Distribution patterns, Diversity, and Endemism of Melochia L. (Byttnerioideae – Malvaceae) in Brazil

Abstract

Melochia (Byttnerioideae – Malvaceae) is widely distributed across tropical and subtropical regions. In Brazil, however, its distribution, species richness, diversity, and endemism remain poorly understood. The present study aimed to analyze the biogeography of Melochia and infer which distribution patterns best reflect its geographic distribution in Brazil and where its centers of diversity and endemism are located. Species occurrence data were compiled from online databases, resulting in 4,136 cleaned records representing 27 species. Richness and diversity analyses were conducted using DIVA-GIS, while QGIS was used to create the distribution maps. Parsimony Analysis of Endemicity was performed with PAUP. Our results indicate that most Melochia species are widely distributed. Seven areas of diversity were identified: Pantanal, Central Brazilian Cerrado, Northern and Southern Espinhaço Range, Serra dos Carajás, Tapajós, and Raposa Serra do Sol Indigenous Land. Two areas of endemism were detected: Chapada Diamantina, and Chapada do Araripe. These patterns highlight the ecological relevance of Melochia across varied environmental gradients and its biogeographical structure, particularly in the Central-West and Espinhaço Range. The study also reveals significant knowledge gaps and underrepresentation of certain species in herbarium collections and databases, stressing the need for increased botanical exploration and habitat protection.

Key words
Biogeography; Brazilian Flora; Conservation; Endemism

INTRODUCTION

Understanding plant diversity and distribution is essential for addressing the challenges of a rapidly changing world. Studies show that global plant biodiversity is largely concentrated in hotspots, areas with high species richness and endemicity and elevated threat levels, such as the tropical forests of Brazil, Africa, and Southeast Asia (Brummitt et al. 2021, Daru 2024). In contrast, globalization, land-use change, and climate change are intensifying pressures on these ecosystems, increasing extinction rates and reducing ecological resilience (Wang et al. 2024).

Studies such as those carried out by Brummitt et al. (2021) and Ondo et al. (2024) highlight the importance of integrating data from different sources, such as herbaria and digital platforms (GBIF, Flora e Funga do Brasil, SpeciesLink) to map diversity patterns, prioritize conservation efforts and guide effective environmental policies in under- represented regions (darkspots), such as the interior of the Amazon and parts of Central Africa, which have significant Linnean (lack of knowledge about species) and Wallacean (lack of knowledge about geographical distributions) deficits.

In addition, surveying species distributions helps to identify ecological and evolutionary processes that sustain biodiversity, as well as understanding patterns of abundance and rarity in order to predict the impacts of anthropogenic changes and direct public policies that promote the sustainable use of natural resources (Callaghan et al. 2023).

Brazil is one of the world’s most megadiverse countries, hosting major biodiversity hotspots such as the Atlantic Forest and the Cerrado, which harbor high levels of endemism and species richness (Myers et al. 2000, Oliveira et al. 2017). Brazil’s continental dimensions, combined with wide latitudinal and climatic variation, favor a high rate of flora diversification, making the country a global center of plant diversity with more than 50,000 species, about 17,000 of them endemic (Vilaça et al. 2024). Yet major gaps in conservation and knowledge persist: roughly 55% of Brazilian species and 40% of evolutionary lineages remain outside protected areas, and most species have less than 30% of their ranges safeguarded (Oliveira et al. 2017).

Brazil harbors over 20% of global Malvaceae diversity (about 890 of 4,250 species), more than half of them endemic and 50 of which are listed as nationally endangered (Christenhusz & Byng 2016, MMA 2022, Flora e Funga do Brasil 2025). The family occurs across all phytogeographic dominions and plays a key ecological role by providing food and shelter for fauna and sustaining ecosystem services such as pollination and nutrient cycling (Torquato et al. 2024).

Melochia L. (Malvaceae) is a genus of mostly herbs and subshrubs, rarely trees, characterized by a gynoecium with a 5-carpellate, uni- to biovulate ovary and five papillate stipes (Goldberg 1967). The genus includes 65 species distributed across tropical and subtropical regions of the Americas, Africa, Asia, and Oceania (POWO 2025). In Brazil, it comprises 24 taxa, six of them endemic, occurring mainly in the Cerrado and in transition zones between the Caatinga and Atlantic Forest (Gonçalez & Coutinho 2020). Although revised by Goldberg in 1967, its diversity, richness, and endemism in Brazil remain poorly documented.

Its species holds economic and ecological importance due to a diverse medicinal and bioactive properties. Several species, including Melochia corchorifolia L., M. hermannioides A.St.-Hil., M. odorata L.f., and M. umbellata (Houtt.) Stapf, show compounds with potential value against cancer, as contraceptives, and in anti-HIV P24 and anti-dengue applications (Arenas & Azorero 1977, Jadulco et al. 2014, Soekamto et al. 2020, Yuan et al. 2019). Despite this relevance, no Brazilian species has had its conservation status assessed, underscoring the limited knowledge of their distributions.

This study aims to investigate the distribution patterns, richness, and diversity of Melochia species in Brazil and to identify areas of endemism with emphasis on regions of high diversity and priority for species conservation. From this perspective, we sought to answer the following questions: I – How many species of Melochia exist in Brazil and which is the distribution of each one? II – Where are the centers of diversity and endemism for the genus in Brazil?

MATERIALS AND METHODS

The geographical coordinates of Melochia specimens were obtained from three main online databases: Global Biodiversity Information Facility (https://www.gbif.org/), CRIA-speciesLink (http://www.splink.org.br/) and Reflora Virtual Herbarium (http://floradobrasil.jbrj.gov.br/). Only specimens identified by specialists in the genus were included. Cleaning and validation were performed manually. The species considered in the analysis were based on the taxonomic revision by Goldberg (1967), supplemented by updates to the Flora e Funga do Brasil (2025) and POWO (2025) databases, the latter providing details on species described after the last revision.

For records without original geographic coordinates, the locality or municipality information from the specimen labels was used, and their coordinates were assigned using the geoLoc tool (accessible from the Environmental Information Reference Center website at SpeciesLink – http://splink.cria.org.br/geoloc). When only the municipality or locality was indicated on the specimen label, the generated coordinates corresponded to the official geographic position of the municipal seat rather than to the geometric centroid of the municipality. This approach is commonly adopted in biodiversity studies when precise locality descriptions are unavailable, as it provides a standardized and reproducible reference point while minimizing spatial uncertainty (Chapman & Wieczorek 2006). Therefore, the inferred coordinates were used exclusively to represent approximate occurrences.

Records that could not be georeferenced to at least the municipal level, along with duplicates, dubious identifications or those without collector numbers, were excluded. After data cleaning, the final dataset comprised 4,136 records belonging to 27 species. Species nomenclature followed the standards of The International Plant Names Index (https://www.ipni.org/), Tropicos (https://www.tropicos.org/home), and Flora e Funga do Brasil (2025).

Recent syntheses indicate that Brazil’s current biogeographic configuration is linked to Cenozoic climatic shifts, particularly the fragmentation of once-continuous tropical forests and the establishment of the so-called “dry diagonal” (Cerrado–Caatinga–Pantanal), which acts both as a barrier and a corridor for species dispersal. Despite this fragmentation, intermittent connections between the Amazon and Atlantic forests likely persisted over time, contributing to lineage exchange and diversification across dominions (Della & Prado 2025).

To classify the distribution patterns of Melochia species in Brazil, we used Area of Occupancy (AOO) as a quantitative metric. We estimated AOO from occurrence records for each individual species with GeoCAT following the IUCN (2025) protocol. AOO offers clear and comparable spatial criteria across species and supports the definition of three distribution categories: (1) widely distributed (WD, AOO > 20,000 km²), when the species occurs homogeneously within the polygon; (2) disjunct (DD, variable AOO), when the distribution shows large gaps (> 200 km) or ecological barriers; and (3) restricted (RD, AOO < 20,000 km²), when the species occurs in a single region or phytogeographic dominion.

The choice of grid cell size plays a crucial role in biogeographic analyses, as it significantly influences the results (e.g., Morrone & Escalante 2002, Contreras-Medina & Luna-Vega 2007, Garraffoni et al. 2017). In this study, grid cell sizes 1° × 1° (approximately 100 km x 100 km) and 2° × 2° (approximately 200 km x 200 km) were chosen for analyses of richness, diversity and PAE. These grids were created using DIVA- GIS software (Hijmans et al. 2001).

The richness (Jackknife1 Index: Hellmann & Fowler 1999) and diversity analysis was carried out using DIVA-GIS software (Shannon Index: Magurran 1988), while QGIS software (QGIS Development Team 2021) was used to build the maps and visualize the distribution of the species. The preliminary conservation status follows the IUCN Red List Categories and Criteria (2025). The GeoCAT tool (Bachman et al. 2011) was used to estimate the extent of occurrence (EOO) and area of occupancy (AOO).

PAE is a method developed by Rosen (1988) and Rosen & Smith (1988) that can be applied statically or dynamically (Nihei 2006). In the static approach used in this study, the method is restricted to a single geological era, representing the present. Smaller grids tend to generate less resolved cladograms of areas, while larger grids favor the detection of broader areas of endemism.

Following Morrone’s methodology (1994), we created grid cells across Brazil and included only those that contained at least one species record. We then generated species-distribution maps in QGIS (QGIS Development Team 2021). Based on these records, we produced a presence–absence matrix in Mesquite (version 3.16; Maddison & Maddison 2021). In this matrix, columns represent species and rows represent grid cells, with each taxon coded as 1 for presence and 0 for absence. We added a hypothetical area with no species as an outgroup to root the cladogram.

In this study, areas of endemism were only considered when the relationships between the grids were supported by the occurrence of two or more exclusive species (synapomorphies) (Garraffoni et al. 2017, Assunção-Silva & Assis 2022). The analysis was carried out in the PAUP* 4.0 software (Swofford 2003), to infer the centers of endemism and reconstruct the most parsimonious tree, using a heuristic algorithm with 1,000 replications from random additions and TBR (tree- bisection-reconnection) branch swapping, saving two trees per replication; the same commands were used for bootstrap analysis, using a traditional search with 1,000,000 replications (Costa et al. 2024). This approach allows the exploration of different levels of endemism in different regions, providing a more comprehensive view of biogeographic distribution.

RESULTS

Geographical distribution patterns, richness and diversity of Melochia L.

The survey of 4,136 occurrence records for 27 Melochia species shows a predominance of widely distributed taxa, such as M. parvifolia, M. pyramidata, and M. spicata. These species occupy all biogeographical dominions, which reflects broad ecological tolerance and effective dispersal. Disjunct distributions likely reflect historical and ecological processes. M. nodiflora, for instance, remains associated with wet environments, including the Amazon, humid high-altitude forests (brejos de altitude) in Ceará, and Atlantic Forest remnants in Rio de Janeiro, which highlights its persistence in mesophytic refuges. In contrast, M. morongii occurs in seasonally dry or flood-prone areas—such as the Pantanal, Cerrado regions of Goiás and Minas Gerais, western São Paulo, and the campos rupestres of the Chapada Diamantina—suggesting an affinity for open habitats with strong fluctuations in water availability.

Restricted species, such as M. argentina and M. lanceolata, which occur in the Cerrado and Amazon, likely show this pattern due to habitat loss, sampling gaps, or limited dispersal. Endemic species, including M. betonicifolia and M. longidentata, tolerate a broader range of environments and occur in the Caatinga as well as in areas adjacent to the Cerrado and Atlantic Forest. In contrast, M. lanata and Melochia cristobaliae, endemic to quartzitic outcrops of the Espinhaço Range and to the Northeastern Atlantic Forest, show marked edaphoclimatic specialization that promotes microendemism. This specialization increases their vulnerability to human impacts and climate change, both common pressures in these ecosystems. The distribution maps for all species are shown in Figures 4 to 8.

For the richness analysis 460 grids 1°× 1° and 171 grids 2°× 2° were generated (Fig. 1) and the areas of greatest richness are represented by the squares A276, A283, and A336 in the smallest grid size (with 14 shared species, five of which are exclusive), and A113, A117, A129, A132, A133, and A143 in the largest grid size (with 18 shared species, seven of which are exclusive). In both analyses, the areas with the highest Melochia richness are the northern (A276 in the smallest grid size and A113 in the largest grid size) and southern (A336 in the smallest grid size and A132, A133 and A143 in the largest grid size) portions of the Pantanal, the Cerrado of Central Brazil (A283 in the smallest grid size and A117 in the largest grid size) and the southern portion of the Espinhaço Mountain Range (A129 only in the largest grid size). The difference between the analyses lies in the appearance of an area of richness in the south of the Espinhaço Range in the larger grid size. We will therefore use the 2nd grid to discuss the results obtained here.

Figure 1
Melochia richness map. a. Analysis with grid 1°× 1°. b. Analysis with grid 2°× 2°. In both analyses, the main centers of Melochia richness in Brazil are the Central-West region and the southern portion of the Espinhaço Range.

Like the richness analysis, the diversity analysis (Figure 2) generated 460 grids in size 1°× 1° and 171 grids in size 2°× 2°. The greatest diversity in grid size 1°× 1° was found in grids A4 and A9, in the Raposa Serra do Sol Indigenous Land (Roraima); A42, in the Tapajós National Forest (FLONA) and A102, near the Serra dos Carajás (Pará); A236, in the northern portion of the Espinhaço Range (Bahia); A265, on the border between the states of Goiás and Bahia; A319 and A334, southern portion of the Espinhaço Range (MG); A277, A293, A294, and A309, northern portion of the Pantanal (Mato Grosso), and A337, A338, A340, A355, and A372, southern portion of the Pantanal and its surroundings (Mato Grosso do Sul). In the grid size 2°× 2° some areas coincided with those of a smaller grid size, such as the Raposa Serra do Sol Indigenous Land (A1); Tapajós FLONA (A22 and A23); Serra dos Carajás (A37, A50, and A51); Northern portion of the Espinhaço Range (A97 and A108); Southern portion of the Espinhaço Range (A129 and A138); Northern portion of the Pantanal (A112 and A113); Southern portion of the Pantanal and its surroundings (A132, A133, A134, A142, A143, and A144) and a distinct area emerged on the larger grid corresponding to the Cerrado of Central Brazil (A93 - on the border between the states of Tocantins and Mato Grosso; A104 - east of Mato Grosso; A107 - on the border between the states of Bahia and Goiás; A115 - on the border between the states of Goiás and Mato Grosso and A126 - on the border between Goiás and São Paulo). All these areas had a Shannon H’ index between 1.599 and 2.000.

Figure 2
Melochia diversity map. a. Analysis with grid 1°× 1°. b. Analysis with grid 2°× 2°. In both analyses, the main centers of Melochia diversity in Brazil are: Northern and Southern Portion of the Espinhaço Range, Pantanal, Raposa Serra do Sol Indigenous Land, Serra dos Carajás, Tapajós FLONA, and Cerrado of Central Brazil (only in the analysis with grid 2°).

The main difference between the analyses was the appearance of an area of diversity in the central portion of the Cerrado, not seen in the smaller grid size. Thus, the results indicate seven centers of diversity: the Pantanal, the main and largest center, the northern portion of the Espinhaço Range, the southern portion of the Espinhaço Range, the Cerrado of Central Brazil, Serra dos Carajás, Tapajós and the Raposa Serra do Sol Indigenous Land. As with the richness analysis, the largest grid will be used for discussion purposes.

Parsimony analysis of endemicity (PAE)

The PAE analysis generated 85 grid cells 1°× 1° and 36 grid cells 2°× 2°, which represent possible areas of Melochia endemism in Brazil.

For the matrix 1°× 1°, the PAE produced 5,000 equally parsimonious trees, with CI (consistency index) = 0.58 and RI (retention index) = 0.81, based on six parsimony informative characters. The strict consensus tree (Figure 3a) revealed great basal polytomy for most areas and of the remaining 20 quadrants only two formed clades: Chapada do Araripe and Eastern of Sergipe (CAES), Chapada Diamantina and Ecotonal Area of the Eastern of Bahia (CDEB). Both clades showed a 62.9% bootstrap and two or more shared species and can be considered areas of endemism.

Figure 3
Strict consensus tree from the parsimony analysis of endemism in grids. a. 1° × 1° and b. 2° × 2°. The numbers below the lines indicate bootstrap proportions. Two areas of endemism were recovered in the PAE: Chapada do Araripe and eastern Sergipe and Chapada Diamantina and eastern Bahia.

For the 2°× 2° matrix, the PAE produced 5,000 equally parsimonious trees, with CI (consistency index) = 0.55 and RI (retention index) = 0.87, based on six parsimony information characters. The strict consensus tree (Figure 3b) also revealed great basal polytomy for most areas and of the remaining 12 quadrants only two formed clades: CAES and CDEB.

In both analyses, the results were very similar with a large basal polytomy and the formation of only two clades that can be considered areas of endemism. The difference is that in the 2nd matrix the areas of endemism are larger. Therefore, considering that there was no significant difference in both analyses, we will discuss the areas of endemism recovered in the 2nd analysis: CAES and CDEB.

The first area corresponds to the ecotonal areas of southern Ceará, in the Chapada do Araripe, and eastern Sergipe, formed by grid groups A25 and A31 (bootstrap = 91.1%), with two exclusive species and the second area is in the Chapada Diamantina and eastern Bahia in the Caatinga and Atlantic Forest contact area, formed by grid groups A16, A17, A23, and A32 (bootstrap = 50.9%), also with two exclusive species.

Figure 4
Distribution maps of Melochia species. a. M arenosa (WD). b. M argentina (RD). c. M betonicifolia (RD). d. M chamaedrys (WD). e. M. cristobaliae (RD). f. M gardneri (RD).

Figure 5
Distribution maps of Melochia species. a. M graminifolia (WD). b. M hassleriana (RD). c. M hermannioides (RD). d. M illicioides (RD). e. M kerriifolia (RD). f. M lanata (RD).

Figure 6
Distribution maps of Melochia species. a. M lanceolata (RD). b. M longidentata (RD). c. M melissifolia (WD). d. M morongii (DD). e. M nodiflora (DD). f. M parvifolia (WD).

Figure 7
Distribution maps of Melochia species. a. M pilosa (WD). b. M pyramidata (WD). c. M sergipana (RD). d. M simplex (WD). e. M siphonandra (DD). f. M spicata (WD).

Figure 8
Distribution maps of Melochia species. a. M splendens (WD). b. M tomentosa (WD). c. M ulmifolia (WD).

DISCUSSION

Geographical distribution analysis and conservation status

Analysis of the distribution patterns of Melochia species in Brazil revealed that ecological factors associated with the heterogeneity of phytogeographic dominions influence spatial distribution, ranging from species adapted to the most diverse environmental conditions to tolerant species. Based on Table I, three main categories of species distribution are observed: widely distributed (WD), disjunct distribution (DD), and restricted distribution (RD).

Table I
Distribution patterns of Melochia species with associated vegetation AF: Atlantic Forest; AM: Amazon; CA: Caatinga; CE: Cerrado; PP: Pampa; PT: Pantanal; DD: Disjunct Distribution; RD: Restrict Distribution; WD: Widely Distribution.

Species	Number of grids	Phytogeographical Domains	Distribution Pattern
Melochia arenosa Benth.	64	AF, AM, CA, CE, PT	WD
Melochia argentina R.E.Fr.	3	EC	RD
Melochia betonicifolia A.St.-Hil.	30	AF, CA, CE	RD
Melochia chamaedrys A.St.-Hil.	14	CE, AF, PP	WD
Melochia cristobaliae F.C.P. Costa	2	AF	RD
Melochia gardneri Sprague	7	CA, CE	RD
Melochia graminifolia A.St.-Hil.	37	AF, AM, CE, PP, PT	WD
Melochia hassleriana Chodat	6	AF, CE	RD
Melochia hermannioides A.St.-Hil.	5	CE, PP	RD
Melochia illicioides K.Schum.	4	AF, CA	RD
Melochia kerriifolia Triana & Planch.	4	CE, PT	RD
Melochia lanata A.St.-Hil.	2	CE	RD
Melochia lanceolata Benth.	3	AM	RD
Melochia longidentata Goldberg	9	CA, CE	RD
Melochia melissifolia Benth.	35	AF, AM, CA, CE	WD
Melochia morongii Britton	15	AM, CA, CE, PT	DD
Melochia nodiflora Sw.	4	AF, AM, CA	DD
Melochia parvifolia Kunth	54	AF, AM, CA, CE, PP, PT	WD
Melochia pilosa (Mill.) Fawc. & Rendle	54	AF, AM, CA, CE, PP, PT	WD
Melochia pyramidata L.	82	AF, AM, CA, CE, PP, PT	WD
Melochia sergipana Monteiro	3	AF, AM, CA	RD
Melochia simplex A.St.-Hil.	39	AF, AM, CE, PT	WD
Melochia siphonandra (Turcz.) Dorr	7	AM, CA	DD
Melochia spicata (L.) Fryxell	116	AF, AM, CA, CE, PP, PT	WD
Melochia splendens A.St.-Hil. & Naudin	28	AF, AM, CA, CE	WD
Melochia tomentosa L.	46	AF, AM, CA, CE, PT	WD
Melochia ulmifolia Benth.	31	AM, CE, PT	WD

Widely distributed species, such as M. parvifolia, M. pyramidata, and M. spicata (Figs. 6f, 7b, f) occurs predominantly in seasonally dry open habitats of the Caatinga and Cerrado dominions, disturbed areas, and forest edges, which shows their tolerance to poor soils and limited water. This pattern matches that found in other Malvaceae genera, including Gaya, Helicteres, and Wissadula, dominant in Brazilian savannas and grasslands (Cristóbal 2001, Bovini & Baumgratz 2016, Takeuchi & Esteves 2017).

On the other hand, disjunct species such as M. morongii and M. nodiflora (Figs. 6d-e) form isolated population nuclei between biomes, which suggests that rivers, mountain ranges, forest gaps, and Quaternary paleoclimatic cycles shaped their current ranges (Kadereit & Abbott 2021, Guayasamin et al. 2024). Similar disjunctions occur in Helicteres baruensis Jacq. and Wissadula contracta (Link) R.E.Fr. (Cristóbal 2001, Bovini & Baumgratz 2016). H. baruensis occupies two main blocks: one in eastern Brazil from the Northeast to the Southeast, and another in Roraima. W. contracta shows the same pattern, with a southern extension into the South region.

Most restricted species, such as M. argentina, M. kerriifolia, and M. lanceolata, occur in the Chacoan and southeastern Amazonian domains, within the Chaco, Cerrado, Pampeana, and Xingu–Tapajós provinces (Morrone et al. 2022). This pattern also appears in other Malvaceae genera, including Gaya and Wissadula, and in unrelated plant groups (Schneider et al. 2013, Bovini & Baumgratz 2016, Takeuchi & Esteves 2017, Bezerra et al. 2019). Hysterionica nidorelloides (Asteraceae) and Rhynchosia lineata Benth. (Fabaceae) show a distribution similar to that of M. hermannioides in southern Brazil; Gaya bordasii Krapov., Rhynchosia balansae Micheli, and Wissadula indivisa R.E.Fr. mirror the range of M. argentina; and Wissadula glechomifolia (A.St.-Hil.) R.E.Fr. overlaps with M. chamaedrys (Bovini & Baumgratz 2016, Takeuchi & Esteves 2017, Bezerra et al. 2019).

Restricted species, endemic to Brazil, such as M. betonicifolia, M. illicioides, and M. longidentata, occur mainly in the nuclei of seasonally dry forests and woodlands (SDTFW) within the Caatinga and in ecotonal zones between the Caatinga-Cerrado and the Caatinga-Atlantic Forest. The Caatinga represents the largest continuous SDTFW area of the Neotropical region and harbors numerous endemic species of plants such as Anadenanthera colubrina (Vell.) Brenan, Amburana cearensis (Allemão) A.C.Sm., Gymnanthes boticario Esser, M. F. A. Lucena & M. Alves, and Pseudalbizzia inundata (Mart.) E.J.M.Koenen & Duno (Moro et al. 2016, Queiroz et al. 2017, Moro et al. 2024). According to Fernandes et al. (2019), despite its diversity and endemism, the Caatinga is neglected, with only 1.2% of its area included in protected areas, making its biodiversity vulnerable. These regions also act as endemism centers for Helicteres (Malvaceae), Petalostelma (Apocynaceae), and Pfaffia (Amaranthaceae), which reinforces the idea that edaphic and topographic gradients promote local speciation and sustain regional endemism (Cristóbal 2001, Marchioretto et al. 2010, Santos et al. 2021).

The microendemic M. lanata and Melochia cristobaliae occur in small areas of the Espinhaço Range and the Atlantic Forest, respectively, which reflects their specialization to micro-habitats such as rocky fields. This pattern is consistent with the levels of plant endemism documented for campo rupestre vegetation, where a large number of species are restricted to isolated mountaintops associated with quartzitic and sandstone outcrops and nutrient-poor soils, such as Vellozia spp. (Velloziaceae), Paepalanthus spp. (Eriocaulaceae), Barbacenia spp. (Velloziaceae), and Lychnophora spp. (Asteraceae) (Colli-Silva et al. 2019). This pattern matches evidence from multiple taxa—including Gaya xiquexiquensis C. Takeuchi & G.L. Esteves, Oxalis diamantinae Knuth, Petalostelma atlanticum A.P.B. Santos, and Pfaffia rupestris Marchioretto—that identify the Atlantic Forest and the Espinhaço Range as microendemism hotspots in Brazil, characterized by species with narrow ranges and high vulnerability (Marchioretto et al. 2010, Takeuchi & Esteves 2017, Santos et al. 2021, Abreu et al. 2024).

The patterns observed in Melochia align with broader biogeographic models proposed for the Brazilian flora. Species distributions result from interactions among environmental heterogeneity, paleoecological history, dispersal ability, and geographic barriers (Antonelli & Sanmartín 2011). These findings highlight the need to prioritize microendemic species, which occupy extremely small areas, face ongoing habitat loss, and therefore carry a markedly higher extinction risk.

It is essential to recognize the value of newly described species, even when known from few records. Melochia cristobaliae, for example, reveals ongoing taxonomic discovery and expands our understanding of the genus’ diversity. Comparable studies in other groups also report species poorly represented in databases, which underscores the need for further field collections and targeted conservation actions (Giulietti et al. 2005).

The preliminary assessment of the conservation status of Melochia based on the International Union for Conservation of Nature (IUCN 2025) criteria indicates that most species exhibit broad Extent of Occurrence (EOO) and Area of Occupancy (AOO) (see Table II), supporting their inclusion in the Least Concern (LC) category. Species such as M. pyramidata, M. spicata, and M. tomentosa, which are widely distributed in Brazil, fall within this category and appear to tolerate a wide range of environments, including degraded areas (Gonçalez & Coutinho 2020). In contrast, a small group of taxa were categorized as Critically Endangered (CR), Vulnerable (VU), and Extinct in the Wild (EW).

Table II
Conservation status of Melochia species in Brazil. AOO (Area of Occupancy); EOO (Extent of Occurrence). EW: Extinct in the Wild; CR: Critically Endangered; LC: Least Concern; VU: Vulnerable.

Species	(EOO)	(AOO)	Status IUCN
M. arenosa	6,905,855,428 km²	272,500,000 km²	LC
M. argentina	320,288,854 km²	10,000,000 km²	LC
M. betonicifolia	1,305,533,151 km²	330,000,000 km²	LC
M. chamaedrys	532,491,661 km²	102,500,000 km²	LC
M. cristobaliae	0,000 km²	8,000 km²	CR B1ab(i,ii,iii) +2ab(i,ii,iii)
M. gardneri	250,272,919 km²	25,000,000 km²	LC
M. graminifolia	5,936,779,100 km²	132,500,000 km²	LC
M. hassleriana	85,960,217 km²	12,500,000 km²	LC
M. hermannioides	41,256,584 km²	15,422,803 km²	LC
M. illicioides	47,547,805 km²	11,906,558 km²	LC
M. kerriifolia	61,012,514 km²	19,777,059 km²	LC
M. lanata	0,000 km²	4,000 km²	EW
M. lanceolata	25,704,376 km²	8,799,400 km²	CR B1ab(i,ii,iii) +2ab(i,ii,iii)
M. longidentata	417,250,145 km²	42,500,000 km²	LC
M. melissifolia	5,165,657,547 km²	127,500,000 km²	LC
M. morongii	2,406,888,932 km²	45,000,000 km²	LC
M. nodiflora	1,890,493,856 km²	17,500,000 km²	LC
M. parvifolia	7,871,466,430 km²	242,500,000 km²	LC
M. pilosa	3,690,867,233 km²	375,000,000 km²	LC
M. pyramidata	6,507,059,356 km²	620,000,000 km²	LC
M. sergipana	626,925 km²	4,563,112 km²	VU B2ab(i,ii,iii)
M. simplex	3,836,744,514 km²	167,500,000 km²	LC
M. siphonandra	1,223,327,011 km²	15,500,000 km²	LC
M. spicata	8,872,880,427 km²	890,000,000 km²	LC
M. splendens	3,842,475,350 km²	115,000,000 km²	LC
M. tomentosa	4,626,435,959 km²	542,500,000 km²	LC
M. ulmifolia	3,689,508,089 km²	137,500,000 km²	LC

Melochia sergipana presents reduced EOO and AOO values and was therefore classified as Vulnerable (VU). This species occurs predominantly within sustainable-use conservation units, a relatively permissive protection category that does not guarantee full protection against resource extraction or habitat conversion. Its preliminary classification as VU is therefore consistent with the current scenario of habitat fragmentation, the existence of few known populations, and the ongoing degradation within its area of occurrence.

Melochiacristobaliae and M. lanceolata exhibit extremely low EOO and AOO values and were classified as Critically Endangered (CR). Moreover, these reduced values meet the IUCN criteria B1 and B2. Melochia cristobaliae is a microendemic species of the Atlantic Forest known from only two highly fragmented localities that are subject to continuous degradation. A field expedition conducted by the first author to these areas, aiming to recollect this species, was unsuccessful and revealed that both localities have experienced a decline in habitat quality, supporting subcriteria b(i-iii). In contrast, M. lanceolata is restricted to a few localities in northern Brazil and, although presenting slightly higher EOO and AOO values, is also subject to environmental pressures, which supports its preliminary classification as CR.

More strikingly, the EW status assigned to M. lanata reflects the absence of recent records combined with an EOO equal to zero and a minimal AOO (4,000 km²). This species is known from only two collections: Saint-Hilaire s.n., 1816, and Vincent 818, 1919. According to Gonçalez (Gonçalez & Esteves 2017), an expedition to the Espinhaço Range conducted in 2015 aimed to recollect the species in the field, but without success. The Espinhaço Range has been continuously affected by human pressure, and many of its original formations are currently fragmented due to logging, agriculture, mining activities, and recurrent fires (Costa et al. 2023, Gladson et al. 2023). This evidence reinforces the preliminary classification of M. lanata as Extinct in the Wild.

Richness analysis

The results of the richness analysis indicate that the Central-West region of Brazil and the southern Espinhaço Range are important centers for the genus Melochia, with emphasis on areas A113, A132, A133, and A143, which together comprise 16 species. These areas are associated with different environmental and ecological contexts. Area A113, located in the northern portion of the Pantanal, is a transition between the Amazon, Cerrado and Pantanal phytogeographic domains, regions known for their high biodiversity and heterogeneous environmental conditions, which favor floristic diversification (Junk et al. 2006).

The campos rupestres of the Espinhaço Range harbor high levels of species richness and microendemism, many of which are restricted to isolated mountaintops (Colli-Silva et al. 2019). Representative taxa include genera such as Lychnophora, Mikania, Lessingianthus, and Baccharis, which are particularly diverse and often endemic to these environments (Campos et al. 2019).

Areas A132, A133, and A143 comprise the southern portion of the Pantanal, which has a variety of phytophysiognomies associated mainly with soil and climate types. This area is influenced by the Chaco, a geographical region with a subtropical semi-arid climate that covers Argentina, Bolivia and Paraguay, and its floristic composition is similar to that found in steppe savannas (Botelho & Clevelário Junior 2016). Uetanabaro et al. (2007) classified this area as being of great biogeographical importance, as it covers the central portion of the dry ‘diagonal of open formations’ in South America that stretches from the Caatinga (in Brazil) to the Chaco (in Argentina), where there are areas of contact between the Pantanal, Chaco and Cerrado. In addition, its proximity to the Paraguay River, a region of great ecological importance due to its seasonal flooding dynamics, creates varied habitats, promoting the coexistence of different species (Harris et al. 2005).

The species that occur in this area have a wide distribution and can be found in other Brazilian phytogeographic domains such as Amazonia (M. arenosa, M. graminifolia, M. simplex, M. spicata, and M. ulmifolia), Caatinga (M. pyramidata and M. tomentosa) and Cerrado (M. parvifolia and M. pilosa). In addition to these, M. morongii is a species that occurs in the Chaquenha region and M. kerriifolia is a disjunct species, occurring in the Pantanal and in Central America, Colombia, and Venezuela.

Melochia argentina, M. chamaedrys, M. hassleriana, and M. kerriifolia were found only in the southern portion of the Pantanal, while M. ulmifolia, an Amazonian species, occurs only in the north, where it meets its southern limit. According to Pott (2000) and Pott et al. (2011), the central position of the Pantanal in South America has allowed it to form a vegetation mosaic, receiving influence from the surrounding areas.

Area A117, in the central portion of the Brazilian Cerrado, stands out as a center of richness for several taxa (Chamaecrista, Stylosanthes, Microlicia, and Paepalanthus), suggesting that its centrality and connectivity with other phytogeographic domains favor the establishment of these taxonomic groups (Giulietti et al. 2005, Goldenberg et al. 2012, Vanni 2017, Mendes et al. 2020).

The occurrence of ten species of Melochia in this region (M. arenosa, M. argentina, M. gardneri, M. parvifolia, M. pilosa, M. pyramidata, M. simplex, M. spicata, M. splendens, and M. tomentosa) reinforces the importance of the Cerrado as a hotspot of floristic richness within Brazil. The presence of widely distributed species, such as M. parvifolia, M. pilosa, and M. spicata, together with endemic or restricted taxa (M. gardneri and M. argentina), reinforces the role of the central Cerrado as a core of diversity for the group.

The concentration of richness in areas of the Central-West region suggests that factors such as habitat diversity, connectivity between the Amazon, Cerrado and Pantanal phytogeographic dominions and the presence of protected areas play important roles in the distribution of the genus Melochia. Previous studies corroborate these findings, highlighting the floristic richness of the Cerrado and its importance for the conservation of tropical plants (Ratter et al. 2003, Borghetti et al. 2023, Santos & Silva 2025).

According to Souza & Eisenlohr (2020), transition zones between biogeographic dominions harbor high biodiversity due to the coexistence of species from different floristic groups and the presence of multiple vegetation types. In addition, phylogenetic analyses indicate that these transition zones not only concentrate species richness but also harbor high phylogenetic diversity, reflecting the coexistence of lineages from different evolutionary origins and emphasizing their importance for conservation planning (Silva-Pereira et al. 2020).

Together, these findings support the interpretation that the high richness of Melochia in the Central-West results from the combined effects of environmental heterogeneity, hydrographic influence, and biogeographical connectivity among dominions.

Diversity analysis

The results of the diversity analysis indicated seven main centers with the highest Shannon diversity indices (H’ between 1.599 and 2.000): Pantanal (PTN), Cerrado of Central Brazil (CBC), Northern Portion of the Espinhaço Range (NPER), Southern Portion of the Espinhaço Range (SPER) Serra dos Carajás (CJM), Tapajós (TPJ), and Raposa Serra do Sol Indigenous Land (RSSIL). This distribution reflects areas of high environmental heterogeneity and great species richness, corroborating the idea that factors such as habitat variation, ecological interactions and evolutionary history are fundamental to maintaining biological diversity in tropical regions.

The Pantanal had the highest number of grids with high diversity, distributed in the north (grids A112 and A113) and south (grids A132, A133, A134, A142, A143, and A144). This concentration of diversity in the domain is associated with its seasonal character and mosaic of habitats, which include wetlands, riparian forests, grasslands and savannahs (Alho 2011). According to Junk et al. (2006), the seasonality of the floods creates a dynamic environment, promoting the coexistence of generalist and specialized species. In the north of the Pantanal, it is possible to observe the occurrence of species adapted to wetter areas due to the connectivity with the Amazon, such as M. splendens and M. ulmifolia, while in the south, species with adaptability to water deficit and low humidity stand out, such as M. argentina, M. chamaedrys, M. kerriifolia, and M. morongii.

Pott et al. (2011) documented a similar pattern. According to the authors, the headwaters of several tributaries of the Paraguay River originate near the transition with the Amazon basin, creating a corridor that allows Amazonian species such as Nectandra amazonum Nees and Victoria amazonica (Poepp.) J.C. Sowerby to reach the Pantanal. Likewise, as observed for Melochia, species from the Chaco—such as Copernicia alba Morong and Aspidosperma quebracho-blanco Schltdl.—occur south of the Pantanal.

The Espinhaço Range (grids A97, A108, A129, and A138) is recognized as a biodiversity hotspot, especially due to the campos rupestres and their high levels of flora and faunal richness and endemism, concentrating almost 15% of Brazil’s flora in less than 1% of its territory (Giulietti & Pirani 1988, Giulietti et al. 1997, Colli-Silva et al. 2019). The heterogeneity of micro-habitats, associated with poor soils and seasonal climate, promotes adaptive processes that sustain this diversity. Similar results were reported by Silveira et al. (2016), who highlighted the importance of the Serra do Espinhaço for the conservation of threatened species.

As described for the Pantanal, in the Espinhaço Range there is a difference in the composition of Melochia species in its northern and southern portions. The NPER is influenced by the Caatinga, with greater water seasonality and high temperatures, while the SPER, especially in the higher portions, has greater humidity and a milder climate (Giulietti et al. 1997, Zappi et al. 2017). This differentiation between NPER-SPER of Espinhaço Range is consistent with broader biogeographic patterns documented for other plants groups in campos rupestres ecosystems (Alves & Buril 2022, Costa et al. 2024).

Studies on Asteraceae demonstrate that the Espinhaço Range flora is structured into distinct floristic groups influenced by surrounding phytogeographical dominions, particularly the Caatinga in the northern sector, where genera such as Lepidaploa, Lessingianthus, and Vernonanthura occur and the Cerrado and Atlantic Forest in the southern, habitat of Lychnophora, Eremanthus, and Richterago, with climate acting as a primary driver of species distribution (Campos et al. 2019). This supports the interpretation that the NPER–SPER differentiation in Melochia reflects not only edaphic constraints but also dominion-level climatic filtering.

In the NPER, the occurrence of species such as M. illicioides and M. longidentata may be related to the quartzite and acidic substrate, which would limit their expansion to the southern portion that has greater lithological variation, including ferruginous outcrops, which support highly specialized floras, where, for example, M. lanata occurs, known only from one locality in the SPER.

Alves & Buril (2022) reported a similar pattern for Convolvulaceae. Ipomoea chapadensis J.R.I. Wood & L.V. Vasconcelos occurs only in the northern sector of Chapada Diamantina, whereas Distimake repens (D.F. Austin & Staples) Petrongari & Sim.-Bianch. is restricted to the southern Espinhaço Range in Minas Gerais.

The exclusive distributions of certain species may be traces of ancient phylogeographic processes. Studies with other plant groups in the Espinhaço Range, such as Vellozia, Barbacenia (Velloziaceae), Orthophytum (Bromeliaceae), and Paepalanthus (Eriocaulaceae), show that populations from the north and south often form distinct lineages and sometimes cryptic species (Giulietti et al. 2005, Mello-Silva et al. 2011, Costa et al. 2024, Magri et al. 2025).

Grids A37, A50, and A51, located in the Serra dos Carajás, in Pará, and northern Tocantins, reflect the role of this region as a center of diversity due to its transitional vegetation between the Amazon Rainforest and the Cerrado and Campinaranas ecosystems. According to Zappi et al. (2019), the “cangas” outcrops of the Serra dos Carajás are characterized by a poor soil, stony, shallow substrate with low water retention, high surface temperatures and water seasonality, acting as ecological filters for the flora, occurring on mountain tops, forming islands of open vegetation within a matrix of dense ombrophilous forest, which results in environmental heterogeneity and the occurrence of endemic taxa. These areas are home to M. ulmifolia, a typically Amazonian species, and the amphibious species: M. arenosa, M. melissifolia, M. simplex, M. spicata, and M. splendens. These species occur within the hydromorphic formations described by Mota et al. (2015), particularly marshy grasslands and temporary lagoons, which are subject to permanent or seasonal flooding. According to these authors, other paludose Malvaceae species, such as Waltheria indica L. and Hibiscus bifurcatus Cav., can also be found in these environments, as well as species from other families, including Aniseia martinicensis (Jacq.) Choisy (Convolvulaceae), Miconia chamissois Naudin (Melastomataceae), and Aeschynomene rudes Benth. (Fabaceae).

The Tapajós National Forest (Grids A22 and A23) showed a high diversity index, which can be attributed to its forest structure, rich in large trees, and its connectivity with other biodiversity corridors. Works such as those by Espírito-Santo et al. (2005) and Gonçalves & Santos (2008) highlight the role of the Tapajós forest in maintaining diversity. The TPJ shares some species with the CJM, such as M. arenosa, M. melissifolia, M. simplex, M. spicata and M. ulmifolia. It is also the habitat of M. lanceolata, a species endemic to the Amazon, occurring in Brazil, Guyana and Suriname. This area is also home to M. sergipana, a Brazilian endemic known from only three records (Ceará, Pará and Sergipe).

The high diversity in grid A1, located in the Raposa Serra do Sol Indigenous Land in northern Roraima, highlights the importance of protected areas for conservation. This region is characterized by mountain forests, savannas, and rivers, forming a mosaic that supports a rich fauna and flora (Ab’Sáber 2009).

In addition, the traditional management of indigenous communities contributes to the preservation of local biodiversity. The species found in the RSSIL are also found in the other areas of the Amazon (CJM and TPJ), especially M. siphonandra, a species with few reports in Brazil, occurring in the Amazon and humid high-altitude forests (brejos de altitude) of Ceará. According to Moro et al. (2024), these humid enclaves are clearly distinguishable in vegetation indices and represent areas of higher moisture within the Caatinga matrix, harboring distinct plant assemblages and increasing regional heterogeneity.

This reinforces the hypothesis that these humid fragments of northeastern Brazil were connected to the Amazon, which allowed for the dispersal of Amazonian lineages (Prado & Gibbs 1993, Werneck 2011). However, new evidence indicates that these connections were not limited to wetter interglacial periods during the Pleistocene, having occurred repeatedly over time, under different climatic conditions and vegetation types, and followed various routes (Ledo & Colli 2017).

The patterns observed in the results of this study corroborate research that highlights areas of high biodiversity in Brazil, such as the Caatinga (Queiroz et al. 2017) and the campos rupestres (Silveira et al. 2016, Rapini et al. 2021). However, unlike previous studies that focused only on well-preserved areas, our data point to the relevance of regions with land use mosaics, such as the Pantanal, in sustaining high levels of diversity.

Endemicity analysis

The Parsimony Analysis of Endemicity (PAE) applied to the genus Melochia revealed two main groupings of areas of endemism: CAES (Chapada do Araripe and eastern Sergipe) and CDEB (Chapada Diamantina and eastern Bahia), both recovered at both 1° × 1° and 2° × 2° grid resolution. Despite the low resolution of the consensus tree, which showed high basal polytomy, these areas stood out for having exclusive species and significant support values (bootstrap ≥ 50%), which corroborates their biogeographic relevance. These findings are in line with patterns identified in other studies of plants and animals in the Neotropical region (Costa et al. 2000, Ippi & Flores 2001, Morrone & Escalante 2002, Chiron 2009).

The Chapada Diamantina region, recovered as an area of endemism for Melochia (CDEB), has been identified as a center of endemism for several botanical families, such as Asteraceae (Campos et al. 2019), Bromeliaecae (Costa et al. 2024, Souza et al. 2025), Eriocaulaceae (Giulietti & Miranda 2009, Giulietti & Silva 2016), Velloziaceae (Conceição & Pirani 2007), and Fabaceae (Simon & Proença 2000). Costa et al. (2024), for example, identified the Chapada Diamantina as one of the three main areas of endemism for Orthophytum, with five exclusive species and bootstrap support of over 60%, a result analogous to that obtained with Melochia. These coincidences reinforce the importance of the region as a historical center of speciation and climatic refuge, probably favored by its topographical complexity, geographical isolation and environmental heterogeneity.

The CAES area, which includes the Chapada do Araripe, suggests that this ecotonal portion of northeastern Brazil is home to historical processes of retention and differentiation of lineages, especially due to its position of contact between humid (Atlantic Forest) and dry (Caatinga) formations. Recent biogeographical syntheses recognized the Chapada do Araripe as an independent district within the Caatinga dominion, highlighting its unique geomorphological and floristic identity, which corresponds to an uplifted sedimentary basin surrounded by crystalline terrains, forming an isolated plateau that rises above the surrounding semiarid landscapes (Moro et al. 2024).

The CAES emerges as a novelty as an endemic area, since there is no reference in the literature citing endemism for plants in these locations. The most comprehensive work carried out in the Chapada do Araripe cites almost 500 species, however it does not report exclusive species (Loiola et al. 2015). Three endemic species of Melochia were recorded for the Chapada do Araripe: M. betonicifolia, M. longidentata, and M. sergipana. This environmental mosaic and the region’s paleoclimatic history, marked by cycles of vegetation fragmentation and reconnection during the Quaternary, are often cited as probable areas of endemism (Prado & Gibbs 1993, Werneck 2011).

The patterns observed in Melochia reinforce the idea that the Espinhaço Range, especially its north-central portion (Chapada Diamantina and adjacent areas), acts as an important biogeographic barrier and center of speciation, as also discussed for Orthophytum by Costa et al. (2024). The occurrence of Melochia lanata only in the Espinhaço Range and of M. illicioides at its eastern limit is consistent with patterns of microendemism related to edaphic specialization and dispersal limitation in rupestrian environments (Giulietti et al. 1987, Silveira et al. 2016). According to Rapini (2010) and Schaefer et al. (2016), campos rupestres formations on quartzite substrate present edaphoclimatic conditions such as shallow, acidic soil with low fertility, favoring niche segregation and the evolution of endemic and specialized taxa.

The similarity of the patterns between phylogenetically distinct groups (Malvaceae, Eriocaulaceae, and Bromeliaceae) highlights the possible structuring role of these landscapes in maintaining endemic biodiversity.

The PAE with 1° and 2° grids produced congruent results, although the finer resolution (1° × 1°) generated greater basal polytomy. This pattern is common in studies with few species and low sample density, as discussed by Morrone & Escalante (2002) and evidenced in the work with Orthophytum (Costa et al. 2024). The methodological limitations are also noticeable in the CI (0.58 and 0.55) and RI (0.81 and 0.87) values obtained in the Melochia analyses. Although the number of informative characters was low (six), the high RI values indicate the coherence of the groupings, even with low overall topological resolution (Morrone 2009).

It is important to note that the comparison with studies with more representative species highlights the limitations of working with poorly sampled taxa. In these cases, the adoption of larger spatial scales (2° × 2°) tends to smooth out sampling noise and recover more comprehensive areas of endemism, as also proposed by Casagranda et al. (2009) and Linder (2001).

The areas of endemism identified for Melochia coincide with regions of high priority for conservation, both because of the presence of endemic species and because of their representativeness in ecotonal and montane formations. The CDEB area, for example, is home to exclusive species that occur in environments under increasing anthropogenic pressure, such as the Caatinga-Atlantic Forest contact and the Chapada Diamantina rupestrian fields, habitats that are already among the most threatened in Brazil (Silva & Barbosa 2017).

The identification of these areas, even with limited data, contributes to the definition of biotic units and will subsidize the selection of target areas for phylogenetic studies and conservation actions. It also reinforces the importance of historical biogeography as a tool for understanding the evolutionary processes that shape the distribution of Neotropical biodiversity (Morrone 2009, Brown & Lomolino 1998).

CONCLUSIONS

Based on the results presented, this study revealed important patterns of geographical distribution, richness, diversity, and endemicity for the genus Melochia in Brazil. The analysis showed a predominance of widely distributed species, highlighting the ecological adaptability and dispersal strategies of the genus. The most abundant species, such as M. pyramidata, M. spicata, and M. tomentosa, proved to be well adapted to diverse environments, while taxa with restricted distribution, such as M. lanata, M. sergipana, and Melochia sp. nov., pointed to specific, often vulnerable habitats.

The Central-West region of Brazil emerged as a center of richness for Melochia, especially in areas such as the Pantanal and its surroundings, reinforcing the role of regions with high environmental heterogeneity in maintaining floristic diversity. The seven main centers of diversity identified—Pantanal, Cerrado of Central Brazil, Northern portion of the Espinhaço Range, Southern portion of the Serra do Espinhaço, Serra dos Carajás, Tapajós and the Raposa Serra do Sol Indigenous Land—corroborate the relevance of environmental mosaics and protected areas for biodiversity conservation.

The endemicity analysis defined two priority areas with exclusive species, indicating the importance of conservation efforts in these regions. The PAE confirmed the existence of well-defined areas of endemism, such as the northern portion of the Espinhaço Range (Chapada Diamantina) and Chapada do Araripe.

The findings of this study provide a solid basis for future research into the genus Melochia, as well as for planning conservation strategies. By highlighting critical regions of diversity and endemicity, the need to expand collection, monitoring and environmental protection efforts is reinforced, especially in areas subject to anthropogenic pressures.

Acknowledgements

We would like to thank all the curators of the herbaria consulted; the Integrative Systematics Laboratory, for the infrastructure used to carry out the work; MTB would like to thank Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the grant (CNPq 311430/2022-8).

• Data availability
  The datasets generated and/or analyzed during the current study are available from the corresponding author upon reasonable request.

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Edited by

• Handling editor
  Vasco Azevedo

Data availability

The datasets generated and/or analyzed during the current study are available from the corresponding author upon reasonable request.

Publication Dates

• Publication in this collection
  21 Aug 2026
• Date of issue
  2026

History

• Received
  19 Sept 2025
• Accepted
  30 Mar 2026