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Lecythidaceae
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| Lecythidaceae | |
|---|---|
| Illustration of Couroupita guianensis by Lansdown Guilding | |
| Scientific 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] | |
| |


The Lecythidaceae (/ˌlɛsɪθɪˈdeɪsiː/ 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]
- Allantoma Miers
- Asteranthos Desf.
- Barringtonia J.R.Forst. & G.Forst.
- Bertholletia Bonpl.
- Brazzeia Baill.
- Careya Roxb.
- Cariniana Casar.
- Chydenanthus Miers
- Chytroma Miers
- Corythophora R.Knuth
- Couratari Aubl.
- Couroupita Aubl.
- Crateranthus Baker f.
- Eschweilera Mart. ex DC.
- Foetidia Comm. ex Lam.
- Grias L.
- Guaiania O.M.Vargas & C.W.Dick
- Gustavia L.
- Imbiriba O.M.Vargas, M.Ribeiro & C.W.Dick
- Lecythis Loefl.
- Napoleonaea P.Beauv.
- Oubanguia Baill.
- Pachylecythis Ledoux
- Petersianthus Merr.
- Pierrina Engl.
- Planchonia Blume
- Rhaptopetalum Oliv.
- Scottmoria Cornejo
- Scytopetalum Pierre ex Engl.
- Waimiria C.W.Dick & O.M.Vargas
Taxonomy
[edit]According to molecular analysis of Lecythidaceae, including work by Mori et al. (2007),[5] subfamilies include:
Barringtonioidea
[edit]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.
- Barringtonia J.R.Forst. & G.Forst. (synonym Abdulmajidia Whitmore)
- Careya Roxb.
- Chydenanthus Miers
- Petersianthus Merr.
- Planchonia Blume
Foetidioideae
[edit]Previously Foetidiaceae[6] from Madagascar is monogeneric:
- Foetidia Comm. ex Lam.
Lecythidoideae
[edit]Genera restricted to the New World tropics.[7]
- Allantoma Miers
- Bertholletia Bonpl.
- Cariniana Casar.
- Chytroma Miers
- Corythophora R.Knuth
- Couratari Aubl.
- Couroupita Aubl.
- Eschweilera Mart. ex DC.
- Grias L.
- Guaiania O.M.Vargas & C.W.Dick
- Gustavia L.
- Imbiriba O.M.Vargas, M.Ribeiro & C.W.Dick
- Lecythis Loefl.
- Pachylecythis Ledoux
- Scottmoria Cornejo
- Waimiria C.W.Dick & O.M.Vargas
Scytopetaloideae
[edit]The APG II system of 2003 included genera from the family Scytopetalaceae and others
- Asteranthos Desf., also as Asteranthaceae[6]
- Brazzeia Verc.
- Oubanguia Baill.
- Pierrina Engl.
- Rhaptopetalum Oliv.
- Scytopetalum Engl.
Napoleonaeoideae
[edit]Previously as family Napoleonaeaceae;[6] species are native to Africa.
- Crateranthus Baker f., incertae sedis according to Takhtajan,
- Napoleonaea P.Beauv.
References
[edit]- ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
- ^ "Lecythidaceae A.Rich., Dict. Class. Hist. Nat. [Bory] 9: 269 (1825), nom. cons". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew. 2025. Retrieved 21 June 2025.
- ^ a b c "Lecythidaceae A.Rich". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 23 October 2025.
- ^ Mori, Scott A.; Tsou, Chi-Hua; Wu, Chi-Chih; Cronholm, Bodil; Anderberg, Arne A. (2007). "Evolution of Lecythidaceae with an emphasis on the circumscription of neotropical genera: Information from combined NDHF and TRNL-F sequence data". American Journal of Botany. 94 (3): 289–301. Bibcode:2007AmJB...94..289M. doi:10.3732/ajb.94.3.289. PMID 21636402.
- ^ Mori, S. A.; Tsou, C. -H.; Wu, C. -C.; Cronholm, B.; Anderberg, A. A. (2007). "Evolution of Lecythidaceae with an emphasis on the circumscription of neotropical genera: Information from combined ndhF and trnL-F sequence data". American Journal of Botany. 94 (3): 289–301. Bibcode:2007AmJB...94..289M. doi:10.3732/ajb.94.3.289. PMID 21636402.
- ^ a b c d e Takhtajan, A. (1997). Diversity and classification of flowering plants. Columbia University Press. ISBN 0-231-10098-1.
- ^ Oscar M. Vargas, Drew A. Larson, Juvenal Batista, Xavier Cornejo, Bruno Garcia Luize, Diana Medellín-Zabala, Michel Ribeiro, Nathan P. Smith, Stephen A. Smith, Alberto Vicentini, and Christopher W. Dick "Reclassification of the Bertholletia Clade of the Brazil Nut Family (Lecythidaceae) Based on a Phylogenetic Analysis of Plastome and Target Sequence Capture Data," Harvard Papers in Botany 29(1), 159-179, (30 June 2024). https://doi.org/10.3100/hpib.v29iss1.2024.n18
External links
[edit]- Lecythidaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants. http://delta-intkey.com
- Barringtoniaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants. http://delta-intkey.com
- Foetidiaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants. http://delta-intkey.com
- Asteranthaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants. http://delta-intkey.com
- Napoleonaeaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants. http://delta-intkey.com
- The Lecythidaceae Pages by Scott A. Mori and Ghillean T. Prance
- Toa Alta, Puerto Rico
- Lecythidaceae in BoDD – Botanical Dermatology Database
- Lecythidaceae Pages
Lecythidaceae
View on GrokipediaDescription
Habit and vegetative characteristics
Lecythidaceae species exhibit a predominantly woody habit, primarily as evergreen trees or treelets, with shrubs occurring rarely. These plants 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.[1][5] 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.[6] 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 illusion in certain species. 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.[1][5] Leaves are typically glabrous and lack glands.[1] The bark of Lecythidaceae is fibrous and often exudes a linseed-like odor 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.[1][6][5]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.[7] 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.[8] 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.[9] The gynoecium includes an inferior ovary with 2–6 locules, each containing 2–many ovules, topped by a simple or divided style.[8] 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.[10] Fruits in the family are diverse but predominantly woody capsules or berries, arising from the multi-locular inferior ovary, and may be dehiscent via an operculum (lid-like structure) or indehiscent.[11] 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.[12] In Couroupita guianensis, fruits are large, spherical, and woody—resembling cannonballs, typically 15–25 cm in diameter—with a hard pericarp that dehisces irregularly upon maturity.[13] Bertholletia excelsa produces rounded, woody pods about 10–15 cm in diameter that split open to reveal multiple seeds.[14] 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.[14] 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.[15]Distribution and habitat
Geographic distribution
The Lecythidaceae family exhibits a pantropical distribution, primarily occurring in the Neotropics, Paleotropics, and parts of Africa, with no representatives in temperate zones or latitudes beyond approximately 23°N or 23°S.[11] The family comprises around 25 genera and 300 species, with the greatest diversity concentrated in humid tropical regions.[1] In the Neotropics, which serve as the primary center of diversity, approximately 180 species are found across 11 genera, predominantly in South America, especially the Amazon Basin where up to 60 species can occur in a single grid square.[11][1] This region spans from Mexico southward to Paraguay, though abundance and species richness peak in the Amazon.[2] 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 Costa Rica to Brazil.[11][1][16] A notable specific range within this center is Bertholletia excelsa, the Brazil nut tree, which is endemic to the lowland rainforests of the Amazon Basin.[11] In contrast, the Paleotropics host around 80 species, mainly in Southeast Asia (including Malesia, with 71 species across 5 genera), Australia, and the Indo-Pacific region.[11][1] Here, Barringtonia stands out with approximately 69 species distributed along mangrove coasts and tropical shores of the Indo-Pacific.[11] The family's distribution shows clear disjunct patterns between New World and Old World clades, reflecting ancient vicariance events with no overlap between continents.[11] In Africa, diversity is lower at about 40 species, concentrated in West and Central regions, including Madagascar.[11] For instance, the genus Foetidia includes 17 species restricted to Madagascar and nearby islands, such as the Mascarene Islands, with one species extending to East Africa.[11] 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.[11]Habitat preferences
Lecythidaceae species predominantly occupy humid tropical rainforests, seasonal forests, and mangroves across their range, thriving in environments characterized by high humidity and structural complexity. These plants exhibit an altitudinal distribution from sea level to approximately 1,500 m, with many species 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 forest architecture.[11][17][18] 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 flooding and even saline conditions in wetland and mangrove margins. This flood tolerance enables Barringtonia to colonize riverine and coastal edges where water levels fluctuate seasonally.[3][19][20] 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 evergreen or semideciduous habits in humid tropics. While most species are sensitive to prolonged dry periods, savanna-edge taxa like Careya arborea display notable drought tolerance, surviving seasonal water deficits through thick bark and deciduous leaf shedding.[21][21][22] 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 biodiversity in non-inundated upland rainforests.[23][24]Taxonomy and phylogeny
Etymology and classification history
The name Lecythidaceae derives from the type genus Lecythis, which was named by Carl Linnaeus in 1753 from the Ancient Greek word lekythos (λήκυθος), referring to an oil flask or jar, in allusion to the distinctive urn-shaped fruits of species in the genus.[25] 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 Myrtaceae, such as placing Lecythis species under Eugenia due to superficial similarities in fruit structure and tropical distribution.[26] This association persisted into the early 19th century until Swiss botanist Augustin Pyramus de Candolle, in his 1828 Prodromus Systematis Naturalis Regni Vegetabilis, separated the group as the tribe Lecythideae within Myrtaceae, primarily on the basis of unique stamen morphology, including the presence of hooded or connate petals and specialized androecial structures not found in typical myrtaceous taxa.[27] 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.[28] 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.[29] 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 Bertholletia clade, revealing polyphyly in genera like Lecythis and Eschweilera, and proposing taxonomic adjustments that refine the boundaries of New World actinomorphic-flowered genera within the subfamily Lecythidoideae.[30]Phylogenetic position
Lecythidaceae is positioned within the core clade of the order Ericales, where it forms a monophyletic group sister to the holoparasitic family Mitrastemonaceae; this combined clade is in turn sister to the remaining core Ericales, excluding the early-diverging balsaminoid lineage comprising Balsaminaceae, Marcgraviaceae, and Tetrameristaceae.[31] This placement is derived from comprehensive molecular analyses incorporating plastid (e.g., rbcL, ndhF, matK) and nuclear (e.g., 18S rDNA) loci across 22 Ericales families.[32] Earlier studies using ndhF and trnL-F chloroplast sequences had suggested alternative relationships, such as proximity to Sapotaceae and the former Myrsinaceae (now subsumed in Primulaceae), but subsequent supermatrix approaches with broader sampling have refined the topology to the current consensus.[33] 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 New World and Old World lineages while underscoring shared ancestral traits like large, actinomorphic flowers.[33] The origins of Lecythidaceae trace to the Late Cretaceous, with a crown age estimated at approximately 83 million years ago based on fossil-calibrated phylogenies of Ericales.[31] The family's pantropical distribution—spanning the Neotropics, Africa (including Madagascar), and Southeast Asia—is explained by vicariance driven by the fragmentation of Gondwana, which separated ancestral populations and promoted the divergence of the major clades during the early diversification of the order.[32] Distinguishing Lecythidaceae from other Ericales 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 ovary with axile placentation and multiple locules.[26] 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 Sapotaceae or Ericaceae.[26]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.[34][33] Barringtonioideae, with 6 genera primarily distributed in the Old World tropics, is characterized by actinomorphic flowers and capsular fruits, with some species adapted to coastal or mangrove environments; notable examples include Barringtonia (73 species), known for its buoyant seeds aiding dispersal in marine settings.[35] Foetidioideae consists of a single genus, Foetidia (18 species), endemic to Madagascar and nearby islands, featuring dioecious trees with unisexual flowers and a distinctive fetid odor from glandular structures.[34] Lecythidoideae, comprising approximately 20 genera and the majority of the family's species in the New World, is distinguished by large, urn-shaped flowers with hooded petals and operculate fruits; it includes the diverse Eschweilera (71 accepted species), a hyperdominant genus in Amazonian forests.[36][33] The 2024 study by Vargas et al. resolved polyphyly in the Bertholletia clade using plastome and nuclear data, erecting new genera such as Guaiania, Imbiriba, Pachylecythis, Scottmoria, and Waimiria, and refining species boundaries in Eschweilera and Lecythis.[30] Scytopetaloideae contains 6 genera restricted to Africa and Amazonia, with zygomorphic flowers and winged fruits adapted to wind dispersal; Scytopetalum (3 species) is representative.[35] Napoleonaeoideae includes 2 genera native to West and Central Africa, marked by plicate (folded) petals in the buds and explosive seed dispersal mechanisms; Napoleonaea (about 10 species) exemplifies this with its intricate floral architecture.[34] 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.[34][37] 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.[36][38][39]| Subfamily | Number of Genera | Accepted Genera |
|---|---|---|
| Barringtonioideae | 6 | Abdulmajidia, Barringtonia, Careya, Chydenanthus, Planchonia, Petersianthus |
| Foetidioideae | 1 | Foetidia |
| Lecythidoideae | ~20 | Allantoma, Bertholletia, Cariniana, Couratari, Couroupita, Corythophora, Eschweilera, Grias, Guaiania, Gustavia, Imbiriba, Lecythis, Pachylecythis, Scottmoria, Waimiria (and others; recent additions from 2024) |
| Scytopetaloideae | 6 | Asteranthos, Brazzeia, Crateranthus, Oubanguia, Rhaptopetalum, Scytopetalum |
| Napoleonaeoideae | 2 | Crateranthus, Napoleonaea |