Research Article Tropical Conservation Science Volume 16: 1–18 Long-Term Perspective on Fishing and © The Author(s) 2023 Article reuse guidelines: Mammal Defaunation in the Atlantic Forest sagepub.com/journals-permissions DOI: 10.1177/19400829231218419 journals.sagepub.com/home/trc Coast of Brazil Using Archaeological Faunal Remains Thiago Fossile1,2 , Krista McGrath1,2, Samantha Presslee3, George Fogarty3, Diego Dias Pavei4,5, Maria Cristina Alves6, Jessica Ferreira7, Tatiane Andaluzia Montes8, Dione da Rocha Bandeira6,9,10, Fernanda Mara Borba10, and André Carlo Colonese1,2 Abstract Background and research aim: Escalating anthropogenic threats to the Atlantic Forest, a renowned biodiversity hotspot, has placed the region in the spotlight for current conservation efforts. Faunal overexploitation and habitat degradation are among key factors driving the region’s recent declines in faunal populations and biodiversity. Assessing the scale of these impacts is complicated by the near-complete absence of historical reference baselines. Methodology: Here, we provide a contribution to bridge this knowledge gap by analysing faunal remains from two historical archaeological sites, Morro Grande 1 (MG1) and Praia Grande Unidade 21 (PG-U21), in Babitonga Bay (Santa Catarina state, Brazil) dated between 1750 to 1950 AD. Results: Our results revealed that fishing and hunting played a crucial role in household consumption and economic livelihoods during the European colonisation of southern Brazil. Native terrestrial mammals made up a significant proportion of faunal remains at both sites. This suggests that species currently undergoing population declines were exploited in the region for at least the past two centuries. Conclusion: This study provides compelling evidence that the selective hunting of medium- and large-bodied native terrestrial mammals has persisted in the region for over 4500 years, and requires us to reconsider the idea of a heavy reliance on domestic animals during early European colonisation of southern Brazil. Our study thus traces the causes of regional terrestrial mammal defaunation back to the Pre-colonial and Historical times. Implication for conservation: We rec- ommend the integration of historical and archaeological data into modern faunal population assessments and conservation initiatives to set more informed reference baselines. 1 Institute of Environmental Science and Technology (ICTA - UAB), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain 2 Department of Prehistory, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain 3 BioArCh, Department of Archaeology, University of York, York, UK 4 Laboratório de Ensino e Pesquisa em Antropologia e Arqueologia (LEPAArq), Universidade Federal de Pelotas (UFPel), Pelotas, Brazil 5 Laboratório de arqueologia Pedro Ignacio Schimitz (LAPIS), Universidade do Extremo Sul Catarinense (UNESC), Criciúma, Brazil 6 Laboratório de Arqueologia e Patrimônio Arqueológico (LAPArq), Universidade da Região de Joinville (Univille), Joinville, Brazil 7 Programa de Pós-graduação em Arqueologia, Universidade Federal do Rio de Janeiro (UFRJ), São Cristóvão, Brazil 8 Programa de Pós-graduação em Antropologia e Arqueologia, Universidade Federal do Paraná (UFPR), Curitiba, Brazil 9 Programa de Pós-graduação em Patrimônio Cultural e Sociedade, Universidade da Região de Joinville (Univille), Joinville, Brazil 10 Museu Arqueológico de Sambaqui de Joinville (MASJ), Joinville, Brazil Received: 11 August 2023; accepted: 19 November 2023 Corresponding Author: Thiago Fossile, Institute of Environmental Science and Technology (ICTA - UAB) and Department of Prehistory, Universitat Autònoma de Barcelona, Campus de la UAB, Cerdanyola del Vallès 08193, Spain. Email: thiago.fossile@uab.cat Creative Commons Non Commercial CC BY-NC: This article is distributed under the terms of the Creative Commons Attribution-NonCommercial 4.0 License (https://creativecommons.org/licenses/by-nc/4.0/) which permits non-commercial use, reproduction and distribution of the work without further permission provided the original work is attributed as specified on the SAGE and Open Access pages (https://us.sagepub.com/en-us/nam/open-access-at-sage). 2 Tropical Conservation Science Keywords Atlantic Forest, Babitonga Bay, faunal analysis, colonial fishing and hunting, mammal defaunation Introduction Archaeology emerges as a discipline with the potential to illuminate the past daily experiences of vulnerable groups and Tropical and subtropical regions of South America are facing minorities, as well as their interactions with the surrounding significant environmental and ecological challenges which environments (Funari, 1994; Symanski, 2009; Symanski & parallel the global trends resulting from the rapid economic Zarankin, 2014). Currently, Historical Archaeology in Brazil expansion of recent decades (Bogoni et al., 2020; Steffen remains heavily biassed towards the analysis of ceramic et al., 2015; Verba et al., 2020). Brazil’s renowned biodi- artefacts, shipwrecks, infrastructure, and housing and set- versity hotspots of the Amazon and the Atlantic Forest tlement patterns (Lima, 1993; Symanski, 2009; Symanski & (Colombo & Joly, 2010; Rezende et al., 2018) are currently Zarankin, 2014), while faunal remains have received only undergoing rapid biodiversity and population loss due to superficial attention (Nobre, 2004). Analysis of terrestrial overexploitation and habitat degradation (Ceballos et al., mammal remains can provide insights into species compo- 2017; de Lima et al., 2020; Galetti et al., 2015, 2017; sition, distribution and relative abundance, as well as their Scarano & Ceotto, 2015; Verba et al., 2020). Recent studies function (food sources) and perceived value (economic, suggest a staggering loss of over 60% of mammals in the symbolic, etc.) to past human societies, and species responses Atlantic Forest (Bogoni et al., 2020); unfortunately, such to anthropogenic activities (Barnosky et al., 2017; Lyman & figures are likely an underestimation if reference baselines Cannon, 2017; Stahl, 2008). This last point is particularly can be extended to include Pre-colonial and historical relevant in conservation debates due to well known effects of periods. subsistence hunting on tropical forest composition, most Studies have shown that certain anthropogenic stressors responsible for impacting animal diversity and abundance in notably through the removal of medium- and large-bodied recent decades have been present for over a century. As early terrestrial mammals with low population densities (Bogoni as the mid-18th century, intellectual elites in colonial Brazil et al., 2020; Dirzo et al., 2014; Galetti et al., 2017; Peres, were already expressing increasing concern regarding the 2000). Yet, little is known about these processes in historical adverse environmental impacts of colonial economies, es- times, nor the extent to which they may have contributed to pecially deforestation (Pádua, 2000). Similarly, in his seminal the increasing rates of defaunation in the Atlantic Forest in work the Destruction of the Brazilian Atlantic Forest, Dean recent decades (Galetti et al., 2017). discussed the persistent human impact on plants and animals To address this knowledge gap, we present the results of in extensive regions of the Atlantic Forest since, at least, the zooarchaeological analyses conducted on faunal remains European colonisation (Dean, 1997). Others have shown that from colonial (prior to 1822) and post-colonial archaeological colonial and post-colonial fisheries were also capable of sites located in Babitonga Bay, Santa Catarina state (southern overexploiting local organisms, causing a noticeable decline Brazil) (Figure 1). The region preserves numerous historical in fish and coral populations (Fogliarini et al., 2022; Sandoval archaeological sites spanning from the 18th to the 20th Gallardo et al., 2021). Our understanding of these historical centuries (Bandeira & Alves, 2012), whose economies de- processes has predominantly relied on administrative docu- pended on manioc, rice and livestock production, along with ments, letters and accounts of European travellers and nat- hunting and fishing in forest and mangrove ecosystems uralists who began visiting the region in the 16th century (da (Saint-Hilaire, 1936). Of these sites, only 15 have been ex- Rocha, 2022; Russell-Wood, 2001; Whitehead, 2012). While cavated to date, five of which presented faunal remains. certain historical sources, such as newspapers and archives, Marine and terrestrial vertebrate remains from the archaeo- provide valuable descriptions of local plants and animals, logical sites of Morro Grande 1 (MG1) and Praia Grande along with their broad societal significance (Herbst et al., Unidade 21 (PG-U21) were analysed herein using an inte- 2023; Sandoval Gallardo et al., 2021), many written docu- grated approach of conventional zooarchaeological (mor- ments produced into the late 18th century lack specific details phological traits) analysis and collagen peptide mass concerning the relative abundance, distribution, and extent to fingerprinting, known as Zooarchaeology by Mass Spec- which several native animals were used (Pádua, 2000). trometry (ZooMS). Both sites were extensively excavated Nevertheless, these studies reinforce the growing consensus and have the largest collection of faunal remains of all the that a comprehensive understanding of current biodiversity historical sites. Changes in hunting strategies through time and environmental threats necessitates information that and their putative ecological impacts were studied by com- predates the anthropogenic impacts of the most recent de- paring the historical fauna with faunal data from Pre-colonial cades (Jackson & Hobbs, 2009; McClenachan et al., 2012). sites in the region (Enseada I, Bupeva II, Forte Marechal Luz, Fossile et al. 3 Figure 1. Location of Babitonga Bay in southern Brazil, including Pre-colonial and Historical sites in the region, and the two historical sites analysed herein, Praia Grande Unidade 21 (PG-U21) and Morro Grande 1 (MG1). Map generated on data publicly available from NASA/JPL- Caltech (adapted from https://www.jpl.nasa.gov/images/pia03388-south-america-shaded-relief-and-colored-height), Brazilian Agricultural Research Corporation - EMBRAPA (Embrapa, 2021), Natural Earth (https://www.naturalearthdata.com/), Brazilian Institute of Geography and Statistics - IBGE (https://www.ibge.gov.br/geociencias/todos-os-produtos-geociencias.html) and National Institute for Space Research - INPE (Assis et al., 2019). Itacoara, Ilha dos Espinheiros II and Cubatão I) and from earliest non-Indigenous colonial settlements in Babitonga contemporary traditional coastal communities in southern Bay (Pereira, 2004). Written and oral historical records in- Brazil, known as Caiçaras. The overarching aim of this study dicate that the region was primarily dedicated to cultivating was to improve our understanding of the ecological footprint manioc for flour production (Santos, 2004; Silva et al., 2001). of colonial and post-colonial societies, and provide a his- Excavations in 2001 covering an area of 374 m2 found one torical perspective to the ongoing debates on vertebrate de- homogeneous stratigraphic deposit with evidence of resi- faunation in the Atlantic Forest. More specifically, we aimed dential spaces, ceramic artefacts and coins dated to the 19th to understand the socioeconomic importance of marine fish century, and faunal remains. On the other hand, PG-U21 is and native terrestrial mammals during the historical occu- considered a coastal site located in Praia Grande which ex- pation in Babitonga Bay. hibits a clear association with fishing. Historical accounts describe fishing and fish processing (drying) in the area, along with manioc cultivation in more recent times (Alves & Methods Oliveira, 2002). Archaeological excavations conducted in 2002 over an area of 166 m2 uncovered abundant faunal Archaeological Contexts remains, ceramic artefacts dating to the 20th century, and MG1 and PG-U21 are situated within the municipality of São other artefacts likely associated with residential activities Francisco do Sul (Figure 1). Located ∼10 km from the coast, (Figure 2). The stratigraphic sequence at the site encom- MG1 holds historical significance as it is located in one of the passed six distinct layers (layers 1 to 6, with 1 being the most 4 Tropical Conservation Science Archaeological Faunal Identification The vertebrate faunal remains were identified through side- by-side comparison with reference collections from the Museu Arqueológico de Sambaqui de Joinville (MASJ, Brazil), the Laboratório de Arqueologia e Patrimônio Ar- queológico at the Universidade da Região de Joinville (LAPArq/Univille, Brazil) and the Laboratory of Archae- ozoology at the Universitat Autònoma de Barcelona (Spain); along with specialised literature (Adams & Crabtree, 2012; France, 2009; McLelland, 1991; Sebben et al., 2019; Tercerie et al., 2022; Wensing, 2009). The scientific names followed the World Register of Marine Species (Horton et al., 2020) and the Animal Diversity Web (Myers et al., 2020). The abundance of species was represented by the Number of Figure 2. Overview of Praia Grande Unidade 21 (PG-U21) Identified Specimens (NISP), a quantitative unit frequently showing the excavation area (bottom left) and specific grid (I10) analysed in this study. Red points in the excavation area (bottom used in zooarchaeological studies (Grayson, 1984; Lyman, left) indicate the distribution of faunal remains at the site (Alves & 2008a, 2018). The Number of Specimens (NSP), which Oliveira, 2002). The quadrants Q1, Q2, Q3 and Q4 have been represents the total of all analysed specimens (identified or depicted in grid H13 for reference to the excavation grid system unidentified), and the Number of Unidentified Specimens used. (NUSP) were also quantified. Fragments of the same spec- imen that could be refitted were counted as one. Species richness (SR) was calculated using the Minimum Level of recent and 6 being the earliest). These layers form over time, Taxonomic Identification, considering only the minimum one superimposed on the other as a result of human activities, hierarchical level for a particular taxa. In order to account for natural processes, or a combination of both, allowing for the variations in the excavation areas and in the volume of ar- establishment of a chronological sequence of events and chaeological deposits between the sites, as well differences in providing insight into the activities of the people who used the number of bones and taphonomic processes that may the site. variably affect distinct taxonomic groups (Lyman, 2008b; Reitz & Wing, 2008), the NISP was standardised for the volume of sediment (NISP/m³). This index provides an in- Sampling Strategy and Site Chronologies dependent density measurement for each taxonomic class. Analysed faunal remains from MG1 were recovered from 77 Fish remains were separated between postcranial (poste- grids, corresponding to 15.4 m3 of sediment, which was rior to the first precaudal vertebrae) and cranial (neuro- sieved using a 3 mm mesh size (Silva et al., 2001). Analysed cranium and viscerocranium) bones for skeletal frequency faunal remains from PG-U21 instead derived from one grid analysis, which provides insight into processing techniques (I10) located in the centre of the excavation, accounting for (e.g. parts removed during fish processing for storage, trade, 1.66 m3 of sediment, which was sieved using a 2.5 mm mesh etc.) (Lyman, 1994; Zohar et al., 2001). Other taphonomical size (Alves & Oliveira, 2002). Although only vertebrate features such as fragmentation, heat exposure, and cut marks remains were analysed from both sites, a significant number were also recorded (Costamagno et al., 2019; Egeland, 2003; of marine mollusc shells (e.g. Anomalocardia flexuosa) were Fernandez-Jalvo & Andrews, 2016; Yravedra, 2013). Re- also recovered at PG-U21 (Alves, 2003; Alves & Oliveira, mains that were slightly broken (e.g. vertebra without spinous 2002) and are currently undergoing analysis. and transverse processes) were considered whole. In order to improve the chronological attributions of MG1 and PG-U21, bones of terrestrial mammals (n = 4) were Faunal Identification Using Collagen Peptide Mass selected for radiocarbon dating (AMS) at CEDAD, Uni- versità di Salento (Italy). Bones from MG1 (one deer and one Fingerprinting (ZooMS) cattle bone) were sampled from layer 1 (grids A12 According to historical records both cattle (Bos taurus) and [LTL22552] and C13 [LTL22553]). Bones from PG-U21 buffalo (Bubalus spp.), as well as peccary (Tayassuidae) and (one armadillo and one cattle bone) were sampled from domestic pig (Suidae), were exploited in the region. Chickens layer 2 (LTL22550) and layer 3 (LTL22551) in grid I10. (Gallus gallus) were also commonly exploited along with Conventional radiocarbon dates were calibrated using OxCal other native birds. The remains of these morphologically v. 4.4 (Ramsey, 2009), using the 100% atmospheric cali- similar species are complex to set apart using conventional bration curve for the southern hemisphere, SHCal20 (Hogg morphological analysis, requiring additional analytical ap- et al., 2020). Calibrated dates were rounded to 10 years. proaches to achieve accurate identifications. As such, the Fossile et al. 5 remains of 57 mammals and birds from MG1 (n = 30) and PG- AD) included species recovered from MG1 and PG-U21, both U21 (n = 27) were selected for analysis with Zooarchaeology located in Babitonga Bay. We also compared the Pre-colonial by Mass Spectrometry (ZooMS) to confirm the morphological and Historical faunal assemblages with hunted animals among taxonomic identification. ZooMS is a method of peptide mass three Caiçara communities on the southeastern coast of Brazil fingerprinting that takes advantage of interspecies differences (∼150 km from Babitonga Bay) between 1998 and 2000 in the amino acid sequences of collagen to make taxonomic (Hanazaki et al., 2009). This Contemporary assemblage (1998- identifications (Buckley et al., 2009; Welker et al., 2015). 2000 AD) derived from interviews with Caiçara groups who Bone samples, ranging from 10-30 mg, were sampled from descend from Indigenous, African and European populations, each of the 57 remains and put in 250 μl of 0.6 M hydrochloric and thus may serve as a modern analogue for assessing patterns acid (HCl) at 4°C to demineralize. The acid was then removed of animal exploitation in historical times (Begossi, 2006; and the samples were rinsed once in 0.1 M sodium hydroxide Begossi & Richerson, 1993; Hanazaki & Begossi, 2003). (NaOH) to remove humic contaminants, followed by three Present day Caiçara communities value native terrestrial washes with 200 μl of 50 mM ammonium bicarbonate buffer mammals such as deer (Mazama spp.), lowland paca (Cuni- (AmBic, NH4HCO3, pH 8). After the final rinse, 200 μl of culus paca), agouti (Dasyprocta azarae), armadillo (Dasypus AmBic was added and the samples were gelatinized for 1 hour novemcinctus), opossum (Didelphis aurita), capybara (Hy- at 65°C. 125 μl of each sample was transferred to a 96 well drochoerus hydrochaeris), tamandua (Tamandua tetra- plate and the samples were digested overnight at 37°C with the dactyla), and peccary (Pecari/Dicotyles tajacu) as dietary addition of 0.4 μg of trypsin. Samples were acidified to 0.1% sources (Begossi & Richerson, 1993; Hanazaki et al., 2009). trifluoroacetic acid (TFA) to stop the trypsin, then purified Certain species, including deer, capybara, peccary and also using C18 resin ZipTip pipette tips (Pierce™ Thermo Sci- ocelot (Leopardus pardalis), are also exploited for medicinal entific) via an Opentrons OT-2 pipetting robot using an in- purposes (Hanazaki et al., 2009). Data on native non-volant house Python script. The in-house Python script used the terrestrial mammals was obtained from surveys performed in Opentrons Gen 1 300 μL multichannel pipette, three 96 well Babitonga Bay between 1909 and 2016 (Carvalho-Junior, plates and a 12 reservoir plate in order to purify peptides using 2022; Cherem et al., 2004; Dornelles et al., 2017; Ima, the C18 Zip Tips. The dimensions of the C18 Zip Tips and 2009) and complemented by the aforementioned archaeo- accompanying tiprack were added to the robot as a custom logical data, resulting in a total of 53 species. labware definition to allow the robot to recognise and use the Mean body weights were determined for each species based C18 Zip Tips. 1 μl of the extracted peptides was spotted onto a on data from the literature (e.g. Myers et al., 2020), and cat- Bruker target plate and combined with 1 μl of matrix solution egorised into small (<1 kg), medium (1-15 kg), and large (α-cyano-hydroxycinnamic acid) then analysed in triplicate mammals (>15 kg) following Galetti et al. (2017) and Vynne along with calibration standards on a Bruker ultrafleXtreme et al. (2022). Population density (individuals/km2) and trophic MALDI TOF/TOF mass spectrometer at the University of groups were compiled from Robinson & Refford (1986) and, York. Spectra were averaged and analysed using mMass when required, complemented with information from other software (Strohalm et al., 2008) and species were determined literature (Duarte et al., 2012; Faria-Corrêa, 2004; Galante & based on published m/z markers (Buckley et al., 2009, 2014, Cassini, 1994; Myers et al., 2020; Reis et al., 2006; Tomas & 2017; Eda et al., 2020, 2023; Kirby et al., 2013; Welker et al., Desbiez, 2004). Density data was obtained for a total of 46 2015) and in some cases using known reference material species. The average density was calculated following Galetti (unpublished data, see SM4 Table 3). et al. (2017), using game species found in each analysed period. Species were divided into trophic categories of Carnivore, Comparing Native Terrestrial Mammal Functional Myrmecophagy, Insectivore-omnivore, Frugivore-omnivore, Frugivore-granivore, Frugivore-herbivore, Herbivore- Traits Across Time Periods browser, Folivore-herbivore, and Herbivore-grazer (Reis Differences in mean body weight classes (body mass), pop- et al., 2006; Robinson & Redford, 1986). The statistical ulation density, and trophic groups of targeted non-volant significance in the proportions of analysed traits across time terrestrial mammal species were compared across three time periods was tested using a two-tailed Binomial test (p < 0.05) periods: Pre-colonial (4500–1150 cal BP), Historical (1750- in RStudio Software (RStudio Team, 2020). 1950 AD) and Contemporary (1998-2000 AD). These dif- ferences were also compared with the current composition of native terrestrial mammals in the region, against which we Results assessed the degree of selection in past hunting activities. Pre- Site Chronologies colonial assemblages (4500–1150 cal BP) were compiled from archaeological sites located in Babitonga Bay: Enseada I, Terrestrial mammal samples from MG1 and PG-U21 pro- Bupeva II, Forte Marechal Luz, Itacoara, Ilha dos Espinheiros vided calendar ages ranging from 1510-1800 to 1690-1950 II, and Cubatão I (Bandeira, 1992, 2004; Benz, 2000; Bryan, AD (68.4% confidence interval), with medians ranging from 1993; Fossile et al., 2019). Historical assemblages (1750-1950 1650 to 1850 AD (Table 1, Figure 3A). Overall, the calibrated 6 Tropical Conservation Science Table 1. Radiocarbon chronology and contextual information of archaeological bone samples. 14 14 Site CEDAD Lab code Material Context C yr. BP C yr. cal AD (2σ) Median cal AD PG-U21 LTL22550 Bovidae Grid I10-Q1, Layer 2 281 ± 45 1510 – 1800 1660 PG-U21 LTL22551 Dasypodidae Grid I10-Q3, Layer 3 139 ± 40 1690 - 1950 1850 MG1 LTL22552 Cervidae Grid A12, Layer 1 282 ± 40 1510 - 1800 1650 MG1 LTL22553 Bovidae Grid C13, Layer 1 175 ± 40 1670 - beyond calibration range 1810 Species Composition, Distribution and Relative Abundance A total of 11262 bone remains (NSP) were analysed from MG1 and PG-U21, of which 10960 (97.32%) could be identified using distinct taxonomic levels (NISP) and 302 (2.68%) could not be identified (NUSP) (SM1), due to pre- and/or post-depositional alterations (SM2). 57.90% of the remains analysed from MG1 were fragmented, and 52.43% of remains were fragmented from PG-U21. Cutmarks were identified on 76.92% of native terrestrial mammal bones (excluding osteoderms and teeth), ranging from 4.92% in Didelphis spp. (opossum) and Cuniculus paca (lowland paca) to 12.30% in Dasypodidae (armadillo). Cutmarks were also found on 80% of domesticated animal bones (excluding teeth) from MG1 and PG-U21, including Bo- vidae (36.81%) and Ovicaprid (1.39%). While Suina (pig and/or peccary) presented cutmarks in 90.91% of the re- mains. In both cases, the cutmarks were mostly located on limb bones (e.g., tibiae, metapodial and phalange), which can be related to carcass skinning, defleshing and disar- Figure 3. A) Radiocarbon chronology and B) associated material ticulation (Costamagno et al., 2019; Egeland, 2003). Cut- culture (fragment of J&G Meakin Ltd. pottery from PG-U21 with marks confirm that native terrestrial mammal remains from the distinctive impression dated to the 1910’s). both sites resulted from anthropogenic activities (e.g. hunting) rather than being a natural death assemblage. radiocarbon dates were older than the associated material Bony fish (NISP = 9002 remains) from MG1 and PG-U21 culture from both sites (coins, tin cans and ceramic artefacts), together accounted for 82.13% of the faunal remains, in- which have been assigned to the 19th and 20th centuries cluding Genidens barbus (white sea catfish), Conodon no- (Alves & Oliveira, 2002; Silva et al., 2001). For example, bilis (barred grunt), Centropomus spp. (snook), Cynoscion fragments of English pottery produced by J&G Meakin Ltd. spp. (weakfish), Cynoscion leiarchus (smooth weakfish), with a distinguishing “SOL” stamp, were found in layers 1 Larimus breviceps (shorthead drum), Micropogonias furnieri and 2 of PG-U21 (Figure 3B). This particular make of pottery (whitemouth croaker) and Pogonias courbina (black drum). appears to have been produced from the 1910’s, thus offering Fish remains were mostly represented by postcranial bones in a terminus post quem for the uppermost part of the site. both MG1 (n = 180, 84.11% fish remains) and PG-U21 (n = Similarly, diagnostic pottery remains and coins found at MG1 3552, 71.35% fish remains) in relation to cranial bones indicate that some occupations took place from 1821 AD. (MG1, n = 34, 15.89%; PG-U21, n = 1426, 28.65%). The Although it is possible that the radiocarbon dates detected cranial/postcranial (C/P) ratio from aggregated archaeolog- earlier occupations (i.e. 1700 AD), their large probabilistic ical layers differed between MG1 (0.19) and PG-U21 (0.40) distributions combined with the plateau-reversals in the 14C (SM3; Figure 4). calibration curve around 1700-1800 AD and 1820-1920 AD Fish remains were followed by mammals (16.45%, (Manning et al., 2018; Taylor et al., 1996), complicate the NISP = 1803), with medium-bodied mammals represented by radiocarbon interpretation and impinge upon our ability to Didelphis spp. (opossum), Dasypodidae (armadillo), Cebidae precisely constrain the site chronologies (Figure 3A). The (capuchin monkey) and Cuniculus paca (lowland paca), combination of radiocarbon and relative material-based while large mammals were represented by Suina (Tayas- chronologies, therefore, led us to assign a parsimonious suidae (peccary) and/or Suidae (domestic pig)), Cervidae age for both sites of 1750-1950 AD. (deer) and Bovidae (cattle). Birds (1.41%; NISP = 154) were Fossile et al. 7 Figure 4. Aggregated relative frequency of cranial and postcranial fish bones from (A) MG1 and (B) PG-U21. Figure generated on data publicly available from ArchéoZoo.org (https://www.archeozoo.org/archeozootheque/). represented by Aves (cf. Gallus gallus [chicken]). The sole markers at m/z 1991 and 2959, and Sus spp. with markers at species of cartilaginous fish (0.01%; NISP = 1) recorded was m/z 1961 and 2987 (SM4 Figures 1, 2 and 3). While these Carcharhinus spp. (shark) (SM1). markers need to be confirmed with further analyses, including LC-MS/MS sequencing to identify the specific peptides, they were identified in the six Suina samples analysed herein (SM4 MG1 Figures 2 and 3), and also in several additional peccary (m/z 1991, 2959) and pig (both domestic and wild boar, m/z 1961, When assemblages were analysed individually MG1 had a 2987) samples from other sites unrelated to this study. Ad- total of 1136 identified remains (74/m3), of which 53.26% ditionally, bulk collagen isotope analysis performed on these (NISP = 605) corresponded to indeterminate mammals Suina samples as part of an ongoing study (data not presented (Mammalia), followed by indeterminate bony fish (Actino- herein) provides further support that these additional peaks pterygii; 16.11%; NISP = 183), then cattle (6.78%; NISP = can be used to distinguish between Sus and Tayassuidae, as 77), pig (peccary and domestic pig) (6.25%; NISP = 71), deer the two identified as domestic pig (one from MG1 and one (2.73%; NISP = 31), rodents (2.73%; NISP = 31), bird (cf. from PG-U21) had nitrogen stable isotope (δ15N) values of Gallus gallus; 2.02%; NISP = 23) and others (<2%) (SM1). 11.03‰ and 8.09‰, respectively, while the samples iden- The highest species richness in MG1 was represented by tified as peccary (all from MG1) had significantly lower mammals (SR = 10), including both native (deer, lowland values, ranging from 1.32‰ to 1.70‰, suggestive of different paca, armadillo, opossum, dolphin and cf. whale) and do- trophic ecologies. mestic (cattle and sheep) species. The remains of marine mammals (Delphinidae and cf. Mysticeti) appear mostly as artefacts, due to indications that they were intentionally polished and/or modified, such as the presence of nails in two PG-U21 Delphinidae remains. ZooMS analysis confirmed the overall A total of 9825 remains (5918/m3) were recovered from PG- morphological identifications at MG1 (86.7% congruence), U21, of which 74.19% were indeterminate bony fish (Acti- with the exception of three Suina which ZooMS identified as nopterygii, NISP = 7288), followed by armadillo (8.48%, two cattle and one brocket deer, and one cattle that ZooMS NISP = 833 including osteoderms), barred grunt (7.99%, identified as brocket deer (SM4 Table 1). Along with con- NISP = 785 including otoliths), whitemouth croaker (3.31%, firming the exploitation of cattle (Bos taurus, n = 11), brocket NISP = 325 including otoliths), and others (<2%) (SM1). deer (Mazama spp., n = 12), and probably chickens (Gallus Several species including cattle (n = 10), chicken (n = 9), gallus, n = 2, although the spectra were of poor quality so a armadillo (n = 9), and domestic pig (n = 1) were all identified confident identification was not possible), ZooMS also al- with ZooMS (SM4 Tables 1 and 2). This represents 96.3% lowed us to verify that both peccary (Tayassu pecari and/or consistency with morphological identifications, with the Pecari/Dicotyles tajacu, n = 4) and domestic pig (Sus do- exception of one sample morphologically identified as ar- mesticus, n = 1) were exploited at the site (SM4 Table 1). madillo but that could not be identified amongst the species Suines can be difficult to distinguish between as different with currently available ZooMS collagen peptide markers. species of Sus, for example, have highly similar collagen The composition, relative abundance and density of faunal sequences meaning it is often impossible to determine if wild remains of PG-U21 varied considerably through the stra- or domestic pigs were being exploited. Using several known tigraphy. Fish remains were recovered from all layers, but a peccary (Tayassu pecari, Pecari/Dicotyles tajacu) reference noticeable increase in the density of remains (NISP/m3) was bones, differences were identified in the MALDI-TOF-MS documented in layers 1 and 2, possibly reflecting fishing spectra compared to Sus spp., with peccary having peptide intensification (Figure 5). These were mostly represented by 8 Tropical Conservation Science Figure 5. Stratigraphic variation in the relative abundance (%NISP), density (NISP/m³) and number of identified remains (NISP) for fish, native terrestrial mammals, and livestock from PG-U21, excluding the generic attributions to Actinopterygii and Mammalia, and remains of Aves. Based on the ZooMS identifications, Suina were grouped with livestock. barred grunt and whitemouth croaker, followed by shorthead classes ranging from 0.6 kg to 200 kg (SM5; Figure 6A). drum and others. Remains of native terrestrial mammals were Among these, 20 species were documented for the Pre- also found in all layers, represented mostly by armadillo and a colonial period (4500–1150 cal BP, SM6), six for the His- few remains of capuchin monkeys. Interestingly, their den- torical period (1750-1950 cal AD), and 10 for the Contem- sities also increased in layer 2. By contrast, livestock (cattle, porary period (1998-2000 AD). In the Pre-colonial period, the sheep, domestic pig) were only documented in layers 3 and 2, species with the largest and smallest mean body weights were with the highest density in layer 3. Finally, bird (mostly Tapirus terrestris (Brazilian tapir, 200 kg) and Cavia aperea believed to be chicken) remains were found in all layers, with (Brazilian guinea pig, 0.6 kg), respectively. In the Historical higher densities in layers 3, 4 and 5. period, MG1 exhibited a broader range of body weight classes (1.3 kg to 30.5 kg), while PG-U21 had a more re- Trait Variation in Native Terrestrial Mammals Across stricted range (2.8 kg to 5.7 kg). Tayassuidae (peccary, 30.5 kg) had the largest mean body weight, while Didelphis Time Periods spp. (opossum, 1.3 kg) was found to have the lowest. In the A total of 21 native terrestrial mammal species were identified Contemporary period, the largest mean body weight was across the three studied time periods, with body weight represented by Hydrochoerus hydrochaeris (capybara, Fossile et al. 9 Figure 6. Mammalian traits and functional groups across studied time periods. The proportion of mammal species differentiated by (A) body weight classes, (B) population density (individual/km2) and (C) trophic group between Pre-colonial (4500–1150 cal BP), Historical (1750- 1950 cal AD) and Contemporary (1998-2000 AD) periods, along with the current native terrestrial mammal composition of Babitonga Bay (BB). 50.5 kg), while the species with the lowest mean body weight exhibited densities ranging from 1.1 to 30 individuals/km2. was Didelphis aurita (big-eared opossum, 1.3 kg). The These species represented 68% of the terrestrial mammals in current composition of terrestrial mammals from Babitonga the Pre-Colonial period, 83% in the Historical period, and Bay includes species with a broader range of body weight 80% in the Contemporary period. By contrast, these species classes, ranging from the Brazilian tapir (200 kg) to Mon- represent only 39% of Babitonga Bay’s current terrestrial odelphis iheringi (Ihering’s short-tailed opossum, 0.1 kg). mammal composition, which shows density distributions Medium-bodied mammals (1.01-15 kg), accounting for ranging from 0.1 to 400 individuals/km2 (SM5; Figure 6B). 41% of the current species in Babitonga Bay, were the most Although no significant differences were observed across the exploited animals during all studied periods, ranging from studied periods, the results suggest that Pre-colonial, His- 60% in the Pre-colonial and Contemporary periods, to 67% in torical and Contemporary hunting practices selectively tar- the Historical period. These were followed by large-bodied geted medium- and large-bodied animals with low to medium mammals (> 15.1 kg) ranging from 30% in the Contempo- population densities. rary, 33% in the Historical, to 35% in the Pre-colonial period; Regarding trophic groups, the categories ranged from while evidence for targeting small-bodied mammals (< Herbivore-grazers to Carnivores, with the majority of targeted 1.0 kg) was only detected in the Pre-colonial period, repre- mammals belonging to the Frugivore-omnivore category sented by Brazilian guinea pig. When considering the current (SM5; Figure 6C). The highest diversity of trophic categories species composition of Babitonga Bay in comparison to the was detected in the Pre-colonial period, with a preference for three time periods and body weight classes studied, signifi- Frugivore-omnivores (30%), followed by Carnivores (15%), cant differences were only observed for the proportion of Frugivore-granivores (15%), Frugivore-herbivores (15%), and small-bodied mammals between the Pre-colonial period and others at less than 5%. In the Historical period, the exploitation the current composition (p = 0.01367). was mainly focused on Frugivore-omnivores (n = 49%), Targeted mammals had population densities ranging from followed then by Insectivore-omnivores, Frugivore-granivores 0.1 to 65.5 individuals/km2, with the highest density repre- and Frugivore-herbivores (all with n = 17%). In the Con- sented by the Brazilian guinea pig (65.5 individuals/km2) and temporary period, the most exploited category was again the lowest by Puma concolor (cougar) (0.1 individuals/km2). Frugivore-omnivores, Frugivore-granivores and Frugivore- In contrast, all native terrestrial mammals had estimated herbivores (all with n = 20%), followed by Carnivores, density values ranging from 0.1 to 400 individuals/km2. Myrmecophages, Insectivore-omnivores and Herbivore- However, the majority of targeted mammals across all periods browsers (all with n = 10%). The current terrestrial 10 Tropical Conservation Science mammal composition of Babitonga Bay exhibits the same Catarina state (Herbst et al., 2023; Sandoval Gallardo et al., trophic groups as those in the Pre-colonial and Historical 2021); as such, this fishing intensification at PG-U21 may periods; however, the most abundant trophic group is reflect a local response to increased market demand. The Frugivore-granivores (28%), followed by Carnivores (17%), relatively low taxonomic diversity of fish remains suggests Insectivore-omnivores (17%), Frugivore-omnivores (15%), specialised fishing practices aimed at supplying both Frugivore-herbivores (7%), Myrmecophages (4%), and household and local markets, which is largely supported by others. Despite the preference for Frugivore-omnivore spe- historical accounts documenting the importance of fish as a cies in the Pre-colonial, Historical and Contemporary periods, source of food and income in Babitonga Bay since the mid- there were no statistical differences between periods and 19th century (Saint-Hilaire, 1936). The most frequently current compositions. caught species (whitemouth croaker and barred grunt) con- tinue to display high economic and subsistence values today (Haimovici et al., 2016; Lima et al., 2019), and the same fish Discussion species are also consumed by contemporary Caiçara com- Radiocarbon dates and artefacts indicate that MG1 and PG- munities along the Atlantic Forest coast (Begossi, 2006; U21 were occupied from approximately 1750 to 1950 AD, Begossi & Richerson, 1993; Hanazaki & Begossi, 2003). one hundred years after the establishment of the village of Extensive research has demonstrated that these communities Nossa Senhora da Graça on the São Francisco River in 1658 carefully consider the costs and benefits associated with (Cabral, 1937; Saint-Hilaire, 1936). In 1660, the village was pursuing and processing food resources based on factors such elevated to the status of town, and in 1665 it was designated as site location, environmental conditions, and the influence as a parish, and then as a city in 1847, the earliest city in the of urban centres and markets (Hanazaki & Begossi, 2000; state of Santa Catarina (Cabral, 1937). Faunal remains from MacCord & Begossi, 2006). According to this theoretical MG1 and PG-U21 offer valuable insights into the livelihoods proposition, there is a stronger focus on fishing as the and the ecological footprints of these early colonial and post- proximity to the coast increases, which explains the higher colonial decades in Babitonga Bay. The information gained dependence on fishing observed in PG-U21 (coastal) com- also allows us to reassess the often neglected role of fisheries pared to MG1 (inland). Overall the results highlight the during the early modernization of Brazil’s southern coast. significant role of fishing in Babitonga Bay between the 18th The zooarchaeological data revealed noticeable differ- and the 20th century, which adds to the growing evidence for ences in faunal remains between MG1 and PG-U21, both in a heavy reliance on fishing for food security for thousands of terms of species composition, relative abundance and density years in the region (Fossile et al., 2019, 2023; Toso et al., (Figure 7A-B). We acknowledge that differences in recovery 2021). This is particularly significant given that fishers were and analytical methods can affect fundamental and derived often overlooked or inadequately represented in regional measurements of archaeological faunal remains (Grayson, historical narratives (Silva, 1988, 2001). 2014; Lyman, 2008a; Reitz & Wing, 1999); however, these The species found at PG-U21 and MG1 suggest that are unlikely to explain the variations seen among the sites. fishing was mainly practised in coastal waters. Surprisingly, Firstly, in both sites, faunal remains were retrieved using other coastal species widely captured nowadays, such as similar mesh sizes (2.5 and 3 mm), which are considered mullets (Hanazaki & Begossi, 2000, 2003; Herbst & adequate for the recovery of small fragments and anatomical Hanazaki, 2014; Sandoval Gallardo et al., 2021), have not parts, such as fish remains (McKechnie & Moss, 2016; Zohar been recorded at either site. Mugil liza (Lebranche mullet), in & Belmaker, 2005). Secondly, taxonomic identification for particular, is abundantly caught along the southeastern coast both sites was performed using the same reference collections of Brazil during austral autumn and winter, from May to July and by the same analyst, therefore reducing analytical biases. (Steenbock, 2019), and this practice has been regionally Moreover, the proportion of fragmented bones was similar documented since the early 16th century (Staden, 2020). It is among sites. The recovered faunal remains thus confiden- possible that the species was not consumed by local residents, tially reflect fundamental differences in subsistence and as observed among some contemporary communities in economic activities between the sites. southern Brazil (Hanazaki & Begossi, 2003). Alternatively, Fish outnumbered other faunal remains at PG-U21, with processing methods involving salting and drying (locally an overall aggregate density (899/m³) comparable to, or known as cambira, (Alves, 2003; Anacleto et al., 2019)) may higher than, some local Pre-colonial shell mounds (known as have prevented the survival of diagnostic bone remains in the sambaquis, with densities from 42/m3 to 7871/m3) produced archaeological record. Cambira involves a longitudinal in- by groups subsisting largely on fishing (Fossile et al., 2019, cision along the dorsal region of the individual, and the re- 2023) (Figure 7B). Fishing intensification, however, appears moval of the head, prior to salting and drying. This method is to have occurred during the later phases of the site dated to the similar to the one observed by Zohar and Cooke (Zohar & early 20th century. This coincides with a time interval of Cooke, 1997) in the northwest of Panama Bay, who dem- substantial policy incentives for commercial fisheries, with onstrated that it causes damage to, and loss of, cranial bones, increased catches and fishing efforts along the coast of Santa precisely those bones which contain most of the diagnostic Fossile et al. 11 Figure 7. Aggregated density (NISP/m³) for fish, native terrestrial mammals, livestock and birds from (A) MG1 and (B) PG-U21, excluding the generic attributions to Actinopterygii and Mammalia. For MG1, remains of Artiodactyla and Canidae were also excluded from the analysis. Based on the ZooMS identifications, Suina were grouped with native terrestrial mammals for MG1 and with livestock for PG-U21. Figure generated on data publicly available from PhyloPic (http://phylopic.org/). elements needed for taxonomic identification. Nevertheless, Long-Term Selective Hunting in Babitonga Bay the differences in cranial/postcranial values between the two sites and the absence of otoliths in MG1 suggest that a large Native terrestrial mammals made up a significant proportion number of fish may have been locally processed in PG-U21, of faunal remains at both sites. This suggests that species while in MG1 more fish were brought to the site already currently undergoing population declines were exploited in processed (with crania removed). As such, processing the region for at least the past two centuries. When comparing methods are unlikely to explain the absence of mullets at least this data to information gathered from Pre-colonial sites, our in PG-U21, unless the species was processed and cranial findings show a consistent pattern of medium- and large- bones disposed of elsewhere, with only the postcranial bodied terrestrial animal species being hunted by both Pre- fraction transported to the site for consumption or trade. The colonial and historical coastal communities in Babitonga Bay absence of mullets, therefore, remains a matter deserving of (Brazilian tapir, and various species of deer and peccary). further study. This hunting trend persists despite changes in the purpose of Our results indicate that livestock did not play a major role hunting over different study periods, such as subsistence, as a food source (e.g. meat), nor as sources of secondary ritualistic, and livelihood pursuits. Such a pattern is also products (dairy, fur/hide) during the colonial and post- observed among some contemporary Caiçara communities, colonial periods in Babitonga Bay (Figure 7A-B). Rather, and has been extensively documented in tropical forests in local food security and livelihood relied on fishing and the South America and beyond (Ceballos et al., 2017; Darimont hunting of native terrestrial mammals, along with crop ag- et al., 2023; de Souza & Alves, 2014; Dirzo et al., 2014; riculture. Fish were exploited for their meat, fat and oil, while Milner-Gulland & Bennett, 2003; Peres, 2000). native terrestrial mammals could have also been pursued for Significantly, several species identified in the Pre-colonial their secondary products (fur/hide), as suggested by the and Historical periods, such as Pampas deer (Ozotoceros number and location of cutmarks. This is supported by bezoarticus) (Bryan, 1993), Brazilian tapir, white-lipped historical records which report that fish and native terrestrial peccary, and red brocket deer are no longer documented in mammals were consumed at the household level, and used as the region (Carvalho-Junior, 2022; Dornelles et al., 2017). sources of income (Ribeiro & Corção, 2013; Saint-Hilaire, The last recorded sightings of tapirs and peccaries in Babi- 1936). Among domesticated animals, chickens are an ex- tonga Bay date back to 1996, while the last record of a red ception as they very likely played a role as a daily source of brocket deer was in 2001 (Cherem et al., 2004). By contrast, dietary protein (meat and eggs) at both sites, as documented these species have been reported between 2015 and 2019 in a among traditional riverine and coastal communities nowa- Protected Area located approximately 45 km away from days (Hanazaki & Begossi, 2003). Fish, native mammals and Babitonga Bay (Hübel et al., 2021), reinforcing the role of chickens appear to have contributed substantially to the conservation strategies in reducing defaunation (Bogoni overall economy of PG-U21. By contrast, the low density of et al., 2020). Despite a long history of hunting, Babitonga faunal remains in MG1 suggests that other economic activ- Bay shows a notable diversity of medium and large terrestrial ities, such as plant cultivation and processing (e.g. manioc mammal species (n = 30, excluding volant mammals), sur- milling), prevailed at the site. passing the average observed in the Atlantic Forest from 1983 12 Tropical Conservation Science to 2017 (n = 14.7 species) (Bogoni et al., 2018). This suggests origins, which if not recognized can lead to the establishment that the remaining forested areas may still be capable of of inappropriate sustainability targets, weak public and sustaining levels of productivity that offset the detrimental stakeholder support for conservation initiatives, and gener- effects of hunting and habitat degradation (Peres, 2000; Peres ational shifts in the acceptance of systems that are considered & Nascimento, 2006). degraded (Lovell et al., 2020; McClenachan et al., 2018; When considering the average density of mammals, our Pauly, 1995; Soga & Gaston, 2018). In order to obtain a more results suggest hunting intensity was lower in the Historical comprehensive understanding of local animal density and its period (22.1 ind./km2), compared to both the Contemporary implications, further studies are needed, including studies on (18.5 ind./km2) and Pre-colonial periods (18 ind./km2). These past faunal assemblages. Archaeological faunal remains are subtle differences could be attributed to the relatively smaller some of the few available sources of information on Pre- human population residing in Babitonga Bay from the 18th to colonial and historical vertebrate diversity, and as such can the 20th centuries, or changes in the socio-economic nature of shed light on the origin and changing nature of defaunation hunting practices through time. For example, the absence of over long timescales. We recommend the integration of carnivores during the Historical period may be explained by historical and archaeological data into modern faunal pop- evolving social factors. Leónce Aubé (Aubé, 1857) docu- ulation assessments and conservation initiatives to set more mented that native animals in Babitonga Bay were classified informed reference baselines. into two groups by local communities in the 19th century: game species, consisting of animals hunted for their nutri- Implications for Conservation tional value; and “ferrous” animals (e.g. carnivores such as Panthera onca, jaguar), which were not hunted. Present-day Influenced by a lack of historical perspective, the widespread Caiçara communities pursue different species for their food perception of low human impact in tropical forests has led to value, as well as for their medicinal value (Hanazaki et al., limited conservation attention until recent decades. This 2009), and both are strongly tied to cultural traditions that can historical amnesia has hindered assessments of the current be traced back to the 16th century (Camphora, 2021). environmental challenges facing the Atlantic Forest and its Long-term selective hunting may have had significant coastal waters, including defaunation and overfishing. A detrimental social and ecological consequences in the At- growing body of evidence indicates that the diversity of lantic Forest, about which our understanding remains limited Neotropical mammals is currently under threat from various (Jorge et al., 2013). Studies have shown that the selective human-induced pressures, and archaeology is emerging as a removal of medium and large terrestrial species with low key discipline for expanding our understanding of these population densities, low breeding cycles, prolonged gesta- anthropogenic pressures over long time scales. Our analysis tion periods, and extended intervals between births, not only of faunal remains from two historical sites highlights the causes a decline in the overall animal biomass, but also has significant role of fishing and hunting of native terrestrial far-reaching implications for forest ecosystem services wildlife during the last two centuries of human colonisation (Galetti et al., 2015, 2017, 2021; Peres, 2000; Peres & of Babitonga Bay. This study provides compelling evidence Nascimento, 2006). Hunting pressure on medium and large that the selective hunting of medium- and large-bodied native frugivorous mammals, considered “habitat shapers” (Rumeu terrestrial mammals has persisted in the region for over 4500 et al., 2020) for their capacity to disperse large seeds, may years, and requires us to reconsider the idea of a heavy re- disrupt the recruitment and distribution of plant species with liance on domestic animals during European colonisation of trophic cascade effects on plant diversity, soil regulation and southern Brazil. structure, and forest carbon storage capacity, among others (Camargo-Sanabria et al., 2015; Galetti et al., 2015, 2017; Acknowledgements Jorge et al., 2013; Peres, 2000; Rumeu et al., 2020; Villar The authors thank Alejandro Sierra Sainz-Aja, Alba Ruiz Cros, et al., 2021). Moreover, native terrestrial animals play a Roger Alcàntara Fors and Maria Saña Segui for their support in the significant role in the livelihoods of tropical and subtropical identification of domestic faunal remains. The authors also thank the rural communities (Alves et al., 2009; Hanazaki et al., 2009; Centre of Excellence in Mass Spectrometry at the University of York Peres & Nascimento, 2006), and their loss risks increasing the for access to the Bruker ultrafleXtreme MALDI-TOF/TOF-MS. socio-ecological vulnerability of local traditional groups, and their perceptions of forest ecosystems (Milner-Gulland & Declaration of Conflicting Interest Bennett, 2003; Ponta et al., 2019). The commoditization of natural resources, coupled with The author(s) declared no potential conflicts of interest with respect significant progress in hunting, processing, and transportation to the research, authorship, and/or publication of this article. technologies in recent decades, have led to unprecedented levels of defaunation in the Neotropics (Bogoni et al., 2018, Funding 2020; Jorge et al., 2013; Peres, 2000). Our study indicates that The author(s) disclosed receipt of the following financial support for these ecological footprints actually have deeper historical the research, authorship, and/or publication of this article: This work Fossile et al. 13 was funded by the ERC Consolidator project TRADITION, which Bandeira, D. R. (2004). Ceramistas Pré-coloniais da Baı́a da Ba- has received funding from the European Research Council (ERC) bitonga - Arqueologia e Etnicidade [Doctor in History]. under the European Union’s Horizon 2020 research and innovation Universidade Estadual de Campinas. programme under Grant Agreement No 817911. This work con- Bandeira, D. R., & Alves, M. C. (2012). Arqueologia histórica no tributes to the “ICTA-UAB Marı́a de Maeztu’’ Programme for Units nordeste de Santa Catarina. Revista Tempos Acadêmicos - of Excellence of the Spanish Ministry of Science and Innovation Dossiê Arqueologia Histórica, 10, 68–87. (CEX2019-000940-M). This work also contributes to EarlyFoods Barnosky, A. D., Hadly, E. A., Gonzalez, P., Head, J., Polly, P. D., (Evolution and impact of early food production systems), which has Lawing, A. M., Eronen, J. T., Ackerly, D. D., Alex, K., Biber, received funding from the Agència de Gestió d’Ajuts Universitaris i E., Blois, J., Brashares, J., Ceballos, G., Davis, E., Dietl, G. P., de Recerca de Catalunya (SGR-Cat-2021, 00527). The funders had Dirzo, R., Doremus, H., Fortelius, M., Greene, H. W., & Zhang, no role in the study design, data collection and analysis, the decision Z. (2017). Merging paleobiology with conservation biology to to publish, or the preparation of the manuscript. guide the future of terrestrial ecosystems. Science, 355(6325), eaah4787. https://doi.org/10.1126/science.aah4787 ORCID iD Begossi, A. (2006). The ethnoecology of Caiçara metapopulations (Atlantic Forest, Brazil): ecological concepts and questions. Thiago Fossile  https://orcid.org/0000-0002-6997-4677 Journal of Ethnobiology and Ethnomedicine, 2, 40. https://doi. org/10.1186/1746-4269-2-40 Supplemental Material Begossi, A., & Richerson, P. J. (1993). Biodiversity, family income Supplemental material for this article is available online. and ecological niche: A study on the consumption of animal foods on Búzios Island (Brazil). 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