Fire frequency restructures culturally important tree communities across forest and savanna landscapes in the Xingu Indigenous Territory

Fire frequency restructures culturally important tree communities across forest and savanna landscapes in the Xingu Indigenous Territory | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Fire frequency restructures culturally important tree communities across forest and savanna landscapes in the Xingu Indigenous Territory Lui Gustavo Carvalho, Hernani Fernandes Oliveira, Divino vicente Silvério, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9053133/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 21 You are reading this latest preprint version Abstract Fire has become an increasingly pervasive disturbance in southern Amazonia, and climate projections indicate that fire frequency is likely to intensify in the coming decades. These changes pose a growing threat to forest biodiversity and to the ecosystem services that sustain Indigenous peoples. Here, we evaluate how fire frequency influences the provision of ecosystem services derived from tree species used by the Kuikuro people in the Xingu Indigenous Territory (XIT), Brazil. We conducted forest inventories across a fire-history gradient ranging from zero to six burn events over the past 40 years, sampling 32 plots (10 × 10 m) distributed across forest (n = 20) and savanna (n = 12) vegetation types. For all trees with diameter ≥ 10 cm, we measured diameter, and recorded ethnobotanical uses reported by the Kuikuro people, grouped into three categories, (1) food, (2) construction/handcrafts, and (3) medicinal use. We also incorporated environmental predictors derived from remote sensing and soil data, including altitude, slope, soil pH, clay content, and bulk density. Across all plots, we recorded 92 tree species (54 in forest and 38 in savanna). In forest environments, 6.3% of species were used for food, 9.1% for medicinal purposes, and 17.3% for construction or handcrafts. In savanna, these proportions were 5.4%, 11.8%, and 3.6%, respectively. Regression analyses revealed that increasing fire frequency had a strong negative effect on the abundance of tree species used for construction/handcrafts and medicinal purposes in forest areas. In contrast, in savanna environments, fire frequency showed weaker negative effects on food- and construction-related species and a positive association with medicinal species. Because forests dominate much of the XIT and are becoming increasingly exposed to fire, our findings indicate that tree species providing materials for construction, handcrafts, and medicine are particularly vulnerable to altered fire regimes. These changes threaten not only forest biodiversity, but also the livelihoods, traditional knowledge, and cultural integrity of the Kuikuro people, underscoring the need for fire-sensitive management strategies that integrate ecological and Indigenous perspectives. ethnobotany floristic composition provisioning services Amazon-Savanna ecotone protected areas Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Indigenous territories (ITs) play a crucial role in global biodiversity conservation, with Indigenous peoples managing approximately 28% of the Earth’s land surface while safeguarding nearly 80% of the world’s remaining biodiversity (Garnett et al. 2018 ). In Brazil, around 208,000 Indigenous people inhabit the Amazonia biome, occupying approximately 387 Indigenous Territories that collectively represent about 23% of the Brazilian Amazon basin (FUNAI 2024 ; Povos Indígenas no Brasil 2024 ). In the Cerrado, 109 Indigenous Territories cover 4.35% of the total area and play a fundamental role in conserving biodiversity and ecosystem processes in this highly threatened savanna system (WWF 2020 ). Within this context, the Xingu Indigenous Territory (XIT), located in the state of Mato Grosso, encompasses approximately 2.8 million hectares and harbors more than 16 Indigenous peoples with deep ecological knowledge and long-standing cultural connections to forest, savanna, and their natural resources (Sanches et al. 2012 ; Povos Indígenas do Brasil 2024). Among these groups, the Kuikuro people are one of the largest populations and have historically relied on a diverse array of tree species for food, construction materials, handcrafts, and medicinal purposes. Their livelihoods and cultural practices are closely tied to the structure, composition, and regeneration dynamics of both forest and Cerrado ecosystems. Fire has long been an integral component of Indigenous land management in the XIT. The Kuikuro employ traditional fire management practices that are spatially controlled, seasonally timed, and focused on small areas, creating heterogeneous vegetation mosaics at different stages of regeneration. These practices enhance food production, maintain landscape diversity, and ensure the sustained availability of plant resources critical for subsistence and cultural continuity (Denevan 2001 ; Pivello 2011 ; Schwartzman et al. 2013 ; Schmidt et al. 2021 ). Importantly, this knowledge is transmitted across generations and embedded within cultural norms that promote ecosystem resilience and resource renewal. However, Indigenous territories across southern Amazonia and the Amazonia–Cerrado transition are increasingly exposed to uncontrolled forest fires driven by climate change, prolonged droughts, and expanding agricultural frontiers (Cochrane 2003 ; Nepstad et al. 2004 ; Brando et al. 2014 ). These fires differ fundamentally from traditional burning practices: they are larger in extent, more frequent, and often more intense, leading to altered forest structure, increased tree mortality, and long-term changes in species composition (Brando et al. 2014 , 2019 ; Feng et al. 2021 ). As fire frequency increases, ecosystems become progressively more vulnerable, with cascading effects on regeneration processes and the persistence of fire-sensitive tree species. Such changes have direct implications for the provisioning ecosystem services upon which Indigenous peoples depend. Uncontrolled fires can reduce the availability of tree species used for construction, handcrafts, medicine, and food, thereby undermining food security, cultural practices, and traditional knowledge systems (Eloy et al. 2021 ). These impacts are expected to be particularly pronounced in forest environments, where tree species are generally less adapted to frequent fire, while savanna systems may exhibit more variable responses due to long-term evolutionary exposure to fire. Despite growing recognition of the importance of Indigenous fire management for conservation and food security, there remains limited quantitative evidence on how increasing fire frequency affects the diversity and availability of tree species that provide provisioning ecosystem services to Indigenous communities across contrasting vegetation types. Addressing this knowledge gap is critical for informing conservation policies and fire management strategies that integrate ecological processes with Indigenous knowledge and governance systems. In this study, we evaluate how fire frequency influences provisioning ecosystem services derived from tree species used by the Kuikuro people in the Xingu Indigenous Territory. Specifically, we test the hypothesis that increasing fire frequency reduces the diversity and availability of tree species used for food, construction/handcrafts, and medicinal purposes, with stronger negative effects in forest environments than in Cerrado systems, reflecting differences in fire sensitivity and ecological resilience. Materials and methods 1. Study area We conducted this study in the Xingu Indigenous Territory (XIT), a 2.8 million-hectares area located in northern Mato Grosso state, Brazil. We focused on Afukuri village (12°59'S, 53°26'W), inhabited by the Kuikuro people, the most populous group in the Upper Xingu region (Franchetto and Heckenberger 2001) (Fig. 1). The region exhibits diverse vegetation physiognomies resulting from gradual variability in climate and soils, characterizing an ecological transition zone between the Amazonia and Savanna biomes (Marimon et al. 2006). Seasonal Evergreen Forest predominates (Ivanauskas et al. 2008), although typical savanna formations are also present (Sanches et al., 2012). The climate at the southern Amazonia frontier is classified as tropical savanna (Aw Köppen, Alvares et al. 2013), with mean temperatures above 18°C and a pronounced dry season from May to September. 2. Sampling 2.1 Vegetation sampling We conducted a forest inventory in 32 plots (10 × 10 m) distributed across two vegetation formations, with 20 plots located in forest areas and 12 in savanna areas. All plant individuals with diameter at breast height (DBH) ≥ 10 cm were recorded. For each individual, we measured DBH and identified species in the field, with botanical nomenclature verified according to REFLORA (Flora e Funga do Brasil 2024). Using a participatory approach, we interviewed a representative from Afukuri village who assisted with botanical identification, provided Indigenous names, described traditional uses of each species, and shared perceptions about changes in species abundance associated with fire occurrence. Additional information on traditional uses was obtained from case reports and video recordings. Each species was classified according to its relevance to the Kuikuro people into three use categories: (1) food, (2) construction/handcrafts, and (3) medicine. Species was assigned to more than one category when applicable. 2.2 Fire frequency mapping We mapped burned areas in the Xingu Indigenous Territory between 1985 and 2017 using data from the MapBiomas Fire platform (MapBiomas Project 2024). Sampling plot coordinates were overlaid with burned-area polygons to calculate fire frequency for each plot. The national-extent raster was cropped and masked using the crop and mask functions from the raster package (Hijmans et al. 2023) to the boundaries of the Xingu Indigenous Territory, restricting analyses to the study area. Sampling coordinates were converted into spatial objects for integration with fire data. 3. Data analysis 3.1 Phytosociological parameters We calculated phytosociological parameters to assess species abundance, dominance, frequency, and ecological importance in each environment. Importance Value (IV) was calculated following Curtis and McIntosh (1950) as the sum of relative abundance, relative dominance, and relative frequency. This index has been widely used to evaluate species' ecological importance based on population density (i.e., abundance), size (i.e., dominance), and spatial distribution (i.e., frequency). We used the fitoR function (Dalagnol et al. 2017) with the dplyr package (Wickham et al. 2023) to estimate phytosociological parameters of absolute and relative density, frequency, dominance, basal area, species richness, Shannon–Wiener diversity index, and Pielou evenness. Floristic similarity between forest and savanna environments was assessed using Jaccard and Sørensen indices, and we quantified the number of shared and exclusive species between environments. 3.2 Ecosystem service categories To visualize overlap among traditional use categories (food, construction/handcrafts, and medicine), we constructed Venn diagrams. Species classification was based primarily on interviews with traditional knowledge holders, complemented by ethnobotanical literature. 3.3 Relationship between species use and ecological importance We evaluated the relationship between species ecological importance and traditional use by correlating Importance Value (IV) with total species use, defined as the sum of all use categories attributed to each species. We fitted simple linear regression models (Galton 1886) and performed data manipulation using the tidyverse package (Wickham et al. 2019). We examined relationships separately for the three main use categories (food, construction/handcrafts, and medicine) within forest and savanna environments. This approach allowed us to identify patterns of species use in relation to ecological importance and biome type. 3.4 Effects of fire on provisioning ecosystem services To assess the effects of fire on provisioning ecosystem services, we fitted simple linear regression models (Galton 1886) relating fire frequency to the number of useful species, number of individuals, biomass, and mean total species use per plot. We also analyzed mean use for each category (food, construction/handcrafts, and medicine) as a function of fire frequency. Data integration and analysis were conducted using the devtools (Wickham et al. 2022), tidyverse, and ggally (Schloerke et al. 2024) packages. Analyses were performed for total species use and separately for each use category in forest and savanna environments, allowing us to evaluate how fire differentially influences species availability and traditional use patterns. In addition to univariate models, we used a multivariate constrained ordination approach to evaluate how fire frequency, vegetation type, and soil conditions jointly influenced the composition of useful species across plots. We performed Redundancy Analysis (RDA) using the vegan package (Oksanen et al. 2025), treating the abundance of useful species per plot (food, construction/handcraft, and medicinal categories) as the response matrix and environmental predictors (vegetation type, fire frequency, slope, clay content, soil bulk density, and soil pH) as explanatory variables. Prior to analysis, species abundance data were Hellinger-transformed using decostand(method = "hellinger") to reduce the influence of double zeros and to meet Euclidean distance assumptions. Continuous environmental variables were standardized (z-scores) to ensure comparability among predictors, and vegetation type was included as a categorical factor. Model significance was assessed using permutation tests (999 permutations) for the overall model, individual predictors, and constrained axes. This approach allowed us to quantify the relative contribution of fire and environmental gradients to variation in the abundance and composition of culturally important species. All spatial and statistical analyses were performed using R software version 4.3.1 (R Core Team 2025) and QGIS version 3.36 (QGIS Development Team 2025). All figures were generated using ggplot2 (Wickham 2016). Results Phytosociological parameters We recorded a total of 396 tree individuals belonging to 92 species, 56 genera, and 44 families across all sampled plots. Of these, 54 species occurred in the forest and 38 in the savanna environment (Table 1). Species diversity was similar between vegetation types, with slightly higher Shannon diversity in the savanna (H′ = 3.35) compared to the forest (H′ = 3.32). Species evenness was also higher in the savanna (J = 0.92) than in the forest (J = 0.83), indicating a more even distribution of individuals among species in savanna communities. In contrast, plots in the forest exhibited higher tree density and basal area compared to savanna plots. These results highlight the natural structural differences between vegetation types, with forests characterized by higher biomass and dominance, and savannas by greater compositional evenness. Tree species providing provisioning ecosystem services Of the 92 species recorded, 42 species (46%) provided at least one provisioning ecosystem service relevant to the Kuikuro people (Table 2). Most useful species (26 species; 62%) were associated with a single type of use (Fig. 2 ). Among these, 12 species were used exclusively for construction or handcrafts (e.g., Abuta grandifolia, Amaioua guianensis, Curatella americana ), 11 species were used exclusively for medicinal purposes (e.g., Aspidosperma macrocarpon, Bowdichia virgilioides, Dacryodes macrocarpa ), and three species were used exclusively for food ( Mouriri elliptica, Inga heterophylla, Caryocar brasiliense ). The remaining 16 species (38%) provided multiple ecosystem services. Four species were used for both construction/handcrafts and food (e.g., Cecropia palmata, Pseudolmedia macrophylla ), seven species combined medicinal, and construction/handcraft uses (e.g., Annona neoinsignis, Emmotum nitens, Qualea grandiflora ), and five species provided both food and medicinal resources (e.g., Byrsonima crassifolia, Byrsonima pachyphylla, Xylopia amazonica ). These multifunctional species represent an important component of the provisioning services supporting Kuikuro livelihoods. Relationship between species abundance and probability of use The relationship between species abundance and probability of use by the Kuikuro people differed markedly between vegetation types (Fig. 3 ; Table 3; Table S1 ). In savanna environments, species with higher IV tended to have higher probabilities of use, particularly for construction/handcrafts and medicinal purposes. This pattern indicates that, in savannas, commonly species contribute disproportionately to provisioning ecosystem services. In forest environments, however, the opposite pattern emerged. Species used for food, construction/handcrafts, and medicine were generally among the least abundant in the community. In these plots, lower IV values were associated with higher probabilities of use across all three categories, indicating that many culturally important species are relatively rare within tree community in the forest environment. This contrast suggests greater vulnerability of provisioning ecosystem services in forests, where useful species are less abundant and potentially more sensitive to disturbance. Effects of fire frequency on provisioning ecosystem services Provisioning ecosystem services were structured by vegetation type and environmental gradients, with fire frequency contributing to compositional differences among plots (Fig. 4 ; Table S2 ). Redundancy analysis indicated that vegetation type accounted for most of the variation in the abundance of useful species, while fire frequency and soil variables explained additional but comparatively smaller portions of variation. In forest environments, higher fire frequency was associated with lower relative abundance of several species used for construction/handcrafts and medicinal purposes. No positive associations between fire frequency and these use categories were detected in forests. In contrast, savanna environments exhibited a different pattern. Food-related species showed lower abundance with increasing fire frequency, whereas medicinal species tended to increase along the same gradient. Construction/handcraft species displayed weaker negative associations with fire frequency in savannas compared to forests. Overall, these patterns indicate that fire frequency is linked to shifts in the composition and relative representation of useful species within plots, but that responses vary strongly between forest and savanna systems. Vegetation type emerged as the primary driver of use-category structure, with fire and soil conditions contributing to secondary adjustments in species composition. These contrasting responses likely reflect differences in species turnover, fire tolerance, and environmental filtering across vegetation types. Discussion Synthesis of hypotheses and main findings This study assessed how fire frequency influences the provisioning ecosystem services supplied by tree species used by the Kuikuro people in the Xingu Indigenous Territory, with a focus on food, medicinal, and construction/handcraft uses. We hypothesized that increasing fire frequency would alter the richness and availability of culturally important tree species across both forest and savanna environments. Our results partially supported this hypothesis and revealed strong context dependency in fire effects. In forest environments, increasing fire frequency was associated with lower abundance and proportional representation of several species used for construction/handcrafts and medicinal purposes. These patterns suggest that forest-based provisioning ecosystem services may be sensitive to altered fire regimes, particularly where repeated burning leads to changes in species composition. In contrast, savanna environments exhibited weaker negative effects of fire on food- and construction-related species and a positive association between fire frequency and medicinal species. This divergence reflects fundamental ecological differences between forest and savanna systems, particularly in their evolutionary exposure and adaptive responses to fire. Because forests occupy most of the Xingu Indigenous Territory and are increasingly exposed to uncontrolled fires, our results suggest that species providing materials for construction, handcrafts, and medicine may become less abundant or be replaced by other taxa under future fire regimes. Such compositional shifts could influence the availability and reliability of culturally important resources, with potential consequences for Indigenous livelihoods and knowledge systems over time. Fire effects on species providing provisioning ecosystem services The negative associations between fire frequency and useful species observed in forest environments are consistent with the low fire tolerance of most Amazonian forest tree species. Forest vegetation lacks key fire-adaptive traits and experiences substantial structural degradation, increased tree mortality, and shifts in species composition following repeated fires (Cochrane and Schulze 1999 ; Barlow et al. 2003 ; Brando et al. 2014 ). These mechanisms likely contribute to the lower abundance of several useful species in forest plots and their stronger responses to fire frequency in this vegetation type. In contrast, savanna species exhibited greater resilience to fire. Many Cerrado tree species possess adaptive traits such as thick bark, underground storage organs, and strong resprouting capacity, which reduce mortality and allow rapid post-fire recovery (Coutinho 1990 ; Durigan and Ratter 2016 ; Chiminazzo et al. 2021 , 2023 ). These traits likely explain the weaker negative effects of fire on construction- and food-related species in savannas, as well as the positive association between fire frequency and medicinal species. Fire may promote the regeneration or increased availability of certain medicinal plants through resprouting or shifts in competitive dynamics, as documented in other savanna systems (Medeiros and Miranda 2008 ; Sartorelli et al. 2007 ; Souchie et al. 2017 ). Nevertheless, fire emerged as an important driver of community composition in both environments, indirectly influencing the distribution of useful species. While savannas display greater fire tolerance, shifts in fire frequency still altered the composition of useful species, particularly those associated with food production. These results highlight that even fire-adapted systems can experience shifts in functional composition under altered disturbance regimes. Multifunctionality and specialization of plant uses Our results show that forest and savanna environments contribute distinct and complementary sets of provisioning ecosystem services, reflecting limited overlap in tree species composition between vegetation types. Nearly 40% of the species used by the Kuikuro provided multiple ecosystem services, underscoring the multifunctional role of individual species in supporting Indigenous livelihoods. Species such as Emmotum nitens, Kielmeyera coriacea , and Byrsonima crassifolia exemplify this multifunctionality, supporting food security, healthcare, and material needs simultaneously (Alves et al. 2010 ; Nery et al. 2023 ). This multifunctionality amplifies the potential consequences of species loss: declines in particular species may affect multiple dimensions of well-being, from nutrition and health to housing and cultural practices. Conversely, species with exclusive uses—such as those restricted to medicinal or construction purposes—reflect cultural and ecological specialization and may be especially vulnerable to disturbance (Ticktin 2004 ; Cavallo 2018 ). Fire-induced declines of these specialized species may therefore lead to disproportionate impacts. The coexistence of multifunctional and specialized species highlights the importance of conserving not only species richness, but also the functional and cultural diversity embedded within Indigenous-managed landscapes. Implications of altered fire regimes for the Kuikuro people The intensification of uncontrolled fires in the Xingu region poses potential risks to the continuity of Kuikuro livelihoods and traditional land-use practices (Silvério et al. 2015 , 2022 ). Declines in certain tree species used for food and medicine may reduce local availability of these resources and increase reliance on substitutes or external sources (Schmidt et al. 2021 ). Additionally, fire-driven vegetation degradation may facilitate the spread of invasive species, further limiting native regeneration and access to culturally important plants (Silveira et al. 2013 ; Paritsis et al. 2015 ; Sampaio et al. 2021 ). Crucially, our findings reinforce the distinction between traditional Indigenous fire management and contemporary uncontrolled fires. Kuikuro fire practices are guided by ecological knowledge, cultural norms, and social regulation, resulting in small-scale, low intensity burns that promote heterogeneity and resource renewal (Pivello 2011 ; Eloy et al. 2021 ). In contrast, externally driven fires—often associated with agricultural expansion and extreme drought—are spatially extensive, more frequent, and ecologically disruptive, leading to cumulative degradation (Ribeiro and Assunção 2002 ; Ribeiro Filho et al. 2018 ; Shepard Jr et al. 2010 ). Recognizing and reinforcing this distinction is essential for developing conservation and fire management policies that protect biodiversity while respecting Indigenous governance systems. Integrating Indigenous fire knowledge with scientific understanding of fire–ecosystem dynamics offers a pathway to reduce fire impacts, sustain provisioning ecosystem services, and support the cultural integrity of the Kuikuro people. Conclusions This study demonstrates that increasing fire frequency alters species composition and the distribution of culturally important tree species, with potential implications for the provisioning ecosystem services relied upon by the Kuikuro people. Fire showed the strongest negative associations in forest environments, where several species used for construction/handcrafts and medicinal purposes declined with increasing fire frequency. In savannas, although many species exhibit fire adaptations, increased fire frequency was still associated with reduced availability of food- and construction-related species, indicating that even fire-adapted systems are vulnerable to altered fire regimes. Because forests dominate much of the XIT and are becoming increasingly exposed to uncontrolled fires, these findings highlight a disproportionate risk to culturally important tree species and suggest potential changes in the composition and availability of ecosystem services that sustain Indigenous livelihoods, health, and cultural practices. Changes in floristic composition driven by recurrent fire may therefore influence both biodiversity conservation outcomes and the resilience of Indigenous socioecological systems. From a policy perspective, our results underscore the need for fire management strategies that are biome-specific and that explicitly distinguish between traditional Indigenous fire practices and externally driven, uncontrolled fires. Public policies aimed at reducing fire impacts in Indigenous territories should recognize and strengthen Indigenous governance systems, support culturally grounded fire management, and prioritize the conservation of tree species that underpin provisioning ecosystem services. Integrating Indigenous knowledge with ecological monitoring and adaptive fire management is essential to safeguarding biodiversity, maintaining ecosystem services, and supporting the long-term autonomy and well-being of Indigenous peoples. Declarations Funding The authors declare that no funds, grants, or other support were received during the preparation of this manuscript. Competing Interests The authors have no relevant financial or non-financial interests to disclose. Author Contributions LGC: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Writing – original draft, Writing – review & editing; HFMO: Supervision, Validation, Writing – review & editing. LAGS: Formal analysis, Software, Validation, Writing – review & editing. LASC: Validation, Writing – review & editing. DVS: Conceptualization, Supervision, Project administration, Writing 1– review & editing. All authors read and approved the final manuscript. Data Availability The datasets generated and/or analyzed during the current study are available from the corresponding author on reasonable request. References Alvares CA, Stape JL, Sentelhas PC, Gonçalves JLM, Sparovek G (2013) Köppen's climate classification map for Brazil. Meteorol Z 22(6):711–728. 10.1127/0941-2948/2013/0507 Alves JJA, Rocha MSP, Souto WMS, Torquato SC, Portela RA (2010) O conhecimento ecológico tradicional no planejamento e na gestão ambiental. Ateliê Geográfico 4(2):44–60. https://doi.org/10.5216/ag.v4i2.9906 Barlow J, Lagan BO, Peres CA (2003) Morphological correlates of fire-induced tree mortality in a Central Amazonian forest. 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Revista Brasileira de Agroecologia 18(1):416–433. https://doi.org/10.33240/rba.v18i1.23690 Oksanen J, Simpson G, Blanchet F, Kindt R, Legendre P, Minchin P, O'Hara R, Solymos P, Stevens M, Szoecs E, Wagner H, Barbour M, Bedward M, Bolker B, Borcard D, Borman T, Carvalho G, Chirico M, De Caceres M, Durand S, Evangelista H, FitzJohn R, Friendly M, Furneaux B, Hannigan G, Hill M, Lahti L, Martino C, McGlinn D, Ouellette M, Ribeiro Cunha E, Smith T, Stier A, Ter Braak C, Weedon J (2025) vegan: Community Ecology Package_. 10.32614 /CRAN.package.vegan https://doi.org/10.32614/CRAN.package.vegan , R package version 2.7-2, . Paritsis J, Veblen TT, Holz A (2015) Positive fire feedbacks contribute to shifts from Nothofagus pumilio forests to fire-prone shrublands in Patagonia. J Veg Sci 26(1):89–101. https://doi.org/10.1111/jvs.12225 Pivello VR (2011) The use of fire in the Savanna and Amazonian Rainforests of Brazil: past and presente. Fire Ecol 7:24–39. https://doi.org/10.4996/fireecology.0701024 Povos Indígenas no Brasil (2024) Povos indígenas no Brasil. https://pib.socioambiental.org/ . Accessed 22 June 2024 QGIS Development Team (2025) QGIS Geographic Information System. Open Source Geospatial Foundation Project. https://qgis.org/ . Accessed 10 January 2025 R Core Team (2025) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. https://www.R-project.org/ . Accessed 14 May 2025 Ribeiro H, Assunção JV (2002) Efeitos das queimadas na saúde humana. Estudos Avançados 16(44):125–148. https://doi.org/10.1590/S0103-40142002000100008 Ribeiro Filho AA, Adams C, Manfredini S, Munari LC, Silva Junior JA, Ianovali D, Barbosa JM, Barreiros AM, Neves WA (2018) Dynamics of the soil fertility in quilombola shifting cultivation communities of the Atlantic Rainforest, Brazil. Boletim do Museu Paraense Emílio. Goeldi - Ciências Naturais 13(1):79–106. https://doi.org/10.46357/bcnaturais.v13i1.370 Sampaio AB, Ribeiro KT, Vieira DM, Silva DCB (2021) Guia de restauração ecológica para gestores de unidades de conservação: versão 1. Instituto Chico Mendes (ICMBio), Brasília Sanches RA, Rossete AN, Rezende ACP, Alves HQ, Villas-Bôas A (2012) Subsídios para a proteção de áreas úmidas da bacia do rio Xingu (Mato Grosso, Brasil). Revista Árvore 36(3):489–498. https://doi.org/10.1590/S0100-67622012000300011 Sartorelli PAR, Silva JMS, Gorenstein MR, Gomes JE, Ávila EQ (2007) Rebrota após fogo de espécies arbóreas de diferentes grupos fenológicos foliares em Savanna stricto sensu. Revista Científica Eletrônica de Engenharia Florestal 10 Schloerke B, Cook D, Larmarange J, Briatte F, Marbach M, Thoen E, Elberg A, Crowley J (2024) GGally: Extension to 'ggplot2'. R package version 2.2.1. https://CRAN.R-project.org/package=GGally/ . Accessed 12 August 2024 Schmidt MV, Ikpeng YU, Kayabi T, Sanches RA, Ono KY, Adams C (2021) Indigenous knowledge and forest succession management in the brazilian Amazon: contributions to reforestation of degraded areas. Front Forests Global Change 4:605925. https://doi.org/10.3389/ffgc.2021.605925 Schwartzman S, Boas AV, Ono KY, Fonseca MG, Doblas J, Zimmerman B, Junqueira P, Jerozolimski A, Salazar M, Junqueira RP, Torres M (2013) Philosophical Trans Royal Soc B 368(1619):20120164. https://doi.org/10.1098/rstb.2012.0164 Shepard GH Jr, Rummenhoeller K, Ohl-Schacherer J, Yu DW (2010) Trouble in paradise: Indigenous populations, anthropological policies, and biodiversity conservation in Manu National Park, Peru. J Sustainable Forestry 29(2–4):252–301. https://doi.org/10.1080/10549810903548153 Silveira ER, Melo ACG, Contiéri WA, Durigan G (2013) Controle de gramíneas exóticas em plantio de restauração do Savanna. In: Durigan G, Ramos VS (eds) Manejo adaptativo: primeiras experiências na restauração de ecossistemas, 1st edn. Páginas & Letras Editora e Gráfica, São Paulo, pp 1–7 Silvério DV, Brando PM, Macedo MN, Beck PSA, Bustamante M, Coe MT (2015) Agricultural expansion dominates climate changes in southeastern Amazonia: the overlooked non-GHG forcing. Environ Res Lett 10(10):104015. https://doi.org/10.1088/1748-9326/10/10/104015 Silvério DV, Oliveira RS, Flores BM, Brando PM, Almada HK, Furtado MT, Moreira FG, Heckenberger M, Ono KY, Macedo MN (2022) Intensification of fire regimes and forest loss in the Território Indígena do Xingu. Environ Res Lett 17(4):045012. https://doi.org/10.1088/1748-9326/ac5713 Souchie FF, Pinto JRR, Lenza E, Gomes L, Maracahipes-Santos L, Silvério DV (2017) Post-fire resprouting strategies of woody vegetation in the Brazilian savanna. Acta Bot Brasilica 31(2):260–266. https://doi.org/10.1590/0102-33062016abb0376 Ticktin (2004) The ecological implications of harvesting non-timber forest products. J Appl Ecol 41(1):11–21. https://doi.org/10.1111/j.1365-2664.2004.00859.x Wickham H (2016) ggplot2: Elegant graphics for data analysis. Springer, Cham. https://doi.org/10.1007/978-3-319-24277-4 Wickham H, Averick M, Bryan J, Chang W, McGowan LD, François R, Grolemund G, Hayes A, Henry L, Hester J, Kuhn M, Pedersen TL, Miller E, Bache SM, Müller K, Ooms J, Robinson D, Seidel DP, Spinu V, Takahashi K, Vaughan D, Wilke C, Woo K, Yutani H (2019) Welcome to the Tidyverse. J Open Source Softw 4(43):1686. https://doi.org/10.21105/joss.01686 Wickham H, François R, Henry L, Müller K, Vaughan D (2023) dplyr: A grammar of data manipulation. R package version 1.1.2. https://CRAN.R-project.org/package=dplyr/ . Accessed 27 November 2025 Wickham H, Hester J, Chang W, Bryan J (2022) devtools: Tools to make developing R packages easier. R package version 2.4.5. https://CRAN.R-project.org/package=devtools/ . Accessed 28 November 2025 WWF (2020) Savanna perde uma cidade de São Paulo a cada três meses. World Wide Fund for Nature. https://www.gov.br/fundaj/pt-br/destaques/observa-fundaj-itens/observa-fundaj/revitalizacao-de-bacias/savanna-perde-uma-cidade-de-sao-paulo-a-cada-tres-meses/ . Accessed 22 August 2024 Tables Tables are available in the Supplementary Files section. Table 3 is not available with this version Additional Declarations No competing interests reported. Supplementary Files tabelas.docx TableS2RDAresults.docx Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 15 Apr, 2026 Reviews received at journal 15 Apr, 2026 Reviews received at journal 14 Apr, 2026 Reviews received at journal 13 Apr, 2026 Reviews received at journal 10 Apr, 2026 Reviews received at journal 02 Apr, 2026 Reviewers agreed at journal 30 Mar, 2026 Reviewers agreed at journal 28 Mar, 2026 Reviewers agreed at journal 28 Mar, 2026 Reviewers agreed at journal 27 Mar, 2026 Reviewers agreed at journal 27 Mar, 2026 Reviewers agreed at journal 27 Mar, 2026 Reviewers agreed at journal 26 Mar, 2026 Reviewers agreed at journal 26 Mar, 2026 Reviewers agreed at journal 25 Mar, 2026 Reviewers agreed at journal 24 Mar, 2026 Reviewers agreed at journal 24 Mar, 2026 Reviewers invited by journal 24 Mar, 2026 Editor assigned by journal 20 Mar, 2026 Submission checks completed at journal 07 Mar, 2026 First submitted to journal 06 Mar, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9053133","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":611735862,"identity":"b694aa64-5144-4a0e-8f10-dfec0aaef639","order_by":0,"name":"Lui Gustavo Carvalho","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABCElEQVRIiWNgGAWjYBACAwbGBgiLnYeBIaHCRg7EPvCAKC3MQC0fzqQZg7Uk4NUCA0AtjDPbDieCTcCrRSK5+cOPmnuJ/c28Bx/zsB1Onx92+CHQFjs53QZcWhLbJHuOFSfOOMyXbMzDk5678XaaAVBLsrHZAdxamBnYEowZDvOYSfNIWOdunJ0A0nIgcRtuLc2fGf4lGMuDtRgwpxvOTv9ASEuDNGNbgpwBUIvkjATnBHnpHAK28Dxsk+ztS5AzPMxjbPDhQJrhBumcggMJBrj9Yt+e/vjDj28JPHLHewwfJP6zkZefnb75w4cKOzlcWrDYC1ZpQEAVCpBvIEX1KBgFo2AUjAQAAFTKXrXKEPZ7AAAAAElFTkSuQmCC","orcid":"","institution":"Mato Grosso State University","correspondingAuthor":true,"prefix":"","firstName":"Lui","middleName":"Gustavo","lastName":"Carvalho","suffix":""},{"id":611735865,"identity":"aab7c4a0-3af4-40b0-9dd3-fe15c5faf5cd","order_by":1,"name":"Hernani Fernandes Oliveira","email":"","orcid":"","institution":"University of Brasília","correspondingAuthor":false,"prefix":"","firstName":"Hernani","middleName":"Fernandes","lastName":"Oliveira","suffix":""},{"id":611735869,"identity":"7ef3b758-e55c-476d-8576-152438f2e213","order_by":2,"name":"Divino vicente Silvério","email":"","orcid":"","institution":"Federal Rural University of Amazonia","correspondingAuthor":false,"prefix":"","firstName":"Divino","middleName":"vicente","lastName":"Silvério","suffix":""},{"id":611735871,"identity":"c540c21a-ca0d-4413-a863-1c249a241bcf","order_by":3,"name":"Leonardo maracahipes Santos","email":"","orcid":"","institution":"Mato Grosso State University","correspondingAuthor":false,"prefix":"","firstName":"Leonardo","middleName":"maracahipes","lastName":"Santos","suffix":""},{"id":611735879,"identity":"5b45d829-989d-45e9-9c18-d1e567341bfc","order_by":4,"name":"Lorrayne aparecida Gonçalves","email":"","orcid":"","institution":"Federal Rural University of Amazonia","correspondingAuthor":false,"prefix":"","firstName":"Lorrayne","middleName":"aparecida","lastName":"Gonçalves","suffix":""},{"id":611735882,"identity":"0eb43744-499f-4f75-8c89-317821696fdc","order_by":5,"name":"Luiz Antônio Cardoso","email":"","orcid":"","institution":"Federal Rural University of Amazonia","correspondingAuthor":false,"prefix":"","firstName":"Luiz","middleName":"Antônio","lastName":"Cardoso","suffix":""},{"id":611735883,"identity":"d5be44f6-95ab-4fc2-b23b-f3ffe2c119c8","order_by":6,"name":"Marcus vinicius Schmidt","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Marcus","middleName":"vinicius","lastName":"Schmidt","suffix":""},{"id":611735887,"identity":"aedbc684-9a8b-4251-8acb-474e66622472","order_by":7,"name":"Airi Kuikuro","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Airi","middleName":"","lastName":"Kuikuro","suffix":""}],"badges":[],"createdAt":"2026-03-06 18:09:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9053133/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9053133/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105441925,"identity":"c08b0024-5800-4271-b99a-f5a0b64edbef","added_by":"auto","created_at":"2026-03-26 06:12:05","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4632941,"visible":true,"origin":"","legend":"\u003cp\u003eStudy area showing Afukuri village location and vegetation sampling plots in the Xingu Indigenous Territory, northern Mato Grosso, Brazil. Forest plots (n=20) and savanna plots (n=12) were established to assess tree species composition and their uses by the Kuikuro people.\u003c/p\u003e","description":"","filename":"Figure11.png","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/ce0f335f16ff9581a34dcf58.png"},{"id":105441897,"identity":"13d8d154-1eeb-4a69-8444-35cbbf4f3db3","added_by":"auto","created_at":"2026-03-26 06:11:56","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":218798,"visible":true,"origin":"","legend":"\u003cp\u003eVenn diagram representing the distribution of tree species according to their uses by Kuikuro people.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/d736095d72420258a2045fdd.png"},{"id":105441873,"identity":"a07b1021-91d3-465a-8042-d8c177532056","added_by":"auto","created_at":"2026-03-26 06:11:55","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":384064,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between proportion of total use of tree species and importance value index (IVI) in forest and savanna environments in the Xingu Indigenous Territory (Mato Grosso, Brazil).\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/5187d201526919071880820e.png"},{"id":105566415,"identity":"61b7789f-c254-413d-b460-36ab95965941","added_by":"auto","created_at":"2026-03-27 12:56:23","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":227467,"visible":true,"origin":"","legend":"\u003cp\u003eRDA ordination of useful tree species abundance in relation to vegetation type, fire frequency, and soil variables. Points represent plots (colored by vegetation type), arrows show environmental gradients, and labels indicate use categories. RDA1 explained most constrained variation and separated forest and savanna plots. Significance assessed by permutation tests (999 permutations).\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/5aeb483541ec1606196cf568.png"},{"id":105728030,"identity":"c066df43-80bc-4464-a1ea-50f69a01e134","added_by":"auto","created_at":"2026-03-30 11:08:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":6578057,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/5d0dbdf5-2b6c-427d-91a7-7f541135c517.pdf"},{"id":105441919,"identity":"7c077c8f-4003-47fd-991a-ae8270737bda","added_by":"auto","created_at":"2026-03-26 06:12:02","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":25944,"visible":true,"origin":"","legend":"","description":"","filename":"tabelas.docx","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/93126875ced0145157673f20.docx"},{"id":105441872,"identity":"72927233-b886-4618-8748-2a21141f8005","added_by":"auto","created_at":"2026-03-26 06:11:54","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":28403,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2RDAresults.docx","url":"https://assets-eu.researchsquare.com/files/rs-9053133/v1/69a140fd281e72b7b6812132.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Fire frequency restructures culturally important tree communities across forest and savanna landscapes in the Xingu Indigenous Territory","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIndigenous territories (ITs) play a crucial role in global biodiversity conservation, with Indigenous peoples managing approximately 28% of the Earth\u0026rsquo;s land surface while safeguarding nearly 80% of the world\u0026rsquo;s remaining biodiversity (Garnett et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In Brazil, around 208,000 Indigenous people inhabit the Amazonia biome, occupying approximately 387 Indigenous Territories that collectively represent about 23% of the Brazilian Amazon basin (FUNAI \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Povos Ind\u0026iacute;genas no Brasil \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the Cerrado, 109 Indigenous Territories cover 4.35% of the total area and play a fundamental role in conserving biodiversity and ecosystem processes in this highly threatened savanna system (WWF \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Within this context, the Xingu Indigenous Territory (XIT), located in the state of Mato Grosso, encompasses approximately 2.8\u0026nbsp;million hectares and harbors more than 16 Indigenous peoples with deep ecological knowledge and long-standing cultural connections to forest, savanna, and their natural resources (Sanches et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Povos Ind\u0026iacute;genas do Brasil 2024). Among these groups, the Kuikuro people are one of the largest populations and have historically relied on a diverse array of tree species for food, construction materials, handcrafts, and medicinal purposes. Their livelihoods and cultural practices are closely tied to the structure, composition, and regeneration dynamics of both forest and Cerrado ecosystems.\u003c/p\u003e \u003cp\u003eFire has long been an integral component of Indigenous land management in the XIT. The Kuikuro employ traditional fire management practices that are spatially controlled, seasonally timed, and focused on small areas, creating heterogeneous vegetation mosaics at different stages of regeneration. These practices enhance food production, maintain landscape diversity, and ensure the sustained availability of plant resources critical for subsistence and cultural continuity (Denevan \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Pivello \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Schwartzman et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Schmidt et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Importantly, this knowledge is transmitted across generations and embedded within cultural norms that promote ecosystem resilience and resource renewal. However, Indigenous territories across southern Amazonia and the Amazonia\u0026ndash;Cerrado transition are increasingly exposed to uncontrolled forest fires driven by climate change, prolonged droughts, and expanding agricultural frontiers (Cochrane \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Nepstad et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Brando et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These fires differ fundamentally from traditional burning practices: they are larger in extent, more frequent, and often more intense, leading to altered forest structure, increased tree mortality, and long-term changes in species composition (Brando et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Feng et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). As fire frequency increases, ecosystems become progressively more vulnerable, with cascading effects on regeneration processes and the persistence of fire-sensitive tree species. Such changes have direct implications for the provisioning ecosystem services upon which Indigenous peoples depend. Uncontrolled fires can reduce the availability of tree species used for construction, handcrafts, medicine, and food, thereby undermining food security, cultural practices, and traditional knowledge systems (Eloy et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These impacts are expected to be particularly pronounced in forest environments, where tree species are generally less adapted to frequent fire, while savanna systems may exhibit more variable responses due to long-term evolutionary exposure to fire.\u003c/p\u003e \u003cp\u003eDespite growing recognition of the importance of Indigenous fire management for conservation and food security, there remains limited quantitative evidence on how increasing fire frequency affects the diversity and availability of tree species that provide provisioning ecosystem services to Indigenous communities across contrasting vegetation types. Addressing this knowledge gap is critical for informing conservation policies and fire management strategies that integrate ecological processes with Indigenous knowledge and governance systems. In this study, we evaluate how fire frequency influences provisioning ecosystem services derived from tree species used by the Kuikuro people in the Xingu Indigenous Territory. Specifically, we test the hypothesis that increasing fire frequency reduces the diversity and availability of tree species used for food, construction/handcrafts, and medicinal purposes, with stronger negative effects in forest environments than in Cerrado systems, reflecting differences in fire sensitivity and ecological resilience.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e\u003cem\u003e1. Study area\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe conducted this study in the Xingu Indigenous Territory (XIT), a 2.8 million-hectares area located in northern Mato Grosso state, Brazil. We focused on Afukuri village (12°59'S, 53°26'W), inhabited by the Kuikuro people, the most populous group in the Upper Xingu region (Franchetto and Heckenberger 2001) (Fig. 1). The region exhibits diverse vegetation physiognomies resulting from gradual variability in climate and soils, characterizing an ecological transition zone between the Amazonia and Savanna biomes (Marimon et al. 2006). Seasonal Evergreen Forest predominates (Ivanauskas et al. 2008), although typical savanna formations are also present (Sanches et al., 2012). The climate at the southern Amazonia frontier is classified as tropical savanna (Aw Köppen, Alvares et al. 2013), with mean temperatures above 18°C and a pronounced dry season from May to September.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e2. Sampling\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e2.1 Vegetation sampling\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe conducted a forest inventory in 32 plots (10 × 10 m) distributed across two vegetation formations, with 20 plots located in forest areas and 12 in savanna areas. All plant individuals with diameter at breast height (DBH) ≥ 10 cm were recorded. For each individual, we measured DBH and identified species in the field, with botanical nomenclature verified according to REFLORA (Flora e Funga do Brasil 2024). Using a participatory approach, we interviewed a representative from Afukuri village who assisted with botanical identification, provided Indigenous names, described traditional uses of each species, and shared perceptions about changes in species abundance associated with fire occurrence. Additional information on traditional uses was obtained from case reports and video recordings.\u003c/p\u003e\n\u003cp\u003eEach species was classified according to its relevance to the Kuikuro people into three use categories: (1) food, (2) construction/handcrafts, and (3) medicine. Species was assigned to more than one category when applicable.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e2.2 Fire frequency mapping\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe mapped burned areas in the Xingu Indigenous Territory between 1985 and 2017 using data from the MapBiomas Fire platform (MapBiomas Project 2024). Sampling plot coordinates were overlaid with burned-area polygons to calculate fire frequency for each plot. The national-extent raster was cropped and masked using the \u003cem\u003ecrop\u003c/em\u003e and \u003cem\u003emask\u003c/em\u003e functions from the raster package (Hijmans et al. 2023) to the boundaries of the Xingu Indigenous Territory, restricting analyses to the study area. Sampling coordinates were converted into spatial objects for integration with fire data.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3. Data analysis\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.1 Phytosociological parameters\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe calculated phytosociological parameters to assess species abundance, dominance, frequency, and ecological importance in each environment. Importance Value (IV) was calculated following Curtis and McIntosh (1950) as the sum of relative abundance, relative dominance, and relative frequency. This index has been widely used to evaluate species' ecological importance based on population density (i.e., abundance), size (i.e., dominance), and spatial distribution (i.e., frequency). We used the \u003cem\u003efitoR\u003c/em\u003e function (Dalagnol et al. 2017) with the dplyr package (Wickham et al. 2023) to estimate phytosociological parameters of absolute and relative density, frequency, dominance, basal area, species richness, Shannon–Wiener diversity index, and Pielou evenness. Floristic similarity between forest and savanna environments was assessed using Jaccard and Sørensen indices, and we quantified the number of shared and exclusive species between environments.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.2 Ecosystem service categories\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTo visualize overlap among traditional use categories (food, construction/handcrafts, and medicine), we constructed Venn diagrams. Species classification was based primarily on interviews with traditional knowledge holders, complemented by ethnobotanical literature.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.3 Relationship between species use and ecological importance\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe evaluated the relationship between species ecological importance and traditional use by correlating Importance Value (IV) with total species use, defined as the sum of all use categories attributed to each species. We fitted simple linear regression models (Galton 1886) and performed data manipulation using the tidyverse package (Wickham et al. 2019). We examined relationships separately for the three main use categories (food, construction/handcrafts, and medicine) within forest and savanna environments. This approach allowed us to identify patterns of species use in relation to ecological importance and biome type.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.4 Effects of fire on provisioning ecosystem services\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eTo assess the effects of fire on provisioning ecosystem services, we fitted simple linear regression models (Galton 1886) relating fire frequency to the number of useful species, number of individuals, biomass, and mean total species use per plot. We also analyzed mean use for each category (food, construction/handcrafts, and medicine) as a function of fire frequency. Data integration and analysis were conducted using the devtools (Wickham et al. 2022), tidyverse, and ggally (Schloerke et al. 2024) packages. Analyses were performed for total species use and separately for each use category in forest and savanna environments, allowing us to evaluate how fire differentially influences species availability and traditional use patterns. \u003c/p\u003e\n\u003cp\u003eIn addition to univariate models, we used a multivariate constrained ordination approach to evaluate how fire frequency, vegetation type, and soil conditions jointly influenced the composition of useful species across plots. We performed Redundancy Analysis (RDA) using the vegan package (Oksanen et al. 2025), treating the abundance of useful species per plot (food, construction/handcraft, and medicinal categories) as the response matrix and environmental predictors (vegetation type, fire frequency, slope, clay content, soil bulk density, and soil pH) as explanatory variables. Prior to analysis, species abundance data were Hellinger-transformed using \u003cem\u003edecostand(method = \"hellinger\")\u003c/em\u003e to reduce the influence of double zeros and to meet Euclidean distance assumptions. Continuous environmental variables were standardized (z-scores) to ensure comparability among predictors, and vegetation type was included as a categorical factor. Model significance was assessed using permutation tests (999 permutations) for the overall model, individual predictors, and constrained axes. This approach allowed us to quantify the relative contribution of fire and environmental gradients to variation in the abundance and composition of culturally important species. All spatial and statistical analyses were performed using R software version 4.3.1 (R Core Team 2025) and QGIS version 3.36 (QGIS Development Team 2025). All figures were generated using ggplot2 (Wickham 2016).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003ePhytosociological parameters\u003c/h2\u003e \u003cp\u003eWe recorded a total of 396 tree individuals belonging to 92 species, 56 genera, and 44 families across all sampled plots. Of these, 54 species occurred in the forest and 38 in the savanna environment (Table\u0026nbsp;1). Species diversity was similar between vegetation types, with slightly higher Shannon diversity in the savanna (H\u0026prime; = 3.35) compared to the forest (H\u0026prime; = 3.32). Species evenness was also higher in the savanna (J\u0026thinsp;=\u0026thinsp;0.92) than in the forest (J\u0026thinsp;=\u0026thinsp;0.83), indicating a more even distribution of individuals among species in savanna communities. In contrast, plots in the forest exhibited higher tree density and basal area compared to savanna plots. These results highlight the natural structural differences between vegetation types, with forests characterized by higher biomass and dominance, and savannas by greater compositional evenness.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eTree species providing provisioning ecosystem services\u003c/h3\u003e\n\u003cp\u003eOf the 92 species recorded, 42 species (46%) provided at least one provisioning ecosystem service relevant to the Kuikuro people (Table\u0026nbsp;2). Most useful species (26 species; 62%) were associated with a single type of use (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Among these, 12 species were used exclusively for construction or handcrafts (e.g., \u003cem\u003eAbuta grandifolia, Amaioua guianensis, Curatella americana\u003c/em\u003e), 11 species were used exclusively for medicinal purposes (e.g., \u003cem\u003eAspidosperma macrocarpon, Bowdichia virgilioides, Dacryodes macrocarpa\u003c/em\u003e), and three species were used exclusively for food (\u003cem\u003eMouriri elliptica, Inga heterophylla, Caryocar brasiliense\u003c/em\u003e). The remaining 16 species (38%) provided multiple ecosystem services. Four species were used for both construction/handcrafts and food (e.g., \u003cem\u003eCecropia palmata, Pseudolmedia macrophylla\u003c/em\u003e), seven species combined medicinal, and construction/handcraft uses (e.g., \u003cem\u003eAnnona neoinsignis, Emmotum nitens, Qualea grandiflora\u003c/em\u003e), and five species provided both food and medicinal resources (e.g., \u003cem\u003eByrsonima crassifolia, Byrsonima pachyphylla, Xylopia amazonica\u003c/em\u003e). These multifunctional species represent an important component of the provisioning services supporting Kuikuro livelihoods.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eRelationship between species abundance and probability of use\u003c/h3\u003e\n\u003cp\u003eThe relationship between species abundance and probability of use by the Kuikuro people differed markedly between vegetation types (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Table\u0026nbsp;3; Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). In savanna environments, species with higher IV tended to have higher probabilities of use, particularly for construction/handcrafts and medicinal purposes. This pattern indicates that, in savannas, commonly species contribute disproportionately to provisioning ecosystem services. In forest environments, however, the opposite pattern emerged. Species used for food, construction/handcrafts, and medicine were generally among the least abundant in the community. In these plots, lower IV values were associated with higher probabilities of use across all three categories, indicating that many culturally important species are relatively rare within tree community in the forest environment. This contrast suggests greater vulnerability of provisioning ecosystem services in forests, where useful species are less abundant and potentially more sensitive to disturbance.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eEffects of fire frequency on provisioning ecosystem services\u003c/h3\u003e\n\u003cp\u003eProvisioning ecosystem services were structured by vegetation type and environmental gradients, with fire frequency contributing to compositional differences among plots (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e). Redundancy analysis indicated that vegetation type accounted for most of the variation in the abundance of useful species, while fire frequency and soil variables explained additional but comparatively smaller portions of variation. In forest environments, higher fire frequency was associated with lower relative abundance of several species used for construction/handcrafts and medicinal purposes. No positive associations between fire frequency and these use categories were detected in forests. In contrast, savanna environments exhibited a different pattern. Food-related species showed lower abundance with increasing fire frequency, whereas medicinal species tended to increase along the same gradient. Construction/handcraft species displayed weaker negative associations with fire frequency in savannas compared to forests.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eOverall, these patterns indicate that fire frequency is linked to shifts in the composition and relative representation of useful species within plots, but that responses vary strongly between forest and savanna systems. Vegetation type emerged as the primary driver of use-category structure, with fire and soil conditions contributing to secondary adjustments in species composition. These contrasting responses likely reflect differences in species turnover, fire tolerance, and environmental filtering across vegetation types.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eSynthesis of hypotheses and main findings\u003c/h2\u003e \u003cp\u003eThis study assessed how fire frequency influences the provisioning ecosystem services supplied by tree species used by the Kuikuro people in the Xingu Indigenous Territory, with a focus on food, medicinal, and construction/handcraft uses. We hypothesized that increasing fire frequency would alter the richness and availability of culturally important tree species across both forest and savanna environments. Our results partially supported this hypothesis and revealed strong context dependency in fire effects.\u003c/p\u003e \u003cp\u003eIn forest environments, increasing fire frequency was associated with lower abundance and proportional representation of several species used for construction/handcrafts and medicinal purposes. These patterns suggest that forest-based provisioning ecosystem services may be sensitive to altered fire regimes, particularly where repeated burning leads to changes in species composition. In contrast, savanna environments exhibited weaker negative effects of fire on food- and construction-related species and a positive association between fire frequency and medicinal species. This divergence reflects fundamental ecological differences between forest and savanna systems, particularly in their evolutionary exposure and adaptive responses to fire.\u003c/p\u003e \u003cp\u003eBecause forests occupy most of the Xingu Indigenous Territory and are increasingly exposed to uncontrolled fires, our results suggest that species providing materials for construction, handcrafts, and medicine may become less abundant or be replaced by other taxa under future fire regimes. Such compositional shifts could influence the availability and reliability of culturally important resources, with potential consequences for Indigenous livelihoods and knowledge systems over time.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eFire effects on species providing provisioning ecosystem services\u003c/h3\u003e\n\u003cp\u003eThe negative associations between fire frequency and useful species observed in forest environments are consistent with the low fire tolerance of most Amazonian forest tree species. Forest vegetation lacks key fire-adaptive traits and experiences substantial structural degradation, increased tree mortality, and shifts in species composition following repeated fires (Cochrane and Schulze \u003cspan class=\"CitationRef\"\u003e1999\u003c/span\u003e; Barlow et al. \u003cspan class=\"CitationRef\"\u003e2003\u003c/span\u003e; Brando et al. \u003cspan class=\"CitationRef\"\u003e2014\u003c/span\u003e). These mechanisms likely contribute to the lower abundance of several useful species in forest plots and their stronger responses to fire frequency in this vegetation type.\u003c/p\u003e \u003cp\u003eIn contrast, savanna species exhibited greater resilience to fire. Many Cerrado tree species possess adaptive traits such as thick bark, underground storage organs, and strong resprouting capacity, which reduce mortality and allow rapid post-fire recovery (Coutinho \u003cspan class=\"CitationRef\"\u003e1990\u003c/span\u003e; Durigan and Ratter \u003cspan class=\"CitationRef\"\u003e2016\u003c/span\u003e; Chiminazzo et al. \u003cspan class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e2023\u003c/span\u003e). These traits likely explain the weaker negative effects of fire on construction- and food-related species in savannas, as well as the positive association between fire frequency and medicinal species. Fire may promote the regeneration or increased availability of certain medicinal plants through resprouting or shifts in competitive dynamics, as documented in other savanna systems (Medeiros and Miranda \u003cspan class=\"CitationRef\"\u003e2008\u003c/span\u003e; Sartorelli et al. \u003cspan class=\"CitationRef\"\u003e2007\u003c/span\u003e; Souchie et al. \u003cspan class=\"CitationRef\"\u003e2017\u003c/span\u003e). Nevertheless, fire emerged as an important driver of community composition in both environments, indirectly influencing the distribution of useful species. While savannas display greater fire tolerance, shifts in fire frequency still altered the composition of useful species, particularly those associated with food production. These results highlight that even fire-adapted systems can experience shifts in functional composition under altered disturbance regimes.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eMultifunctionality and specialization of plant uses\u003c/h2\u003e \u003cp\u003eOur results show that forest and savanna environments contribute distinct and complementary sets of provisioning ecosystem services, reflecting limited overlap in tree species composition between vegetation types. Nearly 40% of the species used by the Kuikuro provided multiple ecosystem services, underscoring the multifunctional role of individual species in supporting Indigenous livelihoods. Species such as \u003cem\u003eEmmotum nitens, Kielmeyera coriacea\u003c/em\u003e, and \u003cem\u003eByrsonima crassifolia\u003c/em\u003e exemplify this multifunctionality, supporting food security, healthcare, and material needs simultaneously (Alves et al. \u003cspan class=\"CitationRef\"\u003e2010\u003c/span\u003e; Nery et al. \u003cspan class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis multifunctionality amplifies the potential consequences of species loss: declines in particular species may affect multiple dimensions of well-being, from nutrition and health to housing and cultural practices. Conversely, species with exclusive uses—such as those restricted to medicinal or construction purposes—reflect cultural and ecological specialization and may be especially vulnerable to disturbance (Ticktin \u003cspan class=\"CitationRef\"\u003e2004\u003c/span\u003e; Cavallo \u003cspan class=\"CitationRef\"\u003e2018\u003c/span\u003e). Fire-induced declines of these specialized species may therefore lead to disproportionate impacts. The coexistence of multifunctional and specialized species highlights the importance of conserving not only species richness, but also the functional and cultural diversity embedded within Indigenous-managed landscapes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eImplications of altered fire regimes for the Kuikuro people\u003c/h2\u003e \u003cp\u003eThe intensification of uncontrolled fires in the Xingu region poses potential risks to the continuity of Kuikuro livelihoods and traditional land-use practices (Silvério et al. \u003cspan class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e2022\u003c/span\u003e). Declines in certain tree species used for food and medicine may reduce local availability of these resources and increase reliance on substitutes or external sources (Schmidt et al. \u003cspan class=\"CitationRef\"\u003e2021\u003c/span\u003e). Additionally, fire-driven vegetation degradation may facilitate the spread of invasive species, further limiting native regeneration and access to culturally important plants (Silveira et al. \u003cspan class=\"CitationRef\"\u003e2013\u003c/span\u003e; Paritsis et al. \u003cspan class=\"CitationRef\"\u003e2015\u003c/span\u003e; Sampaio et al. \u003cspan class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCrucially, our findings reinforce the distinction between traditional Indigenous fire management and contemporary uncontrolled fires. Kuikuro fire practices are guided by ecological knowledge, cultural norms, and social regulation, resulting in small-scale, low intensity burns that promote heterogeneity and resource renewal (Pivello \u003cspan class=\"CitationRef\"\u003e2011\u003c/span\u003e; Eloy et al. \u003cspan class=\"CitationRef\"\u003e2021\u003c/span\u003e). In contrast, externally driven fires—often associated with agricultural expansion and extreme drought—are spatially extensive, more frequent, and ecologically disruptive, leading to cumulative degradation (Ribeiro and Assunção \u003cspan class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ribeiro Filho et al. \u003cspan class=\"CitationRef\"\u003e2018\u003c/span\u003e; Shepard Jr et al. \u003cspan class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRecognizing and reinforcing this distinction is essential for developing conservation and fire management policies that protect biodiversity while respecting Indigenous governance systems. Integrating Indigenous fire knowledge with scientific understanding of fire–ecosystem dynamics offers a pathway to reduce fire impacts, sustain provisioning ecosystem services, and support the cultural integrity of the Kuikuro people.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThis study demonstrates that increasing fire frequency alters species composition and the distribution of culturally important tree species, with potential implications for the provisioning ecosystem services relied upon by the Kuikuro people. Fire showed the strongest negative associations in forest environments, where several species used for construction/handcrafts and medicinal purposes declined with increasing fire frequency. In savannas, although many species exhibit fire adaptations, increased fire frequency was still associated with reduced availability of food- and construction-related species, indicating that even fire-adapted systems are vulnerable to altered fire regimes. Because forests dominate much of the XIT and are becoming increasingly exposed to uncontrolled fires, these findings highlight a disproportionate risk to culturally important tree species and suggest potential changes in the composition and availability of ecosystem services that sustain Indigenous livelihoods, health, and cultural practices. Changes in floristic composition driven by recurrent fire may therefore influence both biodiversity conservation outcomes and the resilience of Indigenous socioecological systems.\u003c/p\u003e\u003cp\u003eFrom a policy perspective, our results underscore the need for fire management strategies that are biome-specific and that explicitly distinguish between traditional Indigenous fire practices and externally driven, uncontrolled fires. Public policies aimed at reducing fire impacts in Indigenous territories should recognize and strengthen Indigenous governance systems, support culturally grounded fire management, and prioritize the conservation of tree species that underpin provisioning ecosystem services. Integrating Indigenous knowledge with ecological monitoring and adaptive fire management is essential to safeguarding biodiversity, maintaining ecosystem services, and supporting the long-term autonomy and well-being of Indigenous peoples.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that no funds, grants, or other support were received during the preparation of this manuscript.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eCompeting Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eLGC: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Writing \u0026ndash; original draft, Writing \u0026ndash; review \u0026amp; editing; HFMO: Supervision, Validation, Writing \u0026ndash; review \u0026amp; editing. LAGS: Formal analysis, Software, Validation, Writing \u0026ndash; review \u0026amp; editing. LASC: Validation, Writing \u0026ndash; review \u0026amp; editing. DVS: Conceptualization, Supervision, Project administration, Writing 1\u0026ndash; review \u0026amp; editing. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eData Availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated and/or analyzed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAlvares CA, Stape JL, Sentelhas PC, Gon\u0026ccedil;alves JLM, Sparovek G (2013) K\u0026ouml;ppen's climate classification map for Brazil. 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Accessed 22 August 2024\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables are available in the Supplementary Files section.\u003c/p\u003e\n\u003cp\u003eTable 3 is not available with this version\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biodiversity-and-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bioc","sideBox":"Learn more about [Biodiversity and Conservation](https://www.springer.com/journal/10531)","snPcode":"10531","submissionUrl":"https://submission.nature.com/new-submission/10531/3","title":"Biodiversity and Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"ethnobotany, floristic composition, provisioning services, Amazon-Savanna ecotone, protected areas","lastPublishedDoi":"10.21203/rs.3.rs-9053133/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9053133/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eFire has become an increasingly pervasive disturbance in southern Amazonia, and climate projections indicate that fire frequency is likely to intensify in the coming decades. These changes pose a growing threat to forest biodiversity and to the ecosystem services that sustain Indigenous peoples. Here, we evaluate how fire frequency influences the provision of ecosystem services derived from tree species used by the Kuikuro people in the Xingu Indigenous Territory (XIT), Brazil. We conducted forest inventories across a fire-history gradient ranging from zero to six burn events over the past 40 years, sampling 32 plots (10 \u0026times; 10 m) distributed across forest (n\u0026thinsp;=\u0026thinsp;20) and savanna (n\u0026thinsp;=\u0026thinsp;12) vegetation types. For all trees with diameter\u0026thinsp;\u0026ge;\u0026thinsp;10 cm, we measured diameter, and recorded ethnobotanical uses reported by the Kuikuro people, grouped into three categories, (1) food, (2) construction/handcrafts, and (3) medicinal use. We also incorporated environmental predictors derived from remote sensing and soil data, including altitude, slope, soil pH, clay content, and bulk density. Across all plots, we recorded 92 tree species (54 in forest and 38 in savanna). In forest environments, 6.3% of species were used for food, 9.1% for medicinal purposes, and 17.3% for construction or handcrafts. In savanna, these proportions were 5.4%, 11.8%, and 3.6%, respectively. Regression analyses revealed that increasing fire frequency had a strong negative effect on the abundance of tree species used for construction/handcrafts and medicinal purposes in forest areas. In contrast, in savanna environments, fire frequency showed weaker negative effects on food- and construction-related species and a positive association with medicinal species. Because forests dominate much of the XIT and are becoming increasingly exposed to fire, our findings indicate that tree species providing materials for construction, handcrafts, and medicine are particularly vulnerable to altered fire regimes. These changes threaten not only forest biodiversity, but also the livelihoods, traditional knowledge, and cultural integrity of the Kuikuro people, underscoring the need for fire-sensitive management strategies that integrate ecological and Indigenous perspectives.\u003c/p\u003e","manuscriptTitle":"Fire frequency restructures culturally important tree communities across forest and savanna landscapes in the Xingu Indigenous Territory","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-26 06:11:22","doi":"10.21203/rs.3.rs-9053133/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-15T08:05:37+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-15T05:57:50+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-14T20:22:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-13T20:10:31+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-10T09:55:54+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-02T10:58:20+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"150769151542194085438449960454256818711","date":"2026-03-30T19:46:36+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"242359346406501239190875636062638454952","date":"2026-03-28T15:26:29+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"144820908891586113402533134330167099523","date":"2026-03-28T06:21:40+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"57043656784880469557271743484075745514","date":"2026-03-27T11:13:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"170161634718667769244298888090797378559","date":"2026-03-27T10:23:13+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"83732849932344854767022384239911377944","date":"2026-03-27T09:02:44+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"9074987614453447733342109003627151149","date":"2026-03-26T20:40:56+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"308480857616373130526577689497227900730","date":"2026-03-26T09:49:05+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"281881968340697774093151614665799160784","date":"2026-03-26T02:43:44+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"268610386259366236676942772074115022129","date":"2026-03-24T09:26:16+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"100210545921654926217855944220387138957","date":"2026-03-24T06:07:22+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-24T04:03:19+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-03-20T10:53:34+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-03-07T11:04:59+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biodiversity and Conservation","date":"2026-03-06T17:54:57+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biodiversity-and-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bioc","sideBox":"Learn more about [Biodiversity and Conservation](https://www.springer.com/journal/10531)","snPcode":"10531","submissionUrl":"https://submission.nature.com/new-submission/10531/3","title":"Biodiversity and Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"03db15e3-8a30-4aa2-860e-b1a13fda10df","owner":[],"postedDate":"March 26th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-14T21:38:06+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-26 06:11:22","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9053133","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9053133","identity":"rs-9053133","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}