Lecythidaceae A.Rich.

Timothy Utteridge & Gemma Bramley (2020). The Kew Tropical Plant Families Identification Handbook, Second Edition. Kew Publishing Royal Botanic Gardens, Kew

Note

Trees, bark fibrous, smelling of linseed. Leaves alternate, simple, distichous, margin serrate (sometimes inconspicuous). Flowers large, with many stamens often fused into a disk, ovary inferior. Fruit large, capsular, many-seeded.

Recognition

Characters of similar families: Ochnaceae: more obvious subparallel veins, ovary superior, fruit drupaceous. Clusiaceae: latex present, veins parallel, ovary superior. Icacinaceae: petiole twisted, flowers small, stamens fewer, ovary superior. Salicaceae: pellucid dots sometimes present, tertiary venation parallel, leaf margins often serrate, ovary superior.

Morphology General Habit

Trees, treelets or rarely shrubs, bark fibrous and smelling of linseed

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves simple, alternate, distichous, margin serrate (sometimes difficult to observe), often chartaceous to leathery

Morphology Reproductive morphology Inflorescences

Inflorescences racemes or panicles, sometimes solitary

Morphology Reproductive morphology Flowers

Flowers actinomorphic or zygomorphic due to morphology of stamens; calyx (2–)4–6(–12)-merous, fused to the hypanthium; corolla 4–8(–18)-merous, petals free, aestivation imbricate; stamens generally numerous, rarely 10, fused in a ring and sometimes forming a cap over the stigma; ovary inferior, 2–6-locular, placentation axile, pendulous or erect, single style and stigma

Morphology Reproductive morphology Fruits

Fruit capsular with an upper lid (pyxidium), sometimes indehiscent (Bertholletia, Couroupita and Barringtonia), usually large and woody, rarely a berry

Morphology Reproductive morphology Seeds

Seed medium to large, sometimes winged (Couratari and Cariniana), endosperm copious, with proteins and oils, embryo straight to coiled, cotyledons variable.

Distribution

Pantropical family with 25 genera and c. 300 species, with a centre of diversity in South American rain forests. Main genera: Eschweilera, Couratari, Lecythis, Cariniana, Couroupita (cannon-ball tree) and Bertholletia (Brazil-nut) in the Neotropics, Barringtonia in the Old World, and a single genus in Madagascar (Foetidia).

Description Author

Daniela Zappi

Timothy M. A. Utteridge and Laura V. S. Jennings (2022). Trees of New Guinea. Kew Publishing. Royal Botanic Gardens, Kew

Distribution

Pantropical family with 25 genera and c. 300 species, with a centre of diversity in South American rain forests including the Brazil nut (Bertholletia Bonpl.). In Malesia, there are five genera with 71 species, with Barringtonia J.R.Forst & G.Forst., Chydenanthus Miers and Planchonia Blume in New Guinea.

Morphology General Habit

Trees, treelets or rarely shrubs, bark fibrous and smelling of linseed

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves simple, alternate, distichous, margin entire, serrulate to serrate (sometimes obscure), often chartaceous to leathery

Morphology Reproductive morphology Inflorescences

Inflorescences terminal, axillary or ramiflorous, racemose or paniculate, sometimes flowers solitary

Morphology Reproductive morphology Flowers

Flowers bisexual, zygomorphic due to stamen arrangement or actinomorphic (in New Guinea); calyx (2–)4–6(–12)-merous, fused to the hypanthium; corolla 4–8(–18)-merous, petals free, imbricate in bud; stamens usually numerous, rarely 10, fused in a ring and sometimes forming a cap over the stigma; ovary inferior, 2–6-locular, placentation axile, style terminal, unbranched and often longer than stamens (in New Guinea), stigma capitate

Morphology Reproductive morphology Fruits

Fruit capsular with an upper lid (pyxidium), sometimes indehiscent (in New Guinea), usually large and woody and crowned by the persistent calyx lobes (in New Guinea)

Morphology Reproductive morphology Seeds

Seed 1 to several (15 or more in Planchonia), endosperm copious, embryo straight to coiled, cotyledons variable, sometimes absent.

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Trees or shrubs, sometimes poorly branched, very rarely woody lianas, exudate absent, all parts smelling of linseed oil, bark fibrous

Morphology Leaves Stipules

Stipules absent or inconspicuous

Morphology Leaves

Leaves alternate, sometimes condensed at the apex of the branch and then looking verticillate, petiolate, often glabrous, venation pinnate, secondary and tertiary venation often parallel, blades entire, margin serrate to crenate, often appearing entire

Morphology Reproductive morphology Inflorescences

Inflorescences generally terminal, sometimes cauliflorous or ramiflorous, racemose or paniculate, rarely reduced to a fascicle, flowers very rarely solitary

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, epigynous, calyx (2-)4-6(-12)- merous, lobes free or fused, corolla (4-)5-8(-18)- merous (corolla substituted by staminodes in Asteranthos Desf.), lobes fused at base or free, white, cream, orange, dull-red, pink, lilac or pale yellow; stamens (10-)100+, generally adnate at base forming a ring and partially fused to the corolla, or numerous staminodes fused into a cap-like structure that covers the style and stigma, anthers dehiscing longitudinally or by pores; ovary inferior, (2-)4-6-locular and with axile, pendulous or erect placentation, many-ovuled, style present, generally stout

Morphology Reproductive morphology Fruits

Fruit a woody circumcissile capsule (pyxidium) or indehiscent capsule, sometimes with fleshy, smelly pulp, many-seeded; seeds generally large, often winged, sometimes with woody testa (Bertholletia Bonpl. ), embryo straight, cotyledons sometimes absent.

Note

Notes on delimitation: Together with the Styracaceae, Sapotaceae and Myrsinaceae, the Lecythidaceae is currently placed within order Ericales (APGII, 2003). Bertholletiaexcelsa is a source of income for local populations in Amazonian Brazil and Bolivia (Brazil-nut, castanha-do-Pará). Couroupitaguianensis Aubl., the cannon-ball tree, is often used as a street-tree in Rio de Janeiro, despite the weight of its fruits that threaten parked vehicles. Many species are good timber sources. Number of genera: Allantoma Miers - Amazonian Venezuela and Brazil, one species, A. lineata Miers Asteranthos Desf. - Eastern Colombia, Southwestern Venezuela and Northwestern Brazil (upper Rio Negro basin), one species, A. brasiliensis Desf. Bertholletia  Bonpl. - Colombia, Peru, Bolivia, Brazil to the Guianas, one species, B. excelsa Humb. & Bonpl. (Brazil nut). Cariniana Casar. - Colombia and Venezuela, Peru, Bolivia and Brazil, around 9 species. Corythophora R.Knuth - Brazil and the Guianas, 4 species. Couratari Aubl. - Costa Rica to French Guiana, Northern Bolivia and Brazil, 19 species. Couroupita Aubl. - Nicaragua to the Guianas, Peru and Brazil, 3 species. Eschweilera Mart. ex DC. - Trinidad-Tobago, Honduras to French Guiana, Bolivia and Brazil, the largest genus in the family, with over 90 described species. Grias L. - Jamaica, Belize to Peru, around 6 species. Gustavia L. - Costa Rica to French Guiana, Northern Bolivia, Ecuador and Brazil, over 40 species. Lecythis S.A.Mori - Costa Rica to French Guiana, Peru and Brazil, 26 species.

Distribution

Represented in the Neotropics by 11 genera with around 180 species. Lecythidaceae has its main distribution centered within tropical, humid forests, hardly reaching Argentina and Paraguay and being absent from Chile. In the Caribbean only Grias L. has been recorded in nature.

Diagnostic

Leaves crenate, often microscopically so. Flowers with numerous stamens basally fused into a ring or staminodes forming a  cap-like structure covering the stigma. Fruit often a woody circumcissile capsule called a pyxidium. Distinguishing characters (always present): Woody plants with fibrousbark and linseed smell. Flowers with numerous stamens. Ovary inferior. Fruits large, woody, 2-many-seeded. 1. Actinomorphic-flowered genera, woody or fibrous berries, elongated woody capsule or winged capsules ... 21. Zygomorphic-flowered genera, rounded or elongated fruits, dehiscent or not ... 5 2. With dehiscent fruits ... 32. With indehiscent fuits ... 4 3. Lacking petals, capsule with wing formed by expanded calyx... Asteranthos 3. Woody elongated pixidium with winged seeds, petals 6 ... Cariniana 3. Woody elongated pixidium, seeds not winged, petals 5 ... Allantoma 4. Fruit rounded woody berry, flowers large with a staminal ring, petals 6-8(-18) ... Gustavia 4. Fruit oblong fibrous or woody berry, petals 4 ... Grias 5. With dehiscent fruits ... 65. With indehiscent fruits ... 7 6. Fruit elongated pixidium, seeds winged, staminal cap fully coiled ... Couratari 6. Fruit rounded to disk-shaped pixidium with few seeds; staminal cap fully coiled ... Eschweilera 6. Large rounded pixidium; staminal cap not coiled ... Lecythis 7. Inflorescences terminal, flowers pale yellow; rounded fruit with hard, ridged large seeds inside ... Bertholletia 7. Inflorescences cauliflorous, to 1 m long, flowers very showy; large indehiscent fruit with small seeds immersed in smelly pulp ... Couroupita Key differences from similar families: May be confused with Myrtaceae because it has inferior ovary and flowers with many stamens, differing in alternate leaves and lack of pellucid dots or glands (vs. Myrtaceae with generally opposite leaves and pellucid dots) Shares some characters with Chrysobalanaceae (hypanthium and flowers with many stamens) but differs in having fruit with many seeds and fibrousbark (vs. one-seeded drupes and bark not fibrous in Chrysobalanaceae).

Lecythidaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:1. 1954

Morphology General Habit

Trees or shrubs; leaves simple, alternate, not gland-dotted but sometimes with large glands on the margin; stipules absent

Morphology Reproductive morphology Flowers

Flowers usually rather large and showy, actinomorphic or zygomorphic, hermaphrodite

Morphology Reproductive morphology Flowers Calyx

Calyx 4–6-lobed, lobes valvate or imbricate

Morphology Reproductive morphology Flowers Corolla

Petals 4–6, free or united into a campanulate tube and then with many ribs

Morphology Reproductive morphology Flowers Androecium

Stamens numerous, in several series, sometimes the outer ones modified into staminodes and resembling a corona; filaments mostly united; anthers basifixed or rarely adnate, opening at the side by slits; staminal disk sometimes lobed

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior or semi-inferior, 2- or more-celled; style mostly simple; ovules 1 to many on axile placentas, sometimes pendulous from near the apex of the cells

Morphology Reproductive morphology Fruits

Fruit woody, fibrous or fleshy, indehiscent or operculate at the apex

Morphology Reproductive morphology Seeds

Seeds without endosperm; embryo divided or entire

Scytopetalaceae, B. Verdcourt. Flora of Tropical East Africa. 1968

Morphology General Habit

Trees or shrubs, sometimes cauliflorous

Morphology Leaves

Leaves simple, exstipulate, alternate, sessile or petiolate, entire or very rarely denticulate

Morphology Reproductive morphology Inflorescences

Inflorescences axillary, terminal or often borne on the old stems, paniculate, racemose or in fascicles; bracts small, soon falling

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, pedicellate, the pedicel sometimes jointed at the apex

Morphology Reproductive morphology Flowers Calyx

Calyx cupuliform or shallowly tray-shaped, entire, lobed or slightly toothed, persistent at the base of the fruit

Morphology Reproductive morphology Flowers Corolla

Petals 3–16, valvate or sometimes the bud bursting somewhat irregularly; lobes free or joined at the base

Morphology Reproductive morphology Flowers Androecium

Stamens numerous, in 3–6 ranks inserted on the annular disk, essentially epipetalous, often falling with the corolla; filaments free or joined at the base; anthers basifixed

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior or inferior, 3–8-locular, each locule with 2-several ovules; style 1, filiform; stigma small, entire or slightly lobed

Morphology Reproductive morphology Fruits

Fruit a woody or crustaceous capsule, or indehiscent but scarcely fleshy, unilocular

Morphology Reproductive morphology Seeds

Seeds 1–8, with abundant endosperm which is often ruminate

Scytopetalaceae, Hutchinson and Dalziel. Flora of West Tropical Africa 1:2. 1958

Morphology General Habit

Trees; branchlets sometimes winged

Morphology Leaves

Leaves alternate, simple, exstipulate

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic, in terminal panicles or axillary racemes or fasciculate on the old wood

Morphology Reproductive morphology Flowers Calyx

Calyx cupular, entire or toothed

Morphology Reproductive morphology Flowers Corolla

Petals 3–10, valvate, free or shortly connate

Morphology Reproductive morphology Flowers Androecium

Stamens numerous, in several series on the margin of or on the disk, free or united towards the base; anthers 2-celled, opening by a pore or slit at the side or towards the top

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 3–6-celled; ovules 2 to several in each cell, axile

Morphology Reproductive morphology Fruits

Fruit woody

Morphology Reproductive morphology Seeds

Seeds with ruminate or uniform copious endosperm and linear embryo

Barringtoniaceae, A. Fernandes. Flora Zambesiaca 4. 1978

Morphology General Habit

Evergreen trees or shrubs

Morphology Leaves

Leaves alternate, exstipulate, usually large, simple, not gland-dotted

Morphology Reproductive morphology Flowers

Flowers showy, actinomorphic or zygomorphic, bisexual, solitary or in fascicles on the old wood or in terminal racemes

Morphology Reproductive morphology Flowers Receptacle

Receptacle ± crateriform

Morphology Reproductive morphology Flowers Calyx

Calyx (2–3)4–5-lobed, the lobes sometimes calyptriform, covering the petals and stamens in bud

Morphology Reproductive morphology Flowers Corolla

Petals 4 or 0 (Napoleonaed), free or somewhat adnate to the staminal ring

Morphology Reproductive morphology Flowers Androecium

Stamens ¥, in several verticils, all fertile or only some fertile and accompanied by numerous antherless staminodes, the outer series of staminodes united and forming a radiate or cup-shaped pseudocorolla; filaments usually united into a ring of varying depth; anthers basifixed, opening by lateral slits; staminal disk sometimes lobed

Morphology Reproductive morphology Flowers Gynoecium

Ovary inferior or semi-inferior, 2(3)–5-locular; style very long and filiform or short and expanded into a broad peltate disk; ovules l-¥ per loculus, anatropous

Morphology Reproductive morphology Fruits

Fruit woody, fibrous or fleshy, indehiscent or operculate at the top, crowned by the persistent calyx-segments

Morphology Reproductive morphology Seeds

Seeds without endosperm; embryo entire or divided