Holocene molluscs from archaeological sites of ... - Wiley Online Library

Holocene Molluscs from Archaeological Sites of the Pampean Region of Argentina: Approaches to Past Human Uses Mariano Bonomo1, 2,* and Marina L. Aguirre1, 3 1

CONICET Departamento Científico de Arqueología, Museo de La Plata, Paseo del Bosque, (1900) La Plata, Argentina 3 Facultad de Ciencias Naturales y Museo, INGEA, Laboratorio 6 (Malacofauna Cuaternaria), 64 N °3, (1900) La Plata, Argentina 2

The aim of this study is to provide a regional review and update our knowledge of molluscan records from Late Quaternary (12.2–0.4 ka B.P.) archaeological sites of the Pampean region. Comparisons with molluscs from the modern littoral, Holocene beach ridges, and estuarine sediments are made, allowing a synthesis of the main taxa recovered and reinterpretations of the probable past human uses. Overall, the biological and archaeological data gathered reinforce the conclusion that most taxa living far from the intertidal zone were not linked to subsistence of Pampean hunter-gatherers. However, sporadic consumption cannot be discarded, considering the sea-level changes during the Holocene. Other uses including ornamentation, containers, tools, or symbolic meaning can be assigned to marine molluscs which were probably collected as dead shells from the fossil deposits or from the contemporary beach. Selection of the shells by humans related mainly to shell size, shape, and color, and not to their nutritional value. © 2009 Wiley Periodicals, Inc.

INTRODUCTION In addition to providing information regarding archaeological concerns like seasonality of human occupations, past subsistence, and technology, molluscs are useful proxies for environmental and climatic conditions. These include substrate nature, water energy levels, sea-level variations, sea surface temperatures, and oceanic–atmospheric circulation patterns (among others, Lutaenko et al., 2007; Razjigaeva et al., 2002; Aguirre, Richiano, & Negro Sirch, 2006; Carré et al., 2006; Martínez et al., 2006; other references therein). Since the early to mid-20th century, marine molluscs have been included among the dispersed data from archaeological records, used to establish cultural relationships between distant geographic regions of Argentina. Similarities regarding economic or ornamental uses of molluscs, typological affinities of shell tools, selection *Corresponding author; E-mail: [email protected] Geoarchaeology: An International Journal, Vol. 24, No. 1, 59–85 (2009) © 2009 Wiley Periodicals, Inc. Published online in Wiley Interscience (www.interscience.wiley.com). DOI:10.1002/gea.20254

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of the same species in different sites, and their Atlantic or Pacific provenance, have been considered in order to identify large-scale human migrations as evidence of pre-Hispanic cultural interchange (Martínez Soler, 1958–59, and references therein). Among the uses assigned to marine molluscs in the ethnographic and archaeological literature for different locations of Argentina (Vignati, 1930; Martínez-Soler, 1958–59; Bórmida, 1969; Orquera & Piana, 1999; Damiani & Alvarez Rodríguez, 2005; Prates, 2008), the most common are the consumption of shellfish, the manufacture of tools, musical instruments, vessels, and ornaments, such as the pendants and necklace beads frequently found as part of the funeral trousseau. Many sites of the Pampean region of Argentina (Figure 1) have marine mollusc remains. Several authors (Ameghino, 1909; Bórmida, 1969; Conlazo, 1983) advanced

Figure 1. Study area of the Pampean coastal region. Archaeological sites sampled and reviewed from the available literature: 1, Fontezuelas; 2, Laguna Sotelo; 3, Cueva El Abra; 4, Cueva Tixi; 5, Bellamar 1; 6, Bellamar 3; 7, Túmulo de Malacara; 8, Nutria Mansa 1 (sup.); 9, Nutria Mansa 2 (sup.); 10, Quequén Chico 1; 11, Paso Otero 3; 12, Necochea; 13, Cueva del Tigre sitio 1; 14, Cueva del Tigre 4; 15, Claromecó 1; 16, Arroyo Seco 1; 17, Arroyo Seco 2; 18, Puente de Oriente; 19, Sur Cueva del Tigre; 20, Cueva del Tigre; 21, Norte Cueva del Tigre; 22, El Palomar 1; 23, El Puente 1; 24, Puente de Fierro; 25, La Olla 1; 26, La Toma; 27, SA 17 Avestruz; 28, Laguna Los Chilenos; 29, San Martín 1; 30, Fortín Necochea; 31, Pintado II; 32, La Primavera; 33, Don Aldo; 34, Laguna de Chillhué 1; 35, L. Tapera Moreira; 36, Chenque I; 37 Casa de Piedra 1.

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HOLOCENE MOLLUSCS FROM ARCHAEOLOGICAL SITES

the hypothesis that human groups occupying the Pampean Atlantic coast showed feeding habits focused on marine resources, mainly molluscs. However, detailed studies of archaeological molluscan shells have not been carried out, and there has been no synthesis of taxonomic, taphonomic, paleoecologic, and distributional aspects of the main taxa recovered. This prevents a complete understanding of the way humans could have obtained and used these resources. Molluscan taxa have been recurrently cited in the Pampean literature (e.g., Gradín, 1984; Politis, 1984; Crivelli Montero, Eugenio, & Silveira, 1987–88; Eugenio & Aldazabal, 1987–88; Austral et al., 1988; Barrientos, Leipus, & Oliva, 1997; Martínez, 1999). Most studies report only brief details and for individual sites, except for a few recent cases in which the shells were transformed into ornaments (Mazzanti & Valverde, 2001; Cimino, Guastavino, & Velardez, 2004). The aim of this article is to review and update our knowledge of the mollusc remains found in Pampean archaeological contexts, bringing together disparate archaeological and paleoecological information beyond single-site analyses. The evidence analyzed includes collections from our own fieldwork as well as a compilation of numerous bibliographic data sources. It must be acknowledged that the reliability of the mollusc records from some archaeological sites is unknown, and some of the data are somewhat limited by being incomplete, undated, not figured, and not quantified. The majority, however, are reliable. The study of the whole dispersed record will allow interpretations regarding the source and possible uses (food, artifacts, adornments) for each taxon, within each hunter-gatherer site. A better understanding of the dietary practices of human populations could suggest an interpretation of cultural differences among past inhabitants of different areas within the Pampean region. An update of the taxonomic status of the species is essential, as subsequent synonymy changes affect both estimations of biodiversity in archaeological contexts and inferences of Late Quaternary paleoenvironmental change. In addition, comparisons are made between the archaeological mollusc shells and those from natural shell concentrations along the Pampas (Figure 2). This is the first stage of a major study focused on exploring the relationship between geologic, paleoecologic, and paleoclimatic evidence and past human uses of marine shellfish, an approach that has shown good results in other areas (Rollins, Sandweiss, & Rollins, 1990; Taborín, 1993; Cannon, 2000; Gutiérrez Zugasti, 2006; Lutaenko et al., 2007). This article reports some of the results included in the Ph.D. thesis of one of the authors (Bonomo, 2004a, 2004b, 2005). GEOLOGICAL AND MODERN REGIONAL SETTING The study area (33–40°S, 56°41⬘–68°04⬘W) includes the La Pampa and Buenos Aires provinces and southern San Luis, Córdoba, and Santa Fe provinces (Figure 1). During the Neogene several higher than present sea-level episodes transgressed the coastal zone of Buenos Aires Province (Del Río & Martínez, 1998; Aguirre & Farinati, 1999a). From the Río de La Plata margin to Bahía San Blas (Figure 2), the most extensively preserved episode is the mid-Holocene transgression (ca. 3–7 ka B.P.; Figure 3). During the Holocene regressive-transgressive processes (i.e., Las Escobas GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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Figure 2. Study area along the Pampean coastal region: Holocene fossiliferous shell concentrations and limits of the Late Quaternary marine transgressions. Modified from Fucks, Aguirre, and Deschamps (2005). 62



Figure 3. Stratigraphical framework for the study area. Holocene continental and littoral deposits, available dates, and archaeological sites. Modified from Aguirre and Whatley (1995) and Fucks, Aguirre, and Deschamps (2005). M: marine sediments; C: continental sediments; SC: Spanish conquest; PL: Pleistocene. *Main references in Politis and Madrid (2001) and Berón (2004). ka: 1000 years.

Formation), large numbers of invertebrate skeletons accumulated, forming typical beach ridges lying at ⫹2–6 masl. These ridges contain ca. 80–90% of well-preserved and varied mollusc shells. By contrast, Late Pleistocene marine records, probably of the last interglacial sea level (MIS 5e), are discontinuous and contain fewer fossil shells of poorer preservation and lower faunal diversity. The innermost extension of the Holocene transgression reached ca. 50 km westward of the modern coastline along central Samborombón Bay, but in most places this transgression covered only a narrow fringe of ca. 2–10 km along the coastal area. Stratigraphic interpretations of the marine units in the Pampean area have been provided elsewhere (Fucks, Aguirre, & Deschamps, 2005; Schnack et al., 2005) as well as descriptions of marine landforms (among others, Fidalgo, 1979; Schnack et al., 2005; Violante, 1988, Isla et al., 2000). The modern littoral includes a fluvial-estuarine margin (Río de La Plata) and a fully oceanic sector (southwestern Atlantic Ocean, Argentine Sea) (see Guerrero & GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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Piola, 1997; Boltovskoy et al., 1999; Mianzan et al., 2001, for a complete description of physical and biological characteristics). The salinity of the Río de La Plata ranges from oligohaline (0–5‰) at Buenos Aires to polyeuhaline (18–30‰) at Punta Rasa. The boundary between brackish and marine waters depends on the season and dynamics of the estuary. Muddy substrates are dominant in the area. The oceanic sector from Punta Rasa to the Colorado River outlet is part of the Argentine Sea. Soft, sandy substrates and euhaline (33–35‰) waters dominate this area. The microtidal regime ranges from 1 to 2 m (Punta Rasa–Río Colorado), with a peak at Bahía Blanca of approximately 3 m (Isla & Bértola, 2005). The shallow Brazilian warm current, characterized by a warm temperate shallow-water mass at the Transition Zone (Subtropical–Subantarctic Convergence), influences this area. The position of the South Atlantic anticyclone center, which brings warm dry air over the ocean, shows a seasonal latitudinal displacement, being southernmost during the summer. This pattern controls the winds and precipitation regime over the area. The region belongs to the Argentine Malacological Province, which extends from Espiritu Santo (Brazil) to Golfo San Matías (Patagonia) (Castellanos, 1967; Aguirre & Farinati, 1999a, 2000). Detailed information on the physical and biological features of the area has been published by Bastida et al. (2007), Guerrero and Piola (1997), Boltovskoy et al. (1999), Boltovskoy, Correa, and Boltovskoy (2005), and Isla and Bértola (2005). Previous ecological and biocenological studies of littoral molluscs from the Argentine Sea have mostly focused on small areas and on particular benthic communities. Recent ecological studies (among others, Penchaszadeh, 2004; Bastida et al., 2007) have focused on molluscs of economic value (Lasta et al., 1998), with available information about molluscs living on the Argentine continental shelf (Bastida et al., 1981) obtained from fisheries and oceanographic expeditions (Vema, Shinkai-Maru, SAO campaigns). Current data sets on taxonomic diversity and distribution are biased. Overall, there are still gaps in our knowledge of the habitat, bathymetric ranges, variability, and geographical distribution of most marine molluscs studied. The details of some species are known, however. These include Mesodesma mactroides Deshayes and Donax hanleyanus Philippi, which are among the typical taxa of high nutritional value living in the Pampean intertidal zone, characterized by soft substrates. Another bivalve used for food is Mytilus edulis Linné, which lives in deeper waters and varied substrates along the Bonaerensian littoral zone. ARCHAEOLOGICAL REMAINS Most of the Pampean archaeological sites reviewed here were excavated and studied previously by other authors (Austral, 1965, 1994; Gradín, 1984; Politis, 1984; Loponte & Acosta, 1986; Crivelli Montero, Eugenio, & Silveira, 1987–88; Eugenio & Aldazabal, 1987–88; Austral et al., 1988; Oliva, Gil, & Roa, 1990; Madrid & Politis, 1991; Barrientos, Leipus, & Oliva, 1997; Martínez, 1999; Petz & Saghessi, 2000; Mazzanti & Quintana, 2001; Madrid et al., 2002; Politis, Bonomo, & Prates, 2003; Berón, 2004). For a complete set of references for all the sites reviewed see Bonomo (2004a: Tables IX.2–IX.5). A synthesis of the Pampean archaeological records shows that 64



more than 1200 marine mollusc remains, including shells with no human modifications as well as artifacts, have been recovered from 37 sites of Late Pleistocene to Late Holocene age (Table I). While most of the sites are found in the plains along the southeastern Pampean region near the Atlantic coast, they show a wide regional distribution, in several cases surpassing 200 km from the current coastline (Figure 1). Fifteen human occupations included in this survey are surface sites without chronological control. The remaining have been assigned to Late Pleistocene–Early Holocene (n ⫽ 1), Middle Holocene (n ⫽ 7), and Late Holocene (n ⫽ 14) (Figure 3). In general, the molluscs occur in very low densities. As far as the available data show, shell mounds as a result of artificial accumulations by past populations have not been recorded in the area. Most sites have no more than 10 molluscan remains. The Arroyo Seco 2 (Politis, 1984) and Chenque I (Berón, 2004) sites show the greatest abundance, including more than 1100 beads, and are linked to human bones in funerary contexts.

RESULTS In the Pampean archaeological sites, 19 taxa (11 gastropods and 8 bivalves; Appendix 1) were identified to generic or specific levels. Among the identified taxa, Adelomelon brasiliana, Adelomelon sp., and Volutidae indet. are the most common (Figures 4a, b). Nearly all these taxa are also recorded in Pleistocene and Holocene beach ridges from the same area (Figure 2), except for Mesodesma sp. from the La Primavera archaeological site (Bayón et al., 2004), which has not been recorded from natural deposits. Most taxa live along the modern coast in the Argentine, Brazilian, and/or Antillean Malacological provinces (Tables IIa, IIb). Three taxa, however—Fissurella sp. and Urosalpinx haneti recorded at Chenque I (Berón, 2004) and Noetia bisulcata from El Palomar 1 (Austral, 1965)—are not recorded today in the Pampean area. Fissurella sp. is a typical Patagonian mollusc (Magellanian Province), and Noetia bisulcata and Urosalpinx haneti are typically warm-water types, at present found northwards in the Uruguay, Brazil, north Argentine, and Brazilian provinces (Ríos, 1994). All the taxa are typically benthic marine molluscs, mostly from the littoral to deep bathyal zones, within a bathymetric range of ca. 4 m to more than 200 m. Noticeably, only a few of the taxa recovered from archaeological sites live in the intertidal zone (0–4 m deep) or in shallow waters just below the low-tide level, including the gastropods Fissurella, Urosalpinx haneti, Olivancillaria, and Buccinanops and bivalves Mesodesma and Glycymeris longior (Figure 5). Three taxa have only been recorded as fossil shells (Buccinanops, Urosalpinx, Fissurella; Cimino, Guastavino, & Velardez, 2004). Many authors (Austral, 1965, 1994; Gradín, 1984; Politis, 1984; Loponte & Acosta, 1986; Crivelli Montero, Eugenio, & Silveira, 1987–88; Eugenio & Aldazabal, 1987–88; Oliva, Gil, & Roa, 1990; Barrientos, Leipus, & Oliva, 1997; Petz & Saghessi, 2000; Mazzanti & Valverde, 2001; Berón, 2004, among others) have previously suggested the GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

65

?

5

6 7

8

9 10

11 12 13 14 15

16

19

20

21 22

23 24

25

26 27 28 29

30

31

32 33 34

35

36 37

Note: The numbers of sites correspond to those of Figure 1. Updated names for the species are as follows: Drupa pilsbry ⫽ Morula necocheana (Pilsbry); Arca bisulcata ⫽ Noetia bisulcata (Lamarck); Pecten sp. ⫽ Aequipecten sp.

Fissurella sp. Urosalpinx haneti Drupa pilsbry Drupa sp. Zidona dufresnei Adelomelon beckii Adelomelon brasiliana Adelomelon ancilla Adelomelon sp. Volutidae indet. Olivancillaria sp. Buccinanops sp. BIVALVIA Arca bisulcata Glycymeris longior Mytilus sp. Pecten sp. Mesodesma sp. Amiantis purpurata Amiantis sp. Protothaca antiqua Bivalvia indet. MOLLUSCA INDET

4

17 18

3

GASTROPODA

1 2

Archaeological Sites Reviewed

Molluscan Taxa

Table I. Molluscan taxa recovered in the Pampean archaeological sites from north–south to east–west.


Figure 4. Mollusc taxa recovered from the archaeological sites. (a) Gastropods: 1, Fissurella sp.; 2, Urosalpinx haneti (Petit); 3, Morula necocheana (Pilsb.); 4, Zidona dufresnei (Don.); 5, Adelomelon ancilla (Lightf.); 6, Adelomelon beckii (Brod.); 7, Adelomelon brasiliana (Lamk.); 8, Olivancillaria urceus (Röding); 9, Olivancillaria carcellesi Klapp.; 10, Buccinanops globulosus (Kiener). (b) Bivalves: 1, Noetia bisulcata (Lamk.); 2, Glycymeris longior (Sow.); 3, Mytilus sp.; 4, Aequipecten sp.; 5, Mesodesma sp.; 6, Amiantis purpuratus (Lamk.); 7, Protothaca antiqua (King).

archaeological use of a considerable number of taxa. These include Fissurella sp., Urosalpinx haneti, Morula sp., Adelomelon brasiliana, Adelomelon sp., Volutidae indet., Buccinanops sp., Amiantis purpuratus, Amiantis sp., Protothaca antique, Bivalvia indet. from 21 of the Pampean sites studied (Table III). These shells were most likely used to make personal ornaments, and probably tools and containers. The most frequent use was ornamental and decorative, mainly manufactured beads for corporal adornments, such as necklaces or bracelets. These beads and some molluscs without human modification were found mainly associated with human burials in ritual contexts. Ten shells showing highly polished surfaces were interpreted as manufactured artifacts such as containers, drillers, or retouched artifacts, while others were reported with adhered red mineral pigments on their surfaces. It must be acknowledged that the polished shell surfaces and shapes of some of the tools described in the literature can also be produced naturally by taphonomic GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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Depth in m

Trophic Type

Stratigraphical range of taxa: MIO: Miocene; PLIO: Pliocene; PL: Pleistocene; H: Holocene; M: modern. Geographical range (modern distribution in Malacological provinces): ANT: Antillean; BR: Brazilian; AR: Argentinean; MAG: Magellanean; ADJ: occurrence in the modern littoral of the study area. Depth for each species refers to the zones where they most typically live in their geographical distribution range.

PPS: Most probable provision source of the taxa found in the Pampean archaeological sites: 䉱 alive in intertidal zone; 䊏 shells deposited on the beach during tidal changes and after storm events (transported from the infralittoral and bathyal zones); * taxa recovered as fossil remains. RA: refers to the relative abundance of each taxon within Holocene beach ridges preserved along the Pampean coast: A: absent; S: scarce; C: common; VA: very abundant.

▲ C ▲* C 4–30 0–40

Olivancillaria sp. Buccinanops sp.

VA VA

* S S

* A

R A

VA VA

P P S

⬎10 10–350

Supral.

Adelomelon sp. Volutidae indet.

Mesolit.

VA VA

Sublit.

10–75 18–70

Bathyal

Adelomelon beckii (Broderip) Adelomelon (P.) brasiliana (Lamarck)

3–55 20–100

0–15

Hard. 15–100 10–350

NO

Soft

Zidona dufresnei (Donovan) Adelomelon ancilla (Lightfoot)

Urosalpinx haneti (Petit) Morula necocheana (Pilsbry)

MIO PLI PL H

GASTROPOD TAXA Fissurella sp.

Free

Life Substrate Mode

Sessile

Zonation (Bathymetry)

Carniv.

Geographical Range

M ANT BR AR MAG ADJ NO

Stratigraphical Range

Distribution and Taxa

Herbiv.

Table II. Ecological requirements of the modern representatives of the marine molluscan taxa recovered in archaeological sites. (a) Gastropods.

Filterf.

MIO PLI PL H

BIVALVE TAXA

Trophic Type

2–100

Protothaca antiqua (King)

Stratigraphical range of taxa: MIO: Miocene; PLIO: Pliocene; PL: Pleistocene; H: Holocene; M: modern. Geographical range (modern distribution in Malacological provinces): ANT: Antillean; BR: Brazilian; AR: Argentinean; MAG: Magellanean; ADJ: occurrence in the modern littoral of the study area. Depth for each species refers to the zones where they most typically live in their geographical distribution range.

PPS Most probable provision source of the taxa found in the Pampean archaeological sites: 䉱 alive in intertidal zone; 䊏 shells deposited on the beach during tidal changes and after storm events (transported from the infralittoral and bathyal zones); * taxa recovered as fossil remains. RA: refers to the relative abundance of each taxon within Holocene beach ridges preserved along the Pampean coast: A: absent; S: scarce; C: common; VA: very abundant.

NO

䊏* A

C C

䊏䊏 ? ?

15–20 ⬎15

15–40

Amiantis purpuratus (Lamarck) Amiantis sp.

NO

S A

Noetia bisulcata (Lamarck)

䊏䉱

R A

20–130 0–20

P P S

Aequipecten sp. Mesodesma sp.

Supral.

S

Mesolit.

C S

Sublit.

䊏䉱

Depth in m

䊏

Hard

10–20 4–80

Soft

Glycymeris longior (Sowerby) Mytilus sp.

Epibis.

Life Substrate Mode Upp. Inf.

Zonation (Bathymetry) Deep Inf.

Geographical Range

M ANT BR AR MAG ADJ

Stratigraphical Range

Distribution and Ecology Suspens.

Table II. Ecological requirements of the modern representatives of the marine molluscan taxa recovered in archaeological sites. (b) Bivalves.

Detritiv.

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Figure 5. Marine vertical biological zonation. Not to scale. s: supralittoral; m: mesolittoral (intertidal; 0–4 m); Sublittoral: in: infralittoral (4–40 m); c: circalittoral (40–200 m). Dominant bathymetric distribution of the marine taxa living along the Bonaerensian littoral recovered in archaeological sites. Partially modified from Boltovskoy (1965).

processes. The taphonomic signatures of the same taxa recovered in natural shell concentrations from the area include abraded shell surfaces, rounded shell margins, loss of the original color and ornamentation, and disarticulation (Aguirre & Farinati, 1999b). Information on the systematics, paleoecology, and distribution of the most common taxa recorded in the sites reviewed are shown in Appendix 1. For the complete source of information for each taxon see Aguirre (1993), Aguirre and Farinati (1999a, 2000) and Aguirre, Hlebzsebitch, and Delatorre (2008). 70


HOLOCENE MOLLUSCS FROM ARCHAEOLOGICAL SITES Table III. Different past uses of the taxa found in the archaeological record. Archaeological Uses of Taxa Archaeological Sites

Ornamental

Artifact

*

䉱 ()

Cueva Tixi El Abra

䉱

Túmulo de Malacara

䉱

Quequén Chico 1

䉱

Necochea

䉱

䉱

Cueva del Tigre sitio 1

䊏

Cueva del Tigre 4

⫻ 䉱丢

El Puente 1

丢*

Puente de Fierro

†*

†

Adelomelon sp.

䉱

Ø Buccinanops sp.

䊏 Volutidae indet. ⫻ Amiantis purpurata /

Amiantis sp. Protothaca antiqua

◊

Bivalvia indet.

䉱 MOLLUSCA INDET *

Probable asignation

䉱

San Martín 1

䉱

Fortín Necochea

䉱* 䉱

Pintado II

Casa de Piedra 1

丢 Adelomelon brasiliana

䊏*

La Toma

Tapera Moreira 1

Urosalpinx haneti

() Drupa sp.

䉱

䉱/

El Palomar 1

Chenque I

♣ Fissurella sp.

䊏

Laguna Sotelo

Laguna Los Chilenos

Adhered Pigments

◊*

Fontezuelas

Arroyo Seco 2

Ajuar

♣* Ø ⫻

♣* Ø ⫻ 䊏

䊏丢

DISCUSSION Some aspects of Pampean archaeological remains are still controversial. Among these issues are (1) whether the representation of the mollusc taxa in archaeological sites can be accounted for by preservation problems or by sampling bias; (2) what are the compositional differences between the molluscan fauna recorded at the sites compared to the records from Holocene-aged natural assemblages and with the modern littoral associations; (3) whether Holocene sea-level changes controlled shellfish availability; (4) the eventual economic (subsistence and technology) or symbolic value of the molluscs recovered; and (5) the ways in which hunter-gatherers could have obtained the molluscs. Given the low number of shells recorded in most sites, with around 1100 from two sites and an average of three per site for the other 35, it can be argued that while a large number of molluscs were used during human occupations, their skeletons may have been missed during the archaeological recoveries due to sampling bias or taphonomic processes. Small taxa are not expected to have been missed from the samples GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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due to their size, as many micromolluscs have been identified in the archaeological sites (e.g., Drupa necocheana and Urosalpinx haneti). On the other hand, the molluscs recovered (Urosalpinx, Morula, Zidona, Adelomelon, Olivancillaria, and Buccinanops among the gastropods; Noetia, Glycymeris, Amiantis, and Protothaca among the bivalves) belong to taxa with high preservation potential, characterized by medium- to large-size shells which are thick (2–5 mm) in form. These taxa are abundant within natural Holocene concentrations, where they show little or no loss of their original features (see below). Besides, the alkaline sediments that predominate along the Bonaerensian littoral would favor preservation of calcareous shells. Also, in other similar coastal areas to the north in Uruguay (Inda et al., 2006) and southwards in North Patagonia (Bórmida, 1969; Borella & Favier Dubois, 2007), abundant shells are well preserved as shell middens. Overall, the available evidence suggests that the low number of shells is most likely linked to infrequent human selection and collection. Regarding the Holocene-aged natural assemblages (beach ridges and estuarine facies) of the Bonaerensian coastal area, the factors controlling the distribution of marine benthic taxa include the temperature of shallow oceanic water masses (SST, sea surface temperature), oceanic currents, substrate type, salinity, water depth, and energy. The only differences between Holocene and modern benthic molluscan faunas are variations in abundance, or geographical shifts of a few taxa as a biotic response to the effect of the mid-Holocene “Hypsithermal” (Aguirre, 1993), suggesting slight changes in SST and currents. In addition, all the taxa recorded in the archaeological sites have modern representatives. This suggests that variations in the finds of shells at sites are not due to changes in availability through time, at least during the Holocene. Interestingly, a number of gastropods and bivalves (Adelomelon brasiliana, Adelomelon ancilla, Volutidae indet., Amiantis purpuratus, Amiantis sp., Mollusca indet.) from the archaeological sites (4 sites; Figure 3) older than the mid-Holocene marine transgressive maximum (ca. 6 ka B.P.; Schnack et al., 2005) also occur in younger Late Holocene archaeological sites (ca. 3 ka B.P.). This would imply that climate and sea-level changes did not alter their availability for human uses. The remaining species (Fissurella, Urosalpinx, Morula, Zidona, Olivancillaria, Buccinanops, Glycymeris, Noetia, Protothaca, Mytilus, Pectinidae, Mesodesma) are present in both the more numerous Late Holocene and in the undated sites. This results either from a bias in the sampling size due to the greater number of contexts assigned to late Holocene age, or environmental changes following the mid-Holocene Hypsithermal (⫽ Climatic Optimum; climatic amelioration). Altogether, the general morphology, the ecological and bathymetric preferences, and geographical distribution of modern representatives of the recorded molluscs provide clues or hypothetical explanations for their selection and use by humans. Bathymetry and body size are among the main factors controlling human exploitation of shellfish for eating. Hunter-gatherers would prefer to collect and consume species living in shallower waters near the coast, in supratidal and intertidal zones (Buccinanops, Olivancillaria, Mesodesma), whereas deeper-habitat molluscs like Adelomelon spp. and Zidona dufresnei (Figure 5) would be difficult to obtain alive from the beach. Curiously, shallow taxa very common in the Bonaerensian littoral 72



at present (Donax, Mesodesma, Brachidontes, among others) are very scarce or absent in the Pampean sites. Additionally, people would perhaps favor species with large amounts of soft tissue for eating. However, those molluscs most suitable for eating (for example, Mesodesma) are also very scarce in the Pampean contexts. The availability of edible molluscs and productivity in modern littoral areas worldwide is linked to the substrate and tidal regime. In general, rocky substrates in the macrotidal regime offer higher productivity conditions. Along the study area, the scarcity or absence of hard and rocky bottoms as found on the Patagonian coast explains the reduced availability or absence of typical epifaunal taxa with high nutritional values (Mytilus, Aulacomya, Patinigera). On the other hand, the predominance of soft substrates explains the occurrence of infaunal edible taxa such as the burrowing bivalves Mesodesma and Donax. The microtidal Pampean regime (1–2 m), in contrast to the macrotidal Patagonian littoral (5 m at Comodoro Rivadavia, 10 m at central Santa Cruz Province), implies much lower productivity. Overall, an environmental pattern characterized by a lack of hard substrates and low tidal range was disadvantageous for a dietary use of molluscs by hunter-gatherers. Among the taxa recovered in the Pampean sites, only Mesodema sp. and Olivancillaria sp. could have been used as feeding resources. These taxa are represented by only two specimens from two sites (one of which is located ca. 60 km inland from the modern shore, Arroyo Seco 2; Politis, 1984), implying that it was not used for food (see also Mazzanti & Valverde, 2001; Politis, Bonomo, & Prates, 2003; Berón, 2004). By contrast, the dominant molluscs found at the sites (i.e., Adelomelon spp.) usually reach the shore via storms and waves but inhabit circalittoral–bathyal zones, far from the coastline, where diving is extremely difficult. Moreover, around 1100 of more than 1200 archaeological shells were retrieved from only two inland sites, and their final use was as beads in funerary contexts. Frequencies of molluscan remains are very low in most of the remaining 35 sites. Carrying such small numbers of shells over considerable distances is unlikely to be the result of subsistence behavior. As an alternative explanation to diet, it must be considered that molluscs are also collected according to the properties of their shells which make them attractive or useful. While a great variety of species occur in the Pampean sites, a preference for species of rather large skeleton dimensions (Volutidae) or attractive ornamentation is apparent. Adelomelon is very abundant, most probably due to the great size (36–400 mm) and thickness (3–4 mm) of the shell and because of its suitability for varied technological functions (Table III). Although much smaller and scarcer, other gastropods (Fissurella, Urosalpinx) and some bivalves (Noetia, Amiantis) could have been selected for their attractiveness in terms of shape, sculpture, brightness, and color. Some mollusc shells recorded in the Pampean sites could have been utilized as containers or as manufactured ornaments or tools. Many shells (Table III) were transformed into beads, in some cases with red pigments adhered, and found associated with human burials. In some sites, including Laguna de Chillhué 1, Tapera Moreira, Chenque I, Casa de Piedra 1 (Berón, 2004; Gradin, 1984), such beads have been recorded far from the coast. This non-utilitarian use of marine molluscs and their inclusion as funeral trousseau suggests a symbolic value attached to these chosen objects. GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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The different ways in which hunter-gatherers may have had to obtain the shells can be understood by analyzing their occurrence and relative abundance along the modern beach and in the marine fossil deposits. In Holocene raised beach deposits, most bivalves and gastropods retain their color, brightness, and ornamentation, appearing very similar to the living forms. This, and the lack of precise chronological control for both the cultural remains and the fossil deposits, makes it difficult to establish the provenance of the shells found, or to assess whether they came from previously established shell deposits older than the sites or from the contemporary beach sediments coeval with the occupations. The nature of shell concentrations has been well documented worldwide (Kidwell, 1986; Parsons & Brett, 1991, Mehdahl, 1994; Kowalewski & Flessa, 1995) and also in the study area (Lahille, 1895; Carcelles, 1944; Farinati & Zavala, 1995; Aguirre & Farinati, 1999b; Farinati, Spagnuolo, & Aliotta, 2006). During daily tidal rises or episodic storm events, mollusc shells are usually deposited along the beach. Local topography, substrate nature, water energy, and tidal regime are linked with the accumulation of empty disarticulated and, less frequently, whole joined shells along the supratidal and intertidal zones. In the study area, less energetic and more protected zones with soft bottoms show highly concentrated molluscan skeletons, well known locally as “shell cemeteries.” Occasionally, during episodic storms or nonperiodic wave events, larger amounts of shells are deposited along the littoral zone. Higher wave energy can transport shells from deeper zones (bathyal, circalittoral; e.g., Adelomelon) to the beach, creating a mixed concentration of shells from different habitats (Figure 5; Tables IIa, IIb). Several permanent skeletal concentrations forming “shell cemeteries” along the modern coast of Buenos Aires Province have been known since the 17th century (Cardiel, 1930). These include Punta Rasa, Punta Médanos, the beach zone between Mar Chiquita Lagoon and Faro Querandí, and “Caracolero,” near Claromecó (Figure 2). Moreover, several taxa deposited in these cemeteries also occur in pre-Hispanic human occupation sites. In the modern “Caracolero” accumulations, several gastropods (i.e., Zidona, Adelomelon, Buccinanops, Olivancillaria) and bivalves (i.e., Glycymeris, Brachidontes, Mytilus, Aequipecten, Amiantis, Mesodesma) (see Appendix 2) are shared with the Pampean archaeological sites reviewed. This kind of natural concentration could have been used as a shell supply in the past. It is most likely that the Volutidae, the dominant taxa recovered in the Pampean sites, came from the nearby fossil ridges and estuarine facies, or that they were found dead on the beach at the time of the occupation. However, to assess whether the molluscs were collected either alive in beach drift, as dead shells from the beach, or from older raised beaches, it is necessary to carry out a detailed taphonomic and geochronological analysis for single specimens and sites. FINAL REMARKS Our regional review of more than 30 archaeological sites from the Argentine Pampean area where marine molluscs have been recorded associated with cultural materials represents a useful tool for different fields of earth sciences. It shows a close

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linkage between archaeological remains, ecological, and paleoecological evidence. In turn, the information gathered here could also be applied in studies of other coastal areas with similar remains and problems. Most archaeological information, together with paleocological and biological data sets, indicate that marine molluscs have no clear relationship with the subsistence of the Pampean hunter-gatherers. In summary, the marine molluscs recorded in the Pampean sites revisited suggest that: 1. 2. 3. 4. 5. 6. 7. 8.

the number of shells in most coastal and inland Pampean archaeological contexts is very low; the low frequencies of archaeological molluscs in most sites cannot be explained taphonomically; most recorded taxa are typically not edible molluscs, and edible species in the sites are extremely scarce; selection of the mollusc shells seems to be determined not by the nutritional value of the taxa, but by their size, shape, and color; it is highly likely that the regional use of marine molluscs was non-dietary; they probably had other alternative uses (ornamentation, containers, tools); their main use could have been the manufacture of beads, frequently included with human bodies in funerary contexts; some marine molluscs could have had a symbolic value for pre-Hispanic hunter-gatherers; they were most probably collected as dead shells from fossil littoral ridges or from the contemporary beach.

Waves, tides, currents, input from the continent, benthic assemblages, and humans interplay through time and space in the littoral area. Our results show that Pampean hunter-gatherers did not directly exploit the molluscs of the complex continental shelf environment, but picked up dead shells. Further work, such as more exhaustive sampling in the Pampean region and new archaeological evidence with more precise chronological control are urgently required in the area to reach a better understanding of the human use of marine molluscs and their influence on, and relevance to, human society. The authors wish to thank Jeff Blackford and two anonymous reviewers for their useful comments, which improved earlier versions of the manuscript; to Natalia Carden and Cecilia Deschamps (Museo de La Plata) for their help with the English text; and to Andrés Boltovskoy (Museo de La Plata) for providing a useful bibliography about the physical and biological characteristics of the South Atlantic. This study benefited from grants PIP 5424 (CONICET, Argentina), PICT 12776 (Agencia Nacional de Promoción Científica, Argentina), and Proyecto N503 (Secretaría de Ciencia y Técnica, Universidad Nacional de La Plata) to Gustavo Politis; and PIP 5077/4 (CONICET), Proyecto N459 (Secretaría de Ciencia y Técnica, Universidad Nacional de La Plata), and PICT 2006-00468 (Agencia Nacional de Promoción Científica, Argentina) to Marina L. Aguirre.

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BONOMO AND AGUIRRE Isla, F.I., & Bértola, G.R. (2005). Litoral Bonaerense. In R. Barrio, R. Etcheverry, M. Caballé, & E. Llambías, Relatorio del XVI Congreso Geológico Argentino (pp. 265–276). La Plata: Asociación Geológica Argentina. Kidwell, S. (1986). Models for fossil concentrations: Paleobiologic implications. Paleobiology, 12, 6–24. Kowalewski, M., & Flessa, K. (1995). Comparative taphonomy and faunal composition of shelly cheniers from northeastern Baja California, Mexico. Ciencia Marina, 21, 155–177. Lahille, F. (1895). Contribución al estudio de las volutas argentinas. Revista del Museo de La Plata, 6, 296–332. Lasta, M., Ciocco, N., Bremec, C., & Roux, A. (1998). Moluscos bivalvos y gasterópodos. In E. Boschi (Ed.), El mar y sus recursos pesqueros, 2: Los moluscos de interés pesquero (pp. 115–142). Mar del Plata: Instituto Nacional de Investigación y Desarrollo Pesquero. Loponte, D., & Acosta, A. (1986). Sitios acerámicos de la costa de Necochea. Estudios de Antropología Pampeana, 1, 3–47. Lutaenko, K.A., Zhushchikhovskaya, I.S., Mikishin, Y., & Popov, A.N (2007). Mid-Holocene climatic changes and cultural dynamics in the basin of the Sea of Japan and adjacent areas. In D. Anderson, K. Maasch, & D. Sandweiss, (Eds.), Climate change and cultural dynamics (pp. 331–406). Amsterdan: Elsevier. Madrid, P., & Politis, G. (1991). Estudios paleoambientales en la región pampeana: Un enfoque multidisciplinario del sitio La Toma. XI Congreso Nacional de Arqueología Chilena (pp. 131–152). Santiago de Chile: Sociedad Chilena de Arqueología. Madrid, P., Politis, G., March, R., & Bonomo, M. (2002). Arqueología microrregional en el sudeste de la región pampeana argentina: El curso del río Quequén Salado. Relaciones de la Sociedad Argentina de Antropología, 27, 327–355. Martínez, G. (1999). Tecnología, subsistencia y asentamiento en el curso medio del Río Quequén Grande: Un enfoque arqueológico. Unpublished doctoral dissertation, Facultad de Ciencias Naturales y Museo, UNLP, La Plata. Martínez, S., Rojas, A., Ubilla, M., Verde, M., Perea, D., & Piñeiro, G. (2006). Molluscan assemblages from the marine Holocene of Uruguay: Composition, geochronology, and palaeoenvironmental signals. Ameghiniana, 43, 385–398. Martínez Soler, B.J. (1958–59). Conchyliología ethnológica. Runa, 9, 267–322. Mazzanti, D.L., & Quintana, C. (Eds.). (2001). Cueva Tixi: Cazadores y recolectores de las sierras de Tandilia Oriental. 1: Geología, paleontología y zooarqueología. Publicación Especial 1. Mar del Plata: Laboratorio de Arqueología, UNMdP. Mazzanti, D.L., & Valverde, F. (2001). Artefactos sobre hueso, asta y valva. In D. Mazzanti & C. Quintana (Eds.), Cueva Tixi: Cazadores y recolectores de las sierras de Tandilia Oriental (pp. 157–180). Publicación Especial 1. Mar del Plata: Laboratorio de Arqueología, UNMdP. Mehdahl, K. (1994). Biofacies and taphofacies of a Holocene macrotidal environment: Bahía La Cholla, northern Gulf of California. Ciencia Marina, 20, 555–583. Mianzan, H., Lasta, C., Acha, E., Guerrero, R., Macchi, G., & Bremec, C. (2001). The Río de La Plata estuary, Argentina-Uruguay. Ecological Studies, 144, 185–204. Oliva, F., Gil, A., & Roa, M. (1990). Recientes investigaciones en el sitio San Martín, 1 (BU/PU/5): Partido de Puán, provincia de Buenos Aires. Shincal, 3,135–139. Orquera, L.A., & Piana, E.L. (1999). Arqueología de la región del canal Beagle (Tierra del Fuego, República Argentina). Buenos Aires: Sociedad Argentina de Antropología. Parsons, K., & Brett, C. (1991). Taphonomic processes and biases in modern marine environments: An actualistic perspectiva on fósil assemblage preservation. In S.K. Donovan (Ed.), The processes of fossilization (pp. 22–65). London: Belthaven Press. Penchaszadeh, P. (2004). Caracoles, almejas y mejillones. In E. Boschi & B. Cousseau (Eds.), La vida entre mareas: Vegetales y animales de las costas de Mar del Plata, Argentina (pp. 253–270). Mar del Plata: Publicación Inidep. Petz, R., & Saghessi, M. (2000). Investigaciones arqueológicas en las lagunas Encadenadas del oeste, partidos de Adolfo Alsina y Guaminí (provincia de Buenos Aires). II Congreso de Arqueología de la Región Pampeana Argentina, Universidad Nacional de Mar del Plata, Mar del Plata. Politis, G. (1984). Arqueología del Area Interserrana Bonaerense. Unpublished doctoral dissertation, Facultad de Ciencias Naturales y Museo, UNLP, La Plata.

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HOLOCENE MOLLUSCS FROM ARCHAEOLOGICAL SITES Politis, G., & Madrid, P. (2001). Arqueología Pampeana: Estado actual y perspectivas. In E. Berberián & A. Nielsen (Eds.), Historia Argentina prehispánica II (pp. 737–814). Córdoba: Brujas. Politis, G., Bonomo, M., & Prates, L. (2003). Territorio y movilidad entre la costa Atlántica y el interior de la Región Pampeana (Argentina). Estudos Ibero-Americanos, 29, 11–35. Prates, L. (2008). Los indígenas del río Negro. Un enfoque arqueológico. Buenos Aires: Sociedad Argentina de Antropología, Colección Tesis Doctorales. Razjigaeva, N.G., Korotky A.M., Grebennikova T.A., Ganzey L.A., Mokhova L.M., Bazarova V.B., Sulerzhitsky L.D., & Lutaenko K.A. (2002). Holocene climatic changes and environmental history of Iturup Island, Kurile Islands, northwestern Pacific. The Holocene, 12, 469–480. Ríos, E., (1994). Seashells of Brazil. Rio Grande: Fundação Universidade do Rio Grande. Rollins, H., Sandweiss, D., & Rollins, J. (1990). Molluscs and coastal archaeology; A review. In N. Lasca & J. Donahue (Eds.), Archaeological geology of North America (pp. 467–478). Centennial Special Volume 4. Boulder, CO: Geological Society of America. Schnack, E., Isla, F., De Francesco, F., & Fucks, E. (2005). Estratigrafia del Cuaternario marino tardío en la Provincia de Buenos Aires. In R. Barrio, R. Etcheverry, M. Caballé, & E. Llambías, relatorio del XVI Congreso Geológico Argentino (pp. 159–182). La Plata: Asociación Geológica Argentina. Taborin, Y. (1993). La parure en coquillage au Paléolithique. Gallia Préhistoire, 29. Paris: Centre National de Recherche Scientifique. Vignati, M.A. (1930). Restos del traje ceremonial de un médico patagón. Notas del Museo Etnográfico, 4, 7–52. Violante, R. (1988). Geología de la “planicie costera” entre Villa Gessell y Faro Querandí, Provincia de Buenos Aires. Unpublished doctoral dissertation, Facultad de Ciencias Naturales y Museo, UNLP, La Plata.

Received 12 March 2007 Accepted for publication 16 September 2008 Scientific editing by Jeff Blackford


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Appendix 1. Synthesis (taxonomy and distribution) of the molluscan taxa recovered from the sites revisited. Phylum MOLLUSCA Class GASTROPODA Cuvier, 1797 Genus FISSURELLA Bruguière, 1789 Subgenus FISSURELLA Bruguière, 1789 Fissurella sp. (Figure 4a, 1) Dimensions: 20–80 mm Stratigraphic Range: Pleistocene–Recent Fossil Records in the Quaternary of Argentina: Patagonia: Pleistocene from Cabo Raso to Bahía Bustamante; Holocene from Cabo Raso to Tierra del Fuego. Modern Geographic Range: Atlantic littoral of South America from Tierra del Fuego to Golfo San Matías, Pacific coast of Chile and Cape Horn (Magellanean and Argentine Malacological provinces). Remarks: The shells of this genus are allochthonous for the Pampean region and must have been transported from the coast of northern Patagonia (Río Negro and/or Chubut provinces). Genus UROSALPINX Stimpson, 1865 Urosalpinx haneti (Petit, 1856) (Figure 4a, 2) Dimensions: 10–20 mm Stratigraphic Range: Miocene?, Pliocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Magdalena, Punta Indio and Bahía Samborombón; Holocene at Bahía Samborombón. Patagonia: Tertiary–Quaternary at Cerro Laciar, Santa Cruz. Modern Geographic Range: Absent along the Pampean littoral at present (Antillean and Brazilian Malacological provinces). Remarks: Often mentioned as Urosalpinx rushi (Pilsbry, 1897) by several previous authors. Genus MORULA Schumacher, 1817 Morula necocheana (Pilsbry, 1900) (Figure 4a, 3) Dimensions: 15–30 mm Stratigraphic Range: Pleistocene-Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Holocene at Bahía Blanca. Modern Geographic Range: Southern Brazil to Golfo San Matías in Patagonia (Argentine Malacological Province). Remarks: Cited under Drupa by previous authors. Genus ZIDONA H. & A. Adams, 1853 Zidona dufresnei (Donovan, 1823) (Figure 4a, 4) Dimensions: 40–250 mm Stratigraphic Range: Pleistocene-Recent.

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Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Punta Indio; Holocene at Bahía Samborombón, Sauce Grande and Bahía Blanca. Patagonia: Pleistocene from San Antonio Oeste to Puerto Lobos; Holocene at San Antonio Oeste. Modern Geographic Range: Rio de Janeiro, Brazil, south to Golfo San Matías (Argentine Malacological Province). Genus ADELOMELON Dall, 1906 (non Pilsbry & Olsson, 1954) Adelomelon ancilla (Lightfoot, 1786) (Figure 4a, 5) Dimensions: 150–166 mm Stratigraphic Range: Pleistocene-Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Faro Querandí; Holocene at Bahía Blanca. Modern Geographic Range: Southern Brazil to Estrecho de Magallanes and Islas Malvinas (Argentine and Magellanean provinces). Adelomelon beckii (Broderip, 1786) (Figure 4a, 6) Dimensions: 200–490 mm Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Miocene at Chubut; Holocene at Bahía Blanca and Bahía Solano. Modern Geographic Range: Southern Brazil to Tierra del Fuego (Argentine and Magellanean provinces). Adelomelon (Pachycymbiola) brasiliana (Lamarck, 1811) (Figure 4a, 7) Dimensions: 36–200 mm Stratigraphic Range: Pleistocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene from Magdalena, Punta Indio, Bahía Samborombón, and Laguna de Sotelo; Holocene at Magdalena, Bahía Samborombón, and Bahía Blanca. Patagonia: Pleistocene-Holocene San Antonio Oeste. Modern Geographic Range: Rio de Janeiro south to Golfo San Matías (Argentine Province). Genus OLIVANCILLARIA d’Orbigny, 1842 Subgenus OLIVANCILLARIA s.s. Olivancillaria spp. (Figure 4a, 8, 9) Dimensions: 31–55 mm Stratigraphic Range: Miocene-Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Magdalena, Bahía Samborombón, and Faro Querandí; Holocene at Magdalena, Bahía Samborombón, and Mar Chiquita. Patagonia: Pleistocene at Puerto Lobos and Camarones; Holocene from San Antonio Oeste, Puerto Lobos, Camarones, and Tierra del Fuego. Modern Geographic Range: Espiritu Santo (Brazil) to Golfo San Matías (Argentine Province). GEOARCHAEOLOGY: AN INTERNATIONAL JOURNAL, VOL. 24, NO. 1

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Genus BUCCINANOPS d’Orbigny, 1841 Buccinanops spp. (Figure 4a, 10) Dimensions: 6–50 mm Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Magdalena, Punta Indio, Bahía Samborombón, Laguna de Sotelo, and Bahía Blanca; Holocene at Magdalena, Bahía Samborombón, Sauce Grande, and Bahía Blanca. Patagonia: Pleistocene from Puerto Lobos to Camarones; Holocene from San Antonio Oeste to Tierra del Fuego. Modern Geographic Range: Southern Brazil to Golfo San Matías (Argentine Province). Class BIVALVIA (Buonanni, 1681) Linné, 1758 Genus NOETIA Gray, 1857 Noetia bisulcata (Lamarck, 1819) (Figure 4b, 1) Dimensions: 10–41 mm Stratigraphic range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean coast: Pleistocene at Magdalena, Punta Indio, Bahía Samborombón, and Laguna de Sotelo; Holocene at Punta Indio and Bahía Samborombón. Earlier records are from Brazil, Surinam, and Central and North America (Aguirre, 1993 and references therein). Modern Geographic Range: Western Atlantic from Antilles to Uruguay. Probably Pacific coast of Panamá. Remarks: Mentioned under the genus Arca Linn. by most previous authors. This species is absent from the Argentine waters at present. Its modern northern shift represents a biotic response to climate change and sea-level fluctuations (Aguirre, 1993). Genus GLYCYMERIS Da Costa, 1778 Glycymeris (Glycymeris) longior (Sowerby, 1832) (Figure 4b, 2) Dimensions: 23–40 mm Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Magdalena, Punta Indio, Faro Querandí, Laguna de Sotelo, and Mar del Plata; Holocene at Punta Indio and Bahía Samborombón. Patagonia: Pleistocene from San Antonio Oeste to Camarones; Holocene from San Antonio Oeste to Cabo Raso. Modern Geographic Range: Espiritu Santo (Brazil) to Golfo San Matías (Argentine Province). Genus MYTILUS Linné, 1758 Mytilus sp. (Figure 4b, 3) Dimensions: 10–85 mm Stratigraphic Range: Miocene–Recent Fossil Records in the Quaternary of Argentina: Pampean region: 82



Pleistocene from Bahía Samborombón and Punta Hermengo. Patagonia: Miocene from Puerto Madryn; Pleistocene: from Península Valdés to Puerto Deseado; Holocene: from San Antonio Oeste to Camarones. Modern Geographic Range: Pandemic. Genus AEQUIPECTEN Fischer, 1887 Aequipecten sp. (Figure 4b, 4) Dimensions: 20–77 mm Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene from Magdalena and Punta Indio; Holocene from Bahía Samborombón and Bahía Blanca. Patagonia: Miocene at Puerto Madryn; Pleistocene from Puerto Lobos to Comodoro Rivadavia; Holocene from San Antonio Oeste to San Julián. Modern Geographic Range: Rio de Janeiro (Brasil) to Golfo Nuevo (Argentina); scarce in Camarones-Bahía Bustamante area. Remarks: Often mentioned as Pecten, a genus which is not recorded from the Quaternary deposits in the studied area. Previously cited under the genus Chlamys Röding (Aguirre & Farinati, 2000). Genus MESODESMA Deshayes, 1832 Mesodesma sp. (Figure 4b, 5) Dimensions: 20–50 mm Stratigraphic range: Holocene. Fossil Records in the Quaternary of Argentina: Holocene from the southeastern Pampean coast. Modern Geographic Range: Southern Brazil to Golfo San Matías (Argentine Province). Remarks: The only species of the genus living in the modern littoral between Punta Rasa and Bahía San Blas is Mesodesma mactroides Deshayes, 1854. This species was very common along the Pampean littoral waters until the end of the 20th century. However, due either to human explotation or to climate change impact, its abundance has remarkably decreased over the last decades. There are no records from littoral deposits (shell ridges, coastal lagoon facies) of the marine Quaternary of Argentina. Genus AMIANTIS Carpenter, 1864 Amiantis purpuratus (Lamarck, 1818) (Figure 4b, 6) Dimensions: 50–85 mm Stratigraphic Range: Miocene–Recent. Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Pampean region: Pleistocene at Magdalena, Punta Indio, Faro Querandí, and Laguna de Sotelo; Holocene at Laguna de Sotelo and Sauce Grande. Modern Geographic Range: Espiritu Santo (Brazil) to Golfo San Matías (Argentine Province).


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Genus PROTOTHACA Dall, 1902 Protothaca antiqua (King, 1832) (Figure 4b, 7) Dimensions: 60–80 mm Stratigraphic Range: Miocene–Recent. Fossil Records in the Quaternary of Argentina: Patagonia: Pleistocene from San Antonio Oeste to south of Caleta Olivia; Holocene from Puerto Lobos to Tierra del Fuego. Modern Geographic Range: Perú to Magallanes in Chile (Pacific Ocean); southern Brazil to Tierra del Fuego (Atlantic Ocean). Remarks: This species is allochthonous for the Pampean region and must have been carried by early humans from northern Patagonia (Río Negro and/or Chubut provinces), either collected from shell ridges or along the modern beach.

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Appendix 2. List of molluscan taxa found in natural shell cemeteries of the modern Bonaerensian littoral (Aguirre, 1989; this paper). Shell Cemeteries MOLLUSCAN TAXA

Punta Rasa

Claromecó

Gastropoda Tegula patagonica (d’Orb.) Crepidula protea d’Orb. Crepidula dilatata patagonica d’Orb. Natica isabelleana (d’Orb.) Epitonium georgettinum (Kien.) Zidona dufresnei (Donovan) Adelomelon ancilla (Sol.) Adelomelon brasiliana (Lamk.) Olivancillaria urceus (Röding) Olivancillaria carcellesi Klapp. Olivancillaria auricularia (Lamk.) Dorsanum moniliferum (Val.) Buccinanops cochlidium (Chemn.) Buccinanops deformis (King) Buccinanops gradatum (Desh.) Bivalvia Glycymeris longior (Sow.) Mytilus edulis Linn. Brachidontes rodriguezi (d’Orb.) Chlamys sp. Plicatula lanceo Lamk. Ostrea sp. Ostrea cf. Equestris Say Mactra isabelleana d’Orb. Raeta plicatella (Lamk.) Donax hanleyanus Phil. Mesodesma mactroides Desh. Tivella isabelleana (d’Orb.) Pitar rostratus (Koch) Lanceola purpuratus (Lamk.) Trachycardium muricatum (Linn.) Cyrtopleura lanceolada (d’Orb.) Corbula patagonica d’Orb. OTHER INVERTEBRATES Polychaetes Balanids Brachyura indet.


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