Abstract:

The National Forest Inventory (Inventário Florestal Nacional-IFN) is a large initiative that uses standardised methods to survey Brazilian forestry resources. One target of the IFN is the Cerrado, which contains one of the richest floras in the world. The aim of this study was to assess the contribution of the IFN to the knowledge of Cerrado woody flora. We analysed data from field-collected vouchers sampled by the IFN Cerrado. We restricted our analyses to IFN collections of native trees and shrubs, including palms, which were identified at the species level. Habitat of each collection was obtained by overlaying specimens’ geographic coordinates with land cover maps available in the Mapbiomas platform. Our final dataset comprised 28,602 specimens distributed in 2,779 sites (conglomerates) in Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Piauí, São Paulo and Tocantins. Collections were located in the following habitats: savannas (40.5%), forests (30.2%), anthropic areas (25.6%), grasslands (3.5%), and water (0.2%). We recorded 1,822 species belonging to 543 genera and 105 families, representing 34% of Cerrado woody species recorded on Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... . Fabaceae had the largest number of species, while Tapirira guianensis and Matayba guianensis were the most collected species. We highlight 60 potentially new records of occurrence for several states and 64 new records for the Cerrado, primarily in riparian forests where species from other biomes occur. In addition, 232 recorded species are Cerrado endemics, while 36 are cited in the CNCFlora’s red list as endangered. The systematic sampling carried out by the IFN enabled vegetation sampling in remote and poorly known areas, which expanded the geographic range of many woody species and contributed to the knowledge of plant diversity in the Cerrado.

Keywords:
conservation; endemism; plant diversity; sampling effort; survey; vegetation types

Resumo:

O Inventário Florestal Nacional (IFN) é uma ampla iniciativa que emprega métodos padronizados para inventariar recursos florestais brasileiros. Um dos alvos do IFN é o Cerrado, o qual possui uma das floras mais ricas do mundo. O objetivo deste estudo foi avaliar a contribuição do IFN para o conhecimento da flora lenhosa do Cerrado. Nós analisamos dados de vouchers coletados em campo pelo IFN Cerrado. Nós restringimos nossas análises a coletas do IFN pertencentes a árvores e arbustos, incluindo palmeiras, identificadas ao nível de espécie. O habitat de cada coleta foi obtido pela intersecção entre as coordenadas geográficas dos espécimes com mapas de cobertura disponíveis na plataforma Mapbiomas. O conjunto final de dados foi composto por 28.602 coletas distribuídas em 2.779 sítios (conglomerados) localizados na Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Piauí, São Paulo e Tocantins. As coletas foram realizadas nos seguintes habitats: savanas (40,5%), florestas (30,2%), áreas antrópicas (25,6%), campos (3,5%) e água (0,2%). Ao todo foram registradas 1.822 espécies pertencentes a 543 gêneros e 105 famílias, representando 34% das espécies lenhosas do Cerrado registradas na Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... . Fabaceae apresentou o maior número de espécies, enquanto que Tapirira guianensis e Matayba guianensis foram as espécies mais coletadas. Destacam-se possíveis novos registros de ocorrência de 60 espécies para diversos estados e de 64 espécies para o Cerrado, predominantemente nas florestas ripárias onde geralmente ocorrem espécies de outros biomas. Além disso, foram registradas 232 espécies endêmicas do Cerrado, bem como 36 espécies citadas na lista vermelha do CNCFlora como ameaçadas. A amostragem sistemática realizada pelo IFN permitiu o inventário da vegetação em áreas remotas e pouco coletadas, permitindo a expansão da distribuição geográfica de diversas espécies lenhosas, e contribuindo para o conhecimento da diversidade vegetal no Cerrado.

Palavras-chave:
conservação; endemismo; diversidade vegetal; esforço amostral; inventário; tipos de vegetação

Introduction

Brazil harbours one of the richest floras in the world, and it is home to a large number of endemics (BFG 2021BFG - The Brazil Flora Group. 2021. Leveraging the power of a collaborative scientific network. Taxon https://doi.org/10.1002/tax.12640
https://doi.org/10.1002/tax.12640... ). Recent advances took place after the compilation of an updated national checklist containing information on species descriptions and geographic distributions (Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... ; http://floradobrasil.jbrj.gov.br) However, large gaps still exist in the documentation of plant diversity in Brazil. Such knowledge gaps are related to incomplete taxonomic knowledge (“Linnean shortfall”; Whittaker et al. 2005WHITTAKER, R.J., ARAÚJO, M.B., JEPSON, P., LADLE, R.J., WATSON, J.E.M. & WILLIS, K.J. 2005. Conservation Biogeography: assessment and prospect. Diversity Distrib 11:3-23. https://doi.org/10.1111/j.1366-9516.2005.00143.x
https://doi.org/10.1111/j.1366-9516.2005... ), as represented by a high number of new species being described every year in the country (BFG 2021BFG - The Brazil Flora Group. 2021. Leveraging the power of a collaborative scientific network. Taxon https://doi.org/10.1002/tax.12640
https://doi.org/10.1002/tax.12640... ). These gaps are also a product of unsatisfactory understanding of the geographic distribution of species (“Wallacean shortfall”), particularly in poorly sampled regions of the country (e.g., Sousa-Baena et al. 2014SOUSA-BAENA, M.S., GARCIA, L.C. & PETERSON, A.T. 2014. Completeness of digital accessible knowledge of the plants of Brazil and priorities for survey and inventory. Diversity Distrib 20:369-381. https://doi.org/10.1111/ddi.12136
https://doi.org/10.1111/ddi.12136... , Oliveira et al. 2016OLIVEIRA, U., PAGLIA, A.P., BRESCOVIT, A.D. & et al. 2016. The strong influence of collection bias on biodiversity knowledge shortfalls of Brazilian terrestrial biodiversity. Divers Distrib 22 (12): 1232-1244. https://doi.org/10.1111/ddi.12489
https://doi.org/10.1111/ddi.12489... ).

Since 2007, the Brazilian Forest Service (Serviço Florestal Brasileiro-SFB) has been coordinating a survey of Brazilian forest resources through the National Forest Inventory (Inventário Florestal Nacional-IFN). The IFN aims to provide data about forest structure, composition, vitality, biomass, wood and carbon stock (SFB 2020SFB - Serviço Florestal Brasileiro. 2020. Inventário Florestal Nacional. Available from: http://www.florestal.gov.br/inventario-florestal-nacional (accessed: 17 April 2020).
http://www.florestal.gov.br/inventario-f... ). This initiative seeks to support development policies and assist in the identification of strategies and opportunities for sustainable use, restoration and conservation of forest resources (SFB 2019SFB - Serviço Florestal Brasileiro. 2019. Boletim do IFN Cerrado: levantamento botânico, 1ª ed., Serviço Florestal Brasileiro, Brasília., 2020SFB - Serviço Florestal Brasileiro. 2020. Inventário Florestal Nacional. Available from: http://www.florestal.gov.br/inventario-florestal-nacional (accessed: 17 April 2020).
http://www.florestal.gov.br/inventario-f... ). The IFN adopts a standardised sampling method applied to each Brazilian biome (Amazon, Atlantic Forest, Caatinga, Cerrado, Pantanal and Pampa; SFB 2017aSFB - Serviço Florestal Brasileiro. 2017a. Manual de campo: procedimentos para coleta de dados biofísicos e socioambientais. Serviço Florestal Brasileiro, Brasília.). IFN systematic sampling generates a vast amount of scientific data, including thousands of plant occurrence records widely distributed across the country, mostly trees and shrubs. The compilation and analysis of such dataset would be a useful source of biodiversity information on Brazilian biomes that would help to increase floristic knowledge and support conservation planning. Although summary reports for some states have been published (https://www.florestal.gov.br/resultados) comprehensive analyses, including the large volume of plant diversity data sampled by the IFN surveys, are still missing for most states (but see Versieux et al. 2017VERSIEUX, L.M., DÁVILA, N., DELGADO, G.C., DE SOUSA, V.F., DE MOURA, E.O., FILGUEIRAS, T., ALVES, M.V., CARVALHO, E., PIOTTO, D., FORZZA, R.C., CALVENTE, A. & JARDIM, J.G. 2017. Integrative research identifies 71 new plant species records in the state of Rio Grande do Norte (Brazil) and enhances a small herbarium collection during a funding shortage. PhytoKeys 86:43-74. https://doi.org/10.3897/phytokeys.86.13775
https://doi.org/10.3897/phytokeys.86.137... and Vibrans et al. 2020VIBRANS, A.C. GASPER, A.L., MOSER, P., OLIVEIRA, L.Z., LINGNER, D.V. & SEVEGNANI, L. 2020. Insights from a large-scale inventory in the southern Brazilian Atlantic Forest. Scientia Agricola 77(1):1-12. https://doi.org/10.1590/1678-992X-2018-0036
https://doi.org/10.1590/1678-992X-2018-0... for analyses of Rio Grande do Norte and Santa Catarina, respectively).

The Cerrado, which originally occupied around 23% of the Brazilian territory, is located in Central Brazil between two areas of wet forests, the Amazon and Atlantic Forest, and forms a dry corridor, together with the Caatinga, in the northeast and the Chaco in the southwest (Oliveira-Filho & Ratter 2002OLIVEIRA-FILHO, A.T. & RATTER, J.A. 2002. Vegetation physiognomies and woody flora of the Cerrado biome. In The Cerrados of Brazil: ecology and natural history of a neotropical savanna (P.S. Oliveira & R.J. Marquis, eds). Columbia University Press, p. 91-120., Werneck et al. 2012WERNECK, F.P., NOGUEIRA, C., COLLI, G.R., SITES JR, J.W. & COSTA, G.C. 2012. Climatic stability in the Brazilian Cerrado: implications for biogeographical connections of South American savannas, species richness and conservation in a biodiversity hotspot. J Biogeogr 39:1695-1706. https://doi.org/10.1111/j.1365-2699.2012.02715.x
https://doi.org/10.1111/j.1365-2699.2012... ). The Cerrado presents notable physiographic variation (e.g., Sano et al. 2019SANO, E.E., RODRIGUES, A.A., MARTINS, E.S., BETTIOL, G.M., BUSTAMANTE, M.M.C., BEZERRA, A.S., COUTO JR., A.F., VASCONCELOS, V., SCHÜLER, J. & BOLFE, E.L. 2019. Cerrado ecoregions: a spatial framework to assess and prioritize Brazilian savanna environmental diversity for conservation. J Environ Manage 232: 818-828. https://doi.org/10.1016/j.jenvman.2018.11.108
https://doi.org/10.1016/j.jenvman.2018.1... ) and an associated number of vegetation types. These include grasslands, wetlands, savannas, and seasonally dry and wet forests (Ribeiro & Walter 2008RIBEIRO, J.F. & WALTER, B.M.T. 2008. As principais fitofisionomias do bioma Cerrado. In Cerrado: ecologia e flora (S.M. Sano, S.P. Almeida & J.F. Ribeiro, eds) 2ª ed., Embrapa, Brasília, p. 152-212.), the occurrence of which depends on ecological factors at the local scale, such as soil fertility, water availability and fire regime (Bueno et al. 2018BUENO, M.L., DEXTER, K.G., PENNINGTON, R.T., PONTARA, V., NEVES, D.M., RATTER, J.A. & OLIVEIRA-FILHO, A.T. 2018. The environmental triangle of the Cerrado domain: ecological factors driving shifts in tree species composition between forests and savannas. J Ecol 106(5):2109-2120. https://doi.org/10.1111/1365-2745.12969
https://doi.org/10.1111/1365-2745.12969... ). Because of this complex mosaic of vegetation types, the Cerrado is a savanna-dominated biome with the richest flora in the world (Klink & Machado 2005KLINK, C.A. & MACHADO, R.B.A. 2005. Conservação do Cerrado brasileiro. Megadiversidade 1(1):147-155.). It has around 12,000 angiosperm species, including 40% endemics (Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... ). However, the Cerrado has been highly threatened by deforestation, which has resulted in the loss of around 50% of its native vegetation (Alencar et al. 2020ALENCAR, A., SHIMBO, J.Z., LENTI, F., MARQUES, C.B., ZIMBRES, B., ROSA, M., ARRUDA, V., CASTRO, I., RIBEIRO J.P.F.M., VARELA, V., ALENCAR, I., PIONTEKOWSKI, V., RIBEIRO, V., BUSTAMANTE, M.C.M, SANO, E.E. & BARROSO, M. 2020. Mapping three decades of changes in the Brazilian savanna native vegetation using Landsat data processed in the Google Earth Engine platform. Remote Sens 12:924. https://doi.org/10.3390/rs12060924
https://doi.org/10.3390/rs12060924... ). Because of its species richness, high levels of endemism and anthropic pressure, the Cerrado is considered one of the 35 global biodiversity hotspots for conservation (Mittermeier et al. 2011MITTERMEIER, R.A., TURNER, W.R., LARSEN, F.W., BROOKS, T.M. & GASCON, C. 2011. Global biodiversity conservation: the critical role of hotspots. In Biodiversity hotspots (F. Zachos & J. Habel, eds) Springer, Berlin, Heidelberg, p 3-22.). Its plant species extinctions are projected to increase as a consequence of habitat loss (Strassburg et al. 2017STRASSBURG, B.N., BROOKS, T., FELTRAN-BARBIERI, R., IRIBARREN, A., CROUSEILLES, R., LOYOLA, R., LATAWIEC, A.E., OLIVEIRA FILHO, F.J.B., SCARAMUZZA, C.A.M., SACARANON, F.R., SOARES-FILHO, B. & BALMFORD, A. 2017. Moment of truth for the Cerrado hotspot. Nat Ecol Evol 1:99. https://doi.org/10.1038/s41559-017-0099
https://doi.org/10.1038/s41559-017-0099... ).

Considering the elevated rates of habitat loss and outstanding species richness, it is essential to increase knowledge about the Cerrado’s rich and endangered flora through large-scale inventories. Such inventories are certain to result in new collections of herbarium specimens and occurrence data. This would, in turn, tackle both Linnean and Wallacean shortfalls (Whittaker et al. 2005WHITTAKER, R.J., ARAÚJO, M.B., JEPSON, P., LADLE, R.J., WATSON, J.E.M. & WILLIS, K.J. 2005. Conservation Biogeography: assessment and prospect. Diversity Distrib 11:3-23. https://doi.org/10.1111/j.1366-9516.2005.00143.x
https://doi.org/10.1111/j.1366-9516.2005... ) on plant diversity by increasing the number of specimens available for taxonomic work. It would also expand the geographic ranges of known species, particularly in poorly collected areas. Information provided by new inventories can help in the development of conservation strategies, identify priority areas for threatened species, and foster the discovery of new species.

Here, we carried out a data compilation of species occurrence records generated by the IFN Cerrado, seeking to assess the project’s contribution to knowledge of the Cerrado’s woody flora. It is expected that the systematic data survey and wide geographic breadth of the IFN will provide useful information on species geographical distribution in the Cerrado. Questions we intended to resolve herein are as follows: (1) How many species were recorded by the IFN? (2) What is the proportion of woody species surveyed by IFN compared to information available in the literature? (3) Were any species found and recorded for the first time for the Cerrado flora or for individual states? (4) How many endemic or endangered species were recorded?

Material and Methods

The Serviço Florestal Brasileiro provided the data collected by the IFN Cerrado, which followed a standardised methodology (SFB 2017bSFB - Serviço Florestal Brasileiro. 2017b. Manual de campo: procedimentos para a coleta de dados biofísicos e socioambientais. Anexo I Procedimentos específicos para o Bioma Cerrado. Serviço Florestal Brasileiro, Brasília.). Data were surveyed across most of the extension of the Cerrado biome (sensu Instituto Brasileiro de Geografia e Estatísticas; https://www.ibge.gov.br) in different habitats, except in indigenous lands, where plant collecting would demand special permissions. Surveys were carried out in sampling sites called conglomerates (0.4 ha) that were systematically distributed on a 20 km x 20 km grid that sampled different habitats, including various types of natural vegetation and also anthropic areas (e.g. pastures, agriculture). Each conglomerate was composed of four crosswise subunits formed by rectangles of 20 m x 50 m located 30 m from the conglomerate’s centre. Sampling included all individuals with diameter at breast height (DBH) ≥ 10 cm, or diameter at 30 cm height from the soil (DSH) ≥ 10 cm in cases of adult individuals with height > 1.5 m, but DBH absent. Individuals with DBH ≥ 5 cm were measured in two smaller subplots (10 m x 10 m each). Individuals taller than 1.3 m, but with DBH/DSH < 5 cm, were sampled in two subplots (5 m x 5 m each). This sampling strategy was particularly designed for the IFN Cerrado in order to maximise sampling of savanna trees, which are normally stunted with twisted trunks. Individuals were assigned to morphotypes in the field and later received more accurate taxonomic identification. Detailed information on IFN Cerrado sampling methodology is provided in SFB (2017b)SFB - Serviço Florestal Brasileiro. 2017b. Manual de campo: procedimentos para a coleta de dados biofísicos e socioambientais. Anexo I Procedimentos específicos para o Bioma Cerrado. Serviço Florestal Brasileiro, Brasília..

The analyses presented here were based on the herbarium vouchers collected during field surveys carried out during the IFN Cerrado. Voucher specimens (sterile or fertile) representing species measured within conglomerates were collected in the field for identification in herbaria. Field teams were required to make a collection of any given species at least once every 15 conglomerates, including species readily identified in the field, as well as individuals with uncertain identification. Thus, specimens collected correspond to a subset of all individuals measured in IFN surveys, and are expected to comprise a representative sample of species surveyed. Conglomerates placed in areas harboring a richer flora (e.g., preserved forest) are expected to be more intensively collected than anthropic areas with fewer species (e.g., pastures). Extra collections not corresponding to sampled individuals within plots were also made. Appropriate information regarding collections, such as sampling unit, location, geographic coordinates, field name, habit and other observations, was annotated in a standardised form and later entered into a database. A total of 52,778 specimens collected were sent to the CEN, IBGE and UB herbaria for identification by generalist botanists and also by specialists in several families (Table S1, supplementary material). Specimens were databased, imaged, and finally incorporated into these collections. Duplicates were sent to other herbaria.

Our analysis included specimens collected in 2,779 conglomerates surveyed in ten states (Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Piauí, São Paulo and Tocantins) from 2011 to 2020. Field surveys and botanical identifications for Minas Gerais and São Paulo were not completed by the finalization of our analyses, and data for these states are partial. We restricted our analyses to only herbarium specimens (sterile and fertile) since these records are more reliable than unvouchered occurrence data that were also available from the IFN Cerrado dataset. We included only woody species, the main focus of the IFN, by retaining records belonging to species with life form reported as “shrub” or “tree”, according to the Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... , including shrubby and arborescent palms. From the initial dataset of 52,778 specimens, we excluded 3071 records corresponding to herbs, subshrubs, lianas and bamboos. We also excluded 20,710 records identified only at family, genus or not determined, as well as those records identified at the species level, but with an ambiguous identification (indicated by “cf.” or “aff.”). Also, 323 records corresponding to exotic or introduced species, according to the Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... , were excluded. Infraspecific categories were treated at the species level. After this filtering process, the final dataset totalled 28,602 records of native woody plants identified at the species level (Table S2, supplementary material).

Species names were checked in the RStudio program, version 3.6.3 (RStudio Team 2020RSTUDIO TEAM. 2020. RStudio: Integrated Development for R. RStudio, PBC, Boston, MA URL http://www.rstudio.com/
http://www.rstudio.com/... ), by using the flora package (Carvalho 2017CARVALHO, G. 2017. flora: Tools for Interacting with the Brazilian Flora 2020. R package version 0.3.0.https://CRAN.R-project.org/package=flora
https://CRAN.R-project.org/package=flora... ), the underlying database of which is the Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... . By the end of name checking, species names identified as synonyms were replaced by their respective accepted names. The geographic distribution of each species was verified based on information provided by Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... (accessed through flora package) in order to identify possible new occurrence records for the Cerrado or for individual states. Species that only occur within the Cerrado in Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... were listed here as Cerrado endemics. To identify endangered species collected by the IFN Cerrado, we consulted the CNCFlora Red list (National Centre for Plant Conservation; CNCFlora 2021CNCFlora - Centro Nacional de Conservação da Flora. Jardim Botânico do Rio de Janeiro. 2021. Available from: http://cncflora.jbrj.gov.br/portal/pt-br/listavermelha (accessed: 7 March 2021).
http://cncflora.jbrj.gov.br/portal/pt-br... ).

To characterise the environmental variation of IFN Cerrado sampling sites, we analysed the different habitats in which each specimen was collected. The habitat of each collection was obtained by overlaying specimens’ geographic coordinates with land cover maps available in the Mapbiomas platform (collection 6.0; Souza et al. 2020SOUZA., C.M., Jr., SHIMBO, J.Z., ROSA, M.R., PARENTE, L.L., ALENCAR, A.A., RUDORFF, B.F.T., HASENACK, H., MATSUMOTO, M.G., FERREIRA, L., SOUZA-FILHO, P.W.M., DE OLIVEIRA, S.W., ROCHA, W.F., FONSECA, A.V., MARQUES, C.B., DINIZ, C.G., COSTA, D., MONTEIRO, D., ROSA, E.R., VÉLEZ-MARTIN, E., WEBER, E.J., LENTI, F.E.B., PATERNOST, F.F., PAREYN, F.G.C., SIQUEIRA, J.V., VIERA, J.L., NETO, L.C.F., SARAIVA, M.M., SALES, M.H., SALGADO, M.P.G., VASCONCELOS, R., GALANO, S., MESQUITA, V.V. & AZEVEDO, T. 2020. Reconstructing three decades of land use and land cover changes in Brazilian biomes with Landsat archive and earth engine. Remote Sens 12(17):2735. https://doi.org/10.3390/rs12172735
https://doi.org/10.3390/rs12172735... ), which have a spatial resolution of 30 x 30 m that is compatible with the size of our sampling units (conglomerates). Original land use classes from Mapbiomes (see terms listed in parentheses) were merged into five major categories: grasslands (“formação campestre”), savannas (“formação savânica”), forests (“formação florestal”), anthropic areas (“pastagem”, “agricultura”, “mosaico de agricultura e pastagem”, “silvicultura”), and water (“água”). A land use map for 2017 was downloaded from Mapbiomes as a shapefile, and habitat classes for each of the 28,602 occurrence records were retrieved using ArcGIS 10.3 (Environmental Systems Resource Institute).

We assessed the completeness of species richness recorded in the IFN Cerrado based on the sampling effort (Martins & Santos 1999MARTINS, F.R. & SANTOS, F.A.M. 1999. Técnicas usuais de estimativa da biodiversidade. Revista Holos, Edição Especial 1:236-267.). We verified sampling efficiency by using the rarefaction/extrapolation curve based on sample size (number of specimens) according to the Hill number (q=0) using the iNext package (Chao et al. 2014CHAO, A., GOTELLI, N.J., HSIEH, T.C., SANDER, E.L., MA, K.H., COLWELL, R.K. & ELLISON, A.M. 2014. Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecol Monogr 84(1):45-67. https://doi.org/10.1890/13-0133.1
https://doi.org/10.1890/13-0133.1... ) in the RStudio program. We also compared the number of species in our dataset with the data available in Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... to assess the representativeness of the woody flora sampled by the IFN Cerrado.

Results

Our clean dataset, composed of 28,602 specimens distributed in IFN’s 2,779 conglomerates, covered most of the Cerrado (Figure 1). The number of specimens collected varied from one to 127 per conglomerate (average 10.3) with 68% of conglomerates presenting less than ten collections (Figure 2). The number of specimens was not equally distributed among states, with a higher incidence of collections in the central region of the Cerrado, especially in Goiás, and also in northern Maranhão (Figure 1). Collections were recorded in the following land use classes: savannas (40.5%), forests (30.2%), anthropic areas (25.6%), grasslands (3.5%), and water (0.2%).

Figure 1
Geographic distribution of 2779 conglomerates (sampling sites) within the Cerrado showing the number of collections of woody plants identified at species level per conglomerate. The state of Minas Gerais was only partially sampled. Indigenous lands were not sampled, which correspond to large areas in Mato Grosso and Tocantins.

Figure 2
Histogram showing the number of collections of woody plants per sampling unit (conglomerate; 0.4 ha) in the IFN Cerrado. Among the 2779 conglomerates sampled, most were represented by less than ten collections.

We recorded a total of 1,822 woody species sampled by the IFN Cerrado. However, the rarefaction curve did not reach stability (Figure 3), suggesting that increasing sampling would result in recording additional species. Species numbers varied among states (Table 1) with the highest numbers in Goiás (735), followed by Mato Grosso (683), Maranhão (647), Tocantins (563), Bahia (513), Mato Grosso do Sul (382), São Paulo (314), Piauí (291), Distrito Federal (190), and Minas Gerais (132). Most species were found in savannas (1,204 species) and forests (1,200), followed by anthropic areas (891), grasslands (373), and water (130).

Figure 3
Rarefaction/extrapolation curve of richness of woody species based on the number of specimens (28,602) collected by the IFN Cerrado. Extrapolation of species richness is represented, considering a doubling of sampling effort.

Species recorded belong to 543 genera and 105 families, including two Podocarpus gymnosperm species (see Table S3 for full species list, authorship, habit, number of collections and distribution). Among families collected, the top ten presenting the highest number of species were Fabaceae (336 species), Rubiaceae (85), Myrtaceae (78), Malvaceae (72), Melastomataceae (65), Euphorbiaceae (63), Asteraceae (60), Sapotaceae (49), Malpighiaceae (47) and Annonaceae (46). These families represented 47.2% of specimens and contributed to 49.6% of species.

The number of specimens collected per species varied widely with 29% of species (528) represented by a single collection (Figure 4). The most collected species were Tapirira guianensis (374 specimens), Matayba guianensis (305), Vatairea macrocarpa (290), Myrcia splendens (269), Diospyros lasiocalyx (249), Machaerium acutifolium (220), Qualea parviflora (215), Terminalia argentea (207), Pouteria ramiflora (206) and Callisthene fasciculata (204). All these species are typically found in savannas and forests within the Cerrado, including the widespread riparian forest species Tapirira guianensis. The IFN Cerrado contributed potentially new occurrence records for the Cerrado (64 species) and individual states (60), such as Tocantins (18), Mato Grosso do Sul (16), Goiás (11), and Maranhão (ten). Furthermore, 233 Cerrado endemics were recorded, as well as 36 endangered species (Table 2).

Figure 4
Histogram showing species frequency per range of number of collections made by the IFN Cerrado. Among the 1,822 species recorded, most were collected only once or twice, while 67 species are represented by more than 100 specimens.

Discussion

Botanical collections made during IFN Cerrado surveys covered a broad geographic range that included poorly collected regions. However, the number of specimens collected varied greatly among sampling units (conglomerates) and states. The unbalanced number of collections per conglomerate could have been influenced by the number of species present within each conglomerate. For example, a conglomerate placed in a diverse, well-preserved forest is likely to produce more collections than a conglomerate situated in pastureland wherein only one or a few tree species occur. In addition, we observed that sampling effort varied among the field teams that conducted surveys in different regions within the Cerrado. Although all field teams are supposed to follow the same sampling protocol, we found that some teams were more likely to produce more collections per conglomerate than others. For example, sampling in Piauí averaged only 5.1 specimens per conglomerate, while in Goiás, this number was more than three times higher (16.6). Therefore, it is likely that differential sampling effort among regions may have biased our results. We recommend for future IFN surveys that field teams should increase the number of collections, assuring that a representative sample of the flora within each conglomerate is represented by a set of herbarium specimens. This could be achieved by demanding field teams to collect a minimum number of vouchers per conglomerate, considering the peculiarities of each vegetation type. Such measures would reduce discrepancies in the number of specimens between conglomerates, resulting in a more even collecting effort across different regions. Increasing collecting effort would be particularly important for highly diverse sites where species identification tends to be more problematic (e.g., wet forests).

We listed a total of 1,822 woody species collected by the IFN Cerrado, which corresponds to 34% of the 5,373 woody species native to the Cerrado in the Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... repository. These statistics indicate that the survey was not able to sample the rich Cerrado plant diversity in its entirety. It is likely that unsampled species are rare, i.e., having restricted geographic range and/or low abundance, since the most frequent species in the various habitats were sampled in the IFN Cerrado. The rarefaction curve clearly showed that additional collections would result in higher species richness. Doubling sampling effort would result in an estimated 2,380 species. However, we must recognise that a considerable number of collected specimens (20,806 or 39.4%) were not identified at species level. Improving the identification of these specimens would certainly result in an increased number of species sampled by the IFN Cerrado. Likewise, the completion of the survey in Minas Gerais would have also contributed to an increase in IFN Cerrado species numbers.

Accurate identification of specimens at the species level is clearly a challenge for many large and taxonomically complex plant families (e.g., Fabaceae, Lauraceae and Myrtaceae), and reliable identifications often require samples of fruits or flowers. The challenge of naming species is particularly difficult when identifications are based on sterile specimens, which make up 86.4% of the IFN Cerrado specimens analysed here. In many cases, we recognise that an accurate determination of sterile material at the species level is not possible, even for experienced taxonomists. Uncertainties underlying plant identifications pose a limitation to forest inventories in species-rich countries, and improving this situation demands innovative approaches (Drapper et al. 2020DRAPER, F.C., BAKER, T.R., BARALOTO, C., CHAVE, J., COSTA, F., MARTIN, R.E., PENNINGTON, R.T., VICENTINI, A. & ASNER, G.P. 2020. Quantifying tropical plant diversity requires an integrated technological approach. Trends Ecol Evol 35(12):1100-1109. https://doi.org/10.1016/j.tree.2020.08.003
https://doi.org/10.1016/j.tree.2020.08.0... ) that could be adopted by the IFN. Compared to our results, sampling efficiency of floristic diversity was apparently higher in the IFN survey carried out in Santa Catarina where 831 species of trees and shrubs were found (Vibrans et al. 2020VIBRANS, A.C. GASPER, A.L., MOSER, P., OLIVEIRA, L.Z., LINGNER, D.V. & SEVEGNANI, L. 2020. Insights from a large-scale inventory in the southern Brazilian Atlantic Forest. Scientia Agricola 77(1):1-12. https://doi.org/10.1590/1678-992X-2018-0036
https://doi.org/10.1590/1678-992X-2018-0... ). However, despite all collecting efforts, a considerable number (150 species) of trees and shrubs reported for Santa Catarina in previous studies remained unsampled by that survey.

In general, the most frequently collected species in the IFN Cerrado are among the most common species cited in the literature for the different vegetation types of the Cerrado. Therefore, we can assume that an elevated number of collections for a given species reflects its high natural abundance. For example, among the 38 species considered the most frequent (oligarchic) in the cerrado sensu lato (Ratter et al. 2003RATTER, J.A., BRIDGEWATER, S. & RIBEIRO, J.F. 2003. Analysis of the floristic composition of the Brazilian Cerrado vegetation III: comparison of the woody vegetation of 376 areas. Edinb J Bot 60:57-109. https://doi.org/10.10M/S0960428603000064
https://doi.org/10.10M/S0960428603000064... ), all were recorded by the IFN Cerrado, including some highly collected (>100 records) species, such as Agonandra brasiliensis, Bowdichia virgilioides, Byrsonima coccolobifolia, Connarus suberosus, Machaerium acutifolium, Myrcia splendens, Plathymenia reticulata, Pouteria ramiflora, Qualea grandiflora, Q. parviflora, Terminalia argentea and Vatairea macrocarpa. Typical species of “cerradão”, a forest formation composed of mostly savanna tree species, such as Callisthene fasciculata, Emmotum nitens, Lafoensia pacari and Magonia pubescens (Ribeiro & Walter 2008RIBEIRO, J.F. & WALTER, B.M.T. 2008. As principais fitofisionomias do bioma Cerrado. In Cerrado: ecologia e flora (S.M. Sano, S.P. Almeida & J.F. Ribeiro, eds) 2ª ed., Embrapa, Brasília, p. 152-212.), were also highly collected in the IFN Cerrado.

Wet forests in the Cerrado are often associated with water courses. Among the species frequently found in Cerrado’s riparian forests, several were highly collected in the IFN Cerrado, such as Tapirira guianensis, which was the most collected species in the whole survey, as well as Calophylllum brasiliense, Chysophyllum marginatum, Copaifera langsdorffii, Coussarea hydrangeifolia, Dendropanax cuneatus, Eugenia florida, Hirtella glandulosa, Protium heptaphyllum, P. spruceanum, Tapura amazonica, Vochysia haenkeana and Xylopia emarginata. These species are widely distributed in Brazilian wet forests, and their shared occurrence between central Brazil riparian forests, Amazon and Atlantic Forest reinforces the floristic links among these domains (Oliveira-Filho & Ratter 1995OLIVEIRA-FILHO, A.T. & RATTER, J.A. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinb J Bot 52(2):141-194. https://doi.org/10.1017/S0960428600000949
https://doi.org/10.1017/S096042860000094... , Oliveira-Filho & Fontes 2000OLIVEIRA-FILHO, A.T. & FONTES, M.A.L. 2000. Patterns of floristic differentiation among Atlantic Forests in southeastern Brazil and the influence of climate. Biotropica 32(4b):793-810. https://doi.org/10.1111/j.1744-7429.2000.tb00619.x
https://doi.org/10.1111/j.1744-7429.2000... , Miranda et al. 2018MIRANDA, P.L.S., OLIVEIRA-FILHO, A.T., PENNINGTON, R.T., NEVES, D.M., BAKER, T.R. & DEXTER, K.G. 2018. Using tree species inventories to map biomes and assess their climatic overlaps in lowland tropical South America. Global Ecol Biogeogr 27:899-912. https://doi.org/10.1111/geb.12749
https://doi.org/10.1111/geb.12749... ).

Seasonally dry forests occur as scattered patches within the Cerrado, often associated with limestone outcrops and high-fertility soils, which stand in contrast to the acidic and nutrient-poor soils that predominate in the Cerrado region. As a consequence, central Brazil dry forests, which are mostly deciduous during the dry season, differ markedly in species composition compared to adjacent savannas and wet forests (Pennington et al. 2000PENNINGTON, R.T., PRADO, D.E. & PENDRY, C.A. 2000. Neotropical seasonally dry forests and Quaternary vegetation changes. J Biogeogr 27:261-273. https://doi.org/10.1046/j.1365-2699.2000.00397.x
https://doi.org/10.1046/j.1365-2699.2000... , Bueno et al. 2018BUENO, M.L., DEXTER, K.G., PENNINGTON, R.T., PONTARA, V., NEVES, D.M., RATTER, J.A. & OLIVEIRA-FILHO, A.T. 2018. The environmental triangle of the Cerrado domain: ecological factors driving shifts in tree species composition between forests and savannas. J Ecol 106(5):2109-2120. https://doi.org/10.1111/1365-2745.12969
https://doi.org/10.1111/1365-2745.12969... ). The characteristic dry forest species Aspidosperma subincanum and Tabebuia roseoalba were among those highly collected species in the IFN Cerrado, which also recorded other typical dry forest representatives mentioned in the literature, such as Aspidosperma pyrifolium, Commiphora leptophloeos, Machaerium scleroxylon and Schinopsis brasiliensis (Scariot & Sevilha 2005SCARIOT, A. & SEVILHA, A.C. 2005. Biodiversidade, estrutura e conservação de florestas estacionais deciduais no Cerrado. In Cerrado: ecologia, biodiversidade e conservação (A. Scariot, J.C. Sousa-Silva & J.M. Felfili, eds) Ministério do Meio Ambiente, Brasília, p. 123-139., Pereira et al. 2011PEREIRA, B.A.S., VENTUROLI, F. & CARVALHO, F.A. 2011. Florestas estacionais no Cerrado: uma visão geral. Pesq Agropec Trop 41(3):446-455. https://doi.org/10.5216/pat.v41i3.12666
https://doi.org/10.5216/pat.v41i3.12666... ). Although occurring in discontinuous patches, Cerrado dry forests share several species in common with the Caatinga, highlighting the floristic link between these seasonally dry vegetation nuclei (Prado & Gibbs 1993PRADO, D.E. & GIBBS, P.E. 1993. Patterns of species distribution in the dry seasonal forests of South America. Ann Mo Bot Gard 80(4):902-927. https://doi.org/10.2307/2399937
https://doi.org/10.2307/2399937... , Neves et al. 2015NEVES, D.M., DEXTER, K.G., PENNINGTON, R.T., BUENO, M.L. & OLIVEIRA-FILHO, A.T. 2015. Environmental and historical controls of floristic composition across the South American dry diagonal. J Biogeogr 42(8):1566-1576. https://doi.org/10.1111/jbi.12529
https://doi.org/10.1111/jbi.12529... ). In addition, a large sample of sites located at the Cerrado/Caatinga boundary, particularly in Bahia, contributed to increased numbers of typical dry forests and shrublands in our list.

It is worth highlighting that the systematic sampling methodology of the IFN generated new plant occurrence records in areas little sampled in the Cerrado (Sousa-Baena et al. 2014SOUSA-BAENA, M.S., GARCIA, L.C. & PETERSON, A.T. 2014. Completeness of digital accessible knowledge of the plants of Brazil and priorities for survey and inventory. Diversity Distrib 20:369-381. https://doi.org/10.1111/ddi.12136
https://doi.org/10.1111/ddi.12136... ), such as Piauí, Maranhão (the second state in number of IFN specimens), Tocantins, Mato Grosso and southwestern Goiás, thereby filling some important gaps in species distributions. Another positive aspect of the IFN Cerrado methodology was the sampling in different physiognomies, which included habitats that are not frequently surveyed by botanists, such as riparian forests, swamps and seasonally dry forests, including remote areas difficult to access. Botanical exploration of these areas contributed to a number of new species occurrence records for the Cerrado and also for individual states, contributing floristic knowledge, particularly to some states, such as Maranhão, Piauí and Tocantins, for which floristic information is limited (BFG 2015BFG - The Brazil Flora Group. 2015. Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66:1085-1113. https://doi.org/10.1590/2175-7860201566411
https://doi.org/10.1590/2175-78602015664... ). The contribution of IFN collections towards new state occurrence records has also been highlighted elsewhere for Rio Grande do Norte where 71 new angiosperm records were reported (Versieux et al. 2017VERSIEUX, L.M., DÁVILA, N., DELGADO, G.C., DE SOUSA, V.F., DE MOURA, E.O., FILGUEIRAS, T., ALVES, M.V., CARVALHO, E., PIOTTO, D., FORZZA, R.C., CALVENTE, A. & JARDIM, J.G. 2017. Integrative research identifies 71 new plant species records in the state of Rio Grande do Norte (Brazil) and enhances a small herbarium collection during a funding shortage. PhytoKeys 86:43-74. https://doi.org/10.3897/phytokeys.86.13775
https://doi.org/10.3897/phytokeys.86.137... ).

A large number of new occurrence records for the Cerrado, such as Aniba hostmaniana, Calycophyllum spruceanum, Elvasia calophyllea, Eugenia cupulata, Jacaranda copaia, Matayba peruviana, Miconia eugenioides, Pouteria anomala and Theobroma speciosum, were mostly collected in wet forests located in northern Mato Grosso and Tocantins along the Cerrado/Amazonia boundary. This ecotone between the Cerrado and Amazonia, a complex transition zone, is composed of interdigitating patches of savannas and various forest types (Marques et al. 2020MARQUES, E.Q., MARIMON-JUNIOR, B.H., MARIMON, B.S., MATRICARDI, E.A.T., MEWS, H.A. & COLLI, G.R. 2020. Redefining the Cerrado-Amazonia transition: implications for conservation. Biodivers Conserv 29:1501-1517. https://doi.org/10.1007/s10531-019-01720-z
https://doi.org/10.1007/s10531-019-01720... ), and it has certainly incremented the number of species sampled by the IFN Cerrado. Coinciding with an area that lacks botanical records overall (Souza-Baena et al. 2014), this ecotone also helps to explain the large number of new occurrences in the Cerrado for species previously reported only to Amazonia. New occurrences for the Cerrado also comprised typical elements from the Caatinga (e.g., Cratylia mollis, Pouteria furcata) and Atlantic Forest (e.g., Dendropanax denticulatus, Myrocarpus frondosus), albeit to a lesser extent compared to Amazonia. These results highlight the contribution of marginal/ecotonal areas between the Cerrado and other biomes to the species richness of the Cerrado’s tree flora, as reported in previous studies (Oliveira-Filho & Ratter 1995OLIVEIRA-FILHO, A.T. & RATTER, J.A. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinb J Bot 52(2):141-194. https://doi.org/10.1017/S0960428600000949
https://doi.org/10.1017/S096042860000094... , Françoso et al. 2016FRANÇOSO, R.D., HAIDAR, R.F. & MACHADO, R.B. 2016. Tree species of South America central savanna: endemism, marginal areas and the relation with others biomes. Acta Bot Brasilica 30(1):78-86. https://doi.org/10.1590/0102-33062015abb0244
https://doi.org/10.1590/0102-33062015abb... , Miranda et al. 2018MIRANDA, P.L.S., OLIVEIRA-FILHO, A.T., PENNINGTON, R.T., NEVES, D.M., BAKER, T.R. & DEXTER, K.G. 2018. Using tree species inventories to map biomes and assess their climatic overlaps in lowland tropical South America. Global Ecol Biogeogr 27:899-912. https://doi.org/10.1111/geb.12749
https://doi.org/10.1111/geb.12749... ).

It is important to mention that new occurrence records should be viewed with caution since most were based on sterile specimens identified by non-specialists. Consequently, they may not represent accurate records. This means that potentially new occurrence records should be further confirmed, preferably based on fertile specimens determined by experts.

Among the 214 species collected by the IFN Cerrado cited in the CNCFlora’s red list, 36 are classified to some degree of threat, including 2 critically endangered (CR), 14 endangered (EN) and 20 vulnerable (VU). Another 179 species are classified as least concern (LC) or near threatened (NT). A total of 232 Cerrado endemic species were collected, representing 12.4% of 1,858 endemic woody species registered in the Flora do Brasil 2020FLORA DO BRASIL 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed: 12 March 2021).
http://floradobrasil.jbrj.gov.br/... for this biome. New occurrence records for threatened, as well as rare and endemic species provide crucial data expanding knowledge of geographic ranges and, hence, enabling updates of species threat status and furthering efforts to subsidise conservation initiatives. In addition to new distribution records, specimens collected by the IFN Cerrado supported the description of a new Harpalyce (Fabaceae) species from western Bahia (São-Mateus et al. 2019SÃO-MATEUS, W.M.B., SIMON, M.F., QUEIROZ, L.P., JARDIM, J.G. & CARDOSO, D.B.O.S. 2019. Two new species of Harpalyce (Leguminosae, Papilionoideae) from the Cerrado hotspot of biodiversity in Brazil. Kew Bull 74(61):1-11. https://doi.org/10.1007/S12225-019-9845-Y
https://doi.org/10.1007/S12225-019-9845-... ). It is likely that ongoing taxonomic work by specialists based on IFN Cerrado collections deposited in herbaria will reveal more new species to science. Also, with progress in specimen identification in herbaria, new species occurrences for the Cerrado and states, as well as records for endemic and threatened species, are expected.

Conclusion

Although initially developed to assess land coverage, forest structure and wood production, national forest inventories have also played a role as a major source of data for monitoring forest biodiversity (e.g., Chirici et al. 2012CHIRICI, G., MCROBERTS, R.E., WINTER, S., BERTINI, R., BRÄNDLI, U., ASENSIO, I.A., BASTRUP-BIRK, A., RONDEUX, J., BARSOUM, N. & MARCHETTI, M. 2012. National Forest Inventory contributions to forest biodiversity monitoring. Forest Sci 58:257-268.). In the present study, focusing specifically on the biodiversity dimension, we showed that the IFN Cerrado has provided a useful source of occurrence data for woody species, spanning wide geographic and habitat coverage. The large number of woody species reported here reflects the floristic variation found in diverse vegetation types found in the Cerrado region, which includes savannas, seasonally dry forests, and wet forests. Our analyses, based on the latest dataset available, showed that about a third of the floristic diversity of the Cerrado woody plants was sampled in the IFN. Our results also show that the typical floristic composition of the different vegetation types of the Cerrado was captured by that survey, and that species cited in the literature as common appear in high numbers among IFN collections. However, differential sampling efforts between field teams and difficulties in naming specimens are expected to influence the floristic diversity reported here. The fact that 35% of conglomerates sampled were located in anthropic areas, which are likely to be less diverse than well-preserved areas, may have also influenced the great variation found in sampling units in terms of species richness. The floristic richness compiled by the IFN Cerrado, which included a number of potential new occurrence records, was greatly influenced by typical woody elements from surrounding biomes, such as Amazonia, Caatinga and Atlantic Forest. Species collected in these marginal/ecotonal zones greatly contributed to the overall number of species recorded here.

Although considered a biodiversity hotspot, it is estimated that the Cerrado will lose 31-34% of its remaining native vegetation by 2050, mostly from agricultural expansion and limited protected areas (Strassburg et al. 2017STRASSBURG, B.N., BROOKS, T., FELTRAN-BARBIERI, R., IRIBARREN, A., CROUSEILLES, R., LOYOLA, R., LATAWIEC, A.E., OLIVEIRA FILHO, F.J.B., SCARAMUZZA, C.A.M., SACARANON, F.R., SOARES-FILHO, B. & BALMFORD, A. 2017. Moment of truth for the Cerrado hotspot. Nat Ecol Evol 1:99. https://doi.org/10.1038/s41559-017-0099
https://doi.org/10.1038/s41559-017-0099... ). Therefore, increasing Cerrado floristic knowledge is critical to support conservation planning in this threatened biome. We expect that the occurrence records derived from IFN collections, particularly those from rare, endemic and endangered species, will contribute to the identification of priority areas for further surveys and conservation of the rich Cerrado flora.

Supplementary Material

The following online material is available for this article:

Table S1 - Specialists that contributed with the identification of selected families, and generalist botanists that identified miscellaneous families (bottom of the table).

Table S2 - Herbarium specimens of woody species collected by the IFN Cerrado. Only records belonging to native species identified at the species level are listed.

Table S3 - List of woody species collected by the IFN Cerrado, including information on states of occurrence, number of specimens collected, habit, vegetation type, and selected voucher specimen. A voucher specimen, either sterile or fertile, was chosen to represent each species.

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