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Lecythidaceae
Lecythidaceae
from Wikipedia

Lecythidaceae
Illustration of Couroupita guianensis by Lansdown Guilding
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Ericales
Family: Lecythidaceae
A.Rich.[1]
Type genus
Lecythis
Genera

See text

Synonyms[3]
  • Barringtoniaceae DC. ex F.Rudolphi
  • Scytopetalaceae Engl.
Barringtonia acutangula (freshwater mangrove) fruits in Kolkata, India
Careya arborea in Narsapur, Medak district, India

The Lecythidaceae (/ˌlɛsɪθɪˈds/ LESS-ith-ih-DAY-see) comprise a family of about 30 genera[3] and 250–300 species of woody plants native to tropical South America, Africa (including Madagascar), Asia and Australia.

Well known members of the family include the cannonball tree (Couroupita guianensis) and the edible Brazil nut (Bertholletia excelsa).[4]

Genera

[edit]

30 genera are currently accepted.[3]

Taxonomy

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According to molecular analysis of Lecythidaceae, including work by Mori et al. (2007),[5] subfamilies include:

Barringtonioidea

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Previously Barringtoniaceae;[6] also sensu Takhtajan 1997;[6] this subfamily was also called Planchonioideae (which included Barringtonia). Genera are restricted to the Old World tropics.

Foetidioideae

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Previously Foetidiaceae[6] from Madagascar is monogeneric:

Lecythidoideae

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Genera restricted to the New World tropics.[7]

Scytopetaloideae

[edit]

The APG II system of 2003 included genera from the family Scytopetalaceae and others

Napoleonaeoideae

[edit]

Previously as family Napoleonaeaceae;[6] species are native to Africa.

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Lecythidaceae, the , is a family of flowering in the order , comprising approximately 25 genera and 300 species of mostly evergreen trees and shrubs that inhabit humid tropical forests. These are distinguished by their simple, alternate, distichous leaves with serrate margins, large and showy hermaphroditic flowers featuring numerous stamens and an inferior , and hard woody fruits that are typically capsular or indehiscent and contain many large seeds. The family exhibits a mix of zygomorphic and actinomorphic flowers across its genera, reflecting evolutionary adaptations for by bees, bats, and other animals. The center of diversity for Lecythidaceae lies in the Neotropics, particularly the Amazonian rainforests of , where about 11 genera and 180 species occur, making it one of the most abundant families in terms of density—ranking third in Amazonian forests based on extensive inventories. The largest genus, Eschweilera, includes over 90 species and dominates in abundance, followed by (around 41 species) and Lecythis (26 species). While predominantly Neotropical, the family extends to other regions, including five genera and 71 species in (subfamily Barringtonioideae) and a single genus in , with overall distribution ranging from southward to but absent from . Ecologically, Lecythidaceae play a key role in tropical ecosystems through their contributions to forest structure, networks, and mechanisms, which include animal-mediated (e.g., via large fruits attracting mammals) and or dispersal in some . Economically, the family is significant for timber production from genera like Cariniana and Couratari, as well as edible nuts from Bertholletia excelsa, the tree, which supports and local livelihoods in the Amazon. Ornamental , such as the cannonball tree (), are also cultivated for their striking flowers and spherical fruits. Ongoing research emphasizes , conservation, and , given threats from in their native habitats.

Description

Habit and vegetative characteristics

Lecythidaceae species exhibit a predominantly woody habit, primarily as trees or treelets, with shrubs occurring rarely. These often form large canopy trees in tropical forests, such as Bertholletia excelsa, which can reach heights of 40–60 m with a straight trunk up to 2–4 m in diameter and an umbrella-shaped crown spanning 10–20 m. Branching patterns vary, typically featuring monopodial or sympodial growth, with branches often plagiotropic and sparsely distributed in some genera, leading to a poorly branched appearance overall. Leaves in the family are simple, alternate, and arranged in a spiral phyllotaxy, though they may appear distichous or condensed at branch apices, giving a verticillate in certain . They are petiolate, exstipulate or bearing small, inconspicuous stipules, and possess chartaceous to leathery blades with entire, serrate, or crenate margins. Venation is pinnate, characterized by parallel secondary and tertiary veins that are often prominent, especially on the abaxial surface, as seen in Bertholletia excelsa where blades measure 17–45 cm long by 6.5–15.5 cm wide. Leaves are typically glabrous and lack glands. The bark of Lecythidaceae is fibrous and often exudes a linseed-like when cut, serving as a diagnostic trait. In mature trees, it is commonly smooth to fissured, with deep vertical ridges developing over time; for example, in Bertholletia excelsa, the grey-brown bark is 1.5 cm thick, resinous, and reddish internally. Larger species frequently develop low, thick buttresses at the base of the trunk for stability.

Flowers, fruits, and seeds

The flowers of Lecythidaceae are typically large and showy, exhibiting either actinomorphic (radially symmetric) or zygomorphic (bilaterally symmetric) forms, and are hermaphroditic. They possess 2–6 sepals that are often valvate or imbricate, and 3–6 (rarely up to 18) free, imbricate petals, though petals are absent in some genera like Foetidia. The androecium is distinctive, featuring numerous stamens—often equal in number to the petals and opposite them—basally connate into a tube or ring, with anthers that are typically porose and thecae of unequal length; staminodes frequently form a hood- or cap-like structure enclosing the stigma. The gynoecium includes an inferior ovary with 2–6 locules, each containing 2–many ovules, topped by a simple or divided style. Inflorescences vary from terminal or axillary racemes and panicles to spikes or solitary flowers, sometimes cauliflorous or ramiflorous, and often include modified androecial structures serving as floral nectaries. Fruits in the family are diverse but predominantly woody capsules or berries, arising from the multi-locular inferior , and may be dehiscent via an operculum (lid-like structure) or indehiscent. Representative examples include the urn-shaped, woody capsules of Pachylecythis pisonis (formerly Lecythis pisonis), which can reach up to 30 cm in diameter and open by a basal operculum to release seeds. In , fruits are large, spherical, and woody—resembling cannonballs, typically 15–25 cm in diameter—with a hard pericarp that dehisces irregularly upon maturity. Bertholletia excelsa produces rounded, woody pods about 10–15 cm in diameter that split open to reveal multiple seeds. Seeds are generally large, with copious endosperm rich in proteins and oils, and an embryo that is straight to coiled with variable cotyledons; they often feature accessory structures such as a basal aril or wing for protection or potential dispersal aid. In Bertholletia excelsa, the seeds are triangular, 4–5 cm long, and packed tightly (10–25 per fruit) within the pod, enclosed in a hard testa. Seeds of Lecythis species, e.g., L. lurida, are fusiform or triangular, up to 6 cm long, with a thin testa and often a white, fleshy aril surrounding the funicle.

Distribution and habitat

Geographic distribution

The Lecythidaceae family exhibits a distribution, primarily occurring in the Neotropics, Paleotropics, and parts of , with no representatives in temperate zones or latitudes beyond approximately 23°N or 23°S. The family comprises around 25 genera and 300 species, with the greatest diversity concentrated in humid tropical regions. In the Neotropics, which serve as the primary center of diversity, approximately 180 species are found across 11 genera, predominantly in , especially the where up to 60 species can occur in a single grid square. This region spans from southward to , though abundance and peak in the Amazon. Neotropical dominance is exemplified by genera such as Eschweilera, with about 84 species mostly confined to Amazonian lowlands, and Couratari, comprising 19 species primarily in Amazonian and Guianan forests from to . A notable specific range within this center is Bertholletia excelsa, the tree, which is endemic to the lowland rainforests of the . In contrast, the Paleotropics host around 80 species, mainly in (including , with 71 species across 5 genera), , and the region. Here, Barringtonia stands out with approximately 69 species distributed along coasts and tropical shores of the . The family's distribution shows clear disjunct patterns between and clades, reflecting ancient vicariance events with no overlap between continents. In , diversity is lower at about 40 species, concentrated in West and Central regions, including . For instance, the genus Foetidia includes 17 species restricted to and nearby islands, such as the , with one species extending to . These patterns underscore the family's evolutionary history tied to tropical Gondwanan fragments, with subclades like Lecythidoideae exclusive to the Neotropics and Barringtonioideae to the Paleotropics.

Habitat preferences

Lecythidaceae predominantly occupy humid tropical rainforests, seasonal , and mangroves across their range, thriving in environments characterized by high and structural complexity. These exhibit an altitudinal distribution from to approximately 1,500 m, with many favoring lowland settings while others extend into premontane zones. Their woody habit contributes to canopy dominance in these biomes, allowing them to form integral parts of the . In terms of soil preferences, Lecythidaceae generally require well-drained, fertile loams typical of upland forests, which support their growth in non-flooded terra firme habitats. However, certain genera demonstrate remarkable adaptability, with Barringtonia species tolerating periodic ing and even saline conditions in and margins. This tolerance enables Barringtonia to colonize riverine and coastal edges where levels fluctuate seasonally. Climatically, the family demands high annual rainfall ranging from 1,500 to 3,000 mm and consistent temperatures between 20°C and 30°C to sustain their or habits in humid . While most species are sensitive to prolonged dry periods, savanna-edge taxa like display notable , surviving seasonal water deficits through thick bark and leaf shedding. Within specific ecosystems, Lecythidaceae achieve prominence in Amazonian terra firme forests, where genera such as Cariniana contribute significantly to canopy composition, often representing a substantial proportion of large emergent trees. This dominance underscores their ecological role in maintaining forest structure and in non-inundated upland rainforests.

Taxonomy and phylogeny

Etymology and classification history

The name Lecythidaceae derives from the type genus Lecythis, which was named by in 1753 from the word lekythos (λήκυθος), referring to an oil flask or jar, in allusion to the distinctive urn-shaped fruits of species in the genus. The family itself was formally established as Lecythideae by French botanist Achille Richard in his 1825 Dictionnaire classique d'histoire naturelle, marking the initial recognition of these plants as a distinct group based on their woody habit and floral characteristics. In the 18th century, Linnaeus classified early described species of what are now Lecythidaceae within the family , such as placing Lecythis species under Eugenia due to superficial similarities in fruit structure and tropical distribution. This association persisted into the early until Swiss botanist , in his 1828 Prodromus Systematis Naturalis Regni Vegetabilis, separated the group as the tribe Lecythideae within Myrtaceae, primarily on the basis of unique morphology, including the presence of hooded or connate petals and specialized androecial structures not found in typical myrtaceous taxa. Significant advancements in classification occurred in the late 20th century through the comprehensive monographs by botanists Ghillean T. Prance and Scott A. Mori, published in the Flora Neotropica series—Part I in 1979 covering actinomorphic-flowered New World genera, and Part II in 1990 addressing zygomorphic-flowered ones—which synthesized morphological data to recognize approximately 25 genera across the pantropical family, emphasizing its diversity in the Neotropics. These works built on earlier revisions and provided a stable framework for delimiting genera based on fruit, seed, and floral traits. The Angiosperm Phylogeny Group III (APG III) classification, published in 2009, further refined the family's position by placing Lecythidaceae firmly within the order Ericales, integrating molecular evidence with traditional morphology to confirm its ericalean affinities. A recent phylogenetic study by Oscar M. Vargas and colleagues in 2024 utilized complete plastome sequences and target capture data from 343 nuclear loci to reclassify the , revealing in genera like Lecythis and Eschweilera, and proposing taxonomic adjustments that refine the boundaries of actinomorphic-flowered genera within the Lecythidoideae.

Phylogenetic position

Lecythidaceae is positioned within the core of the order , where it forms a monophyletic group to the holoparasitic family Mitrastemonaceae; this combined is in turn to the remaining core , excluding the early-diverging balsaminoid lineage comprising , Marcgraviaceae, and Tetrameristaceae. This placement is derived from comprehensive molecular analyses incorporating (e.g., rbcL, ndhF, matK) and nuclear (e.g., 18S rDNA) loci across 22 families. Earlier studies using ndhF and trnL-F sequences had suggested alternative relationships, such as proximity to and the former Myrsinaceae (now subsumed in Primulaceae), but subsequent supermatrix approaches with broader sampling have refined the to the current consensus. Phylogenetic analyses confirm the monophyly of Lecythidaceae, resolving the family into two principal clades corresponding to geographic realms: the predominantly Neotropical Lecythidoideae and the Paleotropical Barringtonioideae (formerly Planchonioideae) along with other Paleotropical subfamilies (Foetidioideae, Napoleonaeoideae, and Scytopetaloideae). This internal structure is robustly supported by chloroplast ndhF and trnL-F sequence data, which highlight the deep divergence between and lineages while underscoring shared ancestral traits like large, actinomorphic flowers. The origins of Lecythidaceae trace to the , with a crown age estimated at approximately 83 million years ago based on fossil-calibrated phylogenies of . The family's distribution—spanning the Neotropics, (including ), and —is explained by vicariance driven by the fragmentation of , which separated ancestral populations and promoted the divergence of the major clades during the early diversification of the order. Distinguishing Lecythidaceae from other families are key synapomorphies, including the fusion of numerous stamens into a tubular or hood-like androecium that envelops the style, and a fully inferior with axile and multiple locules. These morphological innovations, evident across the family's woody habit and large-fruited species, evolved early in the lineage and contrast with the typically free-filamentous stamens and often superior or semi-inferior ovaries found in relatives like or .

Subfamilies and genera

The family Lecythidaceae is currently subdivided into five subfamilies based on molecular and morphological evidence, reflecting their pantropical distribution and diverse floral and fruit characteristics. Barringtonioideae, with 6 genera primarily distributed in the tropics, is characterized by actinomorphic flowers and capsular fruits, with some species adapted to coastal or environments; notable examples include Barringtonia (73 species), known for its buoyant seeds aiding dispersal in marine settings. Foetidioideae consists of a single , Foetidia (18 ), endemic to and nearby islands, featuring dioecious trees with unisexual flowers and a distinctive fetid from glandular structures. Lecythidoideae, comprising approximately 20 and the majority of the family's in the , is distinguished by large, urn-shaped flowers with hooded petals and operculate fruits; it includes the diverse Eschweilera (71 accepted ), a hyperdominant in Amazonian forests. The study by Vargas et al. resolved in the Bertholletia using plastome and nuclear data, erecting new such as Guaiania, Imbiriba, Pachylecythis, Scottmoria, and Waimiria, and refining boundaries in Eschweilera and Lecythis. Scytopetaloideae contains 6 genera restricted to and Amazonia, with zygomorphic flowers and winged fruits adapted to dispersal; Scytopetalum (3 ) is representative. Napoleonaeoideae includes 2 genera native to West and , marked by plicate (folded) petals in the buds and explosive mechanisms; Napoleonaea (about 10 ) exemplifies this with its intricate floral architecture. The family encompasses approximately 30 accepted genera in total, with recent taxonomic updates incorporating molecular data leading to mergers, re-circumscriptions, and new genera, such as the placement of Allantoma within Lecythidoideae following phylogenetic analyses distinguishing it from superficially similar Caryocaraceae allies. Genus-level diversity is highest in Lecythidoideae, where Eschweilera dominates with 71 species, followed by Gustavia (47 species) and Lecythis (26 species), underscoring the subfamily's role in Neotropical forest canopies.
SubfamilyNumber of GeneraAccepted Genera
Barringtonioideae6Abdulmajidia, Barringtonia, Careya, Chydenanthus, Planchonia, Petersianthus
Foetidioideae1Foetidia
Lecythidoideae~20Allantoma, Bertholletia, Cariniana, Couratari, Couroupita, Corythophora, Eschweilera, Grias, Guaiania, Gustavia, Imbiriba, Lecythis, Pachylecythis, Scottmoria, Waimiria (and others; recent additions from 2024)
Scytopetaloideae6Asteranthos, Brazzeia, Crateranthus, Oubanguia, Rhaptopetalum, Scytopetalum
Napoleonaeoideae2Crateranthus, Napoleonaea
This classification aligns with phylogenetic studies supporting monophyly for each subfamily while highlighting intergeneric relationships within Lecythidoideae.

Ecology

Pollination and reproduction

Members of the Lecythidaceae family exhibit predominantly entomophilous pollination, primarily facilitated by large-bodied bees such as euglossine and carpenter bees (e.g., Euglossa spp. and Xylocopa frontalis), which are attracted to the flowers' nectar, scents, and pollen rewards. In species like Bertholletia excelsa, the zygomorphic flowers feature specialized hoods that restrict access to robust pollinators capable of navigating the complex androecium to effect pollen transfer. Bat pollination has been documented in several Neotropical taxa, including Lecythis poiteaui and L. barnebyi, where nocturnal flowers produce strong odors and abundant nectar to accommodate chiropterophilous visitation. Avian pollination occurs sporadically, particularly in Old World species like Barringtonia racemosa, though bees remain the dominant vectors across the family. Breeding systems in Lecythidaceae are largely characterized by , promoting and high , as evidenced by multilocus rates near 1.0 in species such as Bertholletia excelsa and Couratari guianensis. Flowers are typically hermaphroditic with protandrous presentation to minimize , though partial self-compatibility has been observed in some individuals under environmental stress. In the genus Foetidia, populations exhibit variable selfing rates despite hermaphroditic flowers, potentially due to fragmented habitats influencing pollinator availability. The subfamily Lecythidoideae often displays , where bees vibrate the poricidal anthers to release , enhancing efficiency in pollen presentation. Flowering phenology varies with habitat; in seasonal forests, blooms are synchronized to the dry season, as in Bertholletia excelsa (August–November) and Careya arborea (March–April), coinciding with pollinator activity peaks. In everwet tropical regions, flowering can be continuous or suppressive, allowing extended reproductive windows. Post-pollination, syncarpic fruits develop slowly over 12–18 months, with maturation timed to wet seasons for seed viability, as seen in Bertholletia excelsa where fruits ripen from February to April following the prior year's flowering. Reproductive success is bolstered by high pollen-ovule ratios, such as 26,755 in B. excelsa, indicative of obligate xenogamy and adaptations for reliable cross-pollination.

Seed dispersal and role in ecosystems

Seed dispersal in Lecythidaceae varies by genus and reflects adaptations to tropical environments, primarily involving animal-mediated, wind, water, and gravity mechanisms. In the Neotropical subfamily Lecythidoideae, most species rely on zoochory, where seeds serve as diaspores with protective seed coats featuring lignified exotesta, tannin cells for chemical defense, and arils or sarcotesta to attract dispersers. For instance, in Bertholletia excelsa (Brazil nut), heavy woody capsules fall to the ground via gravity, after which agoutis (Dasyprocta spp.) act as primary dispersers by gnawing open the capsules and scatter-hoarding the large seeds, sometimes moving them up to several meters away; secondary dispersal can occur via tapirs or other mammals that consume and excrete intact seeds. In Lecythis species, such as L. pisonis, bats and small mammals disperse seeds attracted to basal arils, while the seed coats provide mechanical protection during handling. Wind dispersal occurs in genera like Cariniana and Couratari, where dehiscent capsules release winged seeds that glide away from the parent tree. In the pantropical genus Barringtonia, particularly mangrove-associated species like B. racemosa and B. asiatica, buoyant, fibrous fruits with air-filled chambers facilitate hydrochory, allowing long-distance dispersal by tides and currents along coastal and riverine habitats. Lecythidaceae play a pivotal role as keystone taxa in ecosystems, particularly in Amazonian rainforests where they contribute significantly to structural diversity and function. Emergent trees in genera like Bertholletia and Lecythis form part of the upper canopy, providing microhabitats such as rough bark that supports epiphytic orchids, bromeliads, and lichens, thereby enhancing overall . Their nuts and seeds integrate into food webs, sustaining populations of dispersers like agoutis, which in turn promote forest regeneration by caching uneaten seeds that germinate into shade-tolerant seedlings; this mutualism is crucial in nutrient-poor soils, where large seeds endow seedlings with substantial reserves for establishment. In Amazonian forests, Lecythidaceae often comprise 5-10% of canopy tree individuals in certain plots, underscoring their dominance and influence on community assembly and nutrient cycling. Evolutionary adaptations in Lecythidaceae further bolster their ecological contributions, with large seeds adapted to oligotrophic soils through enhanced nutrient storage in the and protective coats that deter predators and pathogens. These traits enable effective in low-fertility environments typical of Amazonian terra firme forests. While most thrive in moist rainforests, some, like certain Couroupita in transitional habitats, exhibit traits conferring partial fire tolerance, such as thick bark on adults that protects against low-intensity burns in savanna-forest edges, aiding post-fire recovery and maintenance. Overall, these interactions position Lecythidaceae as foundational elements in dynamics, supporting disperser populations and facilitating succession.

Human uses and conservation

Economic importance

The family Lecythidaceae holds significant economic value primarily through its contributions to , timber, medicinal applications, and other utilitarian products, with species like Bertholletia excelsa driving much of the global trade. of B. excelsa, known as Brazil nuts, are a major harvested from the , with worldwide production estimated at around 33,400 tons in 2021, supporting livelihoods for thousands of smallholder extractivists across , , and . However, production has declined recently due to the 2023-2024 , with global output dropping and prices surging as of 2025. These nuts are notably rich in , with a single nut providing 68-96 micrograms, exceeding the daily recommended intake of 55 micrograms for adults and contributing to their high market demand in health foods and . Additionally, certain species offer fruits and seeds for local consumption; for instance, the fruits of Gustavia augusta contain an , yellowish pulp surrounding the seeds, while those of Grias peruviana yield firm, walnut-like flesh that is gathered wild or cultivated in home gardens for its nutritional value. Timber from Lecythidaceae species is prized for its and versatility in and . Woods from Cariniana species, such as C. legalis (jequitibá), are heavy and resistant to decay, resembling in texture and used extensively for structural beams, , furniture, and in tropical regions. In contrast, Couratari species, including C. guianensis (tauari), produce lighter, fine-textured wood suitable for interior applications like furniture, paneling, and veneers, with a Janka rating of about 1,650, making it harder than red oak yet workable for detailed craftsmanship. Medicinal and cultural uses further underscore the family's importance in traditional practices. Bark infusions from Lecythis pisonis are employed in folk medicine to treat and , with phytochemical analyses confirming the presence of bioactive compounds like that support these applications. Culturally, species like (cannonball tree) are valued as ornamentals, planted in botanic gardens and urban parks for their striking, fragrant flowers and dramatic fruit, enhancing landscapes in tropical areas from the to . Other economic potentials include extracted from Careya arborea bark, which yield natural dyes for textiles and wood staining, offering an eco-friendly alternative to synthetic colors due to their and phenolic content. Emerging interest in highlights Lecythidaceae's role in sustainable ; B. excelsa, for example, shows high survival rates in degraded Amazon sites when integrated with secondary forests, promoting while generating long-term nut yields for smallholders.

Conservation concerns

Lecythidaceae species face significant threats from , primarily driven by in the , where approximately 20% of the original forest cover has been lost since the , severely impacting the habitats of key genera like Bertholletia and Eschweilera. Despite ongoing threats, rates decreased by 11% in the Brazilian for the period ending July 2025, the lowest in over a . Overharvesting for timber and non-timber products, such as Brazil nuts from Bertholletia excelsa, exacerbates these pressures, leading to population declines in exploited areas. In ecosystems, some species like encounter additional risks from coastal development and potential competition with , though this is less documented compared to inland threats. Many Lecythidaceae taxa are assessed as threatened on the , reflecting their vulnerability to these anthropogenic pressures. For instance, Bertholletia excelsa is classified as Vulnerable due to and loss from and . Several Eschweilera species, such as Eschweilera jacquelyniae (Endangered) and Eschweilera subcordata (Critically Endangered), face similar risks from selective and forest fragmentation in the Neotropics. In eastern Brazil's , assessments of 22 native Lecythidaceae species indicate that at least 14 are threatened, with ongoing evaluations highlighting the need for updated conservation priorities. Conservation efforts include the establishment of protected areas that safeguard Lecythidaceae diversity, such as Ecuador's , which harbors high Neotropical richness for genera like Grias and Couratari amid broader Amazon biodiversity hotspots. In , national regulations restrict the harvest and trade of Bertholletia excelsa to promote sustainable management, compensating for the absence of international listing. programs, including living collections and seed banking at institutions like the , support propagation and reintroduction efforts for across the family. Climate change poses emerging risks to Lecythidaceae, with models predicting range shifts in Paleotropical species like those in Barringtonia due to altered rainfall patterns and temperature increases in Southeast Asian habitats. In fragmented Neotropical populations, post-2020 genetic studies reveal reduced diversity and increased inbreeding in Bertholletia excelsa, heightening susceptibility to environmental stressors and limiting adaptive potential. These impacts underscore the urgency of integrating into ongoing protection strategies for the family.

References

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