrations of nitrate (NO−3), sulfate (SO2− −
4 ), chloride (Cl ), fluoride This alluvial succession (33°28′ S, 69°03′ W) crops out on the right
(F−), phosphate (PO3− −2 −
4 ), carbonate (CO3 ), bicarbonate (HCO3), magne- margin of Arroyo La Estacada, a tributary of Río Tunuyán (in the area of
2+ 2+
sium (Mg ), calcium (Ca ), and silica (SiO2). Chemical analyses were modern sites 1 and 2; Fig. 1). The succession was analyzed in detail by
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.
108 C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113
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).
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 species (H. hatcheri and H. aff. parchappii) together with C. mendozana,
and the sphaeriids dominated in less vegetated lotic water bodies with
The PCA clearly discriminated between sites inhabited by molluscs higher current velocity and coarser sediments, as well as lower pH. In
and uninhabited sites. The former was characterized by high addition, these species appeared to be more tolerant to slightly higher
vegetation cover, low conductivity and current speed as well as conductivity. Even though the waters of the Bañado Carilauquen are
predominance of fine sediments (mainly mud). Depth, substrate, flow, clear and variations in pH scarcely significant, Ciocco and Scheibler
and vegetation have been considered important in explaining snail (2008) also observed a predominance of L. viator and B. peregrina at
distributions (see Dillon, 2000 and references therein). Pulmonate higher pH, and in turbid waters, whereas C. mendozana and H. hatcheri
snails are primarily adapted to calmer water, with few species dominated in more clear and oxygenated waters, with high HCO−3. These
inhabiting lotic environments, their occurrence in rivers restricted to results are in agreement with previous studies that suggested the
calm backwaters and eddies (Dillon, 2000). Therefore, molluscs in the occurrence of L. viator and B. peregrina in lentic bodies (shallow
semiarid region of central-western Argentina occur primarily in calm, lakes, ponds) of areas containing submerged aquatic vegetation
vegetated, shallow water bodies. With the exception of a few species (de Castellanos and Landoni, 1981; Rumi, 1991) as well as the presence
that can tolerate high conductivity such as Heleobia parchappii (see of H. hatcheri and C. mendozana in rivers and streams (Gaillard and de
De Francesco and Isla, 2004), assemblages are also confined to low Castellanos, 1976; de Castellanos and Gaillard, 1981). Thus, from a
salinities (below 1.9 mS cm− 1). paleoecological standpoint, we conclude that these two assemblages
Heleobia parchappii was the only mollusc species recorded in saline are indicative of different energetic environments related to the
waters (8–16 mS cm− 1). In similar saline habitats in central-western stability of the flow regime. The assemblage dominated by L. viator, P.
Argentina, Doering (1884) reported Heleobia occidentalis. Based on acuta and B. peregrina represents lentic habitats (shallow lakes) or calm
their similar shell morphology, Gaillard and de Castellanos (1976) backwaters (pools) within lotic systems such as streams and rivers; the
suggested that H. occidentalis could be a geographical variety of assemblage dominated by H. hatcheri, H. aff. parchappii, C. mendozana,
H. parchappii adapted to saline waters. De Francesco (2007) suggested and Sphaeriidae indicates lotic habitats where a directional water flow
that they could be synonyms. Ciocco and Scheibler (2008) found no (of variable velocity) exists.
conchological differences between the two species. We did not find The two main groups observed in the modern dataset were
any significant morphological or anatomical (penial structure) recognized in the fossil record. According to the DCA, the mollusc
difference between them concluding that they may be synonyms. In assemblages preserved in Puesto Vicencio (PV7 and PV4) are included in
the Pampean region, H. parchappii occurs both in fresh and saline the group 2 mentioned above. Consequently, they represent shallow
water. However, when the species occurs in saline waters it is the only vegetated streams characterized by moderate current velocity and low
mollusc species present (De Francesco and Isla, 2004; Peretti, 2005) conductivity. According to the clusters observed, a subtle difference
while in fresh water it is usually accompanied by other mollusc between both levels can be inferred. This is due to the occurrence of
species. Ciocco and Scheibler (2008) found that along a gradient of some L. viator specimens in PV4, which may be consequence of
conductivity in the Bañado Carilauquen, H. parchappii coexisted with taphonomic alteration. According to the poor preservation of these
other species only in the headwaters and middle reaches at low shells (in most cases, the weathered shells simply disintegrate when
salinities (0.9–1.6 mS cm− 1) but was the only species found in the touched), it is probable that they represent a mixed assemblage,
extremely saline waters (8.0–19.5 mS cm− 1) of the outlet into the lake. including specimens that would have been deposited after some
Therefore, although we cannot morphologically distinguish the transport by currents. Kotzian and Simões (2006) found that small
populations of H. parchappii in fresh water from those inhabiting scale lateral transport of shells does occur in fluvial systems. The
saline waters, the composition of the assemblage may be indicative of occurrence of L. viator here suggests shallow water bodies probably
these two sorts of habitats, which has important implications for the connected to the main lotic system. This mixed assemblage probably
value of freshwater mollusc assemblages as indicators of past represents a taphocoenosis deposited during flooding events. Moreover,
continental salinities. the whole PV succession may reflect the variations in water flow that
Although the shell morphology of Heleobia aff. parchappii is different occurred in the floodplain during the Holocene in Río Atuel. An
from typical H. parchappii, the anatomy of the penial complex is similar. important conclusion is that the molluscs preserved in this sort of
We suggest that this species may be a variety of H. parchappii. It is alluvial succession represent mixed assemblages that suffered some
known that Heleobia displays a significant phenotypic plasticity in shell degree of taphonomic alteration. Because of the natural dynamics of the
morphology (De Francesco, 2007). Another possibility is that this taxon fluvial environment, it is expected that molluscs usually be subject to
is an undescribed species, ecologically similar to H. parchappii. A third transport and redeposition.
possibility is that this taxon corresponds to Heleobia kuesteri as recently The LB and PM successions represent lentic habitats with nil or very
suggested by Masi and Ciocco (2008). Heleobia kuesteri is a problematic low water velocity (Group 1). These habitats would have been vegetated
species that was originally described for Mendoza, of which there is no shallow lakes within the floodplain and not connected to the fluvial
updated taxonomic or ecological information. In fact, it has been system. The excellent preservation of the shells from LB supports this
considered as species inquirenda (Cazzaniga, 1981) because of the interpretation (see De Francesco et al., 2007). As L. viator inhabits the
difficulties for assessing its taxonomic identity. Although we cannot be submerged vegetation in very shallow areas where small streams flow
sure of the taxonomic identity of this taxon with the scattered into the lake, shells may be deposited in situ and, thus, represent
observations presented here, the presence of this same morphology parautochthonous assemblages. Some LB levels (LB4–LB6, LB11, LB14),
as fossil in the area (it was published as Heleobia sp. or Heleobia cf. H. dominated by B. peregrina, did not have a modern analogue in the
australis by De Francesco and Dieguez, 2006) allows their utility as a present model. However, B. peregrina is widely distributed in shallow
reliable bioindicator with important consequences for the reconstruc- lakes from La Pampa and Buenos Aires Province (Rumi, 1991; Peretti,
tion of past environments. In other words, despite its taxonomic 2005), regions characterized by higher humidity than the region
identity the ecological characteristics of this taxon are still an studied here. Thus, these LB levels probably indicate relatively wetter
informative indicator of the original depositional environment. conditions, which agrees with the previous interpretation, based on the
The CCA clearly indicated a strong relationship between mollusc whole assemblage together with the isotopic evidence, that the studied
taxa and environmental variables. Two main groups can be recognized: section records a mild interval corresponding to an interstadial (Marine
(Group 1) the species L. viator, P. acuta, and B. peregrina dominated in isotope stage 3) of the last glacial cycle (De Francesco et al., 2007).
highly vegetated sites characterized by high pH and NO2− 3 , and lower Although the results obtained here indicated that mollusc species
current velocity, on fine sediments, and (Group 2) the two Heleobia are useful for inferring past habitats in the semiarid region of
C.G. De Francesco, G.S. Hassan / Palaeogeography, Palaeoclimatology, Palaeoecology 274 (2009) 105–113 113
Mendoza, the presence of species not recorded as fossils in the area (H. de Castellanos, Z.A., Gaillard, M.C., 1981. Mollusca Gasteropoda Chilinidae. In: Ringuelet,
R.A. (Dir.), Fauna de agua dulce de la República Argentina, Vol. 15 (4), Fundación
hatcheri and P. acuta) introduces a complication in using the modern para la Educación, la Ciencia y la Cultura, Buenos Aires, pp. 19–51.
assemblage to infer paleoenvironments. P. acuta (= P. cubensis) is an de Castellanos, Z.A., Landoni, N.A., 1981. Mollusca Gasteropoda Lymnaeidae. In:
introduced species widely distributed in Argentina (Núñez and Ringuelet, R.A. (Dir.), Fauna de agua dulce de la República Argentina, Vol. 15 (5),
Fundación para la Educación, la Ciencia y la Cultura, Buenos Aires, pp. 53–82.
Pelichotti, 2003). This species has a superior reproductive capacity de Cock, K.N., Wolmarans, C.T., 2007. Distribution and habitats of the alien invader
compared to other pulmonate snails, as well as ability to migrate freshwater snail Physa acuta in South Africa. Water S. A. 33, 717–722.
upstream and to quickly recolonize a water-body (see de Cock and De Francesco, C.G., 2007. Las limitaciones a la identificación de especies de Heleobia
Stimpson, 1865 (Gastropoda: Rissooidea) en el registro fósil del Cuaternario tardío y
Wolmarans, 2007 and references therein). Consequently, it may have a sus implicancias paleoambientales. Ameghiniana 44, 631–635.
negative impact on indigenous freshwater molluscs in particular and De Francesco, C.G., Dieguez, S., 2006. Paleoambientes del Cuaternario tardío del sur de
on the biodiversity of freshwater habitats in general. One important Mendoza: estado del conocimiento, problemas y perspectivas. In: Durán, V.,
Cortegoso, V. (Eds.), Anales de Arqueología y Etnología, vol. 61. Mendoza, pp. 69–80.
caveat regarding paleoenvironmental interpretations is the lack of
De Francesco, C.G., Isla, F.I., 2004. Reproductive period and growth rate of the freshwater
knowledge of the ecological role played in the past by other modern snail Heleobia parchappii (d'Orbigny, 1835) (Gastropoda: Rissooidea) in a shallow
coexisting species such as L. viator and B. peregrina. It is likely that, in brackish habitat (Buenos Aires Province, Argentina). Malacologia 45, 443–450.
the absence of P. acuta, these species would have occupied a different De Francesco, C.G., Zárate, M.A., 2006. Moluscos holocenos del Valle de Uco, Mendoza.
Academia Nacional de Ciencias (Córdoba), Abstracts, p. 63.
range of habitats or, eventually, experienced an increase in their De Francesco, C.G., Zárate, M.A., Miquel, S.E., 2007. Late Pleistocene mollusc assemblages
distribution. Similarly, the absence of H. hatcheri in the past suggests and inferred paleoenvironments from the Andean piedmont of Mendoza, Argentina.
that the species colonized Mendoza in recent times (in fact, the Palaeogeogr. Palaeoclimatol. Palaeoecol. 251, 461–469.
Dieguez, S., De Francesco, C.G., Gil, A.F., Neme, G.A., 2004. Paleoecología, paleoambientes y
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deposits of up to ca. 500 years B.P.; De Francesco and Zárate, 2006), so (San Rafael, Argentina). 2° Reunión Binacional de Ecología, Mendoza, Abstracts, p. 421.
the habitats they occupy today would have been absent in the past or Dillon Jr., R.T., 2000. The Ecology of Freshwater Molluscs. Cambridge University Press,
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occupied by a different species, which cannot be ascertained. Even Doering, A., 1884. Apuntes sobre la fauna de moluscos de la República Argentina. Bol.
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Financial support for this study was provided by Agencia Nacional http://folk.uio.no/ohammer/past.
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organic and carbonate content in sediments: reproductibility and comparability of
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(Museo de Historia Natural de San Rafael) for support during field Kotzian, C.B., Simões, M.G., 2006. Taphonomy of recent freshwater molluscan death
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