Systematic Treatment

LECYTHIDACEAE Poiteau, Mem. Mus. Hist. Nat. 13: 141-165. 1825 (sub Lecythidees); Myrtaceae tribus Lecythideae Rich, ex DC., Prodromus 3: 288-291. 1828; Myrtiflorae subord Lecythideae Endlicher, Gen. PI. 1233-1234. 1840; Myrtaceae subtribus Eulecythideae Benth. & Hook., Gen. PI. 1(2): 720-725. 1865; Lecythidaceae subfam Lecythidoideae Niedenzu in Engler & Prantl, Nat. Pflanzenfam. 3(7): 26-41. 1892.

Small to large trees, leaves alternate, simple, estipĂșlate or with minute, caducous stipules, without pellucid punctations. Flowers actinomorphic or zygomorphic; petals usually present, less frequently absent; stamens numerous, connate at the base into a staminal ring, the ring actinomorphic or prolonged on one side into a strap-like structure which arches over the summit of the ovary, some apetalous species with the outermost stamens modified into a petaloid corona; intrastaminal disc present or absent; ovary inferior to half-inferior. Fruits fibrous berries, dry drupes, or woody circumscissile capsules with one to many seeds. Seeds with or without cotyledons.

Type genus. Lecythis Loefl.

Key to Subfamilies of Lecythidaceae

1. Pollen syntricolpatc. Cortical bundles with reversed orientation (i e, xylem outside, phloem inside). Secondary xylem without crystal chains. Principally Asian, with 1 sp in East Africa and 2 spp. in Madagascar. subfam I. Planchonioideae.

1. Pollen tricolpate or tricolporate. Cortical bundles with reversed or normal orientation. Secondary xylem with or without crystal chains. New or Old World.

2. Flowers apetalous, without a corona of staminal origin; filaments weakly fused at the base. Ovules horizontal, situated in a vertical ring on a thick shield-like placenta; funicle and micropyle directed outwards. Cortical bundles with reverse orientation. Secondary xylem with crystal chains. Madagascar, India, Malaysia. subfam II. Foetidioideae.

2. Flowers with or without petals, if without then possessing a corona of staminal origin; filaments fused at base to form distinct staminal ring. Ovules in rows, hanging or ascending from a central columnar placenta, if horizontal then with the micropyle facing inwards. Cortical bundles with normal orientation. Secondary xylem with or without crystal chains. West Africa or the Neotropics.

3. Flowers without petals, actinomorphic; outer row of filaments fused to form a petaloid corona. Fruits indehiscent. Secondary xylem without crystal chains. West Africa (except for Asteranthos brasiliensis). subfam III. Napoleonaeoideae.

3. Flowers with petals, actinomorphic or zygomorphic; outer row of filaments not forming a petaloid corona. Fruits dehiscent in all but Couroupita, Grias, and Gustavia. Secondary xylem with crystal chains. Neotropics. subfam IV. Lecythidoideae.

I. Lecythidaceae subfam Planchonioideae Niedenzu in Engler & Prantl, Nat.

Myrtaceae tribus Barringtoniae DC., Prodromus 3: 288-296. 1828. pro parte; Barrington-iaceae Lindley, Vegetable Kingdom 716-730. 1846. pro parte; Myrtaceae subtribus Bar-ringtonieae Benth. & Hook., Gen. PI. 1(2): 720 725. 1865, pro parte.

Flowers actinomorphic; petals present, corona of staminal origin absent, inner or outer whorl of stamens sometimes staminodial, the base of the stamens fused into a distinct staminal ring except in Petersianthus where they are weakly fused; intrastaminal disk present, pollen syntricolpate; ovary 2-4 locular, placenta columnar. Fruit indehiscent; fibrous berries (Planchonia) or dry drupes which contain from one (all species of Barringtonia) to many (some species of Planchonia) seeds; cotyledons present in Planchonia and Petersianthus, absent in Barringtonia, Careya and Chydenanthus. Secondary xylem without crystal chains in parenchyma, cortical bundles reversely oriented with the xylem on the outside, x = 13.

A subfamily of 55 species in 6 genera (Table III) distributed throughout tropical Asia, Malaysia, northern Australia, and the Pacific Islands. Tropical Asia and the islands to the south and east have the greatest number of species of this subfamily.

Type genus. Planchonia Blume

II. Lecythidaceae subfam Foetidioideae Niedenzu in Engler & Prantl, Nat.

Pflanzenfam. 3(7): 29-30. 1892; Barringtoniaceae tribus Foetidieae Knuth,

Pflanzenreich IV. 219a: 62-65. 1939; Foetidiaceae Airy Shaw in Willis, J.

C. A Dictionary of Fl. PI. and Ferns ed. 8, 465.1973.

Flowers actinomorphic; petals absent, corona of staminal origin absent, staminodes absent, the base of the stamens only weakly fused and not forming a staminal ring; intrastaminal disk present; pollen tricolpate; ovary 4 locular, placenta peltate. Fruit indehiscent; cotyledon type unknown. Secondary xylem with crystal chains in the parenchyma, cortical bundles reversely oriented. Chromosome number unknown.

The 5 species of the single genus, Foetidia, of this subfamily are distributed in Madagascar, India, and Malaysia. Type genus. Foetidia Commerson ex Lamarck

III. Lecythidaceae subfam Napoleonaeoideae Niedenzu in Engler & Prantl, Nat. Pflanzenfam. 3(7): 33-34. 1892; Napoleonaeaceae Beauv., Fl. Oware Benin Afr. 2: 29. 1810 (sub Napoleonees); Myrtaceae subtribus Napol-eoneae Benth. & Hook., Gen. PI. 1(2): 723-727. 1865; Barringtoniaceae tribus Napoleoneae R. Knuth, Pflanzenreich IV. 219a: 67-73.1939.

Belvisiaceae R, Brown, Trans. Linn. Soc. London 13:222-223. 1821. Asteranthaceae R. Knuth, Notizbl. Bot. Gart. Berlin-Dhalem 11(110): 1034-1036. 1934. Barringtoniaceae tribus Craterantheae R. Knuth, Pflanzenreich IV. 219a: 65-67. 1939.

Flowers actinomorphic; petals absent, corona of staminal origin present, staminodes present in Napoleonaea absent in Crateranthus and Asteranthos, the base of the stamens fused into a distinct staminal ring except in Asteranthos where they are weakly fused; intrastaminal disk present in Napoleonaea absent in Crateranthus and Asteranthos; pollen tricolpate; ovary 3 or 5 locular, placenta columnar. Fruit indehiscent, berry in Crateranthus, drupe in Napoleonaea, capsule ? in Asteranthos; cotyledons present. Secondary xylem without crystal chains in parenchyma, cortical bundles normally oriented with phloem on the outside. x= 16 in Napoleonaea.

A subfamily of 12 species in 3 genera of West Africa and the upper Rio Negro region of Amazonia. Crateranthus and Napoleonaea are distributed in western Africa, and Asteranthos is confined to the periodically flooded 'caatinga' forest of the upper Rio Negro in Colombia, Venezuela and Brazil. Type genus. Napoleonaea Beauv.

Key to the Genera of Napoleonaeoideae

1. Calyx distinctly 3 or 5 lobed; pseudocorollas 1 or 3, if 1 then campanulate; seeds several to many per fruit, exalbuminous.

2. Calyx lobes 5; pseudocorolla 3-rowed, rotate, staminodes present; ovary inferior, 5

locular. Napoleonaea.

2. Calyx lobes 3; pseudocorolla 1-rowed, campanulate; staminodes absent; ovary semi-inferior, 3 locular. Crateranthus.

1. Calyx dentate but not distinctly lobed; pseudocorolla 1, radiate; seed one per fruit, albuminous. Asteranthos.

IV. Lecythidaceae subfam Lecythidoideae Niedenzu in Engler & Prantl, Nat. Pflanzenfam. 3(7): 34-41.1892.

Myrtaceae tribus Barringtonieae DC., Prodromus 3: 288-296. 1828, pro parte, typus exclusus; Barringtoniaceae Lindley, Vegetable Kingdom 716-730. 1846. pro parte, typus exclusus; Myrtaceae subtribus Barringtonieae Benth. & Hook., Gen. PI. 1(2); 720-725. 1865 .pro parte, typus exclusus. Lecythidaceae subfam Lecythidoideae tribus Griadeae Pichon, Notul. Syst. (Paris) 12: 195. 1945.

Small, unbranched to huge, much-branched emergent trees. Leaves alternate, exstipulate or with minute caducous stipules, simple, pinnately nerved, the margins usually entire, sometimes crenulate to serrate, very large and clustered at the branch ends in Grias and some species of Gustavia, medium-sized and not clustered in the remaining species. Inflorescences simple racemes, panicles with 2 or 3 orders of racemose or spicate branches or fascicles, suprafoliar (then either terminal or subterminal), axillary, or cauline. Flowers actinomorphic or zygomorphic, inferior or half-inferior, perfect; calyx entire or of 2 to 6 triangular to broadly ovate lobes; petals 4, 6, or 8, infrequently 12 or 18; stamens arising from a connate staminal ring in Gustavia, Grias, and Allantoma, the staminal ring slightly expanded on one side in Cariniana, and markedly expanded into a strap-like ligule with an enlarged hood at the apex in the remaining genera, the hood appendages with or without anthers; ovary usually 2, 4, or 6-locular, with 2-115 anatropous ovules per locule, the axile placentae at the apex, base, or throughout the length of the locule. Fruits indehiscent, then somewhat fleshy and berry-like (Grias, Gustavia) or with a thin, ligneous exocarp (Couroupita), or dehiscent via a cir-cumscissile operculum, then often very large and woody; seeds winged in Cariniana and Couratari, without wings in the remaining genera, with or without arils; embryos undifferentiated, or with fleshy, plano-convex or foliaceous cotyledons. Secondary xylem with prominent crystal chains in the parenchyma, cortical bundles normally oriented, x= 17.

A subfamily of about 212 species in 10 genera endemic to the neotropics.

Type genus. Lecythis Loefl.

Key to the Genera of New World Lecythidaceae (including Asteranthos of the Napoleonaeoideae)

1. Androecium actinomorphic.

2. Flowers solitary in leaf axils, without petals; outer row of stamens fused to form a radiate, petaloid corona. Embryo J-shaped, embedded in ruminate endosperm.

1. Asteranthos.

2. Flowers in multi-flowered inflorescences, infrequently with a single flower, with 4-8(-18) separate petals without a petaloid corona. Embryo not J-shaped, without endosperm. 3. Flower buds globose, the flowers large, 2.5-20 cm in diameter at anthesis; ovules on the upper 1/2 of the septum. Fruits indehiscent, the seeds released via deliquescence of the pericarp or by fragmentation of the opercular region.

4. Petals 6-8(-18); stamens 500-1200, the anthers linear, 2-5 mm long, dehiscing by apical pores; placcntae expanded, the ovules 7-93 per locule, horizontal or slightly descending. Fruits usually with 2 or more seeds; embryo with plano-convex, fleshy cotyledons. 2. Gustavia.

4. Petals 4; stamens 85-210, the anthers globose, less than 1 mm long, dehiscing by longitudinal slits; placentae not expanded, the ovules 2-4, pendulous. Fruits with a single seed; embryo undifferentiated. 3. Grias.

3. Flower buds globose or oblong, the flowers smaller, less than 2,5 cm in diameter at anthesis; ovules on the lower 1/2 of the septum. Fruits dehiscent, the seeds freely falling.

5. Calyx rim-like or with 5 inconspicuous broadly triangular lobes at anthesis; petals

5. Seeds not winged, the embryo undifferentiated. 4. Allantoma.

5. Calyx with 6 triangular lobes at anthesis; petals 6. Seeds uni-laterally winged, the embryo with 2 foliaceous cotyledons. 5. Cariniana.

1. Androecium zygomorphic.

6. All of the hood appendages bearing anthers. 7. Ovary 6-locular; ovules 30-115 per locule, on bilamellar placentae throughout the length of the locule. Fruit globose, indehiscent. Embryo with 2 foliaccous, highly convoluted cotyledons. 6. Couroupita.

7. Ovary 2-3(-5)-locular; ovules 5-8 per locule, placentae not bilamellar, attached towards the base of the locule. Fruit campanulate or cylindric, dehiscent. Embryo undifferentiated or with 2 foliaceous cotyledons, these not highly convoluted. 8. Androecium elongated on one side into a strap-like structure which bends over the summit of the ovary; ovary 2(-5)-locular. Seeds without wings, the embryo undifferentiated. 7. Corythophora.

8. Androecium elongated on one side but not forming a conspicuous strap-like structure which bends over the summit of the ovary; ovary 3-locular. Seeds unilaterally winged, the embryo with foliaceous cotyledons. 5. Cariniana.

. Usually with all of the hood appendages sterile or, iess frequently, with less than half of the hood appendages bearing anthers.

9. Buds enclosed by calyx except for a horizontal slit at the apex; calyx with 2 lobes at anthesis, the style greater than 10 mm long. Fruit appearing indehiscent but with a small inwardly falling operculum, the seeds being retained within the fruit until the pericarp disintegrates. Seeds with a thick, boney integument. 8. Bertholletia.

9. Buds not enclosed by calyx; calyx with 6 lobes at anthesis, the style less than 10 mm long. Fruit dehiscent, with a relatively large, outwardly falling operculum, the seeds being released at maturity. Seeds with a thinner, non-boney integument.

10. Androecium hood coiled inwards, with an outwardly extended flap at the apex of the coil. Fruits cylindric or campanulate. Seeds with a wing around the circumference, the embryo with 2 foliaceous cotyledons. 9. Couratari.

10. Androecium hood flat, or if coiled inwards without an outwardly extended flap at the apex of the coil. Fruits usually globose. Seeds without wings, the embryo undifferentiated.

11. Androecium coiled inwards, with blunt-tipped appendages at the apex of the coil, these differentiated from the more abundant, echinate hood appendages; ovary usually 2-locular. 10. Eschweilera.

11. Androecium flat or expanded at the apex but not coiled inwards, all hood appendages more or less equal; ovary usually 4-locuIar. 12. Ovary 2-locular, the style not differentiated from the summit of the ovary, the summit umbonate. 7. Corythophora.

12. Ovary 4-locular, the style differentiated from the summit of the ovary, the summit truncate. 11. Lecythis.

ASTERANTHOS by g. T. Prance

1. Asteranthos Desfontaines, Mem. Mus. Hist. Nat. 6: 9,13. 1820; R. Brown, Trans. Linn. Soc. London 13: 222. 1821; de Candolle, Prodr. 7: 55. 1828; Endlicher, Gen. PI. 745. 1839; Meisner, PI. Vase. Gen. 125. 1839; Lindley, Veg. Kingdom 716-730. 1846; Bentham, J. Linn. Soc., Bot. 3: 80-81. 1859; Benth. & Hook., Gen. PI. 1: 724. 1865; Miers, Trans. Linn. Soc. London II. 1: 17-19. 1875; Knuth, Notizbl. Bot. Gart. Berlin 11: 1034-1036. 1934, Pflanzenreich IV. 219b: 1-3. 1939.

Asteranthus Spreng., Linn. Syst. Veg. ed. 16. 2: 568. 1825; Reichenbach, Consp. Veg. 137. 1828; Eichler, Mart. Fl. Bras. 12(1): 496-500. 1889; Niedenzu in Engl. & Prantl, Nat. Pflanzenfam. 3(7): 34. 1892.

Medium sized trees. Leaves alternate, entire, appearing glabrous, but with minute clustered hairs on lower surface visible only with high power magnification; with minute caducous stipules visible only on the youngest leaves. Flowers hermaphrodite, actinomorphic, 3.5-4.5 cm diameter, solitary in leaf axils. Receptacle conical-campanulate with a radiate circular accrescent calyx with crenate margin. Corolla absent. The outer whorl of stamens developed into a corona or corolla-like structure with 24-28 conspicuous veins from center to margin, pleated and unfolding like a parasol, the margins irregularly dentate and ciliate; the fertile stamens numerous, free, inserted in several rows attached in a circle around the base of the sterile outer whorl; anthers 2 locular, longitudinally dehiscent, basally attached. Style erect and thin, the stigma 5-8 lobed. Ovary semi-inferior, 5-8 locular with 4 ovules in each loculus. Fruit an oblong-pyramidal capsule; semi-inferior ±2 cm long with enlarged persistent calyx persisting around middle, the fruiting calyx coriaceous, to 3.5 cm diameter, the lower portion of fruit smooth unribbed, 0.5 cm long, upper part above calyx 1.5 cm long deeply 6 ribbed and tapered to a pointed apex, unilocular with a single cone-shaped seed within, with abundant ruminate endosperm; embryo curved on lower portion to form a J-shape, with 2 membranous cotyledons at apex.

Type species. Asteranthos brasiliensis Desf. The name Asteranthos is derived from the Greek compound word aSnjg = aster ( = star) and avdos-andos (= flower) referring to the star-like flower.

Distribution. 1 species confined to the upper Rio Negro region of Colombia, Venezuela and Brazil.

Niedenzu (1892) and Hutchinson (1969) pointed out that the fused outer row of staminodes was mistaken for a corolla by many previous authors. We agree with Hutchinson that Asteranthos is apetalous but has a corolla- or corona-like row of staminodes. This forms a complete fused circle and the inner rows of stamens are fused to it at their bases. This interpretation is important in relating Asteranthos to the Lecythidaceae a family in which many remarkable structures have developed from the staminal tissue.

Taxonomic History

Asteranthos was described by Desfontaines (1820) who related it to the genus Napoleonaea. The genus was based on a single fragmentary collection without collectors information which was sent to Paris from Portugal but was of Brazilian origin. This was one of the A. R. Ferreira Brazilian collections. From the scrappy material Desfontaines reconstructed the plant and described A. brasiliensis and provided a surprisingly accurate illustration. Desfontaines placed Asteranthos in the Jussieuan family Symploceae. In fact Asteranthos had been mentioned in the literature by Palisot de Beauvois (1810) ten years prior to its formal description. Palisot de Beauvois when he described the new African genus Napoleonaea referred to an undescribed and related genus from Brazil which is in fact Asteranthos,

Robert Brown (1821) described the new family, Belviseae, to accommodate the two genera Belvisia and Asteranthos. Belvisia is a later homonym of Napoleonaea and is consequently illegitimate. Brown was thus linking Napoleonaea and Asteranthos together. He stated that he was uncertain of the exact relationship of his family Belviseae which was described in a footnote of a work discussing the Rafflesiaceae.

Sprengel (1825) placed Asteranthos in the Linnaean group Polyandria monogynia. Reichenbach (1828) treated it together with 'Napoleona'4 in his group Asterantheae which was placed between Olacinae and Aquifoliaceae.

Lindley moved the Belvisiaceae around in his different works as his thoughts developed. In the first edition of his Natural System (1830), he placed Belvisiaceae between Styracaceae and Sapotaceae in the second edition of the same work (1836) he moved the Belvisiaceae to his Alliance Campanales and placed it between the Sphenocleaceae and Columelliaceae. In 1846 Lindley was the first worker to suggest that the Belvisiaceae was related to the Myrtaceae and Lecythidaceae. He placed Belvisiaceae in his order Myrtales between Rhizophoraceae and Melastomaceae. The order also contained Myrtaceae, Lecythidaceae, Onagraceae and four other families.

De Candolle (1838) in his Prodromus placed the family 'Napoleoneae' between Columelliaceae and Vaccinieae. The family contained the two genera 'Napoleona' and Asteranthos.

Meisner (1839) considered the Belvisieae as a family which was placed between the Passifloraceae and Loasaceae. Also in 1839, Endlicher, considered Asteranthos as part of his class Petalanthae and in the 'Order' Ebenaceae in the group Symploceae. However, it is placed together with 'Napoleona' in "Genera dubiaeaffinitatis."

Bentham (1859) discussed Asteranthos in light of further material collected by Spruce which ended the doubt that the original material was from Brazil. He suggested affinity with the Myrtaceae (including Barringtoniae and Lecythidaceae) and thus follows Lindley (1846). A few years later Asteranthos was again included in the Myrtaceae by Bentham and Hooker (1865) who placed it in their tribe Lecythidaceae together with Barringtonia, 'Napoleona' and the South American genera of the family. The early confusion about the position of Asteranthos was partially due to the poor type material. Since Bentham drew attention to the Spruce material it has generally been placed near to or within the Lecythidaceae in the more recent systems.

Le Maout & Decaisne (1873) placed Asteranthos and 'Napoleona' in the family 'Napoleoneae' between the families Myrtaceae (including Lecythidaceae) and 'Melastomaceae.' Baillon (1877) treated the 'Napoleoneae' as one of his 'series' of the Myrtaceae.

Miers (1875a) did not agree that Asteranthos is related to Myrtaceae. He studied the Spruce material and suggested a close relationship with Rhododendron, based mainly on the false assumption that Asteranthos had a fully superior ovary. This affinity has not been adopted by later workers. Miers' suggestion that there is more than one species of the genus is certainly unfoun-ed judging by the more rccent material studied here. In the same work Miers placed Napoleonaea in a unigeneric family Belvisiaceae.

Eichler (1889) treated the Napoleonaeaceae for Martius' Flora Brasiliensis as it was not included with the Myrtaceae in Berg's (1858) earlier account of that family which included the Lecythidaceae. Eichler did not agree with a myrtaceous affinity for Napoleonaeaceae and he discussed the various affinities

The correct orthography of the generic name is Napoleonaea (see Liben, 1971). Most later authors have misspelled the name as 'Napoleona.' In this account the orthography used by each author is given, but between quotation marks where is is incorrect.

suggested by previous authors, he rejected them all and rather hesitantly suggested a relationship to Mesembrianthemum or Cactaceae.

Niedenzu (1892) in the first edition of Engler's Pflanzenfamilien treated the genera 'Napoleona' and Asteranthos as the subfamily 'Napoleonoideae' of the Lecythidaceae.

Knuth (1934, 1939c) treated Asteranthos in a separate family Asteran-thaceae. He considered 'Napoleona' as a separate tribe of the family Barringtoniaceae. Hutchinson (1969) followed Knuth and recognized the Asteranthaceae with some hesitation. Other more recent phyllogenetic systems e g Cronquist (1968), Takhtajan (1969) have included Asteranthos within the Lecythidaceae.

The Systematic Position of Asteranthos

The historical review above shows that there has been considerable disagreement about the systematic position of Asteranthos. Table XII summarizes the most important placings of Asteranthos.

The majority of workers starting with Desfontaines (1820), who described the genus, have related it to Napoleonaea, although the exact position and taxonomic status has varied widely from author to author. In the last detailed monograph of the Lecythidaceae Knuth (1939a, b, c) placed Asteranthos in a separate family, and Napoleonaea as a separate tribe in his family Barringtoniaceae.

In the introductory material to the present work it is shown that Asteranthos is undoubtedly most closely related to the African Napoleonaea. The main reason that some authors have been reluctant to relate the two genera is geographical rather than morphological. There are many examples of relationships between African and American genera and species and the geography does not pose any real problems. However, Asteranthos remains as the most isolated and distinct genus within the Lecythidaceae. It is close enough to Napoleonaea to be included in the same subfamily, but is obviously more primitive with only one staminal whorl developed into a fused corolla-like structure and the rest of the stamens free but attached to the outer whorl. In Napoleonaea there are four whorls of staminal origin, the outermost corollalike, the inner 3 coronas are smaller with staminodes on the second and third rows and with the fertile stamens borne on the innermost corona. In Asteranthos the fruit is reduced to a single seeded capsule but as the flower has a multilocular ovary it is easy to relate to the many seeded drupe of Napoleonaea. The endosperm of Asteranthos is its most important difference from other Lecythidaceae, i e no other Lecythidaceae possess endosperm in the mature seeds.

Asteranthos brasiliensis Desfontaines, Mem. Mus. Hist. Nat. 6: 9, t 3. 1820;

de Candolle, Prodr. 7: 5551. 1838; Miers, Trans. Linn. Soc. London II. 1:

17-19. 1875; Eichler; Mart. Fl. Bras. 12(1): 497-500. 1889, sub Asteranthos;

Knuth, Pflanzenreich IV. 219b: 3. 1939. Figs 7, 16A-D, 40, 41.

Tree to 15 m tall, the young branches glabrous. Leaves with laminas coriaceous,-oblong, 4-13 cm long, 2-4.5 cm broad, almost glabrous on both

Table XII

Proposed relationship of Asteranthos

Relationship proposed Anthor

near Napoleonaea family Symploceae

Desfontaines 1820

Fam. Bclviseae with Bch isia = Napoleonaea

R.Brown 1821

Asterantheae with Napoleonae

Reichenbach 1828

Belvisiaceae near Styracaceae & Sapotaceae

Lindley 1830

Belvisiaceae in Alliance Campanales

Lindley 1836

Belvisiaceae in order Myrtales

Lindley 1846

Fam. Napoleoneae between Columelliaceae & Vaccinieae

de Candolle 1838

Fam. Belvisieae between Passifloraccae & Loasaceae

Meisner 1839

Myrtaceae tribe Lecythidaceae

Bentham & Hooker 1865

Fam. Napoleoneae next to Myrtaceae (incl. Lecythidaceae)

Le Maout & Decaisne 1873

Myrtaceae series Napoleoneae

Baillon 1877

near Rhododendron

Miers 1875a

Napoleonaceae with Napoleonaea

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