Pre-colonial Amerindian legacies in forest composition of southern Brazil

Aline Pereira Cruz; Eduardo Luiz Hettwer Giehl; Carolina Levis; Juliana Salles Machado; Lucas Bueno; Nivaldo Peroni

doi:10.1371/journal.pone.0235819

. 2020 Jul 23;15(7):e0235819. doi: 10.1371/journal.pone.0235819

Pre-colonial Amerindian legacies in forest composition of southern Brazil

Aline Pereira Cruz ^1,^*, Eduardo Luiz Hettwer Giehl ¹, Carolina Levis ¹, Juliana Salles Machado ², Lucas Bueno ², Nivaldo Peroni ¹

Editor: André Jasper³

PMCID: PMC7377383 PMID: 32701950

Abstract

Past human societies have left persistent marks on forests worldwide. However, the degree to which pre-colonial Amerindian societies have affected forest structure is still not fully understood, especially in southern Brazil. This study investigated the influence of two distinct Amerindian groups (Southern-Jê and Guarani) over tree composition of forest fragments in the State of Santa Catarina. Vegetation data was obtained from the Santa Catarina Forest and Floristic Inventory (SCFFI): a statewide systematic vegetation sampling project. Archaeological data was collated from literature reviews as well as existing databases for archaeological sites occupied by Guarani and Southern-Jê groups. Using these sites of known Amerindian occupation, and corresponding environmental variables, ecological niche models were developed for each Amerindian group, predicting potential archaeological sites occupied by these groups across southern Brazil. Maps of these potential occupation sites of pre-colonial Amerindian groups were compared with 417 corresponding floristic inventory plots. Redundancy analysis (RDA) was used to identify floristic composition patterns linked to areas with a high probability of Southern-Jê or Guarani presence. Southern-Jê and Guarani pre-colonial occupations overlapped near main rivers; however, Southern-Jê groups generally occupied elevated areas whereas Guarani occupied mostly coastal areas. We observed differences in forest composition associated with the predicted occurrence of these pre-colonial Amerindian groups. Based on these results, we argue there is a relationship between tree species distribution and pre-colonial human occupation by these two Amerindian groups.

Introduction

To understand species distribution, we need to consider many different aspects, both biotic and abiotic, and their interdependence, as well as the historical processes. Historical ecology research seeks to understand past human legacy on present day species composition, community assemblage, and identify positive interactions (eg. mutualism) and process associated with niche modification [1–3]. A place with the appropriate environmental characteristics for a species is how Grinnell [4] defined a niche. The development of the niche concept incorporated new factors: biotic interactions were considered [5], environment was expanded to multiple layers [6], and the effect of negative biotic interactions (e.g., competition) on species niche retraction were intensely evaluated [7–9]. Subsequent evolution of the niche concept recognized that positive interactions (e.g., facilitation) were capable of widening the niche [10]; introducing the idea that some species can modify their environment, and thereby can promote changes in resource availability for another species. These species are called ecosystem engineers [11]. The most effective ecosystem engineers are species that cause the longest-lasting modifications to their environment, and have the largest population size; for example, humans [11]. Once species modify their environment, they are able to change the direction and force of selection. This bi-directional relationship between species and their environment, may affect the species’ niche as well as the niches of other species [12]. The Niche Construction Theory (NCT) predicts that past bi-directional selective processes result in species composition signatures that are perpetuated over time, and can be recognized in contemporary communities, a pattern called ‘legacy’ [12]. Different societies placed in different contexts modify their environments in different ways [1]. The term Cultural Niche Construction (CNC) has been coined to emphasize that cultural factors drive decisions about the changes that humans promote in their environment and, consequently, to the environment available to other species [13, 14]. CNC theory provides an integrative scenario to understand human legacies on natural ecosystems worldwide [14].

Recent historical, archaeological, and ethnographical studies recognize the historical influence of human activities on what has previously been considered primary, untouched, or pristine forests. Many tropical forests previously thought of as pristine were revealed to have been shaped by past human societies [15, 16]. For example, recent studies have shown that Amazonian forests have been modified by indigenous populations for millennia, altering plant species distributions across the region [17, 18].

The Atlantic Forest is likely no exception: the region was also occupied by Amerindians long before and continued after European arrival [19]. Archaeological studies have reconstructed the long-term human history of the region, identifying many Holocene archaeological sites in southern Brazil (at least 1704 sites already mapped in the region). This indicates that the Atlantic Forest, now drastically reduced and fragmented [20], has had a long history of human interaction.

History of human occupation in the State of Santa Catarina (Fig 1) commenced in the highlands ~ 11,500 Before Present (BP) [21] and in the littoral zone ~ 8, 000 BP [22, 23]. Hunters and gatherers were the first human groups to occupy this territory [24]. Current indigenous peoples in the area belong to the Southern-Jê (Xokleng or Laklaño and Kaingang) and Guarani linguistic groups. Jê groups from Central Brazil has started their migration to Southern around 3, 000 years BP [25] [26]. Archaeological dating in southern Brazil indicates The Southern-Jê commenced occupation along highland rivers, and then moved into littoral areas [25]. Highland occupancy is a general distribution pattern of Macro-Jê linguistic groups in Brazil in the central and eastern plateaus [26]. In the southern Brazil highlands, they relied on ‘pinhão’ (Araucaria angustifolia seed) as a key food source [27, 28], associated with another cultivated resources, as Zea mays, Manihot esculenta, and Dioscorea sp., and widespread hunting and fishing. The Guarani migration is related with Tupi expansion from Amazon. The main route for this Guarani migration from Amazon basin to southern Brazil is related to Paraná and La Plata river basin The two principal Guarani suggested centers are the Amazon and La Plata Basins [29]. The Guarani arrived in Southern Brazil only around 1800 years BP [30]. They are recognized as sealers, fishermen and farmers who followed the main rivers and the coast expansion of their territories. Contrasting both migration dynamics, Guarani groups migrated by expanding their territory, whereas the Southern-Jê migrated by leaving old territory and moving into new ones [25, 26].

The Southern-Jê and Guarani migrated to the south of Brazil to a landscape where forests were expanding and at a time of climate transition. In the highlands, Araucaria moist forests were expanding over what were previously grasslands. Grasslands are remnants of a drier climate, whereas forests are favored by the current climatic conditions [31]. Around 3000 years ago, forest expansion over grasslands intensified [32, 33], coinciding with the arrival of the Southern-Jê people in southern Brazil [25, 26]. Together with climate changes, humans may have acted as a complementary driver of forest expansion due to transportation of seeds [27, 28, 34, 35], and maintained grasslands using fire [32, 33, 36]. Recent research has shown that human action was essential for these forests to reach their maximum distribution [34, 35]. There are many evidence of the long-term use of A. angustifolia by past Amerindian societies also suggests they promoted the population expansion of this species, which is the dominant species of Araucaria moist forests in southern Brazil [37]. Araucaria angustifolia is recognized as a nurse plant, that is, its dispersion would favor the expansion of other forest species [38–40]. The role of human action in the assembly of forest communities in the pre-Colombian period as well as legacies in recent communities are still issues to be understood in the historical ecology of southern Brazil.

Globally, studies showing the influence of past cultural groups on species distribution patterns have intensified (see [16]); however, the long-term history of the Atlantic Forest is still poorly understood. Historical ecology studies in this region are hampered by the intensification of human occupation in recent times, which is obscuring past human legacies. This study aims to address this knowledge gap for southern Brazil. The niche modeling approach was used to understand the distribution of past Southern-Jê and Guarani Amerindian groups and then correlate this distribution with modern forest composition. Based on the premises that, (a) Southern Brazil has been occupied by humans for millennia, and (b) humans are cultural niche constructors, our hypothesis is that floristic composition differs in Southern Atlantic Forests with a high probability of past human activities, and that different cultural groups also leave differing floristic composition and abundance legacies notable yet, despite the disruption of these cultural groups and of the intense changes caused by recent populations. We suggest that humans have been shaping these forest communities for at least 10,000 years; and that some differences in floristic composition are the results of the historical process of human occupancy and cultural variability.

Materials and methods

Study area

The study area (Fig 2) covers part of the pre-colonial Southern-Jê and Guarani distribution, in the Southern portion of the Neotropical Region. Located in southern Brazil, the State of Santa Catarina (SC) was selected because floristic data has been systematically collected across the state [41], providing the opportunity to compare spatial patterns with pre-colonial occupation. The entire area of SC is in the Atlantic Forest Domain [42], one of the global hotspots for biodiversity conservation [43]. Vegetation consists of coastal scrubland, mangroves, grasslands, and forests. Variation in forest types is influenced by geomorphology and climate. The entire State is in the subtropical zone and climatic variation is related to latitudinal and altitudinal gradients. The elevation gradient induces variation in air pressure, and consequently, promotes cooling. Landforms also drive atmospheric water movement and influence rainfall regimes [44]. Temperature variability increases with distance from the coast [44]. Santa Catarina has eighteen hydrographical basins, and combined with a moderate to high annual rainfall, has substantial river resources for local populations.

Vegetation data

We investigated whether there is a legacy of past human populations on present-day forest composition. Vegetation data used in this study was collected by the Santa Catarina Forest and Floristic Inventory (SCFFI), which systematically sampled shrubs and trees across the State in a grid system [41, 45] (Fig 1). Each sample unit covered 4000 m² [41, 45]. All forest plots except sand dunes and mangroves (a total of 417 sample units) were used to evaluate the abundance and distribution of all tree and shrub species in the study area.

Archaeological data

Aiming to identify past human effects on forest composition, the first step was to locate archaeological sites. Archaeological sites within the study area were identified from literature reviews, as well as field data collected since 2011, by the Laboratory of Interdisciplinary Archaeology Studies at the Federal University of Santa Catarina. The literature review considered data published by Bonomo et al [30], and Noelli and Souza [25], covering the south of Brazil, and only used sites that were classified as Southern-Jê or Guarani. Another Southern-Jê archaeological sites were identified in LEIA database based on earthworks description. Using this data, we constructed a matrix with geographical coordinates of Southern-Jê and Guarani archaeological sites. Geographical location was the only information available for spatial analysis that could be obtained from all archaeological sites (although not always mentioned in archaeological studies). Only archaeological data with cultural attribution was used in order to identify any association between species and cultural variability. The selected archaeological sites have been occupied over the past 3,000 years.

Topographic, hydrographic and forest type data

In order to understand which landscape features most influenced past Amerindian distribution, and contributed to defining their niche, we selected variables related to human preferences for specific environments; such as, topography, hydrology [46], and river proximity [23, 47]. The Height Above the Nearest Drainage (HAND) was used as a proxy for water table depth, and altitude and slope were used to define terrain (topography) [48]. Water course vectors were separated using Otto Pfafstetter’s watershed coding method [49]. Coastline distance was used, assuming that the sea is a source of food and other services used by the populations [30, 47, 50]. Correlations between geomorphologic and hydrographic variables were evaluated. Forest types were mapped according to the classification of global terrestrial biomes [51] as follows: (1) Rain Forest (locally called "Dense Forest") on the coastal mountain range; (2) Mixed Forest (locally called "Araucaria Forest") on the highlands; and (3) Seasonal Forest found inland, especially in the Uruguay River basin [52].

Data analysis

River classes area were grouped in case of collinearity (other variables did not present autocorrelation). The distance from each river class was rasterized using the Euclidean Distance function in Qis. We also generated a Euclidean raster distance from the coastline.

Occurrence area of each Amerindian group was defined based on the geographical distribution of Southern-Jê and Guarani archaeological sites. Occurrence area was used to limit the spatial area modeled for each cultural group. To understand the past Amerindian distribution, we developed models based on archaeological, topographic, and hydrographic data in Maxent interface (Ecological Niche Model–ENM; using the package ‘ENMeval’ [53]) to fit models and predictions of archaeological sites beyond sampled locations in R [53, 54]. Since we only had presence-only data on the location of archeological sites and because data wasn’t obtained by systematically sampling, we decided to use Maxent, Maximum Entropy Method, suited for analyzing presence-only data [55]. To further minimize sampling bias, we worked only with one archaeological site on every 10 × 10 km pixel. We used random k-fold validation with 4 k-folds, combining Features Class (FC): Linear (L), Linear and Quadratic (LQ), Hinge (H), and Linear Quadratic Hinge (LQH); and Regularization Multiplier (RM) sequence values 0.5, 4, 0.5. Models were sorted in decreasing order by the largest AUC values (Area under receiver operating Curve) and lowest overfitting, estimated by contrasting AUC values of train and test sets, and then, between top ranking models. Only one model was finally chosen by visual inspection. The chosen model was then used to generate predictive maps of the potential distribution of past Amerindian occupation. Estimates of the variables contribution to each model was expressed in terms of percent contribution and permutation importance values. Two separate maps for Southern-Jê and Guarani sites were constructed. Using these maps, the overlap of Southern-Jê and Guarani past distributions was calculated with the similarity statistic ‘I’ [56], where results fall between 0 and 1 (0 indicating no overlap and 1 indicating full overlap).

Vegetation sampling locations were overlaid with the maps of the potential distribution of both cultural groups. Next, we extracted the probability of each vegetation sample location falling over Southern-Jê or Guarani sites, or both. To assess the relationships between past Amerindian groups distributions and floristic composition of current forest fragments, we carried out a Redundancy Analysis (RDA), using the package ‘vegan’ [57] in R. The matrix of species abundances was used as the response variable and the probabilities of Southern-Jê and Guarani sites for each site were used as predictors. Forest type was added to ordination diagrams to aid interpretation but was not used as a predictor in the analysis. QGIS software was used for all geoprocessing procedures and R for multivariate analyses.

Results

Guarani and Southern-Jê distributions were influenced by the distance to first or second order rivers (using Otto Pfafstetter’s hierarchy, Fig 3). The importance of this variable was 42.2% and 40% (contribution and permutation importance values, respectively) in the Guarani model and 22% and 31.9% in the Southern-Jê model. Guarani distribution was also influenced by coastline distance (contribution: 36.9%; importance: 51.8%), which, conversely, was the environmental variable with the lowest influence over Southern-Jê distribution (contribution: 0.2%; permutation importance: 0.6%). Elevation was the environmental variable with the highest influence over Southern-Jê distribution (contribution: 39.6%; permutation importance: 27.1%), being irrelevant to Guarani distribution (contribution: 1.2%; permutation importance: 5.2%). The most suitable environments for Guarani people were those with proximity to the sea or rivers, while Southern-Jê sites were mostly found in elevated areas and near rivers. The final Guarani model (Fig 4A) was generated with FC = H and RM = 2, showing an AUC = 0.83. The final Southern-Jê model (Fig 4B) was generated with FC = LQP and RM = 4, and had an AUC = 0.71. Final images are available in S1 and S2 Files.

Fig 4 — a) Map of the probability of past Guarani distribution; b) Map of the probability of past Southern-Jê distribution; c) Map without scale of South America highlighted areas showed in a and b. These maps were created with ecological niche models that presented the lowest overfitting and the highest AUC values.

Niche overlap between past distributions of Southern-Jê and Guarani was noticeable (I = 0.64; Fig 5). Overlapping areas are located along the major rivers (1^st and 2^nd order) and along main rivers within intermediate-sized basins (3^rd order, Fig 4).

The first two axes of the Redundancy Analysis (RDA) explained 7% of the data variation (Fig 6) and show (i) directional segregation between the two cultural groups and (ii) the links between the distribution of these Amerindian groups and forest typologies. Ordination of sampling units (Fig 6A) suggest Southern-Jê sites relate with Mixed Forests and Guarani sites are more related with Rain Forests. Seasonal forests were likely used by both groups. The scores of the relation between the first axis of the RDA and the past Amerindian groups distribution was -0.57 for Guarani and 0.85 for Southern-Jê: the second axis had positive scores for both groups (Southern-Jê = 0.52 and Guarani = 0.81). RDA ordination analysis highlighted three clear groups (Fig 6): (1) species shared between the Southern-Jê and Guarani groups that are in the positive range of the second axis, (2) species related to the occurrence of the Southern-Jê group in the positive range of the first axis, and (3) species related to the Guarani presence in the negative range of both axes. Some species related to both axes, but were more strongly related to the Southern-Jê (left side of the first RDA axis) or Guarani (right side of the first RDA axis). Twenty-nine species had ordination scores higher than 0.1 or lower than –0.1 and were considered as associated with Amerindian past distribution. These 29 species were listed along a cultural gradient (Fig 7). The listed species belong to 20 families, Lauraceae (n = 6), Sapindaceae (n = 3), Anacardiaceae and Arecaceae (n = 2), with the other families represented by only one species each.

Fig 7 — These species were selected because they had ordination scores higher than 0.1 or lower than –0.1 in the RDA.

Discussion

This study investigated the cultural legacy of pre-colonial Guarani and Southern-Jê in the present floristic composition of forest remnants. Correlation of the distribution of past Amerindian groups and forest species detected species associated with each group. Landscape compartmentalization between the geographical distribution of archaeological sites occupied by Southern-Jê and Guarani groups was identified. Landscapes suitable for Southern-Jê are generally at higher altitudes and close to water courses. Guarani settlements were more likely to be closer to the sea or large rivers. Suitable landscapes for both groups seem to overlap near the larger rivers.

The species that showed association with pre-colonial Amerindian groups could simply be associated with the landscape characteristics that predicted the distribution of these groups. Alternatively, the plant species could be attracted to human occupation or could be favored by humans, by both intentional or unintentional actions [58]. For instance, Amerindians intentionally cleared the land and used fire to produce crops, and unintentionally favored plants by interrupting seeds dormancy, like Mimosa scabrella [59–61], by improvement in light availability to pioneer species, like Solanum mauritianum [60]. In the State of Santa Catarina palynological evidence indicates that the opening of clearings for cultivation started between 3760 and 2430 B.P (Fig 1) [34]. Guarani expanded their occupation area and transplanted useful plants when they migrated, favoring fruit trees that attract fauna, and expanding the territories of species of cultural importance [62]. Amerindians transplanted cassava, beans, maize, and genipap from the Amazon to southern Brazil as part of a cyclic agroforestry production system [62]. The Southern-Jê were probably responsible for Araucaria forest expansion over grasslands [27, 34, 35]. Pre-colonial Southern-Jê and Guarani archaeology reveals evidence of agriculture [30, 62, 63] and pottery production [30, 62–64], and both these activities require fire. Fire was used for many things (e.g., heating, cooking, ceramic preparation, and cremation) and may have involved selective extraction or gathering of species for fuel. Fuel for funeral use must reach very high temperatures, for domestic use it must be easily controllable [65]. The use of fire in rituals can also lead to the selection of woods by symbolic characteristics. Archaeological research has shown that Southern-Jê used preferred species in contexts that indicate the practice of rituals: Araucaria angustifolia, and Jacaranda sp. in a location where Araucária is not available [66]. Fire use could also be unintentionally selective, driving changes in species composition, unintended by those lighting the fires. Agriculture and transplanting fruit trees are examples of intentional activities that modify species composition. Southern-Jê constructed earthworks, that were landscape modifications benefiting some species, in a similar way to the Amazon mounds earthworks [17].

We observed a distribution pattern of Southern-Jê and Guarani groups that was associated with forest types. Southern-Jê sites occur mainly in Mixed Forests (Araucaria forests), where many subterranean structures, earthworks, have been found. Araucaria forests are recognized as an important element in the social organization of the current Southern-Jê (Kaingang and Xokleng/Laklaño) [34, 66]. Southern-Jê controlled their territories by managing the Araucaria forests [64]. Guarani sites have not been found in Mixed Forests but are common in Rain Forest areas. Both groups co-occurred in Seasonal Forests, probably because this forest type is mainly distributed along an important river (the Uruguay River). Rivers are a factor associated with both groups’ distributions, including their ancestral territories. Rivers are especially important in Guarani culture, they are excellent sealers and fishermen [30]. The Southern-Jê arrived before the Guarani in southern Brazil and initially occupied coastal zones and riversides [25]. However, main rivers and the coastline were also migration routes used by the Guarani [30]. The Southern-Jê subsequently intensified the highlands occupation, and this may be due to arrival of the Guarani people [25, 67], or due to ecological identity with their ancestral territories, which are highland headwaters [26].

We found a species partitioning gradient between the two cultural groups in analysis of the species distribution (Fig 7). An important point is that our analyzes do not isolate the causes of the distribution of plant species, we are considering niche construction as a process in which factors of different natures are inseparable. Many of the species listed in this gradient are cited in the ethnobotanical literature as important for Southern-Jê and Guarani. Species such Dicksonia sellowiana (Fig 7A) is associated with Southern-Jê groups, and cited in literature as a culturally valued species that is used to prepare a traditional drink called “Mõg” [68, 69]. Araucaria angustifolia (Fig 7B) appears to be associated with Southern-Jê presence, and archaeological evidence exists of its use by pre-colonial Southern-Jê groups [27, 70, 71]. Dicksonia sellowiana and A. angustifolia are very abundant species in Mixed Forests, as are Ilex paraguariensis (Fig 7C) and Matayba elaeagnoides (Fig 7D). I. paraguariensis is a key cultural species, whose leaves are used to prepare a drink called "chimarrão" or “mate”, that is popular in South America [72]. The practice of consumption and processing of I. paraguariensis leaves are culturally tran;smitted by Amerindians [72]. M. elaeagnoides is described as a good quality burning wood [73]. Lithraea brasiliensis (Fig 7E) has a popular name “bugreiro” (“bugre” with an added suffix to indicate that it is connected to, or used by, the bugre). Although this species is associated with Southern-Jê, L. brasiliensis was also reportedly used by Guarani for construction purposes [74]. Along the entire gradient, we observed species of the Lauraceae family, composed by the genera Ocotea (Fig 6F) and Nectandra (Fig 7G). Among these and linked with Southern-Jê distribution, the species Ocotea porosa is currently listed as vulnerable to extinction [75]. We observed a change in the composition of Lauraceae inside the gradient. From the species are shared by both Southern-Jê and Guarani, the genus Ocotea is replaced by the genus Nectandra. The species shared by the two cultural groups are at the center of the gradient and all of them are abundant in inland Seasonal forests. In addition, all of these shared species are cited by various cultural groups in ethnobotanical literature, reinforcing that they are species of shared cultural use and knowledge. Luehea divaricata (Fig 7H) is used medicinally by Laklaño (Southern-Jê) [69], Guarani [74, 76, 77], and regional farmers [73]. Cupania vernalis (Fig 7I) is used by Laklaño [69], Guarani [74, 76, 77] and regional nowadays farmers [73] for food, construction and artifact production. Species such as Syagrus romanzoffiana (Fig 7J) and Euterpe edulis (Fig 7K) are associated with Guarani presence, but only S. romanzoffiana was common to both Guarani and Southern-Jê. Both species are reportedly used by the Guarani [74] and S. romanzoffiana is known to be cultivated by them [62]. Arecaceae is an important family for human subsistence in the Atlantic Forests, because its species produce many fruits, and in some cases, palm heart [19]. Arecaceae phytoliths have been found in archaeological Southern-Jê pottery, which could suggest consumption and processing or may represent background vegetation [63]. Euterpe edulis is a very abundant species in dense forest (Rain Forest) [78], while S. romanzoffiana also occurs in other forest typologies [78–80]. Other species associated with Guarani presence are Casearia sylvestris (Fig 7L) and Pera glabrata (Fig 7M), both species with Amerindian references in their popular name: “bugre tea” [73, 74, 76] and “bugre heart” [69] respectively (“bugre” is a pejorative name for native people). Casearia sylvestris is a species culturally prominent for present day Guarani [76] and P. glabrata wood is used by Guarani for construction [74, 77] and by Laklaños (Southern-Jê) for artifact production [69].

Our analysis explained only 7% of data variation; however, given the wide spectrum of environmental characteristics that may influence vegetation patterns at the landscape scale, past Amerindian cultures may be an important factor to consider. Our results indicate past Southern-Jê and Guarani distributions are another factor driving differences in present-day forest species composition.

To infer how much of the activities in the pre-colonial period may be registered in today's forest composition, it is important to consider demographic changes. We illustrate the expectation of human impact at different times (Fig 1). Although we still don't have detailed demographic estimates for the pre-colonial period in southern Brazil [24], we have estimate up 3,000 years ago [81]. The population density in the state of Santa Catarina would have been 0,15 inhab/km2 in the period between 11,000 and 7,000 years ago [81]. That density would have remained in the western portion until 3,000 years ago, but would have increased in the east reaching up to 0,75 inhab / km2 [81]. that we don’t have demographic data, but in the period between 1,800 BP and 1,400 BP the Guarani occupation began [30], there was an increase in the Southern-Jê population [34], and as a consequence of this interaction there were cultural changes [67]. In the present year, 2,020, the population density in the state of Santa Catarina varies from 0.7 inhab/km2 (in the largest portion of the territory) to 400/ inhab/km2 (in a few places) [82]. Thus, despite the collapse of the Amerindian populations, there was a population increase in the study area. That pattern contrasts with the Amazon, where there was general depopulation [58]. If on the one hand in the Amazon, populations of domesticated plant species are being lost [58], on the other hand, pre-colonial legacies are evident [18]. In State of Santa Catarina Amerindian populations have drastically reduced, just like in the Amazon, so populations of domesticated species may also be missing. However, if there was intercultural transmission of knowledge, some populations of domesticated species may have perpetuated. In general way, State of Santa Catarina population density increased and that has resulted in intense recent changes in vegetation [83]. Despite this, we have listed here 29 species related to the Southern-Jê and Guarani occupation in the pre-Colonial period. We suggest that traits of domestication of these species be investigated, because some populations of domesticated species in the pre-Colonial period may have been maintained [84], and we need more investigations in this regard.

Past Amerindian distribution

This study found that Southern-Jê and Guarani group distributions are related to different environments. Landscapes suitable for Southern-Jê were likely higher altitude and close to minor water courses, whereas Guarani settlements were more likely located near the sea. Both groups may have shared locations near large rivers.

The past distribution models generated, are an advance on broad analyses, because they reinforce, and spatialize the preferential landscape characteristics for pre-colonial Southern-Jê and Guarani groups. Maps of past Amerindian group distributions are useful for investigating spatial patterns; such as, relationships with vegetation, or landscape partitioning between cultural groups. Distribution models generated with presence data are only potentially usable when they have an AUC value above 0.75 [85]: however, for goodness of fit evaluation of models in general, AUC values above 0.7 can be considered good [86]. Evaluating AUC values in species distribution models, Elith et al (2006, [87]) observed that 64% of the best models presented AUC values above 0.75, and 14% of the best models presented AUC values between 0.7 and 0.75. In this respect, the Guarani distribution model, with a value of 0.83, can be considered adequate. The Southern-Jê model presented an AUC above 0.7, so we consider it acceptable, but understand there are caveats. Our maps explain the spatial dimension of the occupation of southern-Jê and Guarani, but the temporal dimension mixes the entire period of pre-Colonial occupation.

There was a niche overlap between both cultural groups near main rivers. This may be explained by occupation at different times, or by simultaneous occupation involving interactions between the two groups and remains to be further addressed. The Southern-Jê initially occupied diverse territories in Southern Brazil. Around the time the Guarani arrived and began to occupy areas close to large rivers and the coast the Southern-Jê were settled in the highlands. Although there are records of Southern-Jê presence in the highlands since 2000 years BP, it is only around 1000 years BP that these groups started to build their funerary earthworks [64, 67]. This new architectural expression may be a way of controlling their territory, because it coincides with the arrival of Guarani groups [67]. Guarani groups expanded their occupation along the margins of major rivers and along the coastline [30]. The Jê people were distributed over the highlands in their Brazilian territory [25, 26], and their pattern of migration created isolated groups, differing from the Guarani [25]. Following contact with Europeans, Amerindian populations were reduced due to territorial conflicts, diseases, political, and social disruption. There is evidence of Guarani occupation followed by subsequent Southern-Jê occupation in some locations [25], suggesting that the groups competed for the territories. Furthermore, synchronous Guarani sites inside of Southern-Jê areas have been found, suggesting fluid frontiers where these groups have interacted over time [25].

Conclusion

Although there are still many pieces of the puzzle to be put together for a complete understanding of vegetation patterns, this study is the first to assess pre-colonial cultural effects, on the contemporary forest composition, of a large extent of the Atlantic Forest. This study evaluated forests that have been seriously altered post-European colonization [83] and still detected a legacy of pre-colonial cultural groups on forest composition. Forest composition differs in landscapes previously occupied by different Amerindian groups. This difference may be due to the different ways differing cultures manage territory, indicative of Cultural Niche Construction. We conclude that long-term cultural activities may have acted with other biotic and abiotic processes to determine forest compositions in the Southern Atlantic Forest.

Supporting information

S1 Code. R Scripts.

This file contains all the codes wrote in R.

(DOCX)

Click here for additional data file.^{(473.7KB, docx)}

S1 File. ENM Guarani raster.

This file can be opened using a Geographic Information System (Gis Software) or following the script provided in S1 Code.

(CSV)

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S2 File. ENM Southern-Jê raster.

This file can be opened using a Geographic Information System (Gis Software) or following the script provided in S1 Code.

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S3 File

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S4 File

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S5 File

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S1 Data

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S2 Data

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S1 Fig

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Acknowledgments

We thanks to Francisco Noelli and Jonas Gregorio de Souza, for Southern-Jê data. We are also grateful to the Laboratory of Interdisciplinary Studies in Archaeology (LEIA) at the Federal University of Santa Catarina (UFSC) for archaeological data; thanks to Leonardo de Campos who made many helpful suggestions on an earlier draft; thanks to Mário Tagliari and Ramiro Melinski for contributions in ecological niche modeling; thanks to Christopher Graves for English revision; and thanks to Thiago Ehlert for the tabulation of useful plants data.

Data Availability

We have organized the data in order to preserve the third-party data, but to allow the analysis to be reproduced. The following data is in the supporting information files: All steps of the analysis in R, containing the names of the files we are sharing and / or the download paths. Two georeferenced images, with the results of niche modeling, The Guarani model and the Southern-Jê model. Part of the geographical coordinates of the Southern-Jê archaeological sites. Floristic data are third-party Data. We are sharing the abundance matrix with anonymization of species names and sample units. These data belong to the Floristic Forest Inventory of the State of Santa Catarina (IFFSC), and are published in books, available for download at: <https://sites.google.com/view/iffportal/publica%C3%A7%C3%B5es/livros>. We requested the abundance matrix (raw data) from the IFFSC coordination, through an email with a description of our research objective. The project coordinator is Dr. Alexander Christian Vibrans, from the Regional University of Blumenau - FURB. Their emails are: acv@furb.br; iffsc@furb.br; inventarioflorestal.sc@gmail.com. The portion of the Southern-Jê data that is published in Noelli e Souza, 2017, (doi: 10.1590/1981.81222017000100004) was provided by the authors and is third party data and, for that reason we are not sharing. However, we are sharing a set of locations from Southern-Jê archaeological sites. All these locations are in a radius of 200 m. from some archaeological site published by the National Historical and Artistic Heritage Institute of Brazil (IPHAN) in: http://portal.iphan.gov.br/cna/pagina/detalhes/1227 data Guarani can be found in http://dx.doi.org/10.1016/j.quaint.2014.10.050; The authors of this study had no special access privileges others would not have.

Funding Statement

This work was partly supported by the “Territorialidades Ameríndias no Alto Vale do Itajaí” financed in part by the “Elisabete Anderle Award 2017 - Segment material and immaterial heritage / archaeological research" (contract number 0148/2017 and process number 2578/2017) received by JSM, and is supported partly by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES, https://www.capes.gov.br/) - Finance Code 001, including APC doctoral fellowship. NP has received a productivity fellowship from CNPq (National Council for Scientific and Technological Development, http://www.cnpq.br/)(CNPq 310443/2015-6). CL has received from CNPq a post-doctoral fellowship (CNPq 159440/2018-1).

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PLoS One. doi: 10.1371/journal.pone.0235819.r001

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PONE-D-19-33906

Pre-colonial Amerindian legacies in forest composition of southern Brazil

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Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Partly

Reviewer #2: Yes

**********

2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

Reviewer #2: N/A

**********

3. Have the authors made all data underlying the findings in their manuscript fully available?

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Reviewer #1: Yes

Reviewer #2: Yes

**********

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Reviewer #1: Yes

Reviewer #2: Yes

**********

5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: This is a very interesting and original research on historical ecology carried out in Santa Catarina State in Brazil. The paper is fairly well organised, the topic is very important and deserves to be published. The Introduction is well-written and interesting.

However, in other part the text is sometimes not so clear and there are paragraphs that should be read again to become easy understandable.

As a general advice, I suggest to consider some aspects to be incorporated in the paper:

a) The chronological framework and diachronical succession of events seems to me not discussed; please, consider that the international audience can be not informed about what means ‘pre-colonial’ – and when the change from the hunter-gatherer to Neolithic culture (transitions or migrations?) had occurred in Brazilian regions.

b) The demography datum is missed. One should consider that the number of people changed during the different chrono-cultural phases, and therefore when you mention 11,500 BP hunter-gatherers, were they fewer or had a similar impact than the more recent human groups?

c) Are there known differences in the cultural choices (environmental-dependent?) of the human groups involved in the research? (probably they are evident from the Fig. 6 ) Or the differences are also linked to the changes of one culture along the changing millennia?

Line 36 = Araucaria can be added to the keywords

Lines 68-69 = how long is this ‘long-term’ history? It emerges from the next paragraph that it is about 11,500 years long. Therefore, a more precise chronology can be added, such as changing “identifying many archaeological sites” to “identifying many Holocene archaeological sites”, or something similar.

Line 103 = “humans are cultural niche constructors” = this statement is absolutely sharable and there is a general consensus on this vision, as well as “different cultural groups also leave differing floristic composition legacies” = please note, the presence of archaeological sites and archaeological/archaeobotanical findings is mandatory to infer the type of land use. Further information can be obtained from the ethnobotanical review, as in the discussion of this paper.

Line 244 = the first part of the discussion is well centred on geographical and geomorphological characteristics that can explain the observed results and trends. However, the timeline variable – chronology of archaeological sites/events is not reported in the following paragraphs.

Line 253 = ‘Alternatively, the species could’ = Alternatively, the plant species could

Line 255 = as an example on the need of chronological details, in the sentence “Amerindians intentionally cleared the land and used fire to produce crops,”, you should add the culture and/or Holocene period or millennia BP.

Line 264 = “may have involved selective extraction of species for fuel” = is this a speculative argument or can you make an example from a case study?

Lines 279-310 = This is an interesting ethnobotanical part of the text but it is not clear in which order the plants/uses are discussed, and why they are listed.

Line 285 = phytolits in pottery could suggest consumption or, preferably, are they evidence of plant processing?

Line 385 = practical

Citations 31. 32, 33, 77 = put ‘Aracauria’ in italics

Citation 37, 55 = check the format

Fig. 6 = it is an interesting figure (please, quote it in the text) but I am not sure that it is sufficiently clear the sense and aim of this ‘separation of plant species’ – Cultural? Phytogeographical? Current or past different exploitation? The full name of species should be preferred to the abbreviation of the genus, if possible.

Reviewer #2: The article PONE-D-19-33906 “Pre-colonial Amerindian legacies in forest composition of southern Brazil” presents the analyzes on patterns of occupations and settlements of ancient societies in Brazilian lands. The evidence left by Amerindian societies (especially Southern-Jê and Guarani) is persistent in forests around the world. This research presents some important results about how these societies have affected and managed over time the forest structure in southern Brazil. The investigation allows to evaluate the tree composition of forest fragments and the corresponding environmental variables in the state of Santa Catarina. Archaeological data were used to understand ecological niche models thinking of prediction models. The amount of inventory studied is quite significant. The methodology used is correct and can be an excellent tool for further studies. It is noteworthy that the settlements and sites of these follow the pattern already studied since the 1960s by American and Brazilian archaeologists. Which serves as confirmation of some procedural models. As for the results and analyzes on the floristic composition of forest remnants and the relationship of the distribution of ancient Amerindian groups and the species associated with each group, the sites demonstrate the patterns of occupation at altitudes and reliefs already highlighted in other classical studies of Brazilian archaeology. Structurally the manuscript is in accordance with a scientific text following the ethical and methodological rigors. The bibliography is up to date and of excellent quality. The manuscript deserves to be published and will bring important contributions to current studies on ecological and environmental history in southern Brazil, still ephemeral in archaeological studies.

**********

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Reviewer #1: No

Reviewer #2: No

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PLoS One. 2020 Jul 23;15(7):e0235819. doi: 10.1371/journal.pone.0235819.r002

Author response to Decision Letter 0

29 Feb 2020

Dear Editors,

We are grateful to you and reviewers for comments regarding our work. The comments were very positive and contributed to improving the manuscript.

In short, we are sharing several files, including all the code we have written, to improve the reproducibility of our analyzes. Changes were made in the manuscript and figures to address the issues raised in review process.

We hope that with the improvements in the manuscript it can be published in Plos One.

RESPONSE TO JOURNAL RQUIREMENTS

Journal Point 1: “Please ensure that your manuscript meets PLOS ONE's style requirements, including those for file naming.”

Response to Journal Point 1: We check and adjust the style requirements in the manuscript and in the filenames.

Journal Point 2: “We note that you have indicated that data from this study are available upon request. PLOS only allows data to be available upon request if there are legal or ethical restrictions on sharing data publicly.”

Response to Journal Point 2: We organize the data in order to preserve the third-party data, but to allow the analyzes to be reproduced. All steps of the analysis in R, containing the names of the files we are sharing and / or the download paths are in the file “Methods.docx”.

We are sharing two georeferenced images, with the results of niche modeling. The Guarani model is available in the archive “S1A_result.tif” and the Southern-Jê model in file “S1B_result.tif”.

Floristic data are third-party Data. We are sharing the abundance matrix with anonymization of species names and sample units. These data belong to the Floristic Forest Inventory of the State of Santa Catarina (IFFSC), and are published in books, available for download at:

<https://sites.google.com/view/iffportal/publica%C3%A7%C3%B5es/livros>.

We requested the abundance matrix (raw data) from the IFFSC coordination, through an email with a description of our research objective. The project coordinator is Dr. Alexander Christian Vibrans, from the Regional University of Blumenau - FURB. Your emails are:acv@furb.br; iffsc@furb.br; inventarioflorestal.sc@gmail.com.

We are sharing part of the geographical coordinates of the Southern-Jê archaeological sites in the named file “S1B_POINTS.csv”. We do not share the portion of the data that is published in Noelli e Souza, 2017, (doi: 10.1590/1981.81222017000100004) and was provided by the authors (thirdy-part data). To request this data, the authors must be contacted by e-mails:

Jonas Jonas Gregorio de Souza <jonas.gregorio@yahoo.com.br>;

Francisco Silva Noelli <chico.noelli@gmail.com>.

Journal Point 3: “We note that Figures 1, 3 and 4 in your submission contain map images which may be copyrighted.”

Response Journal Point 3: All figures are original and was created by authors. We don’t utilize proprietary data.

RESPONSE TO REVIEWERS

We thank the reviewers for the assessment and comments of our manuscript. We have modified the text and figures according to the referees’ critiques. Below are our comments on each issue raised by Reviewer #1.

Reviewer Point 1: “This is a very interesting and original research on historical ecology carried out in Santa Catarina State in Brazil. The paper is fairly well organized, the topic is very important and deserves to be published. The Introduction is well-written and interesting. However, in other part the text is sometimes not so clear and there are paragraphs that should be read again to become easy understandable.”

Response to Reviewer Point 1: We hope that the text is clearer with the changes. If there is any critical point, please do not hesitate to let us know.

Reviewer Point 2: “As a general advice, I suggest to consider some aspects to be incorporated in the paper: a) The chronological framework and diachronical succession of events seems to me not discussed; please, consider that the international audience can be not informed about what means ‘pre-colonial’ – and when the change from the hunter-gatherer to Neolithic culture (transitions or migrations?) had occurred in Brazilian regions.”

Response to Reviewer Point 2: We agree with the reviewer on this important point about succession of events. We included a timeline (Fig 1, in line 89), with the aim of making the sequence of events in the population of the State of Santa Catarina better understandable.

About culture transitions, research conducted in recent years have pointed to the complexity of the subject due to regional variations, linked to specific cultural histories and the unsuitability of a consolidated basis for the study of European prehistory in the tropics. In general, these studies have shown that the aspects traditionally considered as key elements for this “Neolithic transition”, such as ceramics, sedentary lifestyle, agriculture, do not appear together at once time. There are several examples in tropical regions for a very old presence of ceramics, which, however, does not remain and is constant or in continuous growth, involving, in some regions, production cycles and disappearance. At the same time, there are indications of domesticated items in contexts with dates for the Early Holocene, which are not associated with the presence of ceramics or changes in the pattern of mobility. In other words, these aspects integrate local historical trajectories, with regional variations in chronological and organizational terms. In southern Brazil, what we have identified is a change in the distribution pattern of the sites and their associated archaeological remains from the 3,000 years BP, when a process of cultural interaction is intensified associated with the migration of Jê groups to the southern plateau. However, on the coast, since 7,000 years ago, with intensification between 5-3,000, significant changes involving demography and territoriality are identified and associated with the construction of large shell mounds.

Reviewer Point 3: “(…) I suggest to consider some aspects to be incorporated in the paper: (…) b) The demography datum is missed. One should consider that the number of people changed during the different chrono-cultural phases, and therefore when you mention 11,500 BP hunter-gatherers, were they fewer or had a similar impact than the more recent human groups?”

Response to Reviewer Point 3: We agree with the reviewer, demography dynamics is important to our discussion. We include demography datum in between lines 370-387. Regarding the impact differences between temporal phases, we expect the marks on the vegetation to be more visible when more recent and / or more persistent. Regarding the persistence of the effects is related to larger population, greater cultural diversity, and more advanced technology. We synthesize the expected effects in a gradient illustrated along the timeline (Fig 1, in line 89).

Reviewer Point 4 “(…) I suggest to consider some aspects to be incorporated in the paper: (…) c) Are there known differences in the cultural choices (environmental-dependent?) of the human groups involved in the research? (probably they are evident from the Fig. 6) Or the differences are also linked to the changes of one culture along the changing millennia?”

Response to Reviewer Point 4: Yes, there are cultural differences related to the choice of environments. In general, both cultural groups prefer proximity to rivers. But “rivers are especially important to Guarani culture, they are sealers, fishermen” (added in text in lines 293-294). Southern-Jê prefers highland areas, and “Araucaria forests are recognized as an important element in the social organization of the current Southern-Jê (Kaingang and Xokleng/Laklaño)”, i.e, Araucaria forests are key in their social organization (added in lines 288-289). In this way, at the Guarani contact Southern-Jê building monument to defend highland territories (were Araucaria forest occur), marking a cultural transition: “Although there are records of Southern-Jê presence in the highlands since 2000 years BP, it is only around 1000 years BP that these groups started to build their funerary earthworks [61,65]. This new architectural expression may be a way of controlling their territory, because it coincides with the arrival of Guarani groups [65].” (lines 419-422). In the moment of contact with European immigrants, another change in the distribution pattern occurs. Due to the collapse of populations, areas were opened for re-occupation: “Following contact with Europeans, Amerindian populations were reduced due to territorial conflicts, diseases, political, and social disruption. There is evidence of Guarani occupation followed by subsequent Southern-Jê occupation in some locations [25], suggesting that the groups competed for the territories” (lines 424-427). The migration patterns of each group also influence the choice of occupied areas: Guarani along rivers and forming networks, and Southern-Jê in highlands. In manuscript is as follows: “Guarani groups expanded their occupation along the margins of major rivers and along the coastline [30]. The Jê people were distributed over the highlands in their Brazilian territory [25,26], and their pattern of migration created isolated groups, differing from the Guarani [25].” (lines 421-423).

Reviewer Point 5: “Line 36 = Araucaria can be added to the Keywords.”

Response to Reviewer Point 5: We agree and include Araucaria in the key words.

Reviewer Point 6: “Lines 68-69 = how long is this ‘long-term’ history? It emerges from the next paragraph that it is about 11,500 years long. Therefore, a more precise chronology can be added, such as changing “identifying many archaeological sites” to “identifying many Holocene archaeological sites”, or something similar.”

Response to Reviewer Point 6: We added a timeline (Fig 1) and added “Holocene” in text.

Reviewer Point 7: “Line 103 = “humans are cultural niche constructors” = this statement is absolutely sharable and there is a general consensus on this vision, as well as ‘different cultural groups also leave differing floristic composition legacies’ = please note, the presence of archaeological sites and archaeological/archaeobotanical findings is mandatory to infer the type of land use. Further information can be obtained from the ethnobotanical review, as in the discussion of this paper.”

Response to Reviewer Point 7: We agree that the description of our hypothesis highlighted aspects that are consensus. We have included an important aspect in the text, about the possibility that there are marks of human actions from the past visible on current forests despite the intense modification in the last century, as in our study area (Sevegnani et al, 2019). In manuscript is as follows: “our hypothesis is that floristic composition differs in Southern Atlantic Forests with a high probability of past human activities, and that different cultural groups also leave differing floristic composition and abundance legacies notable yet, despite the disruption of these cultural groups and of the intense changes caused by recent populations.” (lines 112-116).

The gradient parallel to the timeline in Fig 1 illustrates this.

Reviewer Point 8: “Line 244 = the first part of the discussion is well centred on geographical and geomorphological characteristics that can explain the observed results and trends. However, the timeline variable – chronology of archaeological sites/events is not reported in the following paragraphs.”

Response to Reviewer Point 8: In our analysis, we prioritize the spatial dimension. We ignore the dates because this information is not available for all archaeological sites. Including dates would reduce sampling, making it difficult to calibrate the models. Our results indicate the occupation space accumulated between ~ 3,000-300 years BP. We include this information in the manuscript methods and discussion as bellow: “We do not consider the temporal information of archaeological sites, as this would considerably reduce the sample size.” (lines 156-158) and “Our maps explain the spatial dimension of the occupation of southern-Jê and Guarani, but the temporal dimension mixes the entire period of pre-Colonial occupation.” (lines 412-414).

Reviewer Point 9: “Line 253 = ‘Alternatively, the species could’ = Alternatively, the plant species could.”

Response to Reviewer Point 9: We agree and modify the text.

Reviewer Point 10: “Line 255 = as an example on the need of chronological details, in the sentence ‘Amerindians intentionally cleared the land and used fire to produce crops,’, you should add the culture and/or Holocene period or millennia BP.”

Response to Reviewer Point 10: We agree and modify the text. We highlighted in text the moment when fire started to be used to open clearings: “In the State of Santa Catarina palynological evidence indicates that the opening of clearings for cultivation started between 3760 and 2430 B.P (Fig 1) [34]” (lines 269-271).

Reviewer Point 11: “Line 264 = “may have involved selective extraction of species for fuel” = is this a speculative argument or can you make an example from a case study?”

Response to Reviewer Point 11: We agree and include study case examples: “Fuel for funeral use must reach very high temperatures, for domestic use it must be easily controllable [63]. The use of fire in rituals can also lead to the selection of woods by symbolic characteristics. Archaeological research has shown that Southern-Jê used preferred species in contexts that indicate the practice of rituals: Araucaria angustifolia, and Jacaranda sp. in a location where Araucária is not available [64].”(lines 278-281).

Reviewer Point 12: “Lines 279-310 = This is an interesting ethnobotanical part of the text, but it is not clear in which order the plants/uses are discussed, and why they are listed.”

Response to Reviewer Point 12: We rearranged the paragraph. Now the text follows a sequence from left to right along the gradient illustrated in Fig 7 (it was Fig 6). That is, starting with species related to the Southern-Jê occupation and ending with species related to the Guarani occupation. The species are listed in Fig 7. We have included captions in the text and in the figure to facilitate the location of each species. For some of the listed species, we discussed ecological and utilitarian features that can help us understand this gradient. We also highlight those species that have an indigenous reference in their names.

Reviewer Point 13: They “Line 285 = phytolits in pottery could suggest consumption or, preferably, are they evidence of plant processing?”

Response to Reviewer Point 13: They could be evidence of processing. We modify the text.

Reviewer Point 14: “Line 385 = practical”

Response to Reviewer Point 14: We agree and modify the text.

Reviewer Point 15: “Citations 31. 32, 33, 77 = put ‘Aracauria’ in italics.”

Response to Reviewer Point 15: We agree and modify the text.

Reviewer Point 16: “Citation 37, 55 = check the format.”

Response to Reviewer Point 16: We agree and modify the text.

Reviewer Point 17: “Fig. 6 = It is an interesting figure (please, quote it in the text) but I am not sure that it is sufficiently clear the sense and aim of this ‘separation of plant species’ – Cultural? Phytogeographical? Current or past different exploitation? The full name of species should be preferred to the abbreviation of the genus, if possible.”

Response to Reviewer Point 17: We quote Figure 7 (it was 6) in text. The figure shows a list of species with RDA scores above |0.1| and organizes them along a cultural gradient between Southern-Jê and Guarani. Thus, we show species that has its distribution associated with the distribution of archaeological sites Southern-Jê and Guarani. Species can also be related to the environmental characteristics of the archaeological sites’ specific location. They could be yet related with an interaction of environment and occupation. There are many factors that influence the distribution of species and the causes cannot be explained by our analyzes. We highlight this in the manuscript as follow: “An important point is that our analyzes do not isolate the causes of the distribution of plant species, we are considering niche construction as a process in which factors of different natures are inseparable.” (lines 300-301).

Attachment

Submitted filename: Response to Reviewers.docx

Click here for additional data file.^{(93KB, docx)}

PLoS One. doi: 10.1371/journal.pone.0235819.r003

Decision Letter 1

André Jasper

8 May 2020

PONE-D-19-33906R1

Pre-colonial Amerindian legacies in forest composition of southern Brazil

PLOS ONE

Dear Dr. Pereira-Cruz,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it only a few changes are necessary that your manuscript can be accepted for publication in PLOS ONE. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

We would appreciate receiving your revised manuscript by Jun 22 2020 11:59PM. When you are ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

If you would like to make changes to your financial disclosure, please include your updated statement in your cover letter.

Please include the following items when submitting your revised manuscript:

A rebuttal letter that responds to each point raised by the academic editor and reviewer(s). This letter should be uploaded as separate file and labeled 'Response to Reviewers'.
A marked-up copy of your manuscript that highlights changes made to the original version. This file should be uploaded as separate file and labeled 'Revised Manuscript with Track Changes'.
An unmarked version of your revised paper without tracked changes. This file should be uploaded as separate file and labeled 'Manuscript'.

We look forward to receiving your revised manuscript.

Kind regards,

André Jasper, Ph.D.

Academic Editor

PLOS ONE

Additional Editor Comments (if provided):

Dear Dr. Pereira-Cruz and colleagues,

After carefully revising your resubmission, we are quite reaching a final decision that is now conditioned to a few details to be changed in your manuscript.

Both reviewers and I agreed that your manuscript is an interesting contribution for PlosOne and, for final acceptance, we kindly ask for the following (as pointed out by Dr. Mercuri):

Line 80 = instead of “Zea Mays, Manihot Esculenta, and Dioscorea sp,” = “Zea mays, Manihot esculenta, and Dioscorea sp.”

Line 123 = delete comma after ‘ (SC) ‘

Lines 157-158 = “We do not consider the temporal information of archaeological sites, as this would considerably reduce the sample size.” This sentence was added now to reply to comments by the reviewer, however it may introduce other criticisms: the chronology is a must in archaeology. Probably, it may be better changing the sentence by reporting only the information on the range of chronology of the sites studied, that is: “ the archaeological sites have been occupied during the last XX millennia”

Line 269 = “unintentionally favored certain plants and/or crops by…” = ‘certain’ is vague and, if correct, ‘synanthropic’ (or the name of the species) should be preferred

Line 359 = check “and we need”

Fig. 4 = check the green colors as it seems that the color of Brazil and the of S. America are inverted, especially in the part ‘ c ‘ of the figure.

Looking forward for you return,

André

[Note: HTML markup is below. Please do not edit.]

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #1: All comments have been addressed

**********

2. Is the manuscript technically sound, and do the data support the conclusions?

Reviewer #1: Yes

**********

3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

**********

4. Have the authors made all data underlying the findings in their manuscript fully available?

Reviewer #1: Yes

**********

5. Is the manuscript presented in an intelligible fashion and written in standard English?

Reviewer #1: Yes

**********

6. Review Comments to the Author

Reviewer #1: The clarity of the paper was notably improved, Fig. 1 is very useful and well done; also the Fig. 7 is now more useful. This research is a very good contribution to the field and I support its publication on PLOSone.

Few remarks are reported below.

Line 80 = “Zea Mays, Manihot Esculenta, and Dioscorea sp,” = “Zea mays, Manihot esculenta, and Dioscorea sp.”

Line 123 = delete comma after ‘ (SC) ‘

Line 269 = “unintentionally favored certain plants and/or crops by…” = ‘certain’ is vague and, if correct, ‘synanthropic’ (or the name of the species) should be preferred

Line 359 = check “and we need”

Fig. 4 = check the green colours a sit seems that the colour of Brazil and ithe S. America are inverted, especially in the part ‘ c ‘ of the figure.

**********

7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #1: Yes: Anna Maria Mercuri

PLoS One. 2020 Jul 23;15(7):e0235819. doi: 10.1371/journal.pone.0235819.r004

Author response to Decision Letter 1

22 Jun 2020

Response to reviewer:

We thank the Dr. Mercuri for the assessment and comments of our manuscript. We have modified the text and figures according to the referees’ critiques. Below are our comments on each issue raised by Dr. Mercuri.

Reviewer Point 1: “Line 80 = “Zea Mays, Manihot Esculenta, and Dioscorea sp,” = “Zea mays, Manihot esculenta, and Dioscorea sp.””

Response to Reviewer Point 1: We fix it.

Reviewer Point 2: “Line 123 = delete comma after ‘(SC)‘”

Response to Reviewer Point 2: We delete the comma.

Reviewer Point 3: “Lines 157-158 = “We do not consider the temporal information of archaeological sites, as this would considerably reduce the sample size.” This sentence was added now to reply to comments by the reviewer, however it may introduce other criticisms: the chronology is a must in archaeology. Probably, it may be better changing the sentence by reporting only the information on the range of chronology of the sites studied, that is: “the archaeological sites have been occupied during the last XX millennia””.

Response to Reviewer Point 3: We agree about the high importance of chronology in archaeology and modify the text as suggested. We removed the sentence that might sound while minimizing the importance of the chronology: “We do not consider the temporal information of archaeological sites, as this would considerably reduce the sample size.”. And we include the sentence “The selected archaeological sites have been occupied over the past 3,000 years” (line 159-160).

Reviewer Point 4: “Line 269 = “unintentionally favored certain plants and/or crops by…” = ‘certain’ is vague and, if correct, ‘synanthropic’ (or the name of the species) should be preferred”.

Response to Reviewer Point 4: We agree and remove the word “certain”. We include an example: “… and unintentionally favored plants by interrupting seeds dormancy, like Mimosa scabrella [59–61], by improvement in light availability to pioneer species, like Solanum mauritianum [60].”.

Reviewer Point 5: “Line 359 = check “and we need””.

Response to Reviewer Point 5: We fix it.

Reviewer Point 6: “Fig. 4 = check the green colours a sit seems that the colour of Brazil and it he S. America are inverted, especially in the part ‘ c ‘ of the figure.”

Response to Reviewer Point 6: We fix it.

Attachment

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Click here for additional data file.^{(81.3KB, docx)}

PLoS One. doi: 10.1371/journal.pone.0235819.r005

Decision Letter 2

André Jasper

24 Jun 2020

Pre-colonial Amerindian legacies in forest composition of southern Brazil

PONE-D-19-33906R2

Dear Dr. Cruz and Colleagues,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

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PLoS One. doi: 10.1371/journal.pone.0235819.r006

Acceptance letter

André Jasper

8 Jul 2020

PONE-D-19-33906R2

Pre-colonial Amerindian legacies in forest composition of southern Brazil

Dear Dr. Pereira Cruz:

I'm pleased to inform you that your manuscript has been deemed suitable for publication in PLOS ONE. Congratulations! Your manuscript is now with our production department.

If your institution or institutions have a press office, please let them know about your upcoming paper now to help maximize its impact. If they'll be preparing press materials, please inform our press team within the next 48 hours. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information please contact onepress@plos.org.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

S1 Code. R Scripts.

This file contains all the codes wrote in R.

(DOCX)

Click here for additional data file.^{(473.7KB, docx)}

S1 File. ENM Guarani raster.

This file can be opened using a Geographic Information System (Gis Software) or following the script provided in S1 Code.

(CSV)

Click here for additional data file.^{(4.1KB, csv)}

S2 File. ENM Southern-Jê raster.

This file can be opened using a Geographic Information System (Gis Software) or following the script provided in S1 Code.

(CSV)

Click here for additional data file.^{(4.8KB, csv)}

S3 File

(CSV)

Click here for additional data file.^{(39.6KB, csv)}

S4 File

(CSV)

Click here for additional data file.^{(610.2KB, csv)}

S5 File

(CSV)

Click here for additional data file.^{(13.3KB, csv)}

S1 Data

(RAR)

Click here for additional data file.^{(35.5KB, rar)}

S2 Data

(RAR)

Click here for additional data file.^{(56.9MB, rar)}

S1 Fig

(TIF)

Click here for additional data file.^{(512.7KB, tif)}

Attachment

Submitted filename: Response to Reviewers.docx

Click here for additional data file.^{(93KB, docx)}

Attachment

Submitted filename: Response to Reviewers.docx

Click here for additional data file.^{(81.3KB, docx)}

Data Availability Statement

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Pre-colonial Amerindian legacies in forest composition of southern Brazil

Aline Pereira Cruz

Eduardo Luiz Hettwer Giehl

Carolina Levis

Juliana Salles Machado

Lucas Bueno

Nivaldo Peroni

Roles

Abstract

Introduction

Fig 1. Timeline of diachronical succession of human population events and linked changes in vegetation cover, in State of Santa Catarina.

Materials and methods

Study area

Fig 2. Map of South America highlighting the study area that comprises the State of Santa Catarina.

Vegetation data

Archaeological data

Topographic, hydrographic and forest type data

Data analysis

Results

Fig 3. Ridgeline plot of smoothed density estimates of the probability of occurrence of Southern-Jê (gray) and Guarani (orange) archaeological sites in relation to environmental variables.

Fig 4.

Fig 5. Spatial partitioning between archaeological Southern-Jê and Guarani distribution.

Fig 6. The redundancy analysis of species abundance and the probability of occurrence of Southern-Jê and Guarani archaeological sites.

Fig 7. Relationship between tree species and the two past Amerindian groups (Guaraní and Southern-Jê).

Discussion

Past Amerindian distribution

Conclusion

Supporting information

Acknowledgments

Data Availability

Funding Statement

References

Decision Letter 0

André Jasper

Roles

Author response to Decision Letter 0

Decision Letter 1

André Jasper

Roles

Author response to Decision Letter 1

Decision Letter 2

André Jasper

Roles

Acceptance letter

André Jasper

Roles

Associated Data

Supplementary Materials

Data Availability Statement

ACTIONS

PERMALINK

RESOURCES

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