Abstract
The Bromeliaceae family plays an important ecological, environmental, and economic role in Bahia, harboring high levels of diversity and endemism, although the state still lacks an updated species checklist to support conservation efforts. This study aimed to compile and systematize information on the species richness, distribution, life forms, endemism, and conservation status of Bromeliaceae in Bahia. Data were obtained from field collections and databases such as GBIF, Tropicos, Reflora, SpeciesLink, and the Encyclopaedia of Bromeliads, as well as herbarium specimen images from over 40 national and international herbaria. The data were organized into spreadsheets and analyzed in terms of taxonomy, life form, biome, endemism, and conservation status. A total of 433 taxa were recorded, comprising 426 species, six varieties, and one subspecies, distributed across 36 genera. The subfamily Bromelioideae was the most diverse (308 taxa), followed by Tillandsioideae (102), Pitcairnioideae (22) and Navioideae (1), with Aechmea, Hohenbergia, Cryptanthus, Vriesea, and Tillandsia being the most representative genera in the state. Of the total taxa, 241 (55.6%) are endemic to Bahia, highlighting the state as a center of diversification for the family. Only 180 taxa (41.6%) have been formally assessed for conservation status, of which 89 (20.5%) are considered threatened: 21 listed as Critically Endangered (CR), 48 as Endangered (EN), and 20 as Vulnerable (VU), with a predominance of threatened taxa within Bromelioideae. Regarding life forms, epiphytic species were the most common (90), followed by terrestrial (76) and rupicolous (61) taxa; 206 taxa exhibited ecological plasticity, occurring in more than one category. The highest species richness was recorded in the Atlantic Forest (266 taxa), followed by the Caatinga (86) and the Cerrado (14), reflecting the influence of multiple phytogeographic domains, altitudinal variation, and climatic gradients in Bahia. The study also identified inconsistencies in taxonomic databases, underscoring the need for expert scientific curation. It is concluded that the updated systematization of Bromeliaceae diversity in Bahia is essential to guide research, conservation strategies, and the sustainable use of this highly threatened flora.
Keywords
Bromeliad; Bromelioideae; Tillandsioideae; Pitcairnioideae; Genetic diversity
Resumo
A família Bromeliaceae desempenha um importante papel ecológico, ambiental e econômico na Bahia, abrigando elevada diversidade e endemismo, mas carece de uma listagem atualizada de espécies para subsidiar ações de conservação. Este estudo teve como objetivo compilar e sistematizar informações sobre a riqueza, distribuição, formas de vida, endemismo e status de ameaça das espécies de Bromeliaceae no estado. Foram utilizados dados provenientes de coletas de campo, além de plataformas como GBIF, Tropicos, Reflora, SpeciesLink e Encyclopaedia of Bromeliads, bem como imagens de exsicatas de mais de 40 herbários nacionais e internacionais. As informações foram organizadas em planilhas e analisadas quanto à nomenclatura, forma de vida, bioma, endemismo e status de conservação. Foram registrados 433 táxons (426 espécies, seis variedades e uma subespécie), distribuídos em 36 gêneros. A subfamília Bromelioideae foi a mais diversa (308 táxons), seguida por Tillandsioideae (102), Pitcairnioideae (22) e Navioideae (1 táxon), com destaque para os gêneros Aechmea, Hohenbergia, Cryptanthus, Vriesea e Tillandsia, os mais abundantes no estado. Do total dos táxons, 241 (55,6%) são endêmicas da Bahia, reforçando o estado como centro de diversificação da família. Apenas 180 táxons (41,4%) possuem avaliação quanto ao status de conservação, das quais 89 (20,5%) estão ameaçadas: 21 classificadas como ‘Criticamente em Perigo’ (CR), 48 como ‘Em Perigo’ (EN) e 20 como ‘Vulneráveis’ (VU), com predominância de táxons ameaçados na subfamília Bromelioideae. Quanto ao hábito de vida, observou-se predominância de espécies epífitas (90), seguidas por terrícolas (76) e rupícolas (61); 206 táxons demonstraram plasticidade ecológica, ocorrendo em mais de uma dessas categorias. A maior diversidade foi registrada na Mata Atlântica (266 táxons), seguida pela Caatinga (86) e pelo Cerrado (14), refletindo a influência dos diferentes domínios fitogeográficos, variação altitudinal e gradiente climático da Bahia. O estudo também revelou inconsistências nas bases de dados taxonômicos, evidenciando a importância da curadoria científica especializada. Conclui-se que a sistematização atualizada da diversidade de Bromeliaceae na Bahia é fundamental para orientar ações de pesquisa, conservação e uso sustentável dessa flora altamente ameaçada.
Palavras-chave
Bromélia; Bromelioideae; Tillandsioideae; Pitcairnioideae; Diversidade genética
Introduction
The family Bromeliaceae extends from the southern United States, through Central America and the Caribbean, to South America, reaching southern Argentina and Chile (Smith & Downs 1974, 1977, 1979, Benzing 2000). On the African continent, the family is represented by a single species, Pitcairnia feliciana (A.Chevalier) Harms & Mildbraed, which occurs along the west coast of Africa, specifically in Guinea (Smith & Downs 1974, Porembski & Barthlott 1999, Benzing 2000, Jacques-Félix 2000).
The family occurs in a wide variety of phytophysiognomies, from sea level up to altitudes of approximately 4,000 meters, contributing to a high level of biological diversity (Benzing 2000). Currently, Bromeliaceae comprises 81 genera and 3,848 species (Gouda et al. 2025, cont. updated), standing out as one of the largest families among monocotyledons, with an almost exclusively Neotropical distribution (Gilmartin 1973, Benzing 2000).
Bromeliaceae species are herbaceous plants with epiphytic, terrestrial, or rupicolous habits. They possess a poorly developed stem and leaves arranged in a distichous or polystichous manner, either distributed along the stem or grouped in rosettes, allowing the accumulation of water and nutrients and the formation of phytotelmata that serve as habitats for various organisms, such as algae, amphibians, small reptiles, and bacteria (Romero et al. 2020, Lopes Filho et al. 2023). Leaf margins range from entire to serrated (Benzing 2000, Vasconcellos & Oliveira 2022), and their surfaces are covered by specialized peltate trichomes, which absorb moisture and nutrients from the atmosphere, a distinctive feature of the family (Vasconcellos & Oliveira 2019).
Tank-forming or phytotelm-forming bromeliads exhibit morphologies that favor the accumulation of water and debris, with wide and imbricated leaf sheaths (Benzing 2000). The phytotelma not only ensures plant hydration but also allows the absorption of nutrients derived from the decomposition of organic matter and the bodies of small invertebrates (Benzing 1973, 1990, Leme & Marigo 1993). The accumulated water can persist for long periods, depending on rainfall frequency, providing a suitable environment for insect reproduction, the growth of lichens, fungi, and algae, as well as sheltering amphibians and other organisms throughout their life cycles (Benzing 1980, 2000).
With an impressive diversity of forms, sizes, and colors, species of the family possess high ornamental value and attract landscaping interest, as exemplified by species of the genera Aechmea Ruiz & Pav., Ananas Mill., Bromelia L., Guzmania Ruiz & Pav., Puya Molina, Sincoraea Ule, Tillandsia L., and Vriesea Lindl. (Souza et al. 2012, 2014, Souza et al. 2015, Anacleto & Negrelle 2013). However, the ongoing habitat loss places many of these species at risk of extinction, reinforcing the need for conservation actions.
Traditionally divided into three subfamilies based on morphological characteristics (Bromelioideae, Pitcairnioideae, and Tillandsioideae) (Smith & Downs 1974, 1977, 1979), the family Bromeliaceae is currently subdivided into eight subfamilies based on morphological, molecular, and phylogenetic data: Bromelioideae, Brocchinioideae, Hechtioideae, Lindmanioideae, Navioideae, Pitcairnioideae, Puyoideae, and Tillandsioideae (Givnish et al. 2007, 2011).
The subfamily Bromelioideae is the most diverse, with 43 genera and 1,042 species (Gouda et al. 2025, cont. updated), and is characterized by generally serrated leaves, seeds covered by mucilage, the presence of petal appendages, reduced roots, and indehiscent berry fruits (Smith & Downs 1979, Givnish et al. 2007, Sass & Specht 2010).
The subfamily Tillandsioideae is the largest within Bromeliaceae, encompassing 23 genera and approximately 1,589 species (Gouda et al. 2025, cont. updated). Its species are predominantly epiphytic, with entire, spineless leaves, peltate scales, and plumose seeds (Smith & Downs 1977). The gynoecium generally has a superior ovary, with exceptions in Glomeropitcairnia Harms (with indehiscent capsules), Tillandsia (subg. Pseudalcantarea) Mez, and Alcantarea (E. Morren ex Mez) Harms., which possess a semi-inferior ovary (Beaman & Judd 1996, Versieux 2009, Smith & Downs 1977, Givnish et al. 2007, Barfuss et al. 2005, 2016).
The subfamily Pitcairnioideae comprises approximately 711 species distributed across five genera. In Brazil, it is primarily represented by the genera Dyckia Schult. & Schult.f., Encholirium Mart. ex Schult. & Schult.f., and Pitcairnia L’Heritier (Gouda et al. 2025, cont. updated). Species of this subfamily are predominantly terrestrial and rupicolous, with functional roots and spiny leaves. Most are herbaceous plants, although some are subshrubby, such as in Deuterocohnia Mez. Leaves are often spiny-serrated and remain entire in many Pitcairnia species. The gynoecium of these plants generally presents a superior ovary, although some Pitcairnia species have an inferior ovary. The fruit is predominantly a capsule, occasionally indehiscent. Seeds often have intact appendages or, more rarely, lack a covering (Smith & Downs 1974).
The distinction of the subfamily Pitcairnioideae from other Bromeliaceae subfamilies is based on characteristics such as mature seeds with intact or slightly divided appendages, generally dehiscent fruits, thick and serrated leaves, finely divided scale indument, and the predominance of a terrestrial habit (Smith & Downs 1974).
The subfamily Navioideae currently comprises five genera and a total of 111 species (Gouda et al. 2025, cont. updated). Species of this genus are characterized by entire, serrated, or spiny leaves, with capsular fruits, and generally superior ovaries, although some Navia species present inferior ovaries (Smith & Till 1998).
Brazil harbors the greatest diversity of Bromeliaceae in the world, with approximately 1,417 species and 55 genera, representing about 40% of known species and 80% of the family’s genera (Flora e Funga do Brasil 2025). Furthermore, approximately 85.4% of these species are endemic to the country (Flora e Funga do Brasil 2025).
These numbers continue to increase with ongoing taxonomic research, with dozens of new species described annually (Gouda et al. 2025, cont. updated). Over the past five years, several new Bromeliaceae species have been described for Bahia, reinforcing its role as a major center of diversity. This floristic richness, including endemic genera, underscores the need for intensified conservation efforts and further studies on the state’s flora.
In this context, the preparation of an updated list of Bromeliaceae for Bahia is not only necessary but urgent. Floristic inventories and species lists are fundamental tools for documenting plant diversity, supporting research in botany, ecology, conservation, and sustainable use of biodiversity (Martinelli et al. 2008). Furthermore, such compilations provide a solid basis for environmental public policies and native flora management programs.
Therefore, this study aims to fill an important gap in the knowledge of Bahia’s flora by compiling and systematizing updated data on Bromeliaceae diversity in the state, significantly contributing to the conservation and rational use of these natural resources. Accordingly, the objective of this study was to provide information on the richness and geographic distribution of Bromeliaceae species in Bahia, as well as their conservation status, life forms, endemism, and chronology of species descriptions.
Material and Methods
1. Study area
The state of Bahia, with an area of 564,692.669 km2, is located in the southern portion of Brazil’s Northeast region. It borders the states of Alagoas, Pernambuco, Piauí, Sergipe (Northeast), Tocantins (North), Goiás (Central-West), Espírito Santo, and Minas Gerais (Southeast), and has an extensive coastline of approximately 900 km along the Atlantic Ocean (SEI Bahia 2025).
Bahia is part of the Coastal Zone, which includes characteristic ecosystems such as beaches, restingas, lagoons, and mangroves. The state’s vegetation is marked by the presence of rupestrian fields, seasonal forests, transitional areas, and coastal vegetation (CBPM 2020). The state encompasses three important biomes: Caatinga (64% of the territory), Cerrado (18%), and Atlantic Forest (18%), all of great relevance for Bromeliaceae diversity.
2. Data sources and sampling
The data for the preparation of this checklist were obtained through collection expeditions between 2009 and 2025, from the Global Biodiversity Information Facility (GBIF) (2025, continuously updated), Tropicos (https://tropicos.org/) (2025, continuously updated), the Germplasm Bank of the Graduate Program in Plant Genetic Resources at UFRB/ Embrapa (Banco de Germoplasma do Programa de Pós-Graduação em Recursos Genéticos Vegetais da UFRB/ Embrapa), as well as from the Encyclopaedia of Bromeliad database (Gouda et al. 2025, cont. updated). In addition, herbarium images from ALCB, HURB, HCF, HUEFS, HUFU, K, MBM, NY, US, SP, CEPEC, MBML, RB, GH, BM, WEB, UFG, HVASF, CVJBFZB, HbVirtFlBras, UFP, UPCB, HAL, MO, UFRN, R, BR, WU, CVRD, P, B, HBG (acronyms according to Thiers, 2025, continuously updated, Index Herbariorum) were verified through the SpeciesLink (2025, continuously updated), JABOT (2025, continuously updated), and Virtual Herbarium REFLORA (2025, continuously updated) databases to expand data on distribution, phenology, and any available label information.
The research was conducted in accordance with Brazilian legislation, under the authorization of the Biodiversity Authorization and Information System (Sistema de Autorização e Informação em Biodiversidade) (SISBIO) nº 69.870-1, and the activities are duly registered in the National System for the Management of Genetic Heritage and Associated Traditional Knowledge (Sistema Nacional de Gestão do Patrimônio Genético e do Conhecimento Tradicional Associado) (SisGen) under authorization A9E9F8D.
3. Data compilation and taxonomic treatment
From these sources, primary surveys were conducted based on collection records, field expeditions, and data from herbarium platforms of different institutions. Spreadsheets were organized containing all registered taxa. The criteria adopted for the research included threat status, endemism, geographic distribution, and habitat type. The taxonomic nomenclature adopted in this study followed the guidelines of the International Plant Names Index (IPNI 2025). The taxonomic circumscription adopted in this study follows Gouda et al. (2025, cont. updated), in which Encholirium is recognized as a valid genus distinct from Dyckia.
Subsequently, the data were organized into tables and evaluated regarding conservation status, biome of occurrence, presence or absence in the state of Bahia, life form, locality of species description (focusing solely on records from Bahia), year of publication, and the corresponding voucher number.
4. Conservation status assessment
Species extinction risk assessments were conducted based on official lists published by the Ministry of the Environment (Brazil 2022b, SEMA 2017), the Institute of Environment and Water Resources of Bahia (Bahia 2017), and the Red Book of Brazilian Flora (Forzza et al. 2013), which follow the criteria established by the International Union for Conservation of Nature (IUCN 2025). Additionally, scientific studies and articles published by specialists were considered, especially when species were directly assessed by their authors. Thus, the classifications reflect an integrated and updated evaluation of extinction risk, which is fundamental for guiding public policies and conservation strategies.
5. Geographic analysis and mapping
For the preparation of the geographic distribution map, occurrence records of species were collected based on herbarium labels available on the SpeciesLink (2025, continuously updated) and Virtual Herbarium REFLORA (2025, continuously updated) platforms, with locations verified using Google Maps. The map was generated using QGIS v. 3.34 software (QGIS Development Team, 2025), employing vector layers of biomes and political boundaries available in the Geocentric Reference System for the Americas (SIRGAS 2000), with the aim of visualizing the geographic distribution of species in the state of Bahia, as well as the municipality of description of Bromeliaceae taxa.
Results and Discussion
1. Richness and taxonomic composition
A total of 433 taxa were recorded in the state of Bahia, comprising 426 species, six botanical varieties, and one subspecies, belonging to 36 genera (Table 1). It was observed that the species are distributed throughout the state, occurring in the three phytogeographical domains, with a notable concentration in the Atlantic Forest of the southern region, which harbors the highest number of described species of the family, as well as the greatest degree of endemism (Figures 1 and 2).
List of Bromeliaceae species occurring in Bahia with their respective endemism (END), threat status (THR): CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; LC = Least Concern; DD = Data Deficient. Life form (LFO), biome of occurrence (BIO), occurrence in other states (OCC), city of species description (DES), year of publication (PUB), and voucher deposited in different herbaria (Voucher).
Distribution of endemic Bromeliaceae species and the corresponding municipalities of their original description in Bahia, according to the different phytogeographic domains.
Occurrence of Bromeliaceae species across different vegetation types in the state of Bahia. (A) Lymania involucrata, a microendemic species recorded in Dense Ombrophilous Forest at the Reserva Ecológica da Michelin, Igrapiúna. (B) Aechmea ampla, a microendemic species occurring in restinga areas in southern Bahia, recorded in Belmonte. (C) Wittmackia lingulatoides, a species occurring in Caatinga and Seasonal Semideciduous Forest, recorded in an urban area in Amargosa. (D) Pseudaraeococcus serranensis, a microendemic species recorded in Seasonal Semideciduous Forest in Brejões. (E) Aechmea bahiana, a microendemic species from the Chapada Diamantina, recorded in Rio de Contas. (F) Tillandsia usneoides, a widely distributed species in the Americas, occurring in Caatinga in Milagres. (G) Encholirium spectabile, Hohenbergia flava, and Vriesea neoglutinosa occurring on Caatinga inselbergs in Milagres. (H) Tillandsia oliveira and Encholirium spectabile, occurring on Caatinga inselbergs in Itatim. (I) Ananas ananassoides, a widely distributed species in the Brazilian Cerrado, recorded in Barreiras. Photos: E.H. Souza.
The high degree of endemism observed in the Atlantic Forest of southern Bahia can be attributed to a combination of ecological, historical, and anthropogenic factors that make this region a unique center of diversity in Brazil (Ferreira et al. 2024). Humid and stable climatic conditions, with annual precipitation exceeding 2,000 mm and high temperatures throughout the year, favor the maintenance of Dense Ombrophilous Forests, which in turn support highly specialized plant communities (Peres et al. 2020, Ferreira et al. 2024). Furthermore, the historical floristic isolation of the region, separated from other forest formations by areas of Cerrado and Caatinga, has favored allopatric speciation processes over time, resulting in a flora with a high degree of endemism (Mori et al. 1981). Traditional agricultural practices, such as the cabruca system, based on cacao cultivation under the shade of native trees, have also contributed to biodiversity conservation by maintaining forest structure and allowing the persistence of native and endemic species in anthropized landscapes (Schroth & Harvey 2007, Ferreira et al. 2024). According to the Critical Ecosystem Partnership Fund (CEPF 2020), the region hosts several endemic and threatened species, justifying its recognition as one of the highest conservation priority areas within the Atlantic Forest.
The distribution map of endemic Bromeliaceae species reveals that the central-southern region of Bahia harbors the highest diversity of these plants. The municipality of Una stands out with 15 species, followed by Guaratinga with nine species and Uruçuca with eight species. Camacan and Wenceslau Guimarães each host seven species, while Itamaraju, Milagres, Rio de Contas, and Santa Teresinha present six taxa. Andaraí, Itapetinga, Morro do Chapéu, Mucugê, Palmeira, and Vitória da Conquista each contain five species, and the remaining municipalities have distributions of fewer than five species. The highest concentration of these taxa occurs in the Atlantic Forest biome, where diversity is strongly associated with factors such as climate and high temperatures, contributing to the high ecological plasticity of the central-southern Bahia region (Peres et al. 2020, Ferreira et al. 2024).
The subfamily Bromelioideae was the most representative, with 24 genera and 308 taxa, followed by Tillandsioideae with eight genera and 102 taxa, Pitcairnioideae with four genera and 22 species, and Navioideae with one species (Table 1). These numbers confirm that Bahia is one of the main centers of diversity for the Bromeliaceae family in Brazil (Flora e Funga do Brasil 2025).
The most representative genera in Bromelioideae were Aechmea Ruiz & Pav. and Hohenbergia Schult. & Schult.f. with 52 taxa, Cryptanthus Otto & A.Dietr. with 38 species, and Orthophytum Beer and Wittmackia Mez with 27 species (Figure 3). In Tillandsioideae, the most representative genera were Vriesea Lindl. with 55 taxa and Tillandsia L. with 32 taxa. In Pitcairnioideae, the most representative genera were Dyckia Schult. & Schult.f. and Encholirium Mart. ex Schult. & Schult.f., with nine species each. Finally, the subfamily Navioideae comprises only one genus in the state, Cottendorfia Schult. & Schult.f. (Figure 3).
The remarkable diversity of species in the subfamily Bromelioideae can be attributed to an adaptive radiation process, reflected in the wide morphological and ecological variation observed in the group (Smith & Downs 1974, Benzing 2000). Many of these species have developed an epiphytic habit and specialized structures, such as peltate trichomes and leaf rosettes capable of accumulating water and organic debris, forming phytotelmata. These adaptations, combined with crassulacean acid metabolism (CAM), confer remarkable efficiency in nutrient acquisition and water conservation (Smith & Downs 1974, Benzing 2000). Furthermore, the occurrence of species from this subfamily across different edaphoclimatic conditions in the state—from humid environments to dry and rocky regions—demonstrates their ecological plasticity. This adaptive capacity is closely linked to the group’s rich morphological diversity and the high occurrence of endemic species, particularly in ecosystems such as the Bahian Atlantic Forest and rupestrian fields (Smith & Downs 1974, Benzing 2000).
2. Distribution and ecological patterns
The state of Bahia encompasses three important phytogeographical domains: the Atlantic Forest, the Cerrado, and the Caatinga, in addition to distinct phytophysiognomies, whose proximity and transition are facilitated by a wide altitudinal range (from sea level up to over 2,000 meters) and a marked thermal gradient (with mean annual temperatures ranging from approximately 14 °C to 35 °C). These factors contribute to high environmental heterogeneity and considerable floristic diversity (Campos et al. 2023).
The Bromeliaceae species present in Bahia exhibit considerable ornamental and economic potential, primarily due to their morphological diversity and environmental adaptations. Many of them have foliage with intense coloration, long-lasting inflorescences, and architectural forms that make them highly valued for landscaping and ornamental cultivation, both indoors and outdoors (Souza et al. 2014, 2017, 2025, Donato et al. 2018, Mota et al. 2023). Additionally, some species have direct economic relevance, such as Ananas comosus, widely cultivated for fruit production, particularly in the regions of Itaberaba and Coração de Maria (Matos et al. 2016), and Neoglaziovia variegata, known for the traditional use of its fibers in the manufacture of ropes, nets, and handicrafts, mainly in the Valente region (Silveira et al. 2009). The presence of these genera demonstrates the utilitarian potential of Bahia’s flora, with applications ranging from ornamental and food uses to the textile industry, highlighting not only the ecological but also the socioeconomic importance of these species (Souza et al. 2018, Mota et al. 2024, Silva et al. 2025).
The Atlantic Forest harbors the highest species richness, with 266 records (61.43%), followed by the Caatinga with 86 taxa (19.86%) and the Cerrado with 14 species (3.23%) (Figure 4). The occurrence of 67 taxa in more than one phytogeographical domain demonstrates the ecological plasticity and adaptive potential of these species.
Regarding the distribution of subfamilies by phytogeographical domain, Bromelioideae exhibits 212 taxa in the Atlantic Forest, 62 in the Caatinga, three in the Cerrado, and 31 occurring simultaneously across all three domains. For the subfamily Tillandsioideae, 53 taxa are recorded in the Atlantic Forest, 12 in the Caatinga, five in the Cerrado, and 32 occur in all three biomes. In Pitcairnioideae, 11 species are found in the Caatinga, six in the Cerrado, and only one in the Atlantic Forest, with four species occurring across all three phytogeographical domains. Finally, in Navioideae, the single species occurs in the rupestrian fields of the Chapada Diamantina (Figure 4).
The Atlantic Forest is one of the most species-rich biomes and exhibits a high degree of endemism. However, it is also one of the most threatened domains globally, having suffered intense fragmentation of its original coverage over recent decades (SOS Mata Atlântica 2025). This degradation is strongly associated with fires, predatory extractive practices, and the uncontrolled expansion of human activities, compromising ecosystem integrity (Tabarelli et al. 2005). It is estimated that the Atlantic Forest harbors approximately 20,000 plant species, of which around 8,000 are endemic (SOS Mata Atlântica 2025).
Originally, this biome occupied an area of 1,110,182 km2, corresponding to about 15% of the national territory. Currently, only approximately 12.5% of its original forest cover remains (SOS Mata Atlântica 2025). The Atlantic Forest encompasses different native forest formations, such as Dense Ombrophilous Forest, Mixed Ombrophilous Forest (also known as Araucaria Forest), Open Ombrophilous Forest, Semi-deciduous Seasonal Forest, and Deciduous Seasonal Forest, in addition to associated ecosystems such as mangroves, restingas, high-altitude fields, inland wetlands, and forest enclaves in the Northeast (Brasil 2022a).
The predominance of Bromelioideae in the Atlantic Forest can be explained by the structural and environmental complexity of this phytogeographical domain, which presents high relative humidity, elevated annual rainfall, and great topographic heterogeneity, creating multiple microhabitats favorable to the diversification of Bromeliaceae species, especially epiphytic ones (Siqueira Filho & Tabarelli 2006, Martinelli et al. 2008, Ferreira et al. 2024). According to Benzing (2000), the constant humidity and the availability of arboreal niches in the Atlantic Forest are key factors for the proliferation and specialization of epiphytic Bromeliaceae, which rely on humid environments for their reproductive and physiological development. Scarano (2002) and Ferreira et al. (2024) further emphasize that the Atlantic Forest harbors one of the greatest floristic diversities in the world, with a high degree of endemism, favoring the presence of taxa exclusive to Bromelioideae. The variable altitude, which in Bahia can reach over 2,000 meters in the Chapada Diamantina National Park, also expands climatic diversity, resulting in specific adaptations and higher species richness (Giulietti et al. 2009).
The Caatinga, an exclusively Brazilian biome, occupies approximately 862,818 km2, representing 10.1% of the national territory (IBGE 2019). It is rich in biodiversity, with about 4,963 plant species. However, it faces threats such as wildfires, deforestation, and predatory extractive practices. Approximately 80% of its original area has already been altered (Brasil 2022a). In 2024, about 174,511 hectares were deforested for wind and solar energy projects, as well as agricultural expansion (MapBiomas 2024). This biome, characterized by more adverse environmental conditions such as low rainfall, high temperatures, rocky soils, and prolonged dry periods, harbors 84 Bromeliaceae species. This number is highly relevant, demonstrating the family’s adaptive capacity to semi-arid ecosystems (Souza et al. 2021a). Bromeliaceae species in the Caatinga exhibit morphological and physiological adaptations, such as leaves covered with trichomes that reduce water loss and specialized roots for efficient water uptake, reinforcing their ecological role in xerophytic environments (Rodal et al. 2008, Souza et al. 2021a). Additionally, the Caatinga is recognized for its highly endemic flora, which is also reflected in the occurrence of Bromeliaceae adapted to specific ecological niches, contributing to the maintenance of regional biodiversity (Leal et al. 2005).
It is noteworthy that many species of the genus Tillandsia exhibit high ecological specialization, occurring mainly on inselbergs within the Caatinga, as observed in the regions of Itatim and Milagres, which harbor several microendemic species (Hromadnik & Hromadnik 2020, 2021, Souza et al. 2021a). This specialization is due to the unique environmental conditions of these rocky outcrops, characterized by shallow soils, high solar exposure, extreme temperature fluctuations, and limited water availability, promoting geographic and selective isolation, which favors local speciation processes and endemism (Leodegário et al. 2020, Souza et al. 2021a).
Although recognized worldwide as a biodiversity hotspot, the Cerrado exhibits a lower number of Bromeliaceae species in the state of Bahia, covering approximately 1,983,017 km2 (23.3% of the national territory) (Brasil 2022a). It comprises different phytophysiognomies, including grassland, savanna, and forest formations. The biome harbors around 12,829 plant species, many of which have potential for the recovery of degraded areas and soil protection. However, it suffers from deforestation, wildfires, habitat fragmentation, and agricultural pressure. Currently, only 49.9% of its native vegetation remains (Klink & Machado 2005a, Brasil 2022a).
The limited occurrence of Bromeliaceae in this biome may be associated with the predominance of open formations, with acidic soils and edaphic conditions that favor the presence of grasses and legumes to the detriment of Bromeliaceae (Klink & Machado 2005b). Additionally, the lower representation may reflect gaps in sampling effort, as many areas of the Bahian Cerrado still lack detailed floristic inventories (França & Giulietti 2019). Nevertheless, the presence of species typical of the subfamily Pitcairnioideae, generally adapted to drier conditions and rupestrian environments, indicates the family’s capacity for colonization and specialization within this biome.
Therefore, the greater richness and diversity of Bromeliaceae in the Atlantic Forest of Bahia can be attributed to a combination of climatic factors (high humidity and mild temperature), geomorphological factors (broad altitudinal gradient and complex relief), and ecological factors (availability of niches for epiphytic species). These conditions provide a favorable environment for the adaptive radiation of Bromeliaceae, particularly epiphytic species and those with high ecological specialization (Benzing 2000, Scarano 2002). The significant presence in the Caatinga highlights the phenotypic and physiological plasticity of the family, allowing its survival in arid environments subjected to intense water stress. Finally, although the Cerrado exhibits lower Bromeliaceae diversity in Bahia, it represents an important biogeographic corridor and refuge for some specialized species, emphasizing the need for further studies to better understand the distribution and conservation potential of Bromeliaceae in this biome.
3. Endemism
Regarding endemism, 241 taxa were found to be exclusive to the state of Bahia, corresponding to 55.6% of the total recorded species (Figure 5). The subfamily Bromelioideae presents the highest number of endemic species (192 taxa), followed by Tillandsioideae (37 taxa), Pitcairnioideae (13 taxa), and Navioideae (1 taxon) (Figure 5). Among the species with restricted distribution in Bahia, numerous taxa of Hohenbergia, Lymania, Pseudaraeococcus, Tillandsia, Cryptanthus, and Wittmackia stand out.
The high rate of endemism confirms Bahia as an important center of diversification and speciation for the Bromeliaceae family, resulting from the interaction of various environmental, historical, and geographic factors. The presence of three phytogeographical domains, combined with a pronounced altitudinal and thermal gradient, generates a wide variety of ecological niches, promoting population isolation and consequently favoring speciation (Giulietti et al. 2009, Flora e Funga do Brasil 2025). In addition, transition areas between biomes and geologically ancient environments, such as inselbergs and mountain ranges in Chapada Diamantina and northeastern Bahia, provide unique conditions that contribute to the emergence of microendemic species (Martinelli et al. 2008, Versieux & Wendt 2007).
The concentration of species with extremely restricted distribution makes these regions a priority for conservation actions, as these taxa are highly vulnerable to environmental changes, particularly habitat loss and fragmentation, fires, mining, invasive species, and urban expansion (Zappi et al. 2017). Therefore, conservation strategies should include the expansion and strengthening of fully protected conservation units, the promotion of ex situ conservation programs (such as seed banks and living collections), the restoration of degraded habitats, and the continuous mapping and monitoring of threatened species populations (Ferreira et al. 2024). Integration between scientific research, public policies, and environmental education is essential to ensure the preservation of Bahia’s unique biodiversity and to prevent local extinction of endemic species with highly restricted distributions (Rocha et al. 2004).
4. Conservation status and threats
Of the 433 Bromeliaceae species recorded for Bahia, only 180 (41.57%) have been assessed for conservation status. Among these, 21 species (4.84%) are classified as “Critically Endangered” (CR), 20 (4.61%) as “Vulnerable” (VU), while 48 (11.08%) fall under the “Endangered” (EN) category, distributed among the three subfamilies (Table 1, Figure 6). It is noteworthy that the conservation status of these species was determined based on official lists of threatened species (Forzza et al. 2013, Bahia 2017, Brasil 2022b) and recently published risk assessments in species descriptions.
Number of taxa by IUCN Red List threat category. CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened; DD = Data Deficient; LC = Least Concern.
Despite these advances, most species still lack adequate evaluation. Approximately 252 taxa (58.3%) are currently classified as ‘Not Evaluated’ (NE), highlighting Cottendorfia florida Schult. & Schult.f., the only species belonging to the subfamily Navioideae and endemic to Bahia (Flora e Funga do Brasil 2025).
This high number of unevaluated species reflects a significant knowledge gap regarding the conservation of the family in the state (Figure 6). This deficiency compromises the effectiveness of conservation policies, as the lack of information on threat levels hinders the prioritization and protection of key biodiversity areas (Martinelli et al. 2008, Ferreira et al. 2024).
The situation becomes even more critical when considering the high level of endemism in Bahia’s flora. Many of these species occur in restricted areas and under strong anthropogenic pressure, significantly increasing the risk of local extinction. The study by Almeida Júnior et al. (2025) on the genus Quesnelia, endemic to the Atlantic Forest, exemplifies this scenario: prior to assessment, only four species were officially considered threatened; after a detailed analysis of distribution, population size, and threats, the authors identified 15 of the 26 assessed species as belonging to risk categories according to IUCN criteria. This example underscores the importance of in-depth taxonomic and ecological studies as a basis for effective conservation actions.
In the breakdown by subfamily, Bromelioideae concentrates the highest number of threatened species: 16 classified as ‘Critically Endangered’ (CR), 15 as ‘Vulnerable’ (VU), and 43 as ‘Endangered’ (EN). In addition, 39 species were considered ‘Least Concern’ (LC), five ‘Near Threatened’ (NT), and eight ‘Data Deficient’ (DD). In the subfamily Tillandsioideae, four species were identified as ‘CR’, five as ‘VU’, one as ‘EN’, five as ‘DD’, and 28 as ‘LC’. Pitcairnioideae presents one species as ‘CR’, one as ‘DD’, three as ‘EN’, and five as ‘LC’ (Figure 6).
The predominance of threatened species in Bromelioideae can be attributed to the higher vulnerability of their habitats, often located in areas of the Atlantic Forest, one of the phytogeographical domains most impacted by deforestation and urban and agricultural expansion (Ferreira et al. 2024). Illegal collection of ornamental species and extractivism are also relevant factors for the population decline of these taxa (SOS Mata Atlântica 2025). The absence of legal protection and ecosystem fragmentation make it urgent to expand conservation efforts, including the creation of protected areas, implementation of management plans, and promotion of scientific research focused on understudied groups.
5. Life forms and ecological distribution
Regarding growth habits observed in Bromeliaceae, epiphytism is the most representative, with 90 taxa, predominating in the subfamilies Bromelioideae and Tillandsioideae (Table 1 and Figure 7). Species with an exclusively terrestrial habit total 76 records, while 61 species are classified as rupicolous. Additionally, 206 species demonstrate ecological plasticity, occurring in more than one of these categories (Table 1 and Figure 7). The subfamily Bromelioideae, the most diverse in the state, also stands out for the broad range of ecological habits. It has 75 exclusively terrestrial taxa, 58 epiphytic, and 27 rupicolous. Furthermore, it exhibits various combinations of habits: seven species are epiphytic and/or rupicolous, 37 exhibit all three habits (epiphytic, rupicolous, and terrestrial), 76 combine epiphytic and terrestrial habits, and 28 species occur as both rupicolous and terrestrial. This diversity of growth forms reflects the high ecological plasticity of the group, facilitating its occupation of distinct environments, from shaded humid forests to open rocky areas.
In the subfamily Tillandsioideae, epiphytic species predominate (32 taxa), although 20 rupicolous taxa and one terrestrial taxon are also represented. Other combined habits are observed: 19 species occur as epiphytic and rupicolous, 15 as epiphytic, rupicolous, and terrestrial, 12 as epiphytic and terrestrial, three as rupicolous and terrestrial, and one exclusively terrestrial. This distribution reinforces the adaptive tendency of Tillandsioideae to occupy varied microhabitats, especially on arboreal and rocky substrates, often associated with drier or seasonally dry climates.
In Pitcairnioideae, habits are predominantly rupicolous, with 14 species found on rocky outcrops and nine species occurring as both rupicolous and terrestrial. This predominance is linked to morphological and physiological traits adapted to open, dry environments with high solar incidence, such as coriaceous leaves, presence of peltate trichomes, and in many cases, the use of Crassulacean Acid Metabolism (CAM), which contributes to water conservation (Crayn et al. 2004, Ferreira et al. 2024).
Overall, Bromeliaceae species in Bahia exhibit a wide diversity of life forms. Epiphytic species, common in humid forests, perform essential ecological functions, such as maintaining microhabitats, providing water for fauna, and cycling nutrients within the canopy (Romero et al. 2020). Conversely, rupicolous and terrestrial species are frequently associated with more arid and extreme environments, such as rocky outcrops, where they develop specialized adaptations that ensure survival (Benzing 2000). This variety of habits demonstrates the broad adaptive capacity of the Bromeliaceae family and underscores the importance of conserving different environments, from dense forests to rupestrian fields, as a strategy for maintaining regional biodiversity.
6. Taxonomic advances and data limitations
This study showed that 290 of the 433 Bromeliaceae species recorded in Bahia were originally described based on specimens collected within the state, representing 66.9% of the total. This number highlights Bahia as an important center of diversification for the family, particularly due to its diversity of environments and ecological transition zones (Pontes et al. 2020, Leme et al. 2025).
The subfamily Bromelioideae is by far the most representative, accounting for 229 of the 290 species initially described in Bahia (78.9%). Examples include Aechmea marcpignalii Leme, Bromelia boqueiranensis Leme & J.A.Siqueira, Cryptanthus cruzalmensis Leme & E.H.Souza, Pseudaraeococcus sessiliflorus (Leme & J.A.Siqueira) R.A.Pontes & Versieux, and Wittmackia aurantiolilacina Leme, E. Fernandez & Amorim. The subfamily Tillandsioideae contributes 45 described species (15.6%), including taxa such as Alcantarea nahoumii (Leme) J.R.Grant, Tillandsia itatiensis E.H.Souza & Leodegário, and Vriesea bahiana Leme. Pitcairnioideae, with 16 species (5.5%), includes representatives such as Dyckia burle-marxii L.B.Sm. & Read, Dyckia milagrensis Leme, Encholirium fragae Forzza, and Pitcairnia piepenbringii Rauh & E.Gross (Table 1). Navioideae includes only the species Cottendorfia florida Schult. & Schult.f.
During the construction of the database used in this study, significant inconsistencies were identified in the digital platforms Virtual Herbarium Reflora and SpeciesLink, raising concerns about the reliability of these systems when used as primary sources of taxonomic data. One example is the erroneous record of Aechmea huebneri Harms (US 2526828), an Amazonian species listed for Bahia, when in fact the specimen corresponds to Aechmea ampla L.B.Sm. This taxonomic confusion highlights the need for rigorous identification of exsiccates to avoid incorrect interpretations regarding the geographic distribution of species.
Furthermore, several species have not yet been officially incorporated into the Flora e Funga do Brasil database, such as Hohenbergia alba B.P.Cavalcante, E.H.Souza & Versieux, Hohenbergia sparsiflora B.P.Cavalcante, E.H.Souza & Versieux, Hohenbergia amargosensis E.H.Souza & Leme, and Wittmackia aurantiolilacina. The absence of these species in digital platforms limits access to information and compromises analyses related to the total number of taxa, their distribution, life forms, and threat status.
Another challenge is the lack of nomenclatural updates across various databases. For example, the genus Araeococcus, now segregated as Pseudaraeococcus based on phylogenetic studies by Pontes et al. (2020), continues to be listed under its former classification. The same occurs with Wittmackia species, still erroneously included under the genus Aechmea (Aguirre-Santoro 2017). These taxonomic inconsistencies complicate researchers’ work and may negatively impact data compilation and the formulation of conservation policies.
Additionally, the synonymization of Encholirium with Dyckia, proposed in some recent revisions (Gomes-da-Silva et al. 2019), is not widely accepted by the national and international scientific community, reflecting the need for more rigorous studies before adopting significant nomenclatural changes.
Finally, in SpeciesLink, inconsistencies were also observed regarding exsiccate images, many of which are mislabeled or even belong to other botanical families. Such errors not only hinder the correct identification of specimens but also compromise the reliability of data used in scientific studies. To mitigate these problems, a rigorous review of the provided data is recommended, with direct involvement of taxonomic specialists in curating the information.
The temporal analysis of historical records of Bromeliaceae species described in Bahia shows significant growth, although marked by some fluctuations, over nearly three centuries (Table 1; Figure 8). The first milestone in Bromeliaceae taxonomy was the description of the genus Bromelia by Carl Linnaeus in 1753, which laid the foundations for the study and classification of the family (Linnaeus 1753).
From 1753 to 1800, only eight species were recorded for Bahia, corresponding to the earliest collections and scientific descriptions of the local flora. During this period, pioneering naturalists such as Auguste de Saint-Hilaire, together with botanists like Joseph August Schultes, played a fundamental role in cataloging and expanding knowledge of Bromeliaceae, driving botanical exploration in the 18th and 19th centuries (Saint-Hilaire 1825, Schultes 1830).
The 19th century was marked by a significant increase in the number of described species, peaking at 44 species between 1876 and 1900. This growth reflected the intensification of floristic explorations in Brazil and the increasing interest of European botanists, such as James Smith, who played an important role in the classification and systematic organization of the family, enhancing the understanding of its diversity (Smith 1889).
In the 20th century, especially in the first five decades, the number of descriptions fluctuated, highlighting the work of taxonomist Lyman Smith, whose revisions were fundamental for consolidating Bromeliaceae systematics (Smith 1955). A significant advance occurred with the publication of Flora Neotropica (1974, 1977, 1979), organized by Smith and Downs, which represented a milestone in botany by systematizing and synthesizing knowledge about Bromeliaceae genera and species, strengthening the foundation for subsequent research (Smith & Downs 1974, 1977, 1979).
From the 1970s onwards, a new impulse in species descriptions is observed, with consistent growth culminating in a peak of 45 species described between 1986 and 1995. This period coincides with the consolidation of specialized research groups on the family and intensified collections in Bahia, reflecting greater technical and scientific capacity (Leme 1990).
Over the past two decades, although there have been some variations and a slight reduction in certain intervals, the number of descriptions has remained high, totaling around 120 species described between 2005 and 2025, indicating the continuity of scientific effort and collection work. Contemporary researchers, such as Elton Leme, have played a crucial role both in the description of new species and in taxonomic revisions, promoting deeper understanding of Bromeliaceae biodiversity in Bahia. Of the species described in the state, 139 were recorded or reclassified by this author (Leme 2010, 1995, Leme et al. 2010, 2014, 2020, 2022, 2023, 2025, Leme & Kollmann, 2011, 2013). In the past five years, our research group, represented by Everton Hilo de Souza, has described 22 new species from Bahia (Cavalcante et al. 2020, 2022a, 2022b, 2024, Leme et al. 2020, 2023, 2025, Leodegário et al. 2020, Souza et al. 2021b, 2021c, 2022, 2025, Souza & Leme 2021, Pontes et al. 2021, Ferreira et al. 2024).
Among the recently described species, notable examples include Pseudaraeococcus lageniformis (R.A.Pontes & Versieux) R.A.Pontes & Versieux, Cryptanthus cruzalmensis Leme & E.H.Souza, C. santateresinhensis Leme, Orthophytum brejoense Leme, Vidigal & E.H.Souza, Hohenbergia ituberaensis B.P.Cavalcante, E.H.Souza & Versieux, H. amargosensis E.H.Souza & Leme, Alcantarea vanderbergii Versieux, Tillandsia itatiensis E.H.Souza & Leodegário, T. jequiensis L.Hrom. & H.Hrom., Wittmackia guedesiae E.H.Souza & Leme, W. conduruensis Leme & E.H.Souza, and W. aurantiolilacina Leme, E.Fernandez & Amorim (Cavalcante et al. 2020, Hromadnik & Hromadnik 2020, Leme et al. 2020, 2023, 2025a, 2025b, Leodegário et al. 2020, Pontes et al. 2020, Souza et al. 2022, 2025).
In 2025, new species have already been described for Bahia, including Aechmea marcpignalii Leme, Bromelia boqueiranensis Leme & J.A.Siqueira, Cryptanthus cajuitensis Leme & L.Kollmann, C. spathulatifolius Leme & E.H.Souza, Hohenbergia kollmannii Leme & A.P.Fontana, Orthophytum pseudoestevesii Leme, Portea santosiana Leme & E.H.Souza, Vriesea punctatissima Leme & E.H.Souza, V. serraourensis E.H.Souza & Leme, and W. aurantiolilacina (Leme et al. 2025a, 2025b, Souza et al. 2025). Furthermore, new genera with representatives in Bahia have also been proposed, such as Karawata J.R.Maciel & G.Sousa and Pseudaraeococcus (Mez) R.A.Pontes & Versieux (Maciel et al. 2020; Pontes et al. 2020).
This historical progress highlights the gradual evolution of scientific effort and botanical collection and analysis techniques in Bahia. The contributions of early pioneers, followed by the systematization promoted by Smith and Downs, and the continuous work of current specialists, reflect a trajectory of advances that remains ongoing, given that the inventory of Bromeliaceae in the region is still incomplete. Investment in research and the strengthening of conservation initiatives are therefore essential to expand knowledge and ensure the preservation of this important botanical family.
Conclusion
The systematization of Bromeliaceae diversity in Bahia highlights the state as one of the main centers of diversification and endemism for the family in Brazil, with a total of 433 recorded taxa, of which 55.6% are endemic. The subfamily Bromelioideae proved to be the most diverse, followed by Tillandsioideae and Pitcairnioideae, with emphasis on the genera Vriesea, Aechmea, Hohenbergia, Cryptanthus, Tillandsia, and Wittmackia, which are the most representative in the Bahian flora.
The predominance of species in the Atlantic Forest reinforces the ecological importance of this biome, whose environmental complexity, high humidity, and altitudinal variation favor diversification, particularly of epiphytic species. The data also demonstrate the remarkable ecological plasticity of Bromeliaceae, reflected in the diversity of life forms and the capacity to adapt to both humid and arid environments, including the Caatinga and Cerrado. However, despite this richness, there remains a significant knowledge gap regarding the conservation status of most species, with only 41.57% formally assessed, of which 89 are classified as threatened.
This scenario is further aggravated by inconsistencies in major taxonomic databases, which include incorrect records, outdated nomenclature, and the absence of recently described species. Such limitations compromise access to reliable information and hinder the formulation of effective conservation strategies. In this context, specialized scientific curation proves indispensable for the continuous correction and updating of information.
It can be concluded, therefore, that strengthening floristic inventories and taxonomic curation is essential for biodiversity conservation, guiding concrete actions for the management, protection, and sustainable use of Bromeliaceae species in Bahia.
Acknowledgements
The authors would like to thank the support from of Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) (Finance Code 001), Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB) (FIOL Project PPF 0014/2021) and Plano de Ação Nacional para a Conservação de Árvores Ameaçadas de Extinção do Sul da Bahia – PAN Hileia Baiana. We also thank the Reserva Ecológica Michelin (Process no.: 23007.00020318/2021-14) for providing the research permit and logistical and financial support; Parque Estadual da Serra do Conduru (PESC) and Instituto do Meio Ambiente e Recursos Hídricos (INEMA/Bahia: 046.0525.2022.0004974-68) for providing the research permit. FVDS and LYSA thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for a productivity grant (307128/20210 and 314502/2023-8, respectively).
Data Availability
The datasets generated during and/or analyzed during the current study are available at: https://doi.org/10.48331/SCIELODATA.0PEJFS.
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