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Original Article • Acta Bot. Bras. 39 • 2025 • https://doi.org/10.1590/1677-941X-ABB-2023-0210 linkcopy

Open-access Asteraceae from Barra da Estiva: the use of floristic inventories suggests two bioregions in the Chapada Diamantina - Brazil

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

    Abstract

    Asteraceae is one of the most diverse families in the world, especially in mountainous regions such as Chapada Diamantina, Bahia, Brazil. The objectives of this work were to carry out a floristic survey of Asteraceae in the municipality of Barra da Estiva, discuss the distribution of species found, and analyze the similarity between the Asteraceae diversity in Barra da Estiva and other previously evaluated areas. Herbaria were reviewed, and three field trips were carried out. Barra da Estiva houses 98 species of Asteraceae grouped into 46 genera and 13 tribes. The tribes Vernonieae (34 spp.) and Eupatorieae (24 spp.) were the most diverse, especially the genera Baccharis (7 spp.) and Lepidaploa (7 spp.). A matrix of species presence/absence for different phytophysiognomies was elaborated according to the rendered species list and previous Asteraceae surveys in Chapada Diamantina. Although previous research considered the Chapada Diamantina a single phytogeographic unit, our results suggest its possible subdivision into two bioregions: ‘Piemonte da Diamantina’, including areas of the extreme northeast, and Central-South Chapada, encompassing areas from the central-south region.

    Keywords:
    Compositae; Espinhaço Range; Floristic diversity; Floristic inventories

    Introduction

    Asteraceae corresponds to about 10% of the global flora and it is recognized as one of the largest families within angiosperms, with an estimated diversity of 25,000 species (Mandel et al., 2019). Recognized as a monophyletic group, the family originated in South America approximately 83 million years ago, at the end of the Cretaceous, and it is morphologically characterized by its capitulum inflorescence, synantherous anthers, inferior ovary containing a single ovule with basal placentation, a cypsela fruit, and a usually persistent calyx that is modified into a structure known as pappus (Roque et al., 2017; Mandel et al., 2019).

    Recent advances in molecular studies allowed a better understanding of the group. Currently, Asteraceae has a robust taxonomic classification, including 16 subfamilies, 50 tribes, and about 1,700 genera (Mandel et al., 2019; Susana et al., 2020). In Brazil, it is represented by 27 tribes, 341 genera, and 2,259 species, of which 1,371 are endemic (Flora e Funga do Brasil, 2024). The family is particularly diverse in mountainous and savanna areas, partly due to its heliophilous behavior (Esquivel, 2015), with the Cerrado and Atlantic Forest, which are both considered biodiversity hotspots (Myers et al., 2000), having the highest number of Asteraceae species in the country (Siniscalchi et al., 2021).

    Located on the margins of these two hotspots and the Caatinga domain, the Espinhaço Range (ER) is a mountain range that extends throughout eastern Brazil, in the states of Bahia and Minas Gerais (Campos et al., 2019). Due to its high species richness, holding 14% of Brazilian diversity in about 1% of its territory (Silveira et al., 2016), the Espinhaço Range is recognized as an area of great importance for conservation (Rapini et al., 2008; Chaves et al., 2019). In this context, Asteraceae has stood out as one of the most species-rich families in several studies carried out in the ER and adjacent areas (e.g.,Meguro et al., 1996; Conceição & Giulietti, 2002; Viana & Lombardi, 2007; Meyer & Franceschinelli, 2010; Couto et al., 2011; Campos et al., 2016), being particularly representative in campos rupestres (rocky grasslands) (Silveira et al., 2016; Almeida et al., 2023). Throughout the campos rupestres of the ER, Asteraceae is represented by over 500 species, of which 178 are endemic to this phytophysiognomy (Campos et al., 2019).

    Based on its geomorphology, the ER has traditionally been subdivided into Serra do Espinhaço and Chapada Diamantina (CHD), the latter located in the north-central portion of the state of Bahia (Danderfer & Dardenne, 2002). The Serra do Espinhaço, in turn, is further subdivided into Northern Espinhaço, which runs through the states of Bahia and Minas Gerais, and Southern Espinhaço, exclusively located in Minas Gerais (Campos et al., 2019). Currently, 12 inventories of Asteraceae in the ER are reported, eight in Serra do Espinhaço (Almeida, 2008; Borges & Forzza, 2008; Borges et al., 2010; Santana, 2013; Marques & Nakajima, 2015; Reis et al., 2015; Campos et al., 2016; Contro & Nakajima, 2017) and four in the CHD (Hind, 1995; Moura & Roque, 2014; Roque et al., 2016; Staudt et al., 2017).

    The CHD holds a wide variety of plant communities with a high proportion of endemic and micro-endemic species (Giulietti et al., 1987; Conceição et al., 2007; Colli-Silva et al., 2019), which could be associated with its particular edaphic conditions, with soils originated by the exposure of various types of rocks and mainly characterized as clayey, acid, and infertile (da Nóbrega & Boas, 2020). Composed of 53 municipalities (Ribeiro, 2022), the CHD encompasses a large number of plant collections focused on a few municipalities (e.g. Lençóis, Mucugê, Rio de Contas, Morro do Chapéu, and Piatã) (Campos et al., 2019; Colli-Silva et al., 2019; Ribeiro, 2022). This highlights a knowledge gap concerning Asteraceae diversity in the Northern and Northwestern regions of CHD, such as the areas previously recognized by Campos et al. (2019) as ‘Irecê’ and ‘Piemonte da Diamantina’. The same pattern is observed in the center-south region since a large portion of the collections are condensed in the Chapada Diamantina National Park, an area of scientific importance and higher funding investment (Lucresia et al., 2021).

    Among those areas lacking collections throughout the CHD, Barra da Estiva stands out. The municipality is located in Chapada Diamantina’s southern zone, sheltering the spring of the Paraguaçu River, one of the most important rivers in the state of Bahia; it is distant from conservation units, showing a heterogeneous sampling effort (Santos et al., 2020). Before the development of this work, the municipality presented 415 records of Asteraceae on the Reflora virtual herbarium and SpeciesLink (CRIA, 2022). Other municipalities in CHD, like Mucugê and Rio de Contas, had over 2.000 records reported, while Abaíra and Morro do Chapéu presented over 1.400 records (CRIA, 2022). The small sampling effort observed for Barra da Estiva is therefore evident, demonstrating the need to expand botanical work in this area. Taking this into account, the objectives of this work were to carry out a floristic inventory of the family Asteraceae in the municipality of Barra da Estiva, elaborate an identification key, analyze and discuss the occurrence and distribution of the species of Asteraceae throughout the municipality, as well as compare the similarity between the Asteraceae flora of Barra da Estiva with other previously studied localities in CHD.

    Materials and methods

    Study area

    According to the Köppen-Geiger climate classification, the municipality of Barra da Estiva presents two climates within its territory (Figure 1): while the northwestern area is classified as a Tropical Savannah with dry winters (Aw), the southeastern zone has a hot Semi-Arid climate (BSh) (Rubel & Kottek, 2010). The municipality has a total area of 1,346.6 km², an average elevation of 1,005 m, and its centroid is located at 13°36' 47" S, 41°19' 24" W, bordering the municipalities of Ituaçu, Jussiape, and Ibicoara (IBGE, 2024). Barra da Estiva has a population of approximately 20,198 inhabitants and is characterized by significant agricultural activities, including crop cultivation and livestock farming (IBGE, 2012). The municipality is located on the slopes of the southern portion of Serra do Sincorá (Fig. 1 F ), a geological formation that is part of the CHD and extends through several municipalities, such as Lençóis, Palmeiras, Mucugê, and Andaraí, being recognized as an important region for ecological, geological, and paleontological studies as well as presenting high touristic interest ( Neves & Conceição, 2007).

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    Figure 1.
    Map of the municipality of Barra da Estiva showing climate classification according to Köppen-Geiger: Tropical wet and dry (Aw in green) and Hot semi-arid (BSh in orange). Previous collection sites of Asteraceae (white cricles) and the collection points in the present work (black circles). Photographs of the different phytophysiognomies of the region: A - Campo rupestre, Morro do Ouro; B - Cerrado; C - Caatinga, southeast area of the municipality; D - Mata Ciliar, Cachoeira Grande; E - Anthropized area for livestock farming; F - Serra do Sincorá, northern area of the municipality.

    Floristic survey

    The botanical collections housed at the herbaria ALCB, BHCB, CEPEC, HUEFS, HURB, RB, and SPF (acronyms according to Thiers, 2024, updated continuously) were visited and reviewed. Furthermore, botanical collections from ESA, MBM, and UEC were evaluated using online high-resolution images available on the Specieslink (CRIA, 2022) and JABOT (2024) digital platforms. In addition to the herbaria collections, three field expeditions were carried out in August (2021), January, and July (2022), in which a total of 116 specimens were collected in different parts of the municipality. The targeted sampling localities were selected to include areas for which no specimens were found in herbaria, as well as to increase sampling effort in previously sampled locations of the municipality (Fig. 1). The collected material was deposited in the ALCB herbarium, with duplicates sent to HUEFS and RB (acronyms according to Thiers, 2024, updated continuously).

    Species identification was carried out following specific literature as reference and consulting type specimens and their respective protologues. Valid names followed Flora e Funga do Brasil (2024) and the delimitation of tribes followed the classification proposed by Susanna et al. (2020). The result was compiled into a species list (Table 1) including information about habit, endemism, conservation status according to IUCN (2024), phytophysiognomies of occurrence, and specimen reference. Additionally, a taxonomic key for the taxa found in the municipality was constructed.

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    Table 1.
    List of Asteraceae species from Barra da Estiva, Bahia, Brazil, *Endemic species to Bahia; growth habits; species endemicity: not endemic (NO), endemic to Brazil (BR), endemic to the Espinhaço Range (ER), endemic to the state of Bahia (BA); IUCN threat status: Non-evaluated (NE), Data Deficient (DD), Least Concern (LC), Near Threatened (NT), Vulnerable (VU), Endangered (EN) and Critically Endangered (CR); Phytophysiognomies: AA - anthropized area, CD - Cerrado, CR - Campos Rupestres, CT - Caatinga and F - Forest and their respective vouchers.

    Data analysis

    Due to the diversity of terms used to characterize phytophysiognomies, these were grouped into five broad categories, following Ribeiro & Walter (1998) and Rizzini (1997), with modifications. The first category corresponds to Campo rupestre (CR), a savanna formation found above 900 m of elevation, with shrubs growing directly over rock outcrops (Fig. 1 A ). Specimens that presented the following descriptions were included: “Campo rupestre”, “área rupestre”, “formação rupestre”, and rocky sand. The second category, named Cerrado sensu lato (CD), is characterized by the presence of trees with irregular and twisted branching, generally with evidence of fire, and scattered shrubs/subshrubs with perennial underground structures (Fig. 1 B ). These areas generally present acid, red-yellowish clayey soils and, in the present work, specimens cataloged under the following descriptions were included in this category: “Cerrado”, Cerradão”, “Campos gerais”, “Gerais”, “Campo sujo”, and “Campo limpo”.

    The third category is Caatinga (CT), which encompasses the semi-arid region of Northeastern Brazil, exhibiting vegetation adapted to arid and semi-arid environments characterized by seasonal leaf loss (Fig. 1 C ). It is an environment dominated by the shrubby/arboreal stratum with an abundance of C4 species, or plants with cladodes or spines. Specimens collected in Caatinga and “Carrasco” were included in this category. The fourth category is Forest (F), which includes forest formations with the presence of canopy and leaf litter, the predominance of tree species, and perennial or semi-perennial plants (Fig. 1 D ). Specimens collected in areas described as: “florestas”, “mata”, “mata ciliar”, and “mata de galeria” were included. Finally, and considering the heliophilous behavior of Asteraceae, a fifth category named Anthropized Area (AA) was included, which corresponds to areas where the original vegetation has disappeared due to human modification (Fig. 1 E ). In this category were included specimens of plants collected from, but not cultivated at, urban perimeters, edges of main roads, crops, and pasture areas.

    To compare the species composition of the study area with other areas of CHD, a list of Asteraceae species of Barra da Estiva was compiled in presence/absence matrices for each phytophysiognomy observed in the municipality. The attribution of specimens to different vegetation classes followed the collector’s description and field observations. A list of species available in the inventories of Asteraceae previously carried out in the CHD were also grouped in presence/absence matrices following the descriptions of phytophysiognomies cited by Hind (1995) Moura & Roque (2014), Roque et al. (2016), and Staudt et al. (2017) (Table 2). A survey of the Espinhaço Range carried out in the municipality of Licínio de Almeida was included as an outgroup (Campos et al., 2016).

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    Table 2.
    Previous studies focused on Asteraceae carried out in areas of Chapada Diamantina and Serra do Espinhaço in the state of Bahia, and the respective number of tribes, genera, and species observed for each phytophysiognomy in the different localities: AA - anthropized area, CD - Cerrado, CR - Campos Rupestres, CT - Caatinga and F - Forest; (-) areas not evaluated in the present study.

    These presence/absence matrices (species segmented by phytophysiognomies of occurrence in each locality) and a general presence/absence matrix for each locality were subjected to ordering and clustering analysis (Non-Metric Multidimensional Scaling - NMDS; and Unweighted Pair Group Method with Arithmetic Mean - UPGMA), using the Simpson index (Del Rio et al., 2003) as a dissimilarity metric, as this index is less affected by species richness. Two analyses of similarities (ANOSIM) were carried out based on the binary matrices segmented by phytophysiognomy and locality to evaluate the cohesion of the clusters found. To avoid pseudo-grouping, only phytophysiognomies that presented over five records per locality were included. Statistical procedures were performed in R (R Development Core Team, 2021) using the packages betapart (Baselga & Orme, 2012) and vegan (Oksanen et al., 2021).

    Results

    Asteraceae from Barra da Estiva

    A total of 13 tribes, 46 genera, and 98 species of Asteraceae were found in Barra da Estiva (Table 1). Amongst the 98 species registered for the municipality, 22 spp. correspond to new records (22%), of which 19 spp. are new collections made during this work. Aspilia hispidantha H.Rob. was initially reported for the municipality of Barra da Estiva; however, it was not included in the species list because no morphological differences were found with Aspilia subalpestris Baker, reinforcing the previous synonymization of the former, proposed by Alves & Roque (2018).

    Tribes Vernonieae and Eupatorieae were the most diverse, with 34 and 24 species respectively, representing 59% of Asteraceae diversity in the municipality, followed by Heliantheae with 10 species. The remaining nine tribes each presented four or fewer species, collectively constituting only 15% of the total Asteraceae species in the municipality. The genera Baccharis and Lepidaploa were the most diverse with seven and six species respectively, followed by Lessingianthus, Lychnophora, Mikania, and Moquiniastrum, with five species each. Twenty-eight genera presented only one species each, covering 29% of the total diversity of Asteraceae in the study area.

    The most common growth habit was shrubs, including 46 species, followed by subshrubs (25), herbs (17), and trees (9), while only one species of liana (Mikania phaeoclados Mart.) was reported (Fig. 2). Of the 98 species reported in this work, 71 species (72%) are considered endemic to Brazil (Table 1), distributed as follows: 34 exhibit wide distribution in several Brazilian states; 14 endemic to the Espinhaço Range and reported in the states of Bahia and Minas Gerais; and 23 species reported only in the state of Bahia, being mostly from the Eupatorieae (11 spp.) and Vernonieae (8 spp.). The municipality did not present endemic species; however, Acritopappus harleyi and Stylotrichium glomeratum were described having the municipality as their type locality and, despite one report from the municipality of Ibicoara (Fig. 1 F ), all other collections have been registered in Morro do Ouro in the southern area of Barra da Estiva. A similar case occurs regarding Agrianthus almasensis, recorded only in Barra da Estiva, Rio de Contas, and Piatã; and Agrianthus carvalhoi, recorded in Barra da Estiva and Ibicoara.

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    Figure 2.
    Growth habits of Asteraceae species present in the municipality of Barra de Estiva, with the capitulum inflorescence of Lychnophorella morii in the center.

    Barra da Estiva has seven species classified under some IUCN threat category (IUCN, 2024). Two are classified as critically endangered (CR), two as endangered (EN), and three as vulnerable (VU) (Table 1). Regarding the remaining 90 species, five were assessed and listed as Least Concern (LC), two are Data Deficient (DD), and 83 have not yet been assessed. The species included in the threat categories belong to the two most diverse tribes, Vernonieae (5) and Eupatorieae (3), with six species exclusive to campos rupestres (CR) (Agrianthus almasensis, Stylotrichium glomeratum, Lychnophora granmogolensis, L.rosmarinifolia, Paralychnophora harleyi, and Stilpnopappus tomentosus. Among the non-endemic CR species, Paralychnophora bicolor was registered both for CR and F; while Acritopappus harleyi was reported for CD.

    The phytophysiognomies had variable diversity. CR was the richest, with 71 species, 41 of them being exclusive while CD presented 32 species, being the second most diverse phytophysiognomy. On the other hand, CT and F presented the lowest richness of Asteraceae in the municipality, with six and nine species respectively. The phytophysiognomies were also heterogeneously distributed within the municipality, with the western portion of Barra da Estiva concentrating areas of CR, CD, and AA, while the eastern portion with CT and some humid F (Fig. 1).

    Floristic comparison of Asteraceae between areas of Chapada Diamantina

    Among the Espinhaço areas within the state of Bahia that have been previously inventoried, Barra da Estiva ranks as the fourth locality with the highest number of Asteraceae species after Mucugê (167 spp.), Pico das Almas (124 spp.), and Morro do Chapéu (119 spp.), surpassing only Licínio de Almeida (95 spp.) and Jacobina (80 spp.) (Table 2). Regarding the floristic patterns of Asteraceae in these localities, the NMDS results from the dataset that was not segmented by phytophysiognomy (Fig. 3 A , B) showed a separation of the Northern Espinhaço range (Licínio de Almeida, in black) from the areas located in Chapada Diamantina (in blue and orange). Those areas in turn, are subdivided into two groups. The first, here referred as Piemonte da Diamantina (I), as it corresponds to a political and administrative region with the same name (approximately translated into English as ‘Diamantina foothills’), groups the Jacobina and Morro do Chapéu municipalities, both located northeast of the CHD. The second group, here referred as Central-South Chapada (II), included Barra da Estiva, Mucugê and Pico das Almas. The UPGMA based on localities reinforces the clusters recognized by the NMDS, also separating areas of the Northern Espinhaço range (in black) from areas of the CHD, which are subdivided into the same groups: Piemonte da Diamantina (I) and Central-South Chapada (II) (Fig. 3 C ).

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    Figure 3.
    A- Location in the Espinhaço Range of the compared areas: Licínio de Almeida (L), Pico das Almas (P), Barra da Estiva (B), Mucugê (M), Morro do Chapéu (MC) and Jacobina (J). B - NMDS based on localities: Northern Espinhaço (L-Black); Central-South Chapada (II-Blue); Piemonte da Diamantina (I-Orange). C - UPGMA based on localities: Northern Espinhaço (L); Central-South Chapada (II); Piemonte da Diamantina (I).

    The floristic patterns of Asteraceae segmented by phytophysiognomies reveal an important geographical structure, with a latitudinal clustering pattern in a south-north direction, mostly clustered by localities instead of by phytophysiognomies (Fig. 4). The Forest phytophysiognomy of Pico das Almas (FP) presented the highest dissimilarity among the evaluated areas, being placed as an external group from the others. Moreover, the anthropized areas (AA) were grouped (black triangles, Fig. 4) in an external core in comparison with the natural vegetations recognized by both the NDMS (Fig. 4 A ) and the UPGMA (Fig. 4 B ); however, in the NMDS, one of the natural vegetations is grouped with the AA (CDJ). Among the natural vegetation in the UPGMA, the areas belonging to Jacobina (○, Fig. 4) form a separate group from the remaining natural areas.

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    Figure 4.
    A - NMDS analysis for the different vegetations of each locality, with the first two letters indicating phytophysionomies: AA (anthropized area), CD (Cerrado), CR (Campos Rupestres), CT (Caatinga), and F (Forest) and the remaining letters indicating the localities: B (Barra da Estiva), J (Jacobina), L (Licínio de Almeida), M (Mucugê), MC (Morro do Chapéu), and P (Pico das almas/Rio de Contas); Anthropized areas (black triangles), Northern Espinhaço in black (squared plus), Central-South Chapada in blue, and Piemonte da Diamantina in red/orange. B - UPGMA analysis for the different vegetations following the same nomenclature as in Figure 4A: (black triangles) AA, (white circles) Jacobina, (black squares) South Chapada, (squared plus) Northern Espinhaço, (white square) Mucugê, and Morro do Chapéu.

    Regarding the natural areas observed in the UPGMA (Fig. 4 B ), the South Chapada group (black squares, Fig. 4) stands out, including areas of Barra da Estiva and Pico das Almas, reinforcing the floristic similarities within this grouping. Additionally, the areas belonging to the Northern Espinhaço (squared plus, Fig. 4) were also grouped independently in the UPGMA, while the NDMS analysis suggests a similarity between these areas and the Cerrado of Morro do Chapéu (CDMC). The remaining areas, belonging mostly to Morro do Chapéu and Mucugê, were grouped with Jacobina Campo rupestre (CRJ) and Barra da Estiva Caatinga (CTB) areas (white square, Fig. 4).

    The ANOSIM reinforced the NMDS and UPGMA results, indicating a greater cohesion between localities (ANOSIM=0.4709; P=0.001) than between phytophysiognomies (ANOSIM=0.099; P=0.051).

    Discussion

    Asteraceae in Barra Da Estiva

    Barra da Estiva hosts 98 species of Asteraceae (Table 1), with shrub and sub-shrub growth habits being predominant, representing 73% of all species (Fig. 2). This prevalence is linked to collections from the phytophysiognomies of CR and CD (Rizzini, 1997), environments that were the focus of more extensive collections and species richness in the municipality. Although Asteraceae is the second richest family in lianas in the Neotropics (Sperotto et al., 2023), only one record of a climbing plant was reported. This result may be attributed to the direct correlation between the diversity of climbing plants with hyper-humid forest habitats (Sperotto et al., 2023), which exhibited a reduced territorial extension in Barra da Estiva.

    The diversity of Asteraceae in Barra da Estiva follows patterns previously observed along the Espinhaço Range. Notably, the high richness of the Vernonieae and Eupatorieae tribes (58%) has been reported in several studies carried out in this geological formation, where these tribes represent about 50% of the total diversity of Asteraceae (Campos et al., 2019). Furthermore, Heliantheae was the third richest tribe in Barra da Estiva, a result also observed in Morro do Chapéu (Staudt et al., 2017) and Jacobina (Moura & Roque, 2014). The higher species richness of the genera Baccharis, Lepidaploa, Lessingianthus, Lychnophora, and Mikania further reinforce the similarity in the composition of the family along the Espinhaço Range (Campos et al., 2019). However, it is important to emphasize that this similarity may result from collection patterns focused on CR areas, where these taxa are especially diverse (Campos et al., 2019), potentially overlooking other phytophysiognomies where different taxa/groups could be more prominent.

    The greater diversity of Asteraceae in campos rupestres in Barra da Estiva corroborates previously reported patterns of higher species richness in this type of phytophysiognomy (Campos et al., 2019). The lower number of species attributed to the Caatinga was also expected, as Asteraceae traditionally show less diversity in this phytophysiognomy (Roque et al., 2017). However, the low number of records reported in the municipality may be due to limited collection efforts in Caatinga areas (Fig. 1), because of the difficulty in collecting in this phytophysiognomy, which is particularly dependent on rainfall seasonality (Jardim et al., 2022), coupled whit a higher focus on Campos rupestre, areas that traditionally have been recognized as harboring high diversity (Lucresia et al., 2021).

    Along with the Caatinga, forest areas also exhibited low diversity of Asteraceae. This result could be associated with the limited extent and high fragmentation of forest areas (Giam, 2017), leaving Asteraceae species practically restricted to the margins of some water bodies and transition zones between crops or livestock areas. This is particularly significant in Barra da Estiva, the source of the Paraguaçu River (Santos et al., 2020), since deforestation is directly linked to biodiversity loss, ecosystem fragmentation, soil alteration, and changes in the water cycle (Lawrence et al., 2022)

    Floristic comparison of the family Asteraceae between areas of the Espinhaço range in Bahia

    When analyzing the plant composition at the locality level, both NMDS and UPGMA analyses evidenced a greater similarity among the CHD block when compared to the Northern Espinhaço (Fig. 3), contrasting with the results previously reported for the Espinhaço Range using different biological models (Colli-Silva et al., 2019; Alves & Loeuille, 2021; Lucresia et al., 2021), where the region of Licínio de Almeida municipality grouped with southern areas of the CHD.

    Using Asteraceae from CR as a model, the Northern Espinhaço and CHD areas were previously clustered (Campos et al., 2019). The separation observed in the present study between the Northern Espinhaço and the CHD block in both NMDS and UPGMA analyses based on localities (Fig. 3) can be mainly attributed to the inclusion of other phytophysiognomies besides Campos Rupestres. In localities such as Jacobina and Licínio de Almeida, more than 50% of the reported species occur in CD areas, whereas in Morro do Chapéu 35% of the reported species are distributed in CT areas, indicating the relevance of the inclusion of other areas in the observed patterns (Table 2). Another factor that could explain the separation of the blocks is that Licínio de Almeida exhibits conditions that differ from areas belonging to the CHD, as it is located in lower elevation areas with a greater presence of Cerrado and Caatinga phytophysiognomies (Campos et al., 2016). Additionally, the average precipitation and temperature in Licínio de Almeida differ from those reported for both the south-central and northern areas of the CHD (Ribeiro, 2022).

    According to the obtained results, the CHD block presented two divisions: the northern area (I) and the central-south area (II) (Fig. 3 B , C), suggesting a possible floristic subdivision of the CHD. Similar findings were reported by Bitencourt & Rapini (2013) and Assunção-Silva et al., (2021), when assessing centers of endemism for Lauraceae and Apocynaceae, respectively. These studies identified areas of endemism in the central-southern region of the CHD that differ in composition from northeastern regions. Additionally, Lucresia et al. (2021) separated Jacobina from other CHD localities using the family Myrtaceae as a model. This north and central-south pattern may result from a combination of factors. One of these factors is the presence of semi-arid depressions with deep valleys bordering the areas of Campos Rupestres in the north (Lobão & Vale, 2008; Stadnik et al., 2016), which could promote greater species diversity from different phytophysiognomies. Another differentiating factor is the distinct environmental conditions, such as low seasonality in rainfall and little variation in average annual temperature in the northeastern CHD (Ribeiro, 2022). Rainfall and temperature are traditionally reported as critical factors for the distribution of diversity in flowering plants, regulating growth, flowering, and pollination processes (Memmott et al., 2007; Lawson & Rands, 2019).

    Furthermore, a different factor that could explain this association between the northern areas of the CHD is the high levels of anthropization and, especially, their high fragmentation. By 2004, Jacobina had already 64% of its natural landscape undergoing anthropization, mainly due to the expansion of the agricultural frontier and mineral extraction (Pinheiro, 2004; Stadnik et al., 2016). A similar situation is observed in Morro do Chapéu, where fragility and fragmentation of the vegetation have been reported as a consequence of deforestation from extensive livestock farming and sand extraction (Lobão & Vale, 2008; Staudt et al., 2017). Although regions of the Central-South CHD also present areas with significant deforestation, it is concentrated in areas such as the Mucugê agricultural pole (Ribeiro, 2022). In contrast, areas of the Chapada Diamantina National Park, where Asteraceae collections are concentrated, are better preserved and exhibit less fragmentation due to anthropogenic effects. To test this, an increased collection effort is needed in the Piemonte da Diamantina area (Campos et al., 2019) as well as evaluations of other vegetation groups to better understand the distribution patterns of diversity throughout the CHD.

    When observing the groupings based on Asteraceae within the CHD block (Fig. 3 C ), we observed that these groups overlap with two of the administrative regions of the state of Bahia, which despite being administrative divisions, are still delimited based on parameters such as economy and culture, reflecting the adaptation of human communities to their natural environment (Rueda-Rodríguez, 2022). The first region, called Piemonte da Diamantina (I; Fig. 3 B , C), includes the municipalities of Caém, Jacobina, Miguel Calmon, Mirangaba, Morro do Chapéu, Ourolândia, Saúde, Serrolândia, Umburanas, and Várzea Nova (Flores, 2014), and is predominantly utilized for livestock farming and agricultural production of beans, corn, and cassava, alongside mineral extraction activities (CODETER, 2017).

    The second administrative region called Chapada Diamantina encompasses 24 municipalities, including Barra da Estiva, Mucugê, and Rio de Contas. Within this area, the principal agricultural products are coffee and vegetables, complemented by touristic activities (SECULTBA, 2003). These social parameters defined in the geopolitical subdivision adopted by the state of Bahia reflect environmental conditions, which directly impact the distribution of plant species. Therefore, it may be possible to suggest a biogeographical subdivision of CHD into two bioregions: the first located in the northern region, called Piemonte da Diamantina, and the second denominated Central-South Chapada, which extends through the central-southern area of CHD. Nevertheless, further studies are necessary to explore not only other taxonomic groups, but different phytophysiognomies beyond Campos Rupestres, to gain a more comprehensive understanding of plant diversity in the CHD.

    The NMDS analysis by phytophysiognomies (Fig. 4 A ) reinforces the grouping of areas within the Northern Espinhaço (squared plus, Fig. 4 B ). However, notable proximity is observed, particularly between the areas of CD and CR (CDL and CRL) with areas of the CHD, reflecting a previously reported floristic similarity between these regions (Campos et al., 2019; Colli-Silva et al., 2019; Alves & Loeuille, 2021; Lucresia et al., 2021).

    One of the observed groups is AA, which could be mainly explained by the presence of ruderal plants. These species show adaptations that facilitate both their dispersion and their establishment in previously disturbed areas (Esquivel, 2015), such as those that extend throughout the CHD (Stadnik et al., 2015; Staudt et al., 2017; Lucresia et al., 2022). The proximity of CDJ to AA, observed in the NMDS (Fig. 4 A ) reinforces a possible state of anthropization in Jacobina (Stadnik et al., 2016), which could be one of the reasons explaining the non-grouping of different phytophysiognomies of the municipality. Furthermore, Jacobina exhibits the lowest species richness among the surveyed areas (Table 2), which could be an additional contributing factor. Therefore, as highlighted by Campos et al. (2019), it is crucial to emphasize Jacobina as an important area for further collections.

    Reinforcing the results obtained from the data by localities, NMDS analysis indicates the existence of two clusters within the CHD: The Central-South Chapada (in blue) and Piemonte da Diamantina (in orange and red), highlighting the proximity between the CD and CR areas within Central-South Chapada, which could be explained by the influence that phytophysionomies such as CD have on CR areas (Loeuille et al., 2015; Campos et al., 2019), however, this proximity relationship was not observed in the Piemonte da Diamantina areas (Fig. 4 A ).

    The UPGMA analysis based on phytophysiognomies (Fig. 4 B ) reinforces the presence of an AA cluster (black triangles, Fig. 4 B ). However, the internal groupings do not seem to follow the patterns described by NMDS based on phytophysiognomies (Fig. 4 A ), or with the locality-based analyses (Fig. 3), which grouped areas from Northern Espinhaço with regions of the CHD, as previously reported in other studies (Campos et al., 2019; Colli-Silva et al., 2019; Alves & Loeuille, 2021; Lucresia et al., 2021). For a better understanding of these patterns, increased plant collections throughout other municipalities within both CHD and Northern Espinhaço are needed (Alves & Loeuille, 2021). A significant portion of floristic studies beyond Asteraceae has been concentrated in the same localities (Azevedo & Van Den Berg, 2007; Watanabe et al., 2009; Bastos & Van Den Berg, 2012; Silva & Wanderley, 2013; Sousa et al., 2013; Hurbath et al., 2016; Stadnik et al., 2015; 2016; Azevedo & Conceição, 2017; Borges et al., 2017; Sampaio et al., 2021; Santos et al., 2021), highlighting the need for broader sampling to enhance our understanding of these biogeographical patterns.

    Although the NMDS and UPGMA analyses segmented by different phytophysiognomies for each locality (Fig. 4) suggest some associations between them, it is important to highlight that the diversity of Asteraceae in CHD, shows high spatial cohesion in their groupings. There is greater similarity observed across different geographical areas, regardless of the analyzed vegetation type. This pattern could be attributed to variations in climatic conditions along the Espinhaço Range (Ribeiro, 2022). However, further investigation is needed to gain a deeper understanding of these results and how other factors influence observed patterns, particularly the dispersal methods of Asteraceae.

    The current research suggests a pattern of limited dispersal over short distances, a phenomenon that has been linked to increased instances of endemism and micro-endemism (Hooper, 2009), as reported in CHD (Giulietti et al., 1987; Colli-Silva et al., 2019). While limited dispersal over short distances may seem unexpected, considering the typical dispersal mechanisms associated with Asteraceae (Keeley et al., 2007; Jeffrey, 2009; Mandel et al., 2019), studies report low dispersal capacity associated with a high correlation with climatic relicts, as well as modifications of the pappus in Brazilian Vernonieae species (Collevatti et al., 2009; Loeuille et al., 2015). Thus, dispersal limitation could be relevant in the plant composition across the CHD, supporting the findings from previous studies that suggest the importance of this factor for plant composition in Campos Rupestres areas of the Espinhaço Range (Pacífico et al., 2021). Further research into these mechanisms will be crucial for a comprehensive understanding of the biogeographical patterns observed in Asteraceae within the region.

    Multiple factors could be affecting the grouping containing CT and F areas. Firstly, the low endemism of Asteraceae within these phytophysiognomies suggests a prevalence of widely distributed species (Hind, 1995; Moura & Roque, 2014; Roque et al., 2016; Staudt et al., 2017) so that vegetations with a high number of exclusive species show greater floristic similarity. Another factor that would be related to the non-grouping of forest areas, is the heterogeneity regarding their delimitation, as the term ‘forest’ encompasses a broad spectrum of vegetations, grouping semi-decidual, evergreen, gallery, Atlantic Forest, and high-altitude forests, as a single phytophysiognomy. Areas categorized simply as ‘forests’ might consist of deciduous or seasonally dry forests, which share affinities with Caatinga areas (Queiroz et al., 2017). In the case of Barra da Estiva, despite the increased plant collections during this study, a need to intensify collection efforts specifically targeting Caatinga and forest areas remains. A focused approach will help clarify the botanical composition and ecological relationships within these diverse phytophysiognomies.

    In conclusion, Barra da Estiva stands out as an important municipality for the study and conservation of Asteraceae diversity. Pointing out that 23% of its species are reported as endemic to Bahia, seven are included in some category of threat, and 83 species have not yet received any conservation status evaluation (IUCN, 2024). At the same time, the present research reinforces the importance of floristic studies, indicating a high spatial cohesion of the composition of Asteraceae in the CHD and suggests a possible subdivision of the CHD in two bioregions: Central-South Chapada and Piemonte da Diamantina. This proposal must be corroborated in further investigations encompassing different plant groups, as well as new collections in under-evaluated areas belonging to the CHD.

    Identification key for Asteraceae species in the municipality of Barra da Estiva, Bahia.

    1. Disciform head (two types of different tubulose florets in the same head) ………………... 2

    1'. Radiate head (heterogamous head with marginal and liguliform florets and central tubular florets) or discoid head (contains only disc florets) …………………………………………... 4

    2. Herbs, with leaves in rosette and scapose, solitary head …………….... Chaptalia integerrima

    2’. Subshrubs, leaves with evident internodes, heads grouped in capitulescences ………….... 3

    3. Involucral bracts papyraceous, yellow ………………………..…… Achyrocline satureioides

    3'. Involucral bracts chartaceous, green ………………………………….... Conyza bonariensis

    4. Radiate head ………………………………………………………………………………... 5

    4'. Discoid head …………………………………………………………………………….... 18

    5. Internal involucral bracts with brown or orange striations ……………………………….... 6

    5'. Involucral bracts with similar aspect in the different series ……………………………….. 7

    6. Florets with corolla usually white, filament glabrous, cypselae obcompressed …………………………………………………………………………………..... Bidens pilosa

    6'. Florets with corolla usually orange, filament pilose, cypselae cylindrical …………………………………………………………………..... Cosmos sulphureus (Fig.6B)

    7. Ray florets neutral (androecium and gynoecium not functional) …………………………... 8

    7'. Ray florets pistillate …………………………………………………………………...….. 11

    8. Involucral bracts 3-6-seriate, internal bracts gradually larger ……........……………..……. 9

    8'. Involucral bracts 2-3-seriate, bracts subequal in length ………………........……………... 10

    9. Leaf blade linear, 1-5 × 0.2-0.5 cm …………………………………………... Aspilia foliosa

    9'. Leaf blade lanceolate, 6-15 × 3-8 cm …………………………………... Aspilia squarrosa

    10.Leaf blade linear or lanceolate, involucral bracts with apex caudate ……….... Aspilia martii

    10'. Leaf blade elliptic or narrow elliptic, involucral bracts with apex acute ……………………………………………………...……………. Aspilia subalpestris (Fig.5E)

    11. Cypselae baccate (fleshy like a berry) …………………………….. Tilesia baccata (Fig. 8E)

    11'. Cypselae not fleshy …………………………………………………………………….... 12

    12. Cypselae winged ................................................................................................................ 13

    12'. Cypselae cylindrical …………………………………………………………………….. 14

    13. Leaf blade elliptic, adaxial surface strigose ……………………….. Verbesina luetzelburgii

    13'. Leaf blade lanceolate or narrowly elliptic, adaxial sericeous only in the veins ……………………………………………………………….... Verbesina glabrata (Fig. 8G-H)

    14. Ray florets white and disc florets yellow ……………...……………... Galinsoga parviflora

    14'. Ray and disc florets with the same color ……………………………………………….. 15

    15. All florets white, pappus aristate ……………………………………..... Blainvillea acmella

    15'. All florets yellow, pappus paleaceous ………………………………………………….... 16

    16. Head with 1-2 foliose bracts at the base, leaf blade ovate …………... Calea villosa (Fig. 5I)

    16'. Head without foliose bracts at the base, leaf blade oblong to elliptic ………………….... 17

    17. Leaf blade chartaceous, head with 28-32 florets, pappus 0.3-0.5 mm .................................. …………………………………………………...………………. Calea candolleana (Fig.5G)

    17'. Leaf blade membranaceous, head with 45-50 florets, pappus 0.6-1.4 mm ... Calea pilosa

    18. Head with only bilabiate florets ……………………………………………. Trixis vauthieri

    18'. Head with tubulose, tubulose-filiform, or ligulate florets ……………………………….. 19

    19. Stems and leaves with white latex, head with only ligulate florets …….... Sonchus oleraceus

    19'. Stems and leaves without latex, head with tubulose or tubulose-filiform florets .............. 20

    20. Leaves and involucral bracts with black resin glands ......................................................... 21

    20'. Leaves and involucral bracts without black resin glands ………………………………... 22

    21. Leaf blade linear …………………………………………………... Porophyllum obscurum

    21'. Leaf blade elliptic ………………………………………..... Porophyllum ruderale (Fig. 7I)

    22. Dioecious plants (distinct unisexual individuals) ………………………………………... 23

    22'. Monoecious (plants with both male and female florets on the same individual) or gynodioecious plants (some plants with bisexual florets and some with pistillate florets) ….. 29

    23. Leaves reduced to scales …………………………………………..... Baccharis orbignyana

    23'. Leaves conspicuous, leaf blade lanceolate, linear, ovate or orbiculate ………………….. 24

    24. Discolorous leaves, adaxial surface glabrous, abaxial white tomentose ………………………………………………………………………….... Baccharis calvescens

    24'. Concolorous leaves, glabrous or pubescents not white tomentose …………………….... 25

    25. Leaves petiolate (petiole > 1 cm. long) …………………………………………………... 26

    25'. Leaves sessile or subsessile (without petiole or maximum with 0.5 cm large) …………. 27

    26. Stem fistulose, leaf blade chartaceous, slightly discolorous, glabrous .... Baccharis pingraea

    26'. Stem with pith, leaf blade membranaceous, strongly discolored, dark green adaxial surface with strigose texture, light green abaxially, trichomes restricted to veins ..... Baccharis serrulata

    27. Leaf blade orbicular (length/width ratio 1:1) ……………………….... Baccharis orbiculata

    27'. Leaf blade elliptical (length/width ratio 2:1) …………………………………………..... 28

    28. Leaves pinnately veined, margin entire on lower half, and 1-5 teeth on upper half ………………………………………………………………………….... Baccharis reticularia

    28'. Leaves 3-veined, margin toothed along the entire length ……….. Baccharis retusa (Fig.5F)

    29. Unisexual heads, staminate head with receptacle paleaceous and pistillate head generally uniflorous and receptacle epaleaceous ……………………………….. Ambrosia artemisiifolia

    29'. Bisexual heads or unisexual in gynodioecious plants ………………………………….... 30

    30. Heads with the same number of florets and involucral bracts ………………………….... 31

    30'. Heads with different numbers of florets and involucral bracts ………………………….. 37

    31. Heads with 5 florets and 5 involucral bracts …………………………………………….. 32

    31'. Heads with 4 florets and 4 involucral bracts …………………………………………….. 33

    32. Alternate leaves, lustrous on both sides, entire margins, capitulescence corymbiform ………………………………………………………………………... Hoehnephytum trixoides

    32'. Opposite leaves, dull on both sides, crenate margin, capitulescence paniculiform ………………………………………………………………………….. Stevia morii (Fig. 8B)

    33. Liana .................................................................................................... Mikania phaeoclados

    33'. Shrubs ................................................................................................................................ 34

    34. Capitulescence thyrsoid ..................................................................................................... 35

    34'. Capitulescence paniculiform or racemiform ………………………………………….... 36

    35. Branches puberulent and green, leaf blade obovate, entire margins ................ Mikania hagei

    35'. Branches velvety, reddish, leaf blade ovate, entire to dentate margins ………………………………………………………………………….... Mikania luetzelburgii

    36. Capitulescence paniculiform, leaf blade entire, revolute ……………….... Mikania obtusata

    36'. Capitulescence racemiform, leaf blade crenate, flat ……………………... Mikania nelsonii

    37. Heads with uniseriate involucre ……………………………………….... Emilia sonchifolia

    37'. Heads with bi - multiseriate involucre ............................................................................... 38

    38. Leaf blade pinnatifid …………………………………….. Verbesina bipinnatifida (Fig. 8F)

    38'. Leaf blade entire ................................................................................................................ 39

    39. Style-branches truncate penicillate or rounded, short bilobed glabrous …………………. 40

    39'. Style-branches elongate papillose or with trichomes ……………………………………. 46

    40. Florets yellow, style-branches truncate, penicillate, pappus paleaceous ................................................................................................................. Calea harleyi (Fig. 5H)

    40'. Florets cream or pink, style-branches rounded, short bilobed, glabrous, pappus of bristles ……………………………………………………………………………………………….. 41

    41. Monoecious subshrub, uniseriate pappus .............................. Richterago discoidea (Fig. 8A)

    41'. Gynodioecious shrubs or trees, pappus 2-3 seriate ……………………………………... 42

    42. Shrub, leaves tomentose on both sides ...................................... Moquiniastrum paniculatum

    42'. Trees, leaves glabrous or glabrescent on adaxially surface and abaxially tomentose …... 43

    43. Head pedunculate (peduncle > 1 cm long) ………………….. Moquiniastrum polymorphum

    43'. Head sessile to subsessile (peduncle < 0.5 cm) ………………………………………….. 44

    44. Capitulescence paniculiform reduced, shorter than the leaves …………………………………………………………………. Moquiniastrum oligocephalum

    44'. Capitulescence paniculiform longer than the leaves …………………………………….. 45

    45. Leaf blade orbicular, heads with 6-9 florets ……... Moquiniastrum blanchetianum (Fig. 7F)

    45'. Leaf blade elliptic, heads with 10-18 florets …………….…. Moquinastrum densicephalum

    46. Style-branches with apex acute and trichomes extending below bifurcation ……………. 47

    46'. Style-branches clavate and papillose, apex obtuse or rounded .......................................... 80

    47. Receptacle strongly alveolate, surrounding the cypselae ………...... Albertinia brasiliensis

    47'. Receptacle not alveolate, if alveolate not surrounding the cypselae .................................. 48

    48. Herbs, shrubs, or subshrubs, with scapose head or capitulescence spiciform or in cymes scorpioid .................................................................................................................................. 49

    48'. Shrubs, subshrubs, trees, or herbs with solitary heads, or grouped in capitulescence glomeruliform, thyrsoid or corymbiform ................................................................................. 64

    49. Pappus biseriate double-paleaceous ................................................................................... 50

    49'. Pappus uniseriate or biseriate with the external series paleaceous and internal series bristle-barbellate, or outer series setose and inner paleaceous ............................................................. 52

    50. Capitulescence spiciform .............................................................. Stilpnopappus tomentosus

    50'. Heads in scorpioid cymes .................................................................................................. 51

    51. Involucral bracts glabrous ................................................................ Stilpnopappus pratensis

    51'. Involucral bracts with white tomentose pubescence ........................ Stilpnopappus scaposus

    52. Heads in two series (subduplicate) in the branches of the capitulescence .............................................................................................................. Cyrtocymura scorpioides

    52'. Scapose solitary heads or organized in one series in capitulescence ................................. 53

    53. Pappus biseriate, outer series setose and inner paleaceous .................... Chrysolaena simplex

    53'. Pappus biseriate with the external series paleaceous and internal series bristle-barbellate .................................................................................................................................................. 54

    54. Heads sessile, less than 1 cm diam., style-base dilated ...................................................... 55

    54'. Heads pedunculate, peduncle bigger than 1cm diam., style-base cylindrical .................... 60

    55. Leaf blade concolorous, sericeous, heads with 35-40 florets ..................... Lepidaploa nitens

    55'. Leaf blade discolorous, glabrous on the adaxial surface, and sericeous matte beneath, heads with 15-30 florets ..................................................................................................................... 56

    56. Leaf blade slightly revolute, florets white .................................................. Lepidaploa hagei

    56'. Leaf blade flat, florets purple ............................................................................................. 57

    57. Heads with inconspicuous foliar bracts (shorter than the heads) .................................................................................................... Lepidaploa chalybaea (Fig. 6E)

    57'. Heads with conspicuous foliar bracts (longer than the heads) ........................................... 58

    58. Leaf blade papyraceous, white puberulent on abaxial surface ........... Lepidaploa araripensis

    58'. Leaf blade chartaceous leaves, grayish tomentose on abaxial surface ............................... 59

    59. Leaf blade ovate, abaxially tomentose, heads with 20-30 florets .................................................................................................. Lepidaploa cotoneaster (Fig. 6F)

    59'. Leaf blade elliptic, abaxially grayish pubescent, heads with 22-25 florets .................................................................................................................. Lepidaploa salzmannii

    60. Herbs in rosette, scapose head ............................................................................................ 61

    60'. Shrubs, leaves alternate with visible internodes, capitulescence in scorpioid cymes ........ 62

    61. Leaf blade discolorous, adaxially villous, and abaxially densely pubescent .................................................................................................. Lessingianthus santosii (Fig. 7C)

    61'. Leaf blade concolorous, glabrous ................................... Lessingianthus carvalhoi (Fig. 6G)

    62. Leaf blade linear to aciculate ............................................. Lessingianthus linearis (Fig. 7A)

    62'. Leaf blade ovate to elliptic ................................................................................................. 63

    63. Leaf blade discolorous, green on adaxial surface, yellow underside, margins entire and slightly revolute, floccose on both sides (with white trichomes that come off in tufts when rubbed) .................................................................................. Lessingianthus farinosus (Fig. 6H)

    63'. Leaf blade concolorous, adaxially glabrous, and abaxially puberulent, margins dentate and flat ............................................................................................... Lessingianthus morii (Fig. 7B)

    64. Heads with orange pappus.................................................................................................. 65

    64'. Heads with white or stramineous pappus ........................................................................... 67

    65. Leaf blade hyphodromous, margins strongly revolute ....................... Echinocoryne pungens

    65'. Leaf blade camptodromous, margins flat to slightly revolute ............................................ 66

    66. Involucral bracts dark green-vinaceous ........................... Echinocoryne holoserica (Fig. 6C)

    66'. Involucral bracts pale greenish-yellow ................................................. Echinocoryne stricta

    67. Trees, shrubs, and subshrubs with imbricate leaves, grouped at the apex of the branches and the presence of punctate leaf scars ............................................................................................ 68

    67'. Herbs, shrubs, subshrubs, or trees with lax leaves, if imbricate without punctate leaf scars along the branches .................................................................................................................... 75

    68. Stems with pad-like sheath (with a small bulge where the petiole inserts into the stem) .................................................................................................................................................. 69

    68'. Stems without pad-like sheath ........................................................................................... 71

    69. Leaf blade with decurrent base, cypselae strigose ........................... Lychnophorella bishopii

    69'. Leaf blade with subcordate base, cypselae glabrous .......................................................... 70

    70. Leaf blade ovate, heads with 1-3 florets ............................................... Lychnophorella morii

    70'. Leaf blade narrowly lanceolate, heads with 3-4 florets ..... Lychnophorella triflora (Fig. 7E)

    71. Subshrub bromeliform (basal leaves in rosette) .................. Lychnophora uniflora (Fig. 7D)

    71'. Subshrubs or shrub with alternate leaves and visible internodes ....................................... 72

    72. Heads with 4-9 florets ........................................................................................................ 73

    72'. Heads with 1-3 florets ........................................................................................................ 74

    73. Leaf blade linear, longer than 5 cm long, apex obtuse ...................... Lychnophora salicifolia

    73'. Leaf blade deltate or narrow-elliptic, 1-3 cm long, apex mucronate or acute ......................................................................................................... Lychnophora rosmarinifolia

    74. Base of the leaves with white trichome tufts, broquidodromous venation ...................................................................................................... Lychnophora granmogolensis

    74'. Base of the leaves without white trichome tufts, hyphodromous venation .............................................................................................................. Lychnophora phylicifolia

    75. Trees with heads grouped in syncephaly ............................................................................ 76

    75'. Herbs, shrubs, subshrubs, or trees without syncephaly ...................................................... 77

    76. Heads with 2-3 florets ..................................................... Paralychnophora bicolor (Fig. 7G)

    76'. Heads with 5-12 florets .................................................. Paralychnophora harleyi (Fig. 7H)

    77. Solitary heads with one series of foliose involucral bracts at the base .............................................................................................. Centratherum punctatum (Fig. 6A)

    77'. Heads grouped in capitulescence ....................................................................................... 78

    78. Heads with 8-60 florets ................................................................ Vernonanthura polyanthes

    78'. Heads with 1-6 florets ........................................................................................................ 79

    79. Subshrubs, leaves sessile, leaf blade linear ................................... Stenocephalum monticola

    79'. Trees, leaves petiolate, leaf blade elliptic ............................ Eremanthus capitatus (Fig. 6D)

    80. Style-branches villous below the bifurcation ................. Stylotrichium glomeratum (Fig. 8C)

    80'. Style-branches papillose, glandular, or glabrous below the bifurcation ............................ 81

    81. Receptacle paleaceous, pappus absent, when present, pappus aristate, coroniform or coroniform-aristate .................................................................................................................. 82

    81'. Receptacle epaleaceous, pappus bristle-barbellate or paleaceous-aristate ......................... 84

    82. Leaves sessile or subsessile (petiole shorter than 0.5 cm), glabrous ...................................................................................................... Acritopappus harleyi (Fig.5B)

    82'. Leaves petiolate with tomentose or glandular indument .................................................... 83

    83. Leaf blade abaxially tomentose, heads with 14-25 florets ............................................................................................... Acritopappus heterolepis (Fig.5C)

    83'. Leaf blade abaxially glandular, heads with 5-10 florets ..... Acritopappus confertus (Fig.5A)

    84. Heads with 58 or more florets, pappus paleaceous-aristate…………... Ageratum conyzoides

    84'. Heads with up to 52 florets, pappus bristle barbellate …………………………………... 85

    85. Congested leaves (short internodes) grouped towards the apices of the branches ….….... 86

    85'. Leaves lax (not congested) …………………………………………………….………... 90

    86. Leaf blade strongly discolorous, branches, abaxial leaf blade, and involucral bracts sericeous ………………………………………………………………….... Lasiolaena duartei

    86'. Leaf blade concolorous or slightly discolorous, branches, leaf blade, and involucral bracts glabrous ................................................................................................................................... 87

    87. Cypselae with pappus biseriate ....................................................... Agrianthus empetrifolius

    87'. Cypselae whit pappus uniseriate ........................................................................................ 88

    88. Capitulescence umbelliform ............................................. Agrianthus leutzelburgii (Fig.5D)

    88'. Capitulescence corymbiform ............................................................................................. 89

    89. Leaf blade tomentose, margins serrate ............................................... Agrianthus almasensis

    89'. Leaf blade glabrous, margins entire or only serrate towards the apex .................................................................................................................... Agrianthus carvalhoi

    90. Leaves resinous, pappus biseriate ....................................................................................... 91

    90'. Leaves not resinous, pappus uniseriate .............................................................................. 92

    91. Leaf blade lanceolate ................................................................ Symphyopappus compressus

    91'. Leaf blade orbiculate ................................................... Symphyopappus decussatus (Fig.8D)

    92. Heads with 30-36 involucral bracts .................................................................................... 93

    92'. Heads with 7-20 involucral bracts ...................................................................................... 95

    93. Leaves alternate ......................................................................... Chromolaena cinereoviridis

    93'. Leaves opposite ................................................................................................................. 94

    94. Leaf blade lanceolate, margins strongly serrate, base cuneate, lustrous, glabrous ................................................................................................................ Chromolaena laevigata

    94'. Leaf blade ovate, margins serrate, base rounded, opaque, glabrous or with short strigose trichomes ...................................................................................................... Chromolaena morii

    95. Leaves alternate .................................................................................................................. 96

    95'. Leaves opposite ................................................................................................................. 97

    96. Heads with 18-25 florets, corolla lobe pubescent, pappus plumose, receptacle convex ................................................................................................................. Trichogonia salviifolia

    96'. Heads with 25-35 florets, corolla lobe glabrous, pappus of bristles receptacle conical ............................................................................................................. Conocliniopsis prasiifolia

    97. Leaf blade abaxially pubescent, florets white.............................. Koanophyllon adamantium

    97'. Leaf blade glabrous or pubescent in the veins, florets pink-purplish......Ayapana amigdalina

    thumbnail of Figure 5.
    Figure 5.
    A - Acritopappus confertus; B - Acritopappus harleyi; C - Acritopappus heterolepis; D - Agrianthus luetzelburgii; E - Aspilia subalpestris; F - Baccharis retusa; G - Calea candolleana; H - Calea harleyi; I - Calea villosa. (A, C, E, F - D. Morales; B - L. Lucresia; D, E - G.V. Fonseca; G - N. Roque; I - D. Remor)

    thumbnail of Figure 6.
    Figure 6.
    A - Centratherum punctatum; B - Cosmos sulphureus; C - Echinocoryne holosericea; D - Eremanthus capitatus; E - Lepidaploa chalybaea; F - Lepidaploa cotoneaster; G - Lessingianthus carvalhoi; H - Lessingianthus farinosus. (A, B, E - D. Morales; C, G, H - G. V. Fonseca; D - L. Lucresia; F - D. Remor).

    thumbnail of Figure 7.
    Figure 7.
    A - Lessingianthus linearis; B - Lessingianthus morii; C - Lessingianthus santosii; D - Lychnophora uniflora; E - Lychnophorella triflora F - Moquiniastrum blanchetianum; G - Paralychnophora bicolor; H - Paralychnophora harleyi; I - Porophyllum ruderale (A, C - G.V. Fonseca; B, G - D. Morales; D, F - N. Roque; E, H, I - D. Remor).

    thumbnail of Figure 8.
    Figure 8.
    A - Richterago discoidea; B - Stevia morii; C - Stylotrichium glomeratum; D - Symphyopappus decussatus; E - Tilesia baccata; F - Verbesina bipinnatifida; G, H - Verbesina glabrata. (A - D. Remor; B, C - L. Lucresia; D - G. Guerra; E, F, G, H - D. Morales).

    Acknowledgments

    We thank the Universidade Estadual de Feira de Santana for supporting the development of this research. This study was partially funded by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Brasil (CAPES) - Finance Code 001. DM and NR are grateful to CAPES for their granted fellowships (88887.631120/2021-00).

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    Edited by

    • Associate Editor:
      Annelise Frazão
    • Editor Chef:
      Thais Almeida

    Publication Dates

    • Publication in this collection
      07 July 2025
    • Date of issue
      2025

    History

    • Received
      25 Aug 2023
    • Accepted
      20 Apr 2025
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    This is an open-access article distributed under the terms of the Creative Commons Attribution License
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