Abstract
Despite the multiple constraints of using shell mounds for building relative-sea-level (RSL) curves, one of the premises behind this use is still valid in modern archaeological research. This refers to the sites being continuously built near bodies of water rich in fish species and mollusk beds. Studies that combine the evolution of settlement patterns with the geological evolution of coastal areas in Brazil reach analogous results: the distribution of shell mounds in time and space follows the landscape transformations induced by RSL variations through the Holocene. Although shell mounds are not precise indicators of RSL, they provide evidences of the paleo-geographical changes during the Holocene, of which RSL is one of the many control variables. By collecting and transporting mollusks to the shell mounds, humans inevitably carry sediments from the substrate where mollusks live, for example, the beaches and lagoons near the sites. In this work, the geoarchaeological study of three shell mounds located in the southern coast of Santa Catarina State, combined with mollusk taxa identification, show the direct association of shell mound content with the changing landscape. The predominance of Ostrea sp. versus Anomalocardia brasiliana, the presence of colluvial versus lagoonal sediments, and the occurrence of echinoid spines versus muddy aggregates with diatoms, gastropods, and glauconitic clay characterize the distinction between sites built before and after the maximum Holocene transgression, respectively. This attests the potential of geoarchaeological analyses in shell mounds as a complementary proxy for paleo-environmental reconstructions.
Introduction
Shell mounds, or sambaquis, are accumulations of shells and fish bones mixed with natural sediments, organic debris, and other refuse, including artifacts, made by prehistoric hunter–gatherer–fisher populations. On the coast of Brazil, sites date back to c. 8700 cal. yr BP (Lima et al., 2002). The large volume and the frequent concentration of human burials inside the mounds lead researchers to interpret their formation as building endeavors, although function is still a matter of debate (Afonso and DeBlasis, 1994; DeBlasis et al., 1998; Figuti and Klokler, 1996; Gaspar et al., 2008; Klokler, 2008).
Dating carbonate shells collected from the base of the shell mounds has been traditionally used for reconstructing relative-sea-level (RSL) curves on the coast of Brazil (Bigarella, 1965; Fairbridge, 1976; Martin and Suguio, 1976; Suguio et al., 1992). Although many authors have argued the geological and cultural constraints of using shell mounds for building RSL curves and for paleo-shoreline reconstruction (see Angulo and Lessa, 1997; Angulo et al., 2006; Giannini et al., 2005; Scheel-Ybert et al., 2009), at least one of the premises behind this use is still present in modern archaeology. The premise refers to the chosen location for shell mound construction: always near large bodies of water containing a diversity of fish species, the main subsistence item among coastal groups (Figuti, 1992; Klokler, 2008), as well as intertidal and subtidal resources (Barbosa, 2007; DeBlasis et al., 2007; Gaspar et al., 2008; Klokler, 2008; Kneip, 2004).
Furthermore, recent archaeological studies show that the evolution of prehistoric settlement patterns in the states of Rio Grande do Sul, Santa Catarina, São Paulo, and Rio de Janeiro has been directly influenced by changes in coastal landscapes (Barbosa, 2007; Calippo, 2010; DeBlasis et al., 2007; Giannini et al., 2010; Kneip, 2004; Perreti, 2009), which ultimately result from RSL fluctuations after the maximum Holocene transgression, among other factors. Thus, even if shell mounds are not precise indicators of paleo-shorelines and RSL changes, their constant proximity to water sources means that they can still hold evidences of the paleo-geographical changes in coastal areas during the Holocene, for which RSL is only one of the controlling variables.
Mollusks in shell-bearing sites (e.g. shell middens, shell mounds, etc.) do not necessarily reflect the adjacent aquatic habitat (Claassen, 1991; Stein, 1992a; Waselkov, 1987), but the ecology of the predominant mollusk species can be used as a paleo-environmental proxy whenever there are multiple records available for comparison (geological, botanical, etc.; see examples in Cortez-Sanchez et al., 2008; Fa, 2008; Morey and Crothers, 1998; Morey et al., 2002; Sandweiss, 2003; Stein, 1992b; Van der Schriek et al., 2007, 2008). In this respect, the sediments contained in shell-bearing sites have been less used as an environmental proxy. However, considering the basket-load action of collecting and transporting mollusks to the sites, whether for consumption or as prime material for mound building, remnants of the substrate where the mollusks live (i.e. the beaches or lagoons that existed nearby) are inevitably carried to the sites. Likewise, since some Brazilian shell mounds show evidences of being built after reworking of debris deposited close by (Villagran et al., 2009, 2010), natural sediments from the surroundings could also be carried together. Thus, the sediments in the shell mounds can potentially provide information on the paleo-geography near the sites and can, therefore, be used as another environmental proxy. Based on the dynamics of shell mound formation, we present the geoarchaeological study of three Brazilian shell mounds and their potential for reconstructing paleo-geography, by combining the traditional identification of mollusk species with geoarchaeological analyses of shell mound sediments.
Study area
The study area is located on the southern coast of Santa Catarina State (southern Brazil; Figure 1(A)). The Quaternary geology of the region is characterized by four main depositional systems: lagoonal, transgressive barrier, regressive barrier (strandplain), and eolian (Giannini, 1993; Giannini et al., 2007; Figure 1(B)). The maximum Holocene RSL rise was reached in the area 5700–5100 yr BP or even before, according to Angulo et al. (1999, 2006). Thus, the maximum Holocene transgression coincided or happened before this date. Previously, the area was flooded by a large paleo-bay partially isolated by a sandy transgressive barrier forming a bay–lagoon system. This system differs from the valley–lagoon system that also developed in the area during the maximum Holocene transgression, when valleys incised in regressive marine Pleistocene terraces were drowned and transformed into a lagoon. After 5700–5100 BP, the RSL decline and the progradation of the Tubarão river delta reconfigured the bay–lagoon and valley–lagoon systems into a complex of interconnected coastal lagoons, which were progressively silted until the arrangement seen nowadays (Amaral et al., 2011; Fornari et al., 2012; Giannini, 1993; Nascimento, 2010). The eolian system accompanied this process and developed over the lagoonal, transgressive barrier and regressive barrier (strandplain) systems in at least four subsequent stages from the Pleistocene to the present (Giannini, 1993; Giannini et al., 2007; Martinho, 2004; Sawakuchi et al., 2009). The transgressive barrier system is nowadays almost completely covered by eolian deposits.

(A) Study area (Santa Catarina State, southern Brazil). (B) Quaternary depositional systems and location of the three studied shell mounds: Caipora, Morrinhos, and Jabuticabeira-1. The transgressive barrier system does not appear in the map since it is almost completely covered by eolian deposits.
Materials and methods
Three shell mounds were analyzed in this study: Caipora, Morrinhos, and Jabuticabeira-1 (Figure 1(B)). The sites are among the oldest shell mounds in the region and are located on the hillslopes of pre-Cenozoic granite hills, at about 10–15 km from the present coastline. Further information on size, chronology, and UTM coordinates of the sites is given in Table 1. Of the three sites, only Caipora was probably built before the Holocene maximum transgression, when most of the area was covered by a paleo-bay, while Morrinhos and Jabuticabeira-1 were built during the regression phase when the lagoon system developed.
Size, chronology, sampled material, and UTM coordinates of the three shell mounds analyzed in this study. The maximum and minimum dates are from a sample collected near the base and the top of the site, respectively.
Source: Calibrated dates taken from Giannini et al. (2010).
Mollusk samples and bulk and undisturbed sediment samples were collected from the stratigraphic successions at each site (Figure 2). Mollusk sampling was made using standard volumes of 20 L. Material was recovered by dry and water screening using 4- and 2-mm sieves and weighed for determination of frequency of species. Mollusk analyses of Caipora and Jabuticabeira-1 were taken from Ferraz (2010). Morrinhos was sampled and analyzed for this study. For grain-size analyses, only the mineral sand fraction was used to avoid the interference of anthropic components in the results (e.g. bone, shell, and charcoal). The sand fraction was dry sieved in 0.5-phi intervals. For comparison with natural sediments, off-site samples were collected from nine trenches opened near Caipora and Jabuticabeira-1, in order to sample colluvial and lagoonal sediments, respectively. Data on grain-size distributions of lagoon sediments near Morrinhos were taken from Nascimento (2010; see Supplementary Table A-1 (available online) for location of off-site samples). Grain-size results are expressed using box plot diagrams containing descriptive statistics (mean diameter, standard deviation, and skewness) calculated through the moment’s method. For micromorphology, 14 thin sections of 30 µm thickness were made out of impregnated blocks and analyzed at magnifications ranging from 10× to 50×, using Olympus BX51 and Zeiss Axioplan 2 microscopes.

Sedimentary successions and sampling locations in the three shell mounds studied: (A) Caipora, (B and B.1) Morrinhos, and (C and C.1) Jabuticabeira-1 lower profile, and (D and D.1) upper profile. The black boxes indicate the location and type of sampling (m: micromorphology; s: bulk sediment; k: mollusks).
Results
The analyses of mollusk taxa showed two distinct assemblages: one in Caipora, with a majority of Ostrea sp. (>70%) and presence of vermetid tubes (<1%), and the other in Morrinhos and Jabuticabeira-1, where Anomalocardia brasiliana (Gmelin 1971) is frequent (>60%). A. brasiliana is also present in Caipora, although in lower frequency (~15%), and both Ostrea sp. and mussels are recurrent in the upper layers of Morrinhos and Jabuticabeira-1 (~20% and 5–20%, respectively). Mussel fragments are present in Caipora and Jabuticabeira-1 in equivalent proportion (~4%), with higher frequency in Morrinhos (13–17%). Table 2 shows the complete list of mollusk taxa identified in the three sites. The low frequency (<1%) of species smaller than 1.5 cm in length (e.g. Nassarius vivex, Littorina flava, Neritina virginea, among others) suggests their accidental gathering from the mollusk beds where the dominant species were collected.
Distribution of mollusk species in Caipora, Morrinhos, and Jabuticabeira-1.
: dominant species (70–100%); ••••: frequent species (50–70%); •••: common species (10–50%); ••: some individuals or fragments (1–10%); •: occasional appearance (<1%).
Micromorphology identified four micro-components of potential association with the paleo-geography near the sites. These are echinoid spines (Figure 3(A)), only seen in Caipora, and two biological and a mineral component exclusively seen in Morrinhos and Jabuticabeira-1 inside angular muddy aggregates of 5 mm to less than 2 cm size (Figure 3(B)). The muddy aggregates contain diatoms (Figure 3(C) and (D)), a gastropod genus identified as Heleobia sp. (Figure 3(E)), and grains of glauconitic clay minerals (Figure 3(F)). In Jabuticabeira-1, the aggregates show evidences of anthropic heating like carbonization and rubefaction (Figure 4(A)). Besides the changes induced by heating, the diatoms and gastropods are still visible in the aggregates (Figure 4(B) and (C)). The identification of glauconitic clay was done by its optical properties: grains of cryptocrystalline green clay of 20–30 µm, sub-equidimensional and subrounded shape, and first-order interference color (maximum estimated birefringence around 0.020). XRD analysis to confirm this mineralogy could not be made because of the difficulty to concentrate material that happens in such low quantity. Besides shells, bone, and charcoal fragments, the coarse fraction in the three sites is made of variable proportions of quartz grains, feldspars, and rock fragments, with rock fragments being predominant only in Caipora.

Photomicrographs of microscopic components that give information on paleo-geography (PPL): (A) echinoid spines (s) seen in Caipora; (B) muddy aggregates (c) seen only in Morrinhos and Jabuticabeira-1 associated with shells of Mytella sp. (m); (C) detail view of the muddy clay aggregates with diatoms (d), mostly of the species Paralia sulcata; (D) close view of diatoms (d) of the species P. sulcata, commonly found in paleo-lagoonal sediments of the study area; (E) cross-section of Heleobia sp. (h) frequently found inside the muddy clay aggregates; (F) glauconitic clay (g) in the muddy aggregates. Box plot diagrams with grain-size statistics for the terrigenous sand fraction in archaeological and natural sediments: (G) standard deviation, (H) mean diameter, and (I) skewness. Samples were collected from Caipora, colluvial sediments near Caipora (Coll.), Jabuticabeira-1 (Jab. 1), paleo-lagoonal sediments near Jabuticabeira-1 (PL Jab.1), Morrinhos (Mor.), and paleo-lagoonal sediments near Morrinhis (PL Mor.). Note the affinity between the terrigenous sand fraction in archaeological sediments and natural sediments from the proximities of each site.

Photomicrographs of muddy aggregates in Jabuticabeira-1 with signs of heating (PPL): carbonized (ca) and rubefied (ra) aggregates in-between shell fragments. Note (A) Heleobia sp. (h) within the rubefied clay, (B) rubefied muddy aggregate with diatom species (d) Paralia sulcata, and (C) carbonized muddy aggregate with diatoms.
Box plots of grain-size statistics (mean diameter, standard deviation, and skewness; see Supplementary Table A-2 (available online) for grain-size data) showed the affinity of shell mound sediments with the natural sediments in close proximity: colluvial deposits in Caipora and lagoon sediments in Morrinhos and Jabuticabeira 1 (Figure 3(G)–(I)). For the three statistical parameters, the interquartile range for natural sediments (colluvium in the case of Caipora and paleo-lagoonal for Jabuticabeira-1 and Morrinhos) is contained in the variation range for shell mound sediments.
Discussion
The data set allowed differentiating between the sedimentary signatures of shell mounds built before and after the maximum Holocene transgression. This difference expresses the interplay between the changing environment and the preferences of coastal groups.
Pre-maximum Holocene transgression shell mound
The faunal assemblage in Caipora is characterized by the predominance of Ostrea sp. with frequent vermetid tubes attached to the surfaces of the shells (vermetid fragments also appear loose in the sediments). Two species of Oyster sp. are found on the southern coasts of Brazil: Ostrea equestris (Say 1834) and Ostrea puelchana (Orbigny 1841). Both are euryhaline intertidal and subtidal marine species that live on rocky bottoms (De Souza et al., 2011; Rios, 1985), while vermetids are marine gastropods that live on rocky shores exposed to wave action (Laborel and Laborel-Deguen, 1996). The predominance of Ostrea sp. in Caipora and the exclusive presence of vermetid tubes in this site suggest that before the maximum Holocene transgression, coastal populations were gathering mollusks from rocky shores exposed to marine waters. The echinoid spines (625–645 µm, basal section) seen in thin sections corroborate this. Echinoids are marine subtidal invertebrates that make up the biological assemblage in rocky coasts (Laborel and Laborel-Deguen, 1996). Its presence in the shell mound sediments indicates marine influence in the body of water near the site where resources were being captured. Echinoid spines have also been described in thin sections from other shell-bearing sites located on marine shores in South Africa (Goldberg, 2000), the San Juan Islands (United States; Stein et al., 2011), and Rio de Janeiro (Brazil; Corrêa et al., 2013). The body of water near Caipora would correspond to the large paleo-bay that formed in the mid-Holocene transgression flooding. The chronology of Caipora, the predominance of Ostrea sp., and the presence of echinoids suggest that rocky shores were readily exploited for their proximity to the site (Figure 5(A)).

Landscape reconstruction before and after the maximum Holocene transgression with diagnostic macro-mollusks and micro-components: (A) the Caipora was built when the area was covered by a large paleo-bay and (B) the Morrinhos and Jabuticabeira-1 sites were first settled between 5500 and 4000 BP, already in the regression phase near the newly formed lagoonal system.
The comparison of grain-size statistics of Caipora and nearby natural sediments (colluvium) shows close affinity (Figure 3(G)–(I)). This, and the visual evaluation of the mineral coarse fraction in thin section (with coarse angular quartz grains, sub-angular feldspars and granule sized angular rock fragments), confirms the association of shell mound sediments with colluvial deposits. Besides collecting mollusks from the rocky shores nearby, colluvial sediments from the surroundings were also being brought to the mounds, mixed with faunal and other organic components. This supports the hypothesis of shell mounds being built after reworking of debris previously left near the sites. The natural deposition of colluvial sediments on the shell mound is precluded by the higher topography of the mound in relation to the surroundings.
Post-maximum Holocene transgression shell mounds
Both Morrinhos and Jabuticabeira-1 are mainly composed of A. brasiliana, an intertidal euryhaline species that lives in muddy and muddy sand beaches, commonly bays and estuaries (Rios, 1985). It burrows 5 cm deep in quiet waters with little disturbance of the bottom deposits and is easily collected during low tides (Narchi, 1972). This species proliferated in the region with the development of the lagoon system. However, the presence of Ostrea sp. in the upper layers of both sites attests continuity in the exploitation of this resource with a diminishment in intensity. By the time Morrinhos and Jabuticabeira-1 were active, Ostrea sp. could be collected from rocky shores within the lagoon system or brought from the maritime rocky shores located more than 5 km away (Figure 5(B)). The intermixing of layers rich in A. brasiliana and Ostrea sp. is seen in other sites of the region (Klokler, 2008; Kneip, 2004). The switch in species could be indicative of a foraging behavior intended to prevent the over-exploitation of the shell beds (see Mannino and Thomas, 2002).
Besides the shells of Ostrea sp. and A. brasiliana, sediments in Morrinhos and Jabuticabeira-1 contain quartz sands associated with lagoon sediments, as demonstrated by comparison of statistical parameters between natural and cultural sediments (Figure 3(G)–(I)). Muddy aggregates are another common component in the shell mound sediments. Micromorphological analyses showed that the muddy aggregates contain diatoms, gastropods and glauconitic clay, and, in Jabuticabeira-1, the aggregates were the substrate of hearths lit near or over the shell mound.
The most abundant diatom species found in the muddy aggregates is Paralia sulcata. Other species like Triceratium favus, Actinoptychus vulgaris, and fragments of Biddulphia pulchella and Coscinodiscus sp. appear in less abundance. Both P. sulcata and B. pulchella are marine eurytropic species, which means they tolerate variations in water salinity. They have optimal development in eusaline conditions (salinity between 30‰ and 40‰), but can also grow in meso-polysaline (salinity between 5‰ and 30‰) and even metasaline waters (salinity above 40‰). T. favus and A. vulgaris are marine stenotopic species, with low tolerance to variations in salinity that live only in eusaline conditions. Thus, the diatom assembly in the muddy aggregates suggests meso to eusaline conditions (salinity from 5–40‰) associated with estuarine–lagoonal sediments.
This hypothesis is reinforced by the fine grain size of the aggregates and the high frequency of P. sulcata, whose predominance has already been detected in lagoonal sediments of the area (Amaral et al., 2011). Likewise, the muddy aggregates in Morrinhos and Jabuticabeira-1 contain many specimens of Heleobia sp. (Stimpson 1865), a gastropod commonly found in muddy sandy substrate associated with estuarine–lagoonal environments (Rios, 1985). This gastropod was only identified at genus level because shape alone is not sufficient to determine the species. However, the species Heleobia australis is well known in the area and lives nowadays in the Garopaba do Sul lagoon (Netto et al., 2012), as well as in other coastal lagoons of Brazil, Uruguay, and Argentina (Martínez et al., 2006).
The glauconitic clay minerals in the muddy aggregates testify the history of the formation of the lagoon deposits. Glauconite is a 2:1 clay mineral formed in eodiagenesis, under the influence of marine waters in low sedimentation rates (see Odin, 1988; Odin and Fullagar, 1988). Glauconite is commonly found today in the continental platform (Dillenburg et al., 2000). Its presence in the muddy aggregates, associated with estuarine–lagoonal sediments (fresh to brackish waters) according to the diatom and mollusk assembly, suggests the mechanical reworking of marine sediments from the shelf during the formation of the lagoon system.
A clear association of mussel shells and the muddy aggregates was seen in thin section, which could serve as an indirect indicator of the mussel species. Two mussels are common in Brazilian shell mounds: Mytella guayanensis (Lamarck 1819) and Brachiodontes sp. (Swaison 1840; Figuti, 1993). The mussel M. guayanensis is an euryhaline intertidal bivalve that lives in muddy lagoon substrates and intertidal flats, even attached to mangrove roots (Bacon, 1975; De Souza et al., 2011), while Brachiodontes sp. lives only on rocky substrate (De Souza et al., 2011). The lagoon origin of the muddy aggregates and their frequent association with mussel valves suggests that M. guayanensis, collected from the lagoons nearby, is present in the sites.
The muddy aggregates and the predominant mollusk assemblage in Morrinhos and Jabuticabeira-1 indicate the exploitation of lagoon resources. The affinity seen between the paleo-lagoonal sands and shell mound sands confirm this.
Conclusion
Studies on the relationship of coastal groups and the changing landscape of the Holocene have been done in different states of Brazil (Rio Grande do Sul, São Paulo, and Rio de Janeiro). By contrasting site location and chronology with the RSL curves, studies have shown that shell mounds’ distribution in time and space followed the landscape transformations induced by Holocene RSL variations. This is because of the constant construction of shell mounds near bodies of water, sources of fish, and intertidal resources. In several coastal areas of the world, the ecology of the predominant mollusk taxa, in combination with other records, has been used for paleo-environmental reconstructions. In this work, geoarchaeological analyses in three shell mounds of Santa Catarina state show that archaeological sediments can also record the paleo-geographical changes in coastal settings. The accidental carrying of natural sediments from the shell beds to the shell mounds is the human action behind such use of archeological sediments as an environmental proxy.
In the study area, the sediments and mollusk assemblages in sites built prior to and after the maximum Holocene transgression reflect the paleo-geography near the sites, determined partially by RSL changes. The prehistoric occupations near the large paleo-bay before 5100 yr BP (minimum age of the Holocene maximum transgression) and the exploitation of its resources are evidenced in shell mounds containing Ostrea sp., with vermetids, echinoid spines, and colluvial sediments from the hilly surroundings. The development of the lagoon system after the Holocene transgression and the exploitation of its resources is evidenced in the shell mounds made of A. brasiliana and M. guayanensis and containing paleo-lagoonal sediments (from sands to clay aggregates with diatoms and gastropods). The analyses of shell mound sediments show the constant choice of locating settlements near the bodies of water and the continuity in shell mound building practices, besides landscape transformations.
Footnotes
Acknowledgements
The authors thank Professor Paulo DeBlasis (MAE/USP) coordinator of the ‘Sambaquis and Landscape’ research project. The authors also thank Professor Daniela Klokler, Professor Milene Fornari, Dr Paula G. C. Amaral, Kiristen Bright, Julie Boreham, and Tobias Sprafke. Analyses were done at the laboratories of sedimentology and sedimentary petrology of the Institute of Geosciences, University of São Paulo. Two anonymous reviewers are also thanked for their helpful comments.
Funding
The doctoral research of Ximena S. Villagran was supported by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, proc. 08/51264-0). The study was supported by a research grant of the FAPESP (proc. 04/11038-0).
References
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