Dioecious or monoecious (Endocomia) monopodial shrubs or trees with aromatic oil- cells, (2-)5-15(-40) m high (Virola sessilis, Brazil, 30 cm high); stilt-roots sometimes present; branching whorled, the crown often pyramidal. Bark and wood exuding a watery pink or red sap on cutting. Wood white or soon brownish on cutting, soft; rays narrow, tanniniferous tubes present; growth rings usually distinct and visible to the naked eye. Twigs tomentose, glabrescent. Hairs uniseriate, often with armed cells, appearing as stellate or dendroid. Leaves simple, entire, alternate, usually distichous in plagiotropic shoots, exstipulate, often with indumentum on both surfaces when young, glabrescent, or remaining on the lower surface; lateral nerves distinct, interarching at the margins; reticulations sometimes faint; vernation conduplicate (Asian genera); dark dots (corky hair bases) on lower leaf surface present or absent; glands absent. Inflorescences from foliar axils to ramiflorous (cauline), pedunculate, compound or not, with or without bracts at base, the flowers variably arranged at the ends of the branches, or a short, woody (sub)sessile simple or 2-4-fid scar-covered wart- or worm-like brachyblast with the flowers in (sub)umbels at the end; bracts caducous. Female inflorescences similar to male, usually less branched. Flowers unisexual, mostly pedicellate; bracteole present or absent. Perianth gamophyllous, globose to rotate, buds 2- (Horsfieldia, p.p.) to 5-lobed (Endocomia, p.p.), cleft to variable depths, lobes little or much recurved; the female usually urceolate, larger, more swollen than the male; disc rarely present. Androecium with synandrium sessile or stalked (androphore), in Myristica mostly with a short sterile apex; anthers dorsally adnate to the column and laterally to each other, or free, in Knema stellately arranged around a disc, 2-45 in number, ellipsoid to linear, extrorse, opening by longitudinal slits, 2-loculed, tetrasporangiate; immature pollen sacs septate. Ovary monocarpellate, sessile, superior, ovoid; style absent or short, stigma 2-lobed or rarely peltate with few to many laciniations; ovule 1, mostly anatropous, ± basal. Fruits (sub)globose to oblong, rarely transversely elongate, dehiscing by a longitudinal circumferential suture into 2 valves, rarely indehiscent; pericarp leathery, carnose or subligneous, glabrous or tomentose. Aril (orange-)red (or yellow?), completely covering the seed, laciniate to various depths or sometimes (sub)entire, attached to the base of the seed between the hilum and the micropyle. Seeds with testa of three layers; albumen hard, mostly ruminate, containing oil, sometimes starch. Embryo small, near the base of the seed or a little above; cotyledons connate at base (peltate or cup-shaped) or free; radicle basal, cylindric.

^ Footnote 1) With contributions by RW.J.M. van der Ham (palynology), R. Hegnauer (phytochemistry & chemotaxonomy), J. Koster and P. Baas (vegetative anatomy). Most of the original drawings are by J. H. van Os and some by R. van Crevel.

Pantropical with c. 500 species in 20 genera, more or less equally distributed over and restricted to the three main continental areas: 8 genera (with few species) in Africa, of which 5 in Africa proper and 3 in Madagascar, 6 genera in America, and 6 in Southeast Asia, mainly Malesia.

The largest genera are Virola (c. 50 species) in America, and Horsfieldia (c. 100 species), Knema (c. 90 species), and Myristica (c. 170 species) in Asia. The latter three, together with Endocomia and Gymnacranthera, have widespread distributions; Endocomia occurs from S China to New Guinea (Map 1), Gymnacranthera from S Peninsular Thailand to New Guinea (Map 2); Horsfieldia occurs from Sri Lanka and S China east to the Solomon Islands and N Australia, with centres of species richness in W Malesia, mainly Borneo and New Guinea (Map 3); Knema has a distribution from India to W New Guinea, with a centre of diversity in W Malesia, mainly Borneo (Map 4); the genus Myristica has the largest distribution, from India to N Australia and far into the Pacific, with centres of speciation in W Malesia and, particularly, New Guinea (100 species) (Map 5). Paramyristica (1 species) is endemic to Papua New Guinea (Map 6).

In the present treatment the now official name Papua Barat has been used instead of Irian Jaya.

Low or medium (rarely canopy) trees in various types of primary lowland rain forest, including kerangas and marshy forest. Some species of Horsfieldia (in New Guinea) and several of Knema (e.g., Mt Kinabalu area) and Myristica (in New Guinea) occur in montane forest. Occasionally species are 'sciophilous nomads' (fast growing, shade tolerant), notably some Horsfieldias in New Guinea, and few are found in secondary forest. Sometimes Myristicaceae constitute a considerable component of the forest, especially of the middle storey of the lowland rain forest, but they are not gregarious.

According to Koster & Baas (1981) leaf anatomical characters are xeromorphic, which is unexpected in view of the mesic ecology of recent Myristicaceae (see Vegetative Anatomy').

Pollination & flower biology — Flowering and fruiting generally occurs throughout the year. The usually ± carnose, yellow or brown, inside creamy, pink, or red flowers of several genera have repeatedly been reported as being fragrant, e.g. Horsfieldia irya and Myristica fragrans. Anthesis presumably is mainly nocturnal, and small beetles may effect pollination; nectar is not reported for any species. Male plants of Myristica produce over 50 times as many flowers as do females (Armstrong & Drummond 1986; Armstrong & Tucker 1986; Armstrong & Irvine 1989). Myristica subalulata and some related species are myrmecophilous, the ants possibly involved in pollination (De Wilde 1998). Besides the coloured inside of the perianth, in Knema also the staminal disc may be brightly (purple-red) coloured, the contrast with the creamy-white pollen possibly heightening the attraction of pollinators.

Dispersal — The brown-black seeds contrasting with the orange or red aril and the inner surface of the (red, pink, or white) opened pericarp attract birds and suggest bird dispersal (Howe & Vande Kerckhove 1980), in Asia by fruit pigeons and doves, hornbills, and birds of paradise (Sinclair 1958). Monkeys and rodents may disperse fruits or seeds as well. Dissemination by water may occur in Horsfieldia wallichii, of which the seeds float because of air trapped between aril and seed, and in Horsfieldia irya, a widespread riverine species with cavities in the endosperm. The seeds of the marsh nutmeg, Myristica elliptica, are reported to float, also when the aril is missing.

Germination & seedling — Seeds remain viable for a restricted period only, a few weeks, and germinate only in a damp, shady environment and therefore the natural reintegration of Myristicaceae in secondary forest is impossible. Germination is (mostly) hypogeal. The cotyledons remain within the testa, the taproot and hypocotyl emerge, the shoot is erect, initially with reduced leaves (cataphylls), mostly borne spirally: the Horsfieldia type (subtype) of seedling (De Vogel 1979), a common type in tropical woody dicotyledons.

References:
Armstrong, J. E. & B. A. Drummond Biotropica 18 (1986) 32-38
Armstrong, J.E. & A.K. Irvine Amer. J. Bot. 76 (1989) 74-85, 86-94
Armstrong, J. E. & S.C. Tucker Floral development in Myristica (Myristicaceae) Amer. J. Bot. 73 (1986) 1131-1143
De Vogel, E. F. Seedlings of Dicotyledons (1979) 1-203. Pudoc, Wageningen
De Wilde, W. J. J. O. Blumea 43 (1998) 165-182
Howe, H.F. & G.A. Vande Kerckhove Science 210 (1980) 925-926
Sinclair, J. A revision of the Malayan Myristicaceae Gard. Bull. Sing. 16 (1958) 205-472

The family Myristicaceae is homogeneous and clearly belongs (also phytochemically) within the Magnoliales. Phylogenetic analysis of the genera revealed that the family is monophyletic, of African origin (Sauquet 1998), and seemingly most related to the Annonaceae, mainly through the ruminate endosperm. Canellaceae have been suggested to be allied through the monadelphous androecium (Wilson & Maculans 1967), but according to a recent cladistic analysis of all families (APG 1998) this family is as yet not properly placed as it falls beyond the recognized basal orders. The foliage is generally strongly reminiscent of Annonaceae. The 3-lobed perianth reminds of Lauraceae and Annonaceae. Myristicaceae are distinguished, however, by their unisexual flowers with uniseriate perianth, and monocarpellate 1-ovuled female flowers.

Within the family the relationship of the genera is unclear, and initially one single genus, Myristica (divided into sections), was recognized until Warburg (1897) divided it into several genera. The genera of Madagascar possibly retain the most primitive characters, viz. pollen morphological, not or less consolidated stamens, and a little developed aril.

Within the larger Malesian genera, Horsfieldia, Knema, and Myristica, subgenera or series have been recognized (Warburg 1897; Uphof 1959; Sinclair 1968; De Wilde 1979), but as explained under these genera, the distinctions are not sharp and only allow for informal grouping of species, largely reflected in the keys to the species.

Practical taxonomic notes — All members of the family Myristicaceae can be recognized in the field by their general habit, i.e., a slender bole with monopodial crown, the branches more or less verticillate and tiered, and the rather long exstipulate distichous leaves like those of the Annonaceae. The latter family differs in its flower structure, fruits, and lack of the red exudate of the cut bark. Further differences are obvious in the transversely cut twig where the bark has radiating parenchyma in the Annonaceae, and also the pollen is different. The genera of Myristicaceae can be told apart on vegetative characters only with experience, and flower, inflorescence, and fruit characters are necessary for a definite identification. Useful characters are the non-striate, usually finely lenticellate and granulate bark of the twigs in Gymnacranthera, or the dry leaves not readily breaking into pieces in most species of Knema and Gymnacranthera (both have reticulate sclerenchyma in the mesophyll). Certain leaf and wood anatomical characters can be used in genera diagnoses. All six Malesian genera may reach timber size.

Since myristicaceous specimens either have male flowers, or female flowers and/or fruits, keys have been constructed for both sexes, using also vegetative characters. However, specimens with only female flowers may be difficult to assign to a particular genus, and one should use both types of keys, paying particular attention to the inflorescence type.

References:
APG (Angiosperm Phylogeny Group) An ordinal classification for the families of flowering plants. Ann. Missouri Bot. Gard. 85 (1998) 531-553
De Wilde, W. J. J. O. New account of the genus Knema (Myristicaceae) Blumea 25 (1979) 321-478
E. Soepadmo & L.G. Saw (eds.) Tree Flora of Sabah & Sarawak 3 (2000) 335-473
Sauquet, H Phylogénie des Myristicaceae Rapport de stage. Institut Agronomique, Paris-Grignon (1998)
Sinclair, J. The genus Myristica in Malesia and outside Malesia Gard. Bull. Sing. 18 (1968) 1-540
Uphof, J.C.Th. A. Engler & K. Prantl Die natürlichen Pflanzenfamilien ed 2 17a II (1959) 177-199
Warburg, O. Monographie der Myristicaceae, Nova Acta Acad Caes. Leop.-Carol. 68 (1897) 1-680
Wilson, T. K. & L. M. Maculans The morphology of the Myristicaceae: I. Flowers of Myristica fragrans and M. malabarica. Amer. J. Bot. 54 (1967) 214-220

Notes — 1) Sizes given in the descriptions always relate to (measurements in) the dry state. When single measurements are given they indicate length. 2) Comprehensive discussions on the morphology of Asian Myristicaceae have been published by Sinclair (1958, 1961, 1968).

Growth form — Asian Myristicaceae are always trees, though sometimes only a few metres high. On germination the erect shoot, which initially develops into the orthotropic main stem, carries a number of spirally arranged cataphylls, soon passing into normal leaves. Growth is normally monopodial in flushes, and each season at the end of the new flush the leaves are produced ± crowded into a several-leaved pseudowhorl. The plagiotropic lateral shoots hence are ± whorled, ± horizontal, and so are the main branches on the stem. This growth form of the trees is according to the model of Massart (Hallé et al. 1978; De Wilde 1992). In the plagiotropic shoots, usually in the herbarium specimens, the leaves are generally (sub)distichous (in Malesia in a few species of Horsfieldia phyllotaxis is spiral). In some species (e.g. Gymnacranthera ocellata, Paramyristica sepicana), an apical bud with bud scales is formed, the latter leaving ring-shaped scars at the base of the innovations. Buttresses are frequent, stilt-roots occasional. The monopodial crown often is ± pyramidal in outline. Presence of stilt-roots and other characteristics have been summarized in the field-notes of most species.

Bark — The bark of the trunk is smooth or fissured (furrowed), scaly, or dippled, in certain species of Myristica and Knema blackish and gritty (the penarahan arang group of foresters). The inner bark is fibrous, red-brown. When cut the inner bark (and wood) exude a clear, pale to intense dark red sap (kino), usually free flowing, and typical for the family. The generally soft sapwood is pale, darkening brown-red on exposure; the heart- wood often is dark-coloured; the core of old trees is commonly brittle, or reported as being rotten. Field-notes on bark and wood characteristics have been given under most species. Some photographs of bark have been published by Sinclair (1958).

Indumentum — Almost all Asian Myristicaceae have some sort of indumentum (of sparse minute hairs to thickly woolly), composed of uniseriate hairs (see 'Anatomy') which may be scale-like, stellate, or dendroid. Very often the hairs are rust-coloured and early shed, but an indumentum is usually present on the sterile apical leaf bud, the very apex of the twigs, inflorescences, and the flower buds; it may persist on the lower leaf surface, dependent on the species. The length of the hairs, viz. short (0.1 mm) versus longer (0.2 mm or more) is diagnostic. The indumentum of the fruits, if present, is always diagnostic.

The indumentum in Myristicaceae is also used for the distinction of genera, as explained for Malesia below in 'Vegetative anatomy' by Koster and Baas. The hairs are essentially uniseriate, but the cells may be branched to one or two sides, forming sessile (scale-like) to long-dendroid hairs.

Twigs — The thickness (diameter) of the twigs, measured at the apex in the distal 10 cm, and whether they are terete (as usual), (blunt) triangular, ridged (mostly at both sides in between the insertion of the petiole), or ± flattened, can be used as diagnostic characters. The bark of the twigs may be longitudinally grooved (striate) to various degrees, and in older twigs may become characteristically longitudinally cracked and later on flaking. Colour of the bark of the twigs is usually some shade of brown, straw, or greyish (pale) and contrasting with the dark drying colour of the petiole. Only when colours are contrasting it is mentioned in the descriptions, being characteristic for certain species, especially in Horsfieldia, and for the whole genus Endocomia. The bark of the twigs may be lenticellate to various degrees, according to the species. In Gymnacranthera the twigs are always ± flattened and strongly lenticellate; in Knema and Endocomia lenticels are (almost) absent. Some New Guinean Myristicas are characteristically myrmecophilous with part of the twigs thickened and ant-inhabited (De Wilde 1998).

Leaves — The leaves are simple, exstipulate, pinninerved, and spirally inserted (dispersed) on orthotropic axes. However, in the plagiotropic fertile twigs, i.e. in herbarium specimens, the leaves are usually distichous; rarely the phyllotaxis is spiral, as in some species of Horsfieldia. The blade varies between elliptic to lanceolate, often being broadest at or slightly above, sometimes below the middle. The margin is occasionally revolute on drying, and only then it is mentioned in the descriptions. The lower leaf surface usually is pale and may be papillose or not, or covered with alveolar material (Koster & Baas 1981, 1982), characteristic for Knema or, e.g., Horsfieldia iryaghedhi and certain species of Myristica. The indumentum may be persistent on the lower leaf surface, but in most species of all genera it is early falling. Very characteristic for certain species in Horsfieldia, Knema, and Myristica is the presence or absence of dark-coloured, red-brown or blackish dots and/or dashes, especially on the lower leaf surface, i.e., corky warts developed from the bases of fallen hairs, visible with a strong lens. Much finer dark spots representing tannin-conglomerations are often present. The presence or absence of dots (and dashes) is important for species distinction and for this purpose, to a lesser extent, the presence or absence of microscopic papulation on the lower leaf surface is used (to be seen with a magnification of x60). In general, one should always inspect the lower leaf surface when determining a myristicaceous specimen. For the distinction of the many similar species of Myristica in New Guinea the size of the leaf blades is used; as arbitrary demarcation smaller or larger than about 15 cm is chosen for the leaf length. Whether the midrib and lateral nerves are raised or sunken above (they are always raised beneath) are useful taxonomic characteristics, as is the number of lateral nerves. The distinctness of the veins connecting the laterals (in the descriptions ‘lines of interarching’) near the blade margin, as well as the nature of the tertiary venation (in the descriptions simply 'venation') are of taxonomic importance at the species level. The size of the ultimate areoles formed by the veinlets is important for the distinction of some Knema species. In some species of all genera the colour of the blade on drying is used for species delimitation, i.e. green in Knema viridis. The angle of the lateral nerves to the midrib (in the descriptions indicated by degrees) may be diagnostic.

The sterile apical leaf bud, of a typical elongate-conical shape, has a characteristic indumentum (hairs always appressed in Myristica) and more or less characteristic shape and size. It consists of a single leaf only, and is present and visible as soon as the previous leaf in the flush has developed and expanded. In Asian species the vernation is conduplicate. In a few species, especially those from higher elevations, some bud scales may be present on the apical bud which ends the flush, and this is also rather characteristic for lowland taxa such as Paramyristica and some Myristica and Gymnacranthera species, e.g. G. ocellata, where these bud scales leave two distichous rows of closely set scars at the transition between innovations. In species of a (presumably) more or less seasonal environment (drought, or cold in the mountains), ± ellipsoid or ovoid, sterile or fertile (inflorescence) buds, normally 10 mm long or much less, composed of several cataphylls, can be found axillary to leaves; in these buds the first two scales are minute and essentially placed transversely and opposite (De Wilde 1992), as is common in dicotyledons.

References:
De Wilde, W. J. J.O. The genera of Myristicaceae as distinguished by their inflorescences, and the description of a new genus, Bicuiba Beitr. Biol, der Pflanzen 66 ('1991', 1992) 95- 125
Census of Myristica (Myristicaceae) in New Guinea anno 1994. Blumea 40 (1995) 237-344
Halle, F.et al. Tropical trees and forests. An architectural analysis. Springer, Berlin etc. (1978)
Koster, J. & P. Baas Comparative leaf anatomy of the Asiatic Myristicaceae Blumea 27 (1981) 115-173
Alveolar material in the Myristicaceae. Linn. Soc. Symp. Series, No. 10 Suppl. (1982) 131-138
Sinclair, J. Gard. Bull. Sing. 16 (1958) 205
Sinclair, J. Gard. Bull. Sing. 18 (1961) 102-327
Sinclair, J. Gard. Bull. Sing. 33 (1968) 1-540.

Photographs 1-9:

Photographs nos. 1-8 by W. J. J.O. de Wilde; no. 9 by R. Geesink

Inflorescences — Myristicaceae are practically always dioecious, except Endocomia, which is monoecious, female flowers being mixed within the more numerous male flowers in each paniculate inflorescence. The inflorescences are useful for the recognition of the genera; in detail, inflorescences are also characteristic of species (see De Wilde 1992). They are always axillary (rarely somewhat supra-axillary) amongst or below the leaves, and provided with bracts. They are polythetic, which means that their branches are never terminated by a flower. The male inflorescences are often larger and with many more flowers than the female, and show more interspecific differences. Endocomia, Gymna- cranthera, Horsfieldia, Paramyristica, and part of Myristica have non-woody, paniclelike inflorescences of short duration, while Knema and Myristica, partly, have woody, condensed, knob-like, scar-covered brachyblasts producing at the apex flowers over several seasons. These two types of inflorescence belong to basically different types, a single and a plural (multiple) type (De Wilde 1992).

The architecture of the basic single type of inflorescence corresponds with the typical mode of vegetative branching in the family. This type is axillary, compound, provided with a smooth, non scar-covered, common peduncle; the first lateral branches are opposite, but with branches higher up essentially dispersed. The knob-like inflorescences of Knema, and those of Myristica, partly, can be regarded as derived from these by reduction of branching and clustering of flowers. The superficially similar paniculate inflorescences of the remaining genera appear to be derived from a number of the basic type inflorescences, arranged in a way again comparable to the mode of vegetative branching.

In the multiple-type inflorescence the common peduncle is always provided with the scars of basal prophylls. Clustering of the flowers into flower heads or (sub)umbels adds to the variation in appearance of the inflorescences, but the presence or absence of scars of prophylls at and towards the base of the main peduncle is an essential and easily seen criterion. Knema and Myristica (both those with knob-like as well as panicle-like inflorescences) have the single type, Endocomia (partly), Horsfieldia, and Gymnacranthera have the plural type. In Paramyristica the inflorescences are essentially as in Myristica, but they are panicle-like and arranged in a short-shoot, ending in a vegetative bud.

The two highly distinctive forms of inflorescences in the genus Myristica, discriminating the two sections Myristica and Fatua, both belong to the single type; that of sect. Fatua, as that of Knema, being a strongly condensed form of the panicle-like inflorescences of sect. Myristica. As could be expected, there are quite a number of intermediate forms in Myristica.

Survey of Malesian genera with description of their inflorescences (Fig. 1)

1. Endocomia — Inflorescences predominantly of the plural type, panicle-like, resembling those of most Horsfieldias; only in one species, E. rufirachis, a few single-type inflorescences have been found. Flowers either faintly clustered, or clustered into loose umbels or umbel-like racemes, often with the apical flowers the youngest. Fig. 1a, b.
2. Gymnacranthera — Inflorescences of the plural type, panicle-like, resembling those of Horsfieldia but branching also from the axils of the lower cataphylls, so that the common peduncle is short or absent. Flowers solitary, or grouped in small few-flowered racemes or small clusters. Fig. 1c.
   Similar inflorescences with basal branching are prevalent in the Horsfieldia clavata- group (New Guinea).
3. Horsfieldia (with 3 sections) — Inflorescences of the plural type, panicle-like, variable in size and shape. Main peduncle with scars of cataphylls (bracts) and basal (cataphyll-like) prophylls present; ramification into partial inflorescences predominantly higher up in the inflorescence, but mainly basally in the H. clavata-group (with deviating androecium, New Guinea); see also under Gymnacranthera. Flowers (in most species, sect. Irya and Pyrrosa) on the ultimate branches, solitary or in few- flowered loose clusters, or in H. iryaghedhi (sect. Horsfieldia, Sri Lanka) flowers numerous, clustered into dense capitula. Fig. 1d-g.
4. Knema — Inflorescences of the single type. They consist of persistent scar-covered brachyblasts (of long duration), simple or ± digitately 2-4-branched, producing flowers at the apex each flowering season. Flowers each season few to many, closely set, often appearing as an umbel, spirally arranged, axillary to minute caducous bracts. Common peduncle absent, or up to 10 mm long, smooth. The Knema-type of inflorescence also occurs in Myristica sect. Fatua and is thought as having originated by suppression of most of the common peduncle and by contraction of the flower-bearing raceme. Fig. 1h-k.
5. Myristica — Inflorescences of the single type. The inflorescences of the two sections as recognized by Sinclair (1968: 30,41), sect. Myristica and sect. Fatua, have different designs based on the same plan, taking into account the necessary reductions and corresponding mode of clustering of the flowers. Fig. 1l-o.
   Sect. Myristica: inflorescences branched, panicle-like, generally lasting only one flowering season; peduncle (hypopodium) distinct, often flattened; the proximal pair of main branches and the central branch may be developed variously; sometimes the central branch is lacking, or all three hardly may be developed. Within sect. Myristica all kinds of intermediates between the situations of Fig. 4l-n can be found, sometimes within one species.
   Sect. Fatua: inflorescences as in Knema, i.e., simple or digitately few-branched, with woody scar-covered axes, producing flowers for several seasons; peduncle absent or short, not flattened (Fig. 1o). Also in the Fatua-type inflorescences the two opposite basal side branches are generally clearly visible, with the central branch either well developed or represented by only a single flower or the scar of its pedicel, e.g. as in M. fatua.
   Although Sinclair's sections differ in inflorescence type, it is also true that for many species their assignment to a section is arbitrary. For instance, in sect. Myristica, the type species M. fragrans has male inflorescences which may be judged as short-lived with a few flowers, but generally they grow quite old and last for many flowering seasons, each season producing one or a few flowers at the ends of one or two slender scar-covered raceme-like partial inflorescences, usually with a single middle-flower; this is the dichasial inflorescence as described by Warburg (1897: 42) and later authors. Recently these inflorescences have been described accurately by Armstrong & Tucker (1986). The Knema-like inflorescences of sect. Fatua can be related easily to a variant of the single-type inflorescence occurring in sect. Myristica.
6. Paramyristica — Here the inflorescences are truly of a mixed nature. The true (partial) inflorescences are similar to the single-type, condensed inflorescences of Myristica, but these are distichously arranged on a lateral short-shoot which ends, significantly, in a small vegetative bud. This arrangement seems also predominant in a few New Guinean species of Myristica, for instance M. markgraviana (see De Wilde 1995).

Flowers — The unisexual flowers are campanulate or urceolate, waxy-creamy or yellow outside, greenish, creamy, yellow, red, or white (Knema galeata) inside. They can be glabrous or brownish hairy on both surfaces. Flowers are frequently fragrant (e.g., Horsfieldia iryaghedhi, Myristica fragrans). The uniseriate perianth is (hard) carnose, and cleaves at anthesis along previously developed lines of suture into 2-5 lobes, to various depths, ± according to the genus. The perianth splits deepest, nearly to the base, in Endocomia and part of Knema; in some species of Horsfieldia it opens only inconspicuously at the very apex. The perianth lobes usually curve back at anthesis (especially or only in female flowers), except for Horsfieldia and possibly Paramyristica. In the descriptions the size of the dry mature buds is given, and the length of the lobes by stating the depth of the cleft by fractions. In Knema the full-sized flower buds remain closed for a long time before opening. According to the genus the short or long pedicels may or may not have one bracteole (rather large in Myristica, small in Knema). In some species of Horsfieldia the pedicel (best to be seen in male flowers) is more or less distinctly jointed at the base; this feature can be used in species delimitation. The female flowers (generally somewhat larger than the male flowers) have a single monocarpellate ovary (hairy or glabrous), with sessile or short-stipitate, usually bilobed stigma, the lobes being simple or variously lobulate again; they are conspicuously many-lobulate especially in species of Knema, style and stigma are conspicuously small in Myristica. The androecium of the male flowers is most distinctive for the genera, as explained below (Fig. 2).

Note on the construction of the androecium, and its naming in the six genera - The male organ in the unisexual flowers is formed by the fusion of a variable number of stamens. Together they form the androecium, situated in the centre of the flowers. Each anther consists of a pair of bisporangiate lobes or thecae (which are usually septate when immature). When fully fused, the number of anthers may be difficult to count, but it is half that of the often closely appressed thecae. Genera in which the filaments of the individual stamens have remained partly free (e.g. in the Madagascan genus Maloutchia), can be regarded as comparatively primitive in this respect. The Malesian genera all have (almost) completely fused stamens, forming an androecium of which the shape is characteristic for the genus. For convenience's sake, schematic drawings with a brief explanation of the construction of the androecium and the terms used in the taxonomic text are given for each genus. The (partly) connate or free anthers are shown in solid black. In the case of connate anthers these form the synandrium and the fused filaments below form a stalk called androphore. When the androphore is absent or inconspicuous, the synandrium coincides with the androecium. In Knema the anthers are mostly free, and the androecium is described in a different way. In the descriptions the number of thecae is given, but in Knema only the number of anthers. In Figure 2 the letters correspond with the explanation below.

• Endocomia — The synandrium consists of short anthers adnate by their backs to a short slender column, carried on a stalk or androphore. In the descriptions the following terms are used: synandrium, androphore.
• Gymnacranthera — The androecium practically entirely consists of elongate anthers which are tightly connate into an elongate synandrium by most of their dorsal and lateral sides. The stalk or androphore is so short that the synandrium appears to be sessile. Therefore solely the all-comprising term androecium is used in the descriptions.
• Horsfieldia — Here the synandrium is formed by elongate anthers which are completely or for a lower part attached by their backs to a swollen, solid or usually hollow column. The synandrium is either subsessile (as in Gymnacranthera), or carried on a comparatively short stalk or androphore. In the description either only the general term androecium is used when an androphore is practically absent, or the terms androecium and androphore when the latter is obvious.
• Knema — The androecium is formed by an apical disc-like structure to which the sessile or free anthers are attached radially. Below the disc there is a cylindrical or tapering stalk. In the descriptions the terms staminal disc and staminal column are used.
• Myristica — Tightly set elongate anthers are completely adnate by their backs to a rather thick central column, together forming the elongate synandrium. Above this, the column is usually produced into a sterile portion, called the sterile apex, whereas the synandrium is carried on a distinct cylindrical stalk or androphore. The terms used in the descriptions are sterile apex, synandrium, and androphore.
• Paramyristica — The subglobose synandrium is formed by (tightly set) elongate anthers which are completely fused by their backs to a swollen but largely hollow column, as in most species of Horsfieldia, and is carried on a comparatively long stalk or androphore. Terms used in the taxonomic descriptions are synandrium and androphore.

Fruits — The fruits are ellipsoid or oblong, more rarely (sub)globose, and they vary strongly in size (1-12 cm long); only in Gymnacranthera all species have rather small fruits. Fruits are essentially similar in construction in all genera, when fully ripe a firmly fleshy or ± coriaceous unicarpellate capsule, circumferentially opening at ventral and dorsal side, at the latter, though not completely to the base. Usually the fruits are variously rusty pubescent or glabrous, pale yellow or creamy, pinkish, or salmon, or in Endocomia canarioides glossy dark purple. The colourful unit of brown or black seed with bright orange or red aril remains attached to the inner base of the pericarp (which often is brightly coloured inside). In Endocomia the colour of the aril possibly is not always red, probably in some species yellow, but this needs further observation. The showy open fruits with contrasting colours supposedly attract frugivorous birds.

The fruit (pericarp) usually shrinks considerably on drying, and shape and size information given in the descriptions concerns dried specimens.

Seeds — The single seed is generally similar in shape to that of the fruit. The endosperm is ruminate, the embryo small, the seed coat ligneous, (blackish) brown, or grey, and covered by the firm-fleshy aril. According to Corner (1976) the construction of the seed coat is anatomically characteristic for the various Asian genera. The tegmen is massive (Corner 1976; Van Heel 1982) and causes the characteristic rumination of the seed (rumination in Annonaceae and some other families is of both testal and tegmic origin). The testa in Endocomia is (mostly) variegated, in general an infrequent feature of seeds.

The rather thin (sometimes thick) hard-fleshy aril is a true aril, originating from funicular as well as exostomal tissue. It is always well developed and completely covering the seed in Asian Myristicaceae (reduced in Myristica ingens from New Guinea) and either entire or shallowly to deeply laciniate, according to the genus. The aril is entire or only shallowly lobed in Knema and Horsfieldia, in the latter sometimes ± elongate above the seed into a short folded tube; the aril is incised to about halfway in Endocomia and deeply cleft (nearly) to the base in Gymnacranthera, Myristica, and Paramyristica. The embryo is small and shows variation in the position of the cotyledons and whether or not they are partially connate, according to the genera (Warburg 1897; Sinclair 1958). The endosperm (albumen) is hard and contains fat and/ or fixed (not volatile) oil, and some essential (volatile) oil (3-8% in seeds of M. fragrans, which contains a narcotic); starch may be present, is abundant in Myristica, and absent in Gymnacranthera and Horsfieldia.

References:
Armstrong, J.E. & S.C. Tucker Floral development in Myristica (Myristicaceae) Amer. J. Bot. 73 (1986) 1131-1143
Corner, E. J.H. The seeds of Dicotyledons Cambridge Univ. Press (1976)
De Wilde, W. J. J.O. The genera of Myristicaceae as distinguished by their inflorescences, and the description of a new genus, Bicuiba Beitr. Biol. der Pflanzen 66 ('1991’, 1992) 95-125
Census of Myristicca (Myristicaceae) in New Guinea anno 1994. Blumea 40 (1995) 237-344
Sinclair, J. Gard. Bull. Sing. 16 (1958) 205
33 (1968) 1-540
Van Heel, W. A. Notes on the structure of developing seeds of Knema and Horsfieldia (Myristicaceae) Blumea 28 (1982) 53-60
Warburg, O. Monographie der Myristicaceae Nova Acta Acad. Caes. Leop.- Carol. 68 (1897) 1-680

Little is known about fossil Myristicaceae. A leaf fragment, Myristicophyllum, is described from E Borneo (Geyler 1887); fossil wood, Myristicoxylon princeps, has been described from the Cretaceous in the Sahara (Boureau 1950).

References:
Geyler, H.T. Über fossile Pflanzen von Labuan aus Vega Exped. Vetensk., Jakttagelser, Stockholm 4 (1887) 499, t. 33, f. 3-6
Boureau, E. Étude paléoxylologique du Sahara (IX) Sur un Myristicoxylon princeps, n. gen, n. sp., du Danien d’Asselar. Bull. Mus. Hist. Nat. Paris II, 22 (1950) 523-528

According to the summary presented by Kühn & Kubitzki (1993), based on Marawetz (1986), and Plant Resources of South-East Asia 5 (2, 1995 & 3, 1998), chromosome numbers are high and interpreted as (paleo)polyploid. Known are for Knema: 2n = 42 (K. intermedia: n = 21), Gymnacranthera: 2n = 44 (G. farquhariana var. zippeliana: n = 21), Horsfieldia: 2n = 50 (H. iryaghedhi: n = 25), Myristica: 2n = 42 or 44 (M. elliptica: n = 21, M. fragrans 2n = 42). In the New World much higher numbers have been found (Osteophloeum: 2n = c. 280).

References:
Kühn, U. & K. Kubitzki K. Kubitzki, F.G. Rohwer & V. Bittrich (eds.) The families and genera of vascular plants vol. 2 (1993) Myristicaceae: 461
Marawetz, W. PL Syst. Evol. 152 (1986) 49-100

In most myristicaceous species the heartwood is poorly differentiated from the sapwood, and the wood is of minor commercial importance. The timber is suitable for temporary light constructions. The wood, mainly from the blackish-stemmed group (including Myristica lowiana), is mostly soft or moderately hard or heavy; perishable, but easily treated with preservatives; it is easy to work, but sometimes splits soon. The sapwood is pale, sometimes not well defined, but often the heartwood is dark reddish brown.

The seeds of some species may be used for their fat content or their fragrance, also as medicine. Myristica fragrans is most important, yielding nutmeg (seeds), mace (aril), and the spicy pericarp can be candied. The seeds of M. argentea (W New Guinea) is of minor importance.

Myristicaceae are rarely used in silviculture. Some data are given in PROSEA 5 (2, 1995 & 3, 1998). Propagation is by seed, and shade should be provided for germination and growth. A few species are ornamental (e.g., Horsfieldia iryaghedhi), or may be introduced as such (e.g., H. sylvestris)

Kino — This substance, in the field-notes called exudate or sap, oozes from freshly cut bark in larger or lesser quantities according to individual species. It is also present in the wood, twigs, and to a lesser extent in petioles and inflorescence axis. Its presence is an excellent field test when one suspects a tree to belong to the Myristicaceae. The colour varies from dark red to pink; less often it has an orange tint. Kino is not so obvious in very young trees. The amount probably varies within a species with time. It contains tannin and gum and has left many an indelible stain on the clothes of plant collectors. Warburg (1897) stated that the kino of one species has been used in America as a styptic. Its function is not known, but it may help the wounds of a damaged tree to heal. It has been described as bloody and gruesome and Malays have aptly given Myristicaceae names including darah, the Malay name for blood.

References:
De Wilde, W.J. J.O. M.S.M. Sosef et al. (eds.) PROSEA: Plant Resources of South-East Asia 5 3 (1998) 214-216 (Endocomia W.J. de Wilde), 292-296 (Horsfieldia Willd.)
De Wilde, W.J. J. O.et al. Gymnacranthera (A. DC.) Warb. R. H.M.J. Lemmens et al. (eds.) PROSEA: Plant Resources of South-East Asia 5 2 (1995) 255-260
Sambas, E.N. Knema Lour. M.S.M. Sosef et al. (eds.) PROSEA: Plant Resources of South-East Asia 5 3 (1998) 317-320
Sangat-Roemantyo, H. et al. Myristica Gronov. R. H.M.J. Lemmens et al. (eds.) PROSEA: Plant Resources of South-East Asia 5 2 (1995) 346-356
Warburg, O. Monographie der Myristicaceae Nova Acta Acad. Caes. Leop.-Carol. 68 (1897) 1-680

(J. Koster, leaf anatomy & P. Baas, wood anatomy)

Leaf anatomy — A detailed description of the leaf anatomy of the Asian Myristicaceae was given by Koster & Baas (1981). For short leaf anatomical accounts see also Schouten (1986) and De Wilde (1994). Metcalfe (1987) summarized the vegetative anatomy of the whole family. For the present survey more specimens were examined, including the two new genera Endocomia and Paramyristica.

About 65 of the species belonging to the six Asian genera have been examined leaf anatomically. Individual species will not be mentioned in this synopsis, although many of the species examined can be distinguished by their leaf anatomical characters. Genera will be mentioned when a character has diagnostic value on the genus level.

Hairs are present, at least in young leaves, on both surfaces or only on the abaxial surface (in Gymnacranthera and some Myristica species). A hair is composed of one row of short to tall cells, having one or two arms each, one or two (rarely more) cells nearest the epidermis (the so-called stalk cells) excepted. In Gymnacranthera, Myristica, and Paramyristica the cells have two arms, of unequal length in Gymnacranthera; the cells in Endocomia, Horsfieldia, and Knema have one arm. In older leaves the hairs have often been shed, but the upright walls of the most proximal parts remain as cutinized rings on the epidermis. These rings are subtended by one to numerous small cells, arranged in a circle or oval.

Alveolar material, as an irregularly structured cutinaceous layer overlying the cuticle proper, is present on the abaxial side in many species. The thickness of the cuticle proper measures up to 18 µm adaxially and to 11 µm abaxially. The cuticular flanges on the ad- axial surface are usually more or less sinuous at high focus and more or less straight at lower focus; thin areas of cuticle are present in the loops of the undulations. The cuticular flanges usually show inconspicuous pitting.

Abaxial papillae sometimes occur. Large empty idioblasts (partially) with a thin cuticle or without a cuticle, probably secretory cells, are often present. Some species abound in regular cork warts; groups of basal cells of hairs are probably the origin of some of these structures.

Stomata are usually confined to the abaxial epidermis; the stomatal type is paracytic. The dimensions of the guard cell pairs range from 8 to 21 µm for the width and from 15 to 39 µm for the length. The guard cells are often embedded in the subsidiary cells, which are dome-shaped in Gymnacranthera. In Knema, Myristica, and Paramyristica the stomatal complex is (strongly) sunken; the bordering epidermal cells show papillae. In Knema and some species of Myristica these papillae are more or less horizontally directed, leaving a star-shaped opening above the stomatal complex. In most species of Myristica and in Paramyristica the more or less upright papillae form a ring above the stomatal complex.

An adaxial hypodermis is sometimes present, either as a continuous layer or only locally. An inconspicuous abaxial hypodermis has been recorded for a few species.

The mesophyll is dorsiventral (rarely isobilateral), with mostly two or three, sometimes up to four adaxial layers of palisade parenchyma. In the leaf margin the cells adjacent to the epidermis often have sclerified walls.

The midrib is abaxially prominently raised, and adaxially raised in most Horsfieldia species, in Endocomia, Knema, Myristica, and Paramyristica. There is a more or less straight adaxial vascular bundle (sometimes strongly interrupted) and an arc-shaped abaxial bundle, sometimes joined together. The phloem is arranged in separate strands, often in two layers. One to numerous phloem bundles are interspersed in the ground tissue between the main bundles, often accompanied by xylem elements; in some Knema species there is a complete collateral bundle in the pith. In Gymnacranthera the adaxial bundle is absent. The whole system is surrounded by groups of sclerenchyma fibres, which also occur in the pith, often even in the centre of the phloem bundles. The ground tissue is from centre to periphery parenchymatous to collenchymatous, often interspersed with cells with sclerified walls; in Gymnacranthera and Knema there are often several layers of these cells at the periphery of the midrib. Sometimes adaxial chlorenchyma is continuous in the midrib.

The veins are supplied with collateral bundles; the major veins may have a more complex vascular system, not unlike that of the midrib. Sclerenchyma caps are present at the abaxial and adaxial sides; in Knema a sclerenchymatous bundle sheath is found. A usually poorly differentiated parenchymatous bundle sheath, in Knema sometimes continuous to the epidermides, surrounds the bundle and the sclerenchyma. In Knema strands are present, consisting of sclerenchyma fibres only, in position and distribution not unlike the vein bundles.

The petiole at its basal end has a vascular system consisting of three more or less arc- shaped collateral bundles with free phloem bundles adaxially. The sclerenchyma is usually confined for the greater part to the abaxial sides. The vascular system of the distal end is intermediate between that of the basal end and the midrib.

Crystals may be present in various types. Large druses in enlarged mesophyll idioblasts frequently occur, often adjacent to epidermal cells, which may be extremely flattened and have a thin cuticle and thin and short cuticular flanges (in Myristica adaxially and in Endocomia, Gymnacranthera, and Horsfieldia adaxially and abaxially). Small druses have also been found, but not in Gymnacranthera. Small spindle-shaped particles often occur, usually grouped in cells of the mesophyll and the ground tissue of the midrib. Other crystal types have been found in one or a few species only.

Large, more or less spherical cells frequently occur in the mesophyll and the ground tissue of the midrib. Usually they contain oil, in some species probably tannin- or muci- lage-like substances. The large empty idioblasts in the epidermis have been mentioned above. Fairly thick-walled tubule-shaped cells have sometimes been found, adaxially and abaxially of the sclerenchyma caps of the vein bundles. The content of these cells is probably tannin.

Sclereids are often present as brachy- to astrosclereids in the ground tissue of the midrib. Filiform, rarely branched sclereids have been recorded for Gymnacranthera. Astrosclereids and thick filiform, branched sclereids are present in a few species only. The genera Gymnacranthera and Knema can be distinguished by a combination of leaf anatomical characters. Leaf anatomy provides no means to discriminate between Endocomia and Horsfieldia and between Myristica and Paramyristica.

Wood anatomy — The wood anatomy of the Myristicaceae is fairly uniform. For a detailed family description and literature survey see Metcalfe (1987). Wood anatomy of the main Malesian genera is summarized in the Prosea Handbooks 5: 2 & 3 (Lemmens et al. 1995; Sosef et al. 1998) and pictured by Ilic (1991). The wood is diffuse-porous with vessels medium-sized and in low density (usually 3-12/sq.mm), solitary and in radial multiples. Perforations mixed simple and scalariform, but one of the types dominant or (virtually) exclusive in some species. Intervessel pits ranging from scalariform to opposite and alternate. Vessel-ray pits often coarse and with reduced borders to simple. Fibres thin- to medium thick-walled, with minutely bordered to simple pits, often septate around the vessels. Parenchyma scanty paratracheal to narrowly vasicentric and often also in zonate bands. Rays typically l-2(-3)-seriate, heterocellular and composed of fairly large cells. Crystals present in ray cells or absent. Tanniniferous tubes, usually in very low frequency, present in all species studied, and virtually unique to the family Myristicaceae (except sporadic occurrence in some members of the Ulmaceae). Oil and/ or mucilage cells present among the axial and ray parenchyma in some species.

References:
De Wilde, W. J. J. O. Paramyristica, a new genus of Myristicaceae Blumea 39 (1994) 341-350
Ilic, J. CSIRO Atlas of Hardwoods (1991) 331-334
Koster, J. & P. Baas Comparative leaf anatomy of the Asiatic Myristicaceae Blumea 27 (1981) 115-173
Lemmens, R.H.M. J., I. Soerianegara & W.C. Wong (eds.) Plant Resources of South-East Asia 5 2 Timber Trees: Minor commercial timbers (1995) 255-260
Metcalfe, C.R. Anatomy of the Dicotyledons, Ed. 2, Vol.3 (1987) 56-71
Schouten, R.T.A. Revision of the genus Gymnacranthera (Myristicaceae) Blumea 31 (1986) 451-486
Sosef, M.S.M., L.T. Hong & S. Prawirohatmodjo (eds.) Plant Resources of South-East Asia 5 3 Timber Trees: Lesser-known timbers (1998) 292- 296,317-320, 603-662

(R.W. J.M. van der Ham)

The pollen morphology of the Myristicaceae has been poorly known for a long time. The earliest more extensive account is that by Wodehouse (1937), who dealt with 36 species of the American genera, providing detailed descriptions and drawings. A more inclusive treatment is the light microscopic study by Agababian (1970) of 10 genera from America, Africa and Asia. Further, pollen of a limited number of species is described in pollen floras, of which Tissot et al. (1994) stands out by informative light and scanning electron micrographs (Gymnacranthera, Knema, Myristica). Generic accounts are those by Siddiqi & Wilson (1975; 8 spp. of Knema) and Medeiros Carreira (1985; 36 spp. of Virola, incl. Bicuiba). A preliminary paper by Walker (1976) contains a short family description based on light and scanning electron microscopic data. Comprehensive descriptions of all American, African and Madagascan genera, including scanning and transmission electron micrographs, are in a series of papers by Walker & Walker (1979, 1980, 1981, 1983). The Asian genera, among which the largest in the family (Horsfieldia, Knema, Myristica), are still in need of elaborate palynological study. To date the pollen of the Asian Endocomia (4 spp.) and Paramyristica (1 sp.) and the African Staudtia (2 spp.) is entirely unknown.

Pollen grains of Myristicaceae are usually subspherical to slightly boat-shaped monads with a single, probably always distal aperture. Occasional chance tetrads, observed, for instance, in Iryanthera, are tetragonal. The outline in polar view is subspherical to elliptic, or sometimes obtusely rectangular. Outline in equatorial view is subspherical to elliptic, or often obtusely triangular with a straight to slightly convex apertural side and a more or less strongly convex nonapertural side. Pollen grain size (largest equatorial diameter) is mostly between 20 and 40 µm. Pollen of Brochoneura is smaller (14-21 µm). Some other genera have larger pollen grains: Gymnacranthera (up to 49 µm), Knema (up to 57 µm), Myristica (up to 59 µm), and Mauloutchia (up to 69 µm).

Aperture morphology is relatively simple and not much diverse. It ranges from distinctly sulcate via indistinctly sulcate (sulcoidate) to more or less ulcerate or ulceroidate. The aperture margins are often not clearly defined, which in ulceroidate groups may lead to a superficially inaperturate condition (cryptoaperturate). Wodehouse (1937) observed that an apertural area, even in pollen with a hardly recognizable aperture in the exine, shows a distinctly thickened intine. Sometimes such intine parts seem to be acetolysis-resistant (Walker & Walker 1983).

Exine thickness is from 0.5 to 5 µm. Rather thin exines are found in Brochoneura (0.5 µm), Pycnanthus (0.5 µm, exclusive echinae) and Scyphocephalum (0.8 µm). Fairly thick exines (3-5 µm) occur in the coarsely reticulate Myristica pollen. Exine stratification is usually distinct, with a thin to thick infratectal layer, which is mostly columellate. In Brochoneura the infratectum is thin, so that the granulae observed by Walker & Walker (1979) might actually represent short columellae. In Mauloutchia pollen the verrucate/ scabrate sexine elements seem to stand directly on the nexine, although irregular columella-like structures occur as well. The allegedly primitive granular infratectum reported by Walker & Walker (1979) seems to be part of a granular sexine structure, which is rather a derived feature. Distinct infratectal granules were found so far only in Otoba, more or less adhering to the inner tectum surface and mixed with columellae. In view of the other pollen characters (see below) this is probably also a derived condition. The tectum as well as the nexine can be relatively thin to rather thick. In a few genera most or only the inner part of the nexine may be lamellate. Because of the absence of any contrast in the nexine in transmission electron micrographs, the whole exine is considered to be ectexinous.

Exine ornamentation is the most diverse character in Myristicaceae pollen: from psilate/ perforate via finely fossulate to coarsely reticulate, with several derivations. Pollen of Brochoneura (Madagascar) has a simple massive psilate/perforate tectum. Pollen of the American genus Otoba is psilate/imperforate with a proximal, ± protruding (coarsely) reticulate area. In Compsoneura and Virola (both American) the ornamentation is finely fossulate to coarsely reticulate. In both genera reticulate pollen with scabrate (± banded) muri is found. Such ornamentation occurs also in Iryanthera (America) and Coelocaryon (Africa), while the finely fossulate type with vaguely banded muri of Haematodendron (Madagascar) and the crotonoid type of Scyphocephalum (Africa) can be easily joined. The finely fossulate type of Compsoneura and Virola is also known from Gymnacranthera (Asia), and the (more) coarsely reticulate type from the American Bicuiba and Osteo- phloem, and the Asian Horsfieldia, Knema and Myristica. In Horsfieldia the reticulum is sometimes interrupted, so that an intectate condition remains. Ornamentation in the Madagascan genus Mauloutchia is diverse: scabrate, verrucate (verrucae scabrate or smooth) or scabrate/echinate. Scabrate verrucae occur also in Cephalosphaera (Africa). The pollen of Pycnanthus (Africa) is finely reticulate/echinate.

Concluding, the pollen of the Myristicaceae is diverse. Virtually every genus in the family is palynologically distinct. A rigid subdivision based on pollen morphology, however, is difficult, but may be attempted after a more extensive study of the Asian genera.

The family has a modest fossil pollen record (Muller 1981). Pycnanthus type pollen is known from the Upper Eocene and Lower Miocene of Africa, and pollen grains of the Virola type from Pliocene and Quaternary sediments in Guyana.

References:
Agababian, VS. Biol. Zh. Armenii 23 5 (1970) 58-69
Medeiros Carreira, L.M. Bol. Mus. Paraense E. Goeldi, Bot. 2 (1985) 29-76
Muller, J. Bot. Rev. 47 (1981) 1-142
Siddiqi, M.R. & T. K. Wilson Pak. J. Bot. 7 (1975) 197-200
Tissot, C., H. Chikhi & T. S. Nayar Publ. Dép. Écol. Inst. Fr. Pondichéry 35 (1994)
Walker, J.W. Linn. Soc. Symp. Ser. 1 (1976) 251-308
Walker, J.W. & A.G. Walker Ann. Missouri Bot. Gard. 66 (1979) 731-755
Amer. J. Bot. 67 (1980) 603-611
Grana 20 (1981) 1-17
Amer. J. Bot. 70 (1983) 315-326
Wodehouse, R.P Brittonia 2 (1937) 397-402

(R. Hegnauer)

General remarks — Chemical characteristics of the family were discussed in a number of reviews in recent time (Hegnauer 1969, 1989, 1990; Gottlieb 1979). Many references and structural formulae are given in these surveys. Therefore a compact résumé of presently known facts and some references to most recent chemical investigations of myristicaceous plants, with emphasis on Asian taxa, seem to be appropriate here. Most members of the family are locally used in traditional medicine. This is one of the reasons why we are relatively well informed about its secondary metabolites. Moreover, Myristica fragrans yields the famous spices nutmeg and mace. There is plenty of literature about this plant, its cultivation and its products (e.g., Brticher 1977, Purseglove et al. 1981, Delaveau 1987, Flach & Tjeenk Willink 1989). Because nutmeg, if taken in large amounts, is toxic and causes among other symptoms hallucinations, pharmacologists and ethno- botanists interested in psychotropic plants became also involved in nutmeg research (Efron 1967).

Chemistry of the family — Essential oils, lignans and neolignans, flavonoids in the widest sense and biogenetically related phenolic compounds, peculiar acetogenins based on long-chain fatty acids, and large amounts of a special type of triglycerides in seeds are outstanding myristicaceous features. Moreover, tryptamine-derived alkaloids were reported in several genera, and diterpenoids and triterpenoids were detected only erratically hitherto.

Essential oils — Belonging to woody polycarps (i.e. Magnoliidae-Magnolianae sensu Takhtajan 1980) Myristicaceae have oil cells in most of their parts and usually are aromatic plants. So far only essential oils of nutmeg and mace were investigated thoroughly (Purseglove et al. 1981). There are scarcely qualitative differences between the oils of nutmeg and mace, but rather marked quantitative differences. The same is true between nutmeg oils of different production centres (West India [Grenada], Indonesia and East India) and even between individual trees of the island Grenada. Roughly nutmeg yields 7-16% and mace 4-17% essential oil. Nutmeg oil contains 61-66% mono- terpene hydrocarbons (85-93% individual trees of Grenada), mainly pinenes and sabinene, 5-15% (6.6-12%) monoterpene alcohols and their esters, such as geraniol and linalool, and their acetates, and 2-18% (0-3.5%) phenylpropanoid ethers, among which myristicin, elemicin and safrole predominate. These aromatic allylphenolic ethers are assumed to represent the main toxic and psychotropic constituents of nutmeg and mace. The essential oil of leaves of M. fragrans also contains mainly pinene and myristicin.

Lignans and neolignans — These plant constituents are dimers of phenylpropanoids (C₁ ... C₉ + C_1’ ... C_9’). According to Gottlieb and Yoshida (1989) lignans and neolignans should be defined biogenetically not purely on chemical arguments. They consider lignans as C₈-C_8’-linked dimers of phenolic derivatives of cinnamyl alcohol (C₆H₅-⁷CH = ⁸CH⁹CH₂OH) or phenolic derivatives of cinnamic acid (C₆H₅-⁷CH = ⁸CH-⁹COOH). On the other hand neolignans are dimeric derivatives of allylbenzenes (C₆H₅-⁷CH₂-⁸CH = ⁹CH₂; e.g. eugenol) or propenylbenzenes (C₆H₅-⁷CH = ⁸CH-⁹CH₃;e.g. isoeugenol), and different types of linkage between the two monomers occur, e.g. 8-8', 8-1', 8-3', 8-5', 8-7', 5-5', 1-5', 8-O-4', 4-O-5' etc. In both, lignans and neolignans, one or two additional linkages between the two units are often present. In lignans C₉ and C_9' carry oxygen and in neolignans they do not. A botanical argument which favours such a lignan-neolignan distinction is their distribution in seed-plants. Lignans occur everywhere in gymnosperms and angiosperms, and neolignans are mainly (not wholly!) restricted to Magnoliidae-Magnolianae (Takhtajan 1980) which correspond with woody Polycarpicae + Piperales of Wettstein (1935). In Myristicaceae both true lignans and neolignans occur frequently. Conserva et al. (1990) call neolignans "the most conspicuous constituents of Myristicaceae." Examples of neolignans occurring in the family are dehydroguaiaretic acid and 1,2-dihydrodehydroguaiaretic acid of the stem bark of Knema furfuracea (Pinto et al. 1990), and lignans are represented, e.g. by asarinin, horsfieldin and dihydrocubebin, from leaves, bark, wood and seeds of Horsfieldia iryaghedhi (Gunatilaka et al. 1982; Tillekeratne et al. 1982). By condensations with chalcones or dihydrochalcones neolignans can give rise to still more complex phenolic compounds such as the lignoflavonoids iryantherin A to J of South American Iryanthera taxa (Conserva et al. 1990; Silva et al. 1995). Finally it should be mentioned that not all phytochemists follow the lignan-neolignan-definition of Gottlieb and Yoshida. Many chemists prefer the older definition which considers all 8-8'-linked dimeric phenylpropanoids as lignans, and dimers with other linkages, e.g. the 5-5'-linked dehydrodieugenol and the 8-5'-linked carinatone of Virola carinata, as neolignans.

Flavonoids and bio genetic ally related phenolic compounds — If flavonoids are defined as phytoconstituents derived from a cinnamic acid and three acetates (malonates) which yield the phloroglucinol- or resorcinol-type A-ring, this class of natural products comprises many chemical subclasses. Myristicaceae are outstanding producers of flavonoids sensu lato. At present the following types of flavonoid phenolics are known from the family: Flavonols (e.g. kaempferol, quercetin), flavones (e.g. apigenin, luteolin, 7,4’- dimethoxyflavone, a 5-desoxyflavonoid), flavanones (pinocembrin), several chalcones and dihydrochalcones, several diarylpropanes, isoflavones (e.g. 2’-hydroxyformonon- etin [also a 5-desoxyflavonoid]), pterocarpans (demethylhomopterocarpin, maackiain), flavan-3-ols (catechins, e.g. fisetinidol) and flavans. Moreover, the family is rich in condensed tannins which are based on catechins and leucoanthocyanidins (flavan-3,4- diols). Leucoanthocyanidins have not yet and catechins only rarely been isolated from Myristicaceae. Nevertheless the production of adstringent kino-like exudates by most species and the demonstration of the presence of procyanidins in a few species render possible the conclusion that myristicaceous tannins are of flavanoid nature. Still another type of natural products is represented by stilbenes. In contrast to flavonoids which are C₆-C₃-C₆ compounds, stilbenes are C₆-C₂-C₆ products, because during the biosynthesis of usual stilbenes one CO₂ is lost. Stilbenes occur in Myristicaceae, e.g., as mono-, di- and trimethylethers of resveratrol (= 3,5,4’-trihydroxystilbene).

Peculiar acetogenins based on long chain fatty acids — Two main types of such acetogenins or polyketides occur in the family, (a) The anacardic acid-cardol-type of alkyl- or alkenylphenols which is based on ordinary polyketides. If polyketide synthesis starts with a cinnamic acid molecule this pathway yields ω-phenylalkanyl- and -alkenylphenols. The phenolic part of such acetogenins originates from cyclization of the last three or four acetyl units of the polyketide chain and bears one, two or three phenolic hydroxy Is (= phenol-, resorcinol- and phloroglucinol-type compounds). Additionally this aromatic ring may carry a carboxyl group (anacardic acids sensu stricto) or an acetyl group (acetophenone derivatives). Phenolic alkanones, such as the malabaricones, have an oxo group in the side chain next to the aromatic ring. In some species 3-alkyl- or 3- ω-phenylalkylisocoumarins occur; these metabolites are lactones of an anacardic acid carboxyl with a 2'-hydroxyl in the side chain. Thus this type of biogenetically related phenolic polyketides is extremely diverse in Myristicaceae. (b) The second class of acetogenins bears a methyl- or methylene-butanolide or -butenolide structure which is probably formed by a reaction of the carboxyl group terminating the polyketide chain with a pyruvate unit or its enolic form. Examples of this type of acetogenins are iryellip- tin, grandinolide and the juruenolides of Iryanthera elliptica, grandis, jururensis and ulei and Virola surinamensis (Lopes et al. 1994, 1996).

Most recent phytochemical investigations treat mainly lignanoids, flavonoids and acetogenins. Examples are: Horsfieldia iryaghedhi (Gunatilaka et al. 1982; Tillekeratne et al. 1982). Knema austrosiamensis (Gonzales et al. 1993, 1996), K. elegans (Spencer et al. 1980), K furfuracea (Pinto et al. 1990; Zahir et al. 1993), K. glomerata (Lu Zeng et al. 1994), K. laurina (Kijjoa et al. 1991; Gonzalez et al. 1996), K. tenuinervia subsp. setosa (Kijjoa et al. 1991). Myristica dactyloides (Herath & Priyadarshini 1996, 1997). Pycnanthus angolensis (Omobuwajo et al. 1992). For 1,3-diarylpropanes and 1,3-diarylpropan-2-ols and catechins Virola elongata and minutiflora are noteworthy (Kijjoa et al. 1981). Virola venosa (Kato et al. 1992) and Virola aff. pavonis (Martinez & Torres 1997) represent a notable example for the vicarious occurrence of lignans and neolignans in the family.

Seed fats (oils) — Myristicaceae store large amounts of triglycerides in seeds; they are accompanied by proteins and in some species by starch. The triglycerides of the family contain saturated fatty acids as main acids, usually 14:0 (myristic) and 12:0 (lauric) and sometimes 16:0 (palmitic) or 18:0 (stearic). The presence of large amounts of 14:0 in several seed fats explains the fact that trimyristin could be isolated from the seeds of a number of species. Often the seed lipids contain a large portion of unsaponifiable matter, i.e. non-triglycerides. The non-triglyceride part consists of essential oils, lignans, acetogenins and other resinous matters.

Alkaloids — As already mentioned (Efron 1967) tryptophan-derived protoalkaloids and β-carboline alkaloids occur in several species of Virola. Recently 5-methoxy- N,N-dimethyltryptamine, 6-methoxy-2-methyl-l,2,3,4-tetrahydro-β-carboline and horsfiline, C₁₃H₁₆N₂O₂, a new oxindole alkaloid, were isolated from leaves of Horsfieldia superba (Jossang et al. 1991). Leaves of Osteophloeum platyspermum contain the methylether of N-methyltryptophan. Thus a special metabolism of tryptophan yielding psychotropic tryptamines and simple indolic alkaloids seems to be present in New World (Virola and Osteophloeum species) and Old World (Horsfieldia species) members of the family. Bennett and Alarcón (1994) published recently a remarkable ethnobotanical paper about Amazonian Myristicaceae and about hallucinogenic uses of Osteophloeum platyspermum and Virola duckei in Ecuador.

Meroterpenoids (= compounds of partly terpenoid origins) — Fruits (seed kernels, arilli, pericarps) yield lipid fractions which often contain besides triglycerides appreciable amounts of ‘resinous matter’ of varying composition (essential oils, lignanoids and flavonoids, acetogenins and, in some instances, meroterpenoids). Such a meroterpenoid is komboic acid of seeds of Pycnanthus kombo. It amounts to ca. 23% of total seed 'fat' and was characterized as 16(2’,5’-dihydroxy-3’-methylphenyl)-2,6,10,14-tetramethyl- 2,6,10,14-hexadecatetraenoic acid, i.e. a 2-geranylgeranyl-substituted 6-methylhydroquinone with one of the last methyl groups in the geranylgeranyl side chain oxidated to COOH. Biogenetically related tocotrienols (vitamin E group), of which 2,8-dimethyl- 2-(4,8,12-trimethyl-3,7,l l-tridecatrienyl)-6-chromanol was the main product, were isolated from fruits of Iryanthera grandis, and seeds of Otoba parvifolia yielded a series of farnesylated aromatic to semiaromatic and ring-constricted compounds which probably all derive from the same biogenetic pathway, i.e. farnesylation of gentisic acid and consequent modifications of the resulting farnesylgentisic acid (Ferreira et al. 1989, 1995). Gentisic acid may also be involved in the biosynthesis of komboic acid and the tocotrienols in which an aliphatic diterpene in place of the sesquiterpene farnesol is combined with an aromatic ring.

Diterpenes, triterpenes and phytosterols — Small amounts of phytosterols and tetra- and pentacyclic triterpenes are ubiquists in unsaponifiable matters of plant lipids. Accumulation of diterpenes and triterpenes seem to be much more restricted and rather rare in the family. The tetracyclic triterpenes cycloeucalenol and 24-methylenecycloartanol were isolated from wood of Cephalosphaera usambarensis, and nutmegs yielded a saponin with oleanolic acid as sapogenin. Diterpenoids were isolated from leaves and twigs of Osteophloeum platyspermum (a kaurane and three eperuane derivatives), and recently stem bark of Staudtia kamerunensis yielded staudtienic acid, C₂₀H₂₆O₂ with a rearranged abietane structure (Noumbissie et al. 1992).

Summary and chemotaxonomic remarks — Myristicaceae are chemically characterized by a number of metabolic features.

1. They store large amounts of triglycerides of buttery or fatty consistency because 12:0 and 14:0 (16:0, 18:0) are their main fatty acids.
2. They deposit variable amounts of essential oil in oil cells which occur in practically all plant parts.
3. They have an extremely diverse flavonoid metabolism which results in the production of rather characteristic compounds in a number of their taxa, e. g. isoflavones, ptero- carpans, 1,3-diarylpropanes and their 2-ols, chalcones and dihydrochalcones, flavanones, flavones and catechins.
4. Lignans and/or neolignans occur in large numbers and often in large amounts in all investigated plant parts of practically every member of the family.
5. Two types of polyketides (acetogenins) occur widely in the family: the anacardic acid-type with several variants and the methylbutanolide-type with a smaller number of variants.
6. Myristicaceae are relatively tannin-rich plants, but the precise nature of their tannins is not yet known. All available facts indicate a universal presence of condensed tannins, i.e. oligo- and polymeric proanthocyanidins.

Idioblasts storing essential oils characterize woody polycarps or Magnolianae (Takhtajan 1980). Within this taxon Myristicaceae resemble Lauraceae mostly in their seed fats and in their secondary metabolism. Both families are virtuous producers of neolignans and lignans and of polyketides of the butanolide-type. The typical alkaloids of Magnolianae are benzyltetrahydroisoquinoline-type bases. They are widespread in Lauraceae, but seem to be totally absent in Myristicaceae, but also in Winteraceae, Calycanthaceae and some other, mostly small relictic families. Myristicaceae, however, are not an alkaloid-free taxon. Some of its members produce tryptamine derivatives including β-carboline and the oxindole horsfiline. The tendency to replace benzylisoqui- nolines by tryptamine derivatives does also occur in some members of Lauraceae (Aniba p.p., Nectandra p.p. and Umbellularia p.p.) and is characteristic of Calycanthaceae. Myristicaceae could represent a member of Magnolianae in which accumulation of benzylisoquinolines was totally replaced by intensifying the production of lignanoids and flavonoids s.l. Finally it should be mentioned that Myristicaceae resemble strikingly Papilionoideae (many 5-desoxyflavonoids, isoflavones, pterocarpans, 1,3-diarylpropanes, flavans and fisetinidol-type catechins).

References:
Bennett, B.C. & R. Alarcón Osteophloeum platyspermum and Virola duckei (Myristicaceae): Newly reported as hallucinogens from Amazonian Ecuador Econ. Bot. 48 (1994) 152- 158
Brücher, H. Tropische Nutzpflanzen (Ursprung, Evolution und Domestikation) (1977) 426- 427, Myristica fragrans Springer-Verlag Berlin
Conserva, Lucia M.et al. Phytochemistry 29 (1990) 3911-3918, Dihydrochalcones and the dihydrochalcone-based flavonolignans iryantherin A to E from fruits and barks of Iryanthera laevis, paraensis and ulei
Delaveau, P. Les épices: histoire, description et usage des différents épices: aromates et condiments (1987) Ed. Albin Michel S.A., Paris Part IV Passez muscade (Myristica fragrans) 154-164
Efron, D.H. (Editor-in-chief) Ethnopharmacologic search for psychoactive drugs Public Health Publ No. 1645 (1967) U.S. Government Printing Office, Washington, D.C. Part III: 185-229: Myristica fragrans, with 3 contributions treating mainly psychoactivity, toxicology, pharmacology and chemistry of nutmeg. Part IV. South American snuffs with 6 contributions of which one treats botanical origins of snuffs known as Yakee and Paricá (prepared by some tribes with barks of Virola calophylla, calophylloi- dea and elongata), one treats the snuff Epéna prepared from the bark of Virola calophylloidea, and one treats the identification of N-methyltryptamines and β-carbolines in Epéna and Paricá-snuff preparations and in the bark of Virola calophylla, and an other one treats the psychoactive action of tryptamine derivatives. The harmala alkaloids (= β-carbolines) are treated in part V which is devoted to Malpighiaceae which yield the snuffs Ayahuasca, Caapi and Yagé
Ferreira, A.G.et al. Phytochemistry 28 (1989) 579-583
Ferreira, A.G.et al. Phytochemistry 40 (1995) 1723-1728
Flach, M. & M. Tjeenk Willink E. Westphal & P.C.M. Jansen (Eds.) PROSEA A selection (1989) 192-196, Myristica fragrans. Pudoc Wageningen
Gonzalez, M. J.T.G. et al. Phytochemistry 32 (1993) 433-438, Two resveratrol methyl ethers, a diarylpropane, a flavan, two resorcinol- and two phloroglucincol-type alkanones and two corresponding alkenones and the two lignans episesamin and xanthoxylol from wood of Knema austrosiamensis
Gonzalez, M. J.T.G. et al. Phytochemistry 43 (1996) 1333-1337, Saturated and monounsaturated anacardic acids, alkylphenols and alkanoylphenols from wood of Knema austrosiamensis and bark of K. laurina and 5-hydroxy-3’,4’-methylene-dioxyflavan from bark of K. laurina
Gottlieb, O.R. Chemical studies on medicinal Myristicaceae of Amazonia J. Ethnopharmacol. 1 (1979) 309-323
Gottlieb, O.R. & M. Yoshida Lignans J.W. Rowe (Ed.) Natural products of woody plants (1989) 439-511 Springer-Verlag, Berlin
Gunatilaka, A. A.L. et al. Phytochemistry 21 (1982) 2719-2723, Lignans asarinin, dihydrocubebin and horsfieldin and trimyristin from seeds of Horsfieldia iryaghedhi
Hegnauer, R. Chemotaxonomie der Pflanzen, Vol. V (1969) 144-153,435-437,457
Hegnauer, R. Chemotaxonomie der Pflanzen Vol. VIII (1989) 108-118, Comments on the chemistry of Polycarpicae (= Magnoliidae sensu Cronquist), including fatty acid-derived acetogenins of Myristicaceae
Hegnauer, R. Chemotaxonomie der Pflanzen Vol. IX (1996) 101-111
Herath, H.M.T.B. & A.M. A. Priyadarshini Phytochemistry 42 (1996) 1439-1442
44 (1997) 699-703, Neolignans and arylalkanones (malabaricones) from bark
Jossang, A.et al. J. Org. Chem. 56 (1991) 6527-6530
Kato, M. J.et al. Phytochemistry 31 (1992) 283-287, Two 5-desoxyflavones from flowers, pericarp and leaf and ten lignans from pericarp, aril, seed and root and a special type of w-phenylalkanoylphenol (1-[11- phenylundecanoyl]-3-hydroxycyclohexan-2,6-dione) which occurs in all parts and was already known from Virola elongata and sehifera
Kijjoa, A. et al. Phytochemistry 20 (1981) 1385-1388, Virolane and virolaflorine, two 1,3-diarylpropanes from wood of Virola minutiflora and virolanol, virolanol-B and -C, three l,3-diarylpropan-2-ols and (-)-fisetinidol from wood of V. elongata
Kijjoa, A. et al. Planta Medica 57 (1991) 575-577, Bark of both species yielded only several acetogenins of type (a)
Lopes, N.P et al. Phytochemistry 35 (1994) 1469-1470
43 (1996) 1089-1092, 17 Neolignans from leaves and seeds and juruenolide C and D and 3 propiophenones from seeds and a new juruenolide from seedlings
Lu Zeng et al. J. Natural Products 57 (1994) 376-381, Stem bark yielded 10 (a)-type polyketides among which kneglomeratanol and the two acetophenones kneglomeratanone A and B and the isoflavones fromononetin, biochanin-A and 8- O-methylretusin
Martinez V., J.C. & R. Torres Ch. Phytochemistry 44 (1997) 1179-1182, Leaves yielded nine neolignans among which otobaphenol
Motter Magri, Fatima M. et al. Phytochemistry 43 (1996) 669-671, Unripe pericarps yielded an 1,4-dioxane-type neolignan (linkages 7-0-7' and 8-0-4') and 4 butanolide-type polyketides
Noumbissie, B.E. et al. J. Natural Products 55 (1992) 137-139
Omobuwajo, O.R. et al. Phytochemistry 31 (1992) 1013-1014, 2'- Hydroxyformononetin and its 7-methyl ether from wood
Pinto, Magdalena M.M. et al. Phytochemistry 29 (1990) 1985-1988, (a)-Type acetogenins among which an isocoumarin and two neolignans (dehydroguaiaretic acid and its 1,2-dihydro derivative) from bark
Purseglove, J.W. et al. Spices Vol. 1 (1981) Logman Group Ltd., London Nutmeg and mace p. 174-228: History — Botany — Ecology — Cultivation — Diseases — Pests — Improvement — Products and end-uses — Processing and manufacture — Chemistry — Standard specifications — Production, trade and markets — 3 pp. of References
Silva, Dulce H.S. et al. Phytochemistry 38 (1995) 1013-1016, Iryantherin G to J from fruits of Iryanthera grandis
Spencer, G.F. et al. J. Natural Products 43 (1980) 724-730, Three neolignans and several anacardic acid- and resorcinol-type acetogenins from seeds
Takhtajan, A. J. Outline of the classification of flowering plants (Magnoliophyta) Bot. Rev. 46 (1980) 225-359
Tillekeratne, L.M.V. et al. Phytochemistry 21 (1982) 467-476, Bark, wood and leaves yielded the lignans asarinin and dihydrocubebin and dodecanoylphloroglucinol
Wettstein, R. Handbuch der systematischen Botanik 4 Aufl. 1935 Franz Deuticke, Leipzig- Wien
Zahir, A. et al. J. Natural Products 56 (1993) 1634-1637, Leaves yielded two new phenyl-acylphenols, knerachelin A and B

The preferred trade name applicable for Myristicaceae in general is penarahan. Peninsular Malaysian names are: chendarah, chendarahan, darahan, penarah, penarahan, pendarah, andpendarahan; in Sabah: darah-darah (preferred name); in Sarawak: binarah (Murut), bindara (Ke\abit)Jela bala (Kenyah), kayo bela (Kayan), kayo raha (Berawan), kumpang (preferred name), pang (Bidayuh), pendarahan, pumpu (Bidayuh Sadong), raha meban (Punan Tutoh); the names balun ijok, darah-darah, penaharan, and pianggu are also used in Sarawak; in the Philippines: duguan (Filipino language).

The group of Myristicas with black, gritty bark is called penarahan arang in Malaya and parts of Indonesia. Some Knema species in Indonesia are called ki-mokla (Sundanese). Specific names are, e.g., swamp nutmeg for Myristica elliptica (W Malesia) or mangrove nutmeg for M. hollrungii (New Guinea); thepapua nutmeg is M. argentea, a species locally cultivated in SW Papua Barat. Some other Malay names for Myristicaceae may be mentioned here: the nutmeg, Myristica fragrans, is called buah pala, while wild nutmegs are called pala bukit or pala hutan. Horsfieldia irya is pianggu and H. crassifolia is jangkang paya. Jangkang means stilt-roots and paya is swamp. In Sarawak and Brunei, Myristicaceae are generally called kumpang. In New Guinea the family has many different names in the local languages. This information about the vernacular names comes largely from Sinclair (1958).

Reference:
Sinclair, J. A revision of the Malayan Myristicaceae Gard. Bull. Sing. 16 (1958) 205-472

(based on male flowering specimens)

(based on female flowering and fruiting specimens, also using vegetative characters)

Trees, monoecious (always?). Twigs never angular or ridged, without lenticels, sometimes flaky. Leaves membranous or chartaceous, slightly brittle when dry, lower surface not pale, not papillose, dots absent; reticulation lax, distinct or not. Inflorescences with male and female flowers mixed (always?), sometimes pseudoterminal (terminal bud of twig abortive), paniculate, branched from near the base; basal cataphylls caducous; bracts caducous; flowers in small clusters, buds either in the same or in different stages of development. Flowers short or long pedicellate, at base not articulated; bracteole absent. Male flowers: perianth rotate, thinly leathery, inside pubescent, yellowish or (in New Guinea) bright red; buds small, ellipsoid, ovoid, or subglobose, cleft to c. 3/4 to nearly the base, lobes 3-5, spread or recurved in anthesis; androecium with androphore narrow, glabrous, synandrium small, (depressed) globose or short-ellipsoid, with 2-6 completely fused, short (0.5 mm or less) elliptic anthers. Female flowers: similar to male; ovary ovoid, glabrous, stigma sessile, minute, 2-lobed, lobes narrow or broad and 2-5-lobu- late. Infructescences small or large panicles, to 30 cm long. Fruits ellipsoid or (ob)ovoid, glabrous, yellow or purplish; pericarp thin or thick, fleshy-leathery; aril yellowish (?) or red, subentire or to halfway laciniate; seeds usually pointed at apex, variegated; albumen ruminate, without (?) starch; embryo incompletely known.

Distribution The genus has 4 species, ranging from South China through Southeast Asia and Malesia east to New Guinea; not in Central & East Java, Lesser Sunda Islands, Solomon Islands or Australia. Map 1(see p. 3).

Note In most species female flowers have not been seen; they should be found in the large, paniculate, predominantly male-flowerd inflorescences, or in purely female inflorescences.

Tree 10-35 m. Twigs (grey-)brown, 1.5-4(-10) mm diameter, with minute grey-brown hairs 0.1 mm long, early glabrescent, bark striate, on older twigs sometimes longitudinally cracking, lenticels very inconspicuous. Leaves membranous or chartaceous, elliptic to oblong, 8-30 by 4-13.5 cm, base subcordate or rounded to short-cuneate, apex acute-acuminate; upper surface dark brown, glabrous, lower surface early glabrescent; midrib on both surfaces early glabrescent, above flat or slightly raised; nerves 11-19 pairs, lines of interarching distinct or not; venation lax, distinct or not on both surfaces; petiole 8-30 by 1.5-4 mm; leaf bud 6-25 by 1-3 mm, with dense (grey-)brown hairs 0.1-0.2 mm long or less. Inflorescences predominantly with male flowers, generally behind the leaves, (sub)glabrescent or with sparse greyish hairs 0.1 mm or less, variable in shape, loose or condensed, many-flowered, 6-25 by 3-15 cm, peduncle 0.5-6 cm long, bracts caducous, not seen. Flowers in loose or dense clusters of 3-8, all in about the same stage of development; perianth outside with ± sparse greyish hairs 0.1 (-0.2) mm, or glabrous, lobes inside with pale hairs 0.2-0.3 mm, often in longitudinal rows. Male flowers: pedicel 1.5-2.5 mm; bud broadly ellipsoid-obovoid, 1.5-1.6 by 1.3-1.4 mm, apex broadly rounded or somewhat depressed, base ± tapering, subcircular in cross section, cleft 3/4-4/5, lobes 3 or 4 (or 5), at sutures 0.2(-0.3) mm thick; synandrium depressed globose, 0.3-0.5 by 0.5-0.6 mm, anthers 4-6, androphore 0.3- 0.4 by 0.3 mm. Female flowers not seen; ovary in very immature fruits glabrous. Fruits (solitary or) 2-6 together in panicles of 15-25 cm long, ellipsoid-oblong, (4.5-)5-7 by 2.5-3.5 cm, apex (narrowly) rounded, base rounded or narrowed for 3-5 mm, glabrous, drying dark brown or blackish, finely granulate; pericarp 2-10 mm thick; fruiting pedicel 15-20 mm long; aril laciniate for c. 1/3 to nearly halfway; seeds ellipsoid-oblong, 4-5 cm long, apex subacute or bluntly beaked for up to 3 mm long, testa purplish brownish variegated.

Field-notes Bark smooth, grey- or dark-brown, flaking in small thin pieces, or finely fissured or cracked; slash inner bark brownish white, yellowish brown, or red, with watery reddish exudate; slash wood white or pale yellow. Leaves glossy on both surfaces. Flowers (pale) green. Fruits green turning yellow (Peninsular Malaysia) or purple brown (N Sumatra), ellipsoid-oblong, very large, to 10-12 cm long, mature aril yellow.

Distribution Peninsular Thailand; Malesia: Sumatra, Peninsular Malaysia, Singapore.

Habitat & Ecology Evergreen, open bamboo forest, lowland rain forest, on flat land or hillsides; 0-300 m altitude; fl. Jan.-June; fr. June-Sept.

Note The fruits are very variable in the thickness of the pericarp. When more material becomes available, two varieties may be distinguished through this character. Fruiting specimens with thin pericarp may be difficult to distinguish from large-fruited specimens of Endocomia macrocoma subsp. prainii.

Trees 5-50 m. Twigs (pale) grey brown to dark brown, 2-6(-10) mm diameter, with grey-brown to rusty hairs 0.1-0.5 mm, early to late glabrescent, bark (coarsely) striate or longitudinally cracking, sometimes flaking, lenticels absent or inconspicuous. Leaves membranous or chartaceous, elliptic or obovate to oblong-oblanceolate, (12-) 15-35 (-40) by (4-)5-12 cm, base (narrowly) rounded to attenuate, apex acute-acuminate; upper surface olivaceous to brown, glabrous, lower surface glabrous or (early) glabrescent; midrib above slender, flat or slightly raised; nerves 11-24 pairs, lines of interarching generally indistinct; venation lax or fine, sometimes slightly trabeculate, on both surfaces distinct or not; petiole 10-25 by 1.5-3.5 mm, glabrous to late glabrescent; leaf bud 8-25 by 1.5-3.5 mm, with dense grey brown to rusty hairs 0.1-0.5 mm. Inflorescences (see note 1) slender to stout, much or little branched, glabrescent or with persistent greyish or rusty hairs 0.1-0.5 mm long, in male 6-30 by 2-25 cm, moderately to many-flowered, peduncle 0-5 cm long, bracts ± elliptic, 1.5-3 mm, ± thinly pubescent, caducous. Flowers 4-10 together, buds usually in different stages of development, in ± umbelliform clusters (3-)5-30 mm spaced along the main branches of the inflorescence; perianth outside early glabrescent or thinly with hairs 0.1-0.5 mm long, lobes inside towards the apex with few to many (pale) brownish red hairs 0.2-0.4 mm long, usually in rows in spaces between the anthers, sometimes only near the lobe sutures. Male flowers: pedicel 1.5-7 mm long; buds broadly ellipsoid or broadly ovoid, 1.5-2.3 by 1.3-2 mm, apex rounded to subacute, base rounded to attenuate, in cross section sub- circular or 3- or 4-angular, cleft 2/3-4/5, lobes 3 or 4 (or 5), at sutures 0.2-0.3(-0.5) mm thick; synandrium (depressed) globose to ellipsoid, 0.2-0.5 by 0.3-0.5 mm, anthers 3-6, androphore slender, longer to shorter than the androecium, 0.3-1 mm long. Female flowers: pedicel 2-6 mm; buds ovoid-ellipsoid, 2.2-2.6 by 1.7-2 mm, cleft 2/3-5/6, lobes 0.2-0.3 mm thick; ovary (narrowly) ovoid, 1.3-1.8 by 0.8-1.2 mm, glabrous, stigma 0.2-0.3 mm high, narrowly to broadly 2-lipped, in the latter case the lips minutely 2-5- lobulate. Fruits 3-12 in a pendent loose panicle (10-) 15-30 cm long, ovoid, (narrowly) ellipsoid, or obovoid, 1.5-4.5(-5.5) cm long, glabrous; pericarp 1-3 mm thick, blackish, finely granulate, not or sparingly warted; the aril almost entire to laciniate to about halfway; seeds usually with pointed apex, testa variegated (not always in New Guinea).

Distribution Widespread, continental Southeast Asia and Malesia; see under the subspecies.

Notes 1 The inflorescences often have mixed male and female flowers, the latter possibly most frequent towards the end of the branches; other species may have separate male and female inflorescences on the same twig.

2 The flowers in a subumbel or cluster are generally in different stages of development, as in Endocomia virella, hence open flowers are found together with closed ones and usually with much smaller, still growing flower buds.

Leaf bud and twigs rusty pubescent, hairs 0.2-0.5 mm long, twig apex sometimes early glabrescent. Inflorescences rather condensed, much branched, 6-15 cm long, the flowers densely rusty pubescent, hairs 0.2-0.5 mm long. Pedicels of male and female flowers 3-4 mm long. Male flowers: buds 3-lobed, cleft to c. 4/5, inside greenish yellow; synandrium subglobose to ellipsoid, 0.5 by 0.4-0.5 mm, anthers 4, androphore 0.4-0.6 mm long. Female flowers: ovary ± narrowly ovoid, stigma rather broadly 2-lipped and very finely lobulate. Infructescences 10-16 cm long. Fruits fusiform, ellipsoid, ± obovoid, or pear-shaped, 1.7-3.5 by 1.2-1.9 cm, apex acute to broadly rounded, base rounded or narrowed; pericarp 1-3 mm thick; fruiting pedicel 6-12 mm long; aril at apex laciniate 1/3-1/2; seeds ± ellipsoid, 1.5-2 cm long, apex ± acute, testa blotched and variegated.

Field-notes No buttresses. Outer bark 0.2-0.7 mm thick, slightly fissured or not, little peeling; inner bark 9-18 mm, pale red to pink-ochre, with some pale reddish watery exudate; sapwood whitish to yellowish tinged red, gradually passing into the darker heartwood. Perianth (inside) greenish yellow. Fruits yellow or orange, aril bright red, at apex incised to 1/3-1/2; seeds mottled brown.

Distribution Malesia: Moluccas (Talaud, Morotai, Halmahera, Bacan, Obi, Buru, Seram, Ambon); possibly Sulawesi (sterile coll.) and Philippines (NE Luzon); see notes.

Habitat & Ecology Forest with little undergrowth, on alluvial flats locally with stagnant water, or hill slopes; on porous volcanic soil, or loam soil with stones; also in disturbed forest 0-400 m altitude; ; fl. & fr. throughout the year.

Notes 1 Subsp. macrocoma is arbitrarily separated from subsp. prainii, the latter being very variable in many features, and mainly differing by a much less developed indumentum on leaf buds and inflorescences, and by its generally larger fruits. Subsp. macrocoma has a remarkable variation in fruit shapes. Its fruits are generally ± ellipsoid, but fruits from Bacan and Obi are small, 1.7-2.2 cm long and either broad-ellipsoid or obovoid; in Buru fusiform fruits 28 mm long are found.

2 Ridsdale c. s. ISU 317, from Palanan, NE Luzon, strongly deviates in its very hairy appearance, and may represent a separate taxon.

Leaf bud with hairs dull brown, 0.1 mm long or less, twig apex early glabrescent. Inflorescences lax, little to much branched, 6-30 cm long, with the flowers glabrous or sparingly pubescent, hairs grey-brown, 0.1 mm long; male and female flowers with slender pedicels (2-)4-7 mm long. Male flowers: buds 3- (or 4-)valved, cleft to 4/5-5/6, inside greenish to yellowish; synandrium (depressed) globose, 0.2-0.3 by 0.2-0.4 mm, anthers 2 or 3, androphore slender, (0.6-)0.7-l mm long. Female flowers: ovary narrowly ovoid, stigma shortly and narrowly 2-lipped. Infructescences 15-30 cm long. Fruits narrowly ovoid-ellipsoid, 3.5-4(-5.5) by 1.4-2 cm, apex blunt or subacute, base rounded or narrowed; pericarp 1-2 mm thick; fruiting pedicel 15 mm long; aril laciniate to 1/5; seeds 3 cm long, the apex slightly acute or to 2 mm beaked, testa variegated with longitudinal blotches.

Field-notes Bark of trunk greyish or chocolate brown, somewhat fissured or not and little to profusely scaling in small thin pieces; outer bark 2 mm thick, brown; inner bark 7-17 mm, cream, light brown(-red), or yellowish, with pale orange or reddish watery exudate; sapwood pale yellow or pale brown. Perianth inside yellowish or pale green. Fruits hanging from the branches, green.

Distribution Malesia: E Sumatra and Borneo (Sarawak, Sabah, E and S Kalimantan).

Habitat & Ecology Primary forest, mixed dipterocarp forest, riverside forest, on stream banks or alluvial flats, on deep clay soil, low ridges with sandy or sedimentary soil 0-1000 m altitude;fl. & fr. throughout the year.

Note Related to Endocomia virella, with similar flowers, but the latter has leaves drying distinctly greenish, and much larger fruits.

Leaf bud pubescent, hairs greyish to dull brown, 0.1-0.2 mm long or less, twig apex early glabrescent. Inflorescences condensed to lax, usually much-branched, 8-25 cm long, with the pedicels and perianths sparingly to densely short-pubescent, hairs grey(-brown), 0.1-0.2 mm long; pedicels of male and female flowers slender, 1.5-2.5 mm long. Male flowers: buds 3- or 4- (or 5-)lobed, cleft to (2/3-)3/4-5/6, inside greenish to yellowish, or red; synandrium (depressed) globose or ellipsoid, (0.3-)0.4-0.5 by 0.3- 0.5 mm, anthers (3 or) 4, or 5 or 6 (New Guinea, see note 1), androphore slender, (0.2-)0.3-0.6 mm long. Female flowers: ovary ovoid, with broad 2-lipped stigma. Infructescences 10-30 cm long. Fruits narrowly to broadly ellipsoid to ovoid, (2.2-)3- 4.5(-5.5) by (1—)1.2—2.5 cm, apex obtuse or often subacute, base rounded, or with an up to 5 mm long narrowed part; pericarp 1-2.5 mm thick; fruiting pedicel 5-15 mm long; aril almost entire (Philippines, New Guinea) or laciniate c. 1/5; seeds ellipsoid, 2-3.2 cm long, apex acute or shortly beaked, testa variegated by longitudinal markings, in New Guinea not or only indistinctly so.

Field-notes Without or with short buttresses up to 200 x 30 x 4 cm, branches often horizontally spreading, or drooping. Bark grey to blackish brown, smooth, without or with shallow fissures, shallowly irregularly peeling or not; exudate watery, colourless or pale red to brownish, once recorded as slightly milky; blaze pale brown to salmon; wood white or straw or salmon-cream. Perianth inside greenish to yellow (W Malesia), once purple (Thailand) or dark red to deep maroon (New Guinea), anthers creamy to pale yellow, ovary green with brown or blackish stigma; flowers with sweet scent. Fruits glossy green, turning yellow, in the Philippines and New Guinea orange, aril bright red.

Distribution South China (Yunnan), India (Assam, Andaman I.), Bangladesh, Burma, Indochina; Malesia: W Sumatra, W Java, Philippines, New Guinea.

Habitat & Ecology Lowland and hillside forest, riverine or swamp forest; by streams, on alluvial or clayey soil, in limestone country, or on copper-rich soil; 0-500 m altitude; fl. & fr. throughout the year, but fl. in W Malesia mainly July, Aug., in the Philippines mainly Mar.-May, in New Guinea throughout the year.

Uses The wood is used for house-building (Sepik area).

Notes 1 A variable subspecies, mainly in the indumentum, some features of the male flowers, the shape and size of the fruits, and the colour of the seeds. This variation is more or less correlated with the geography. Specimens from the Philippines differ in inflorescences and indumentum, especially that of the flowers, composed of grey-white rather long hairs 0.1-0.2 mm. Specimens from New Guinea stand apart. They are characterized by generally shorter inflorescences and infructescences, androecium with 5 or 6 anthers, perianth dark red inside, fruits orange and small, sometimes only 2.2-2.5 cm long, testa non-variegated or only faintly so. Specimens from outside New Guinea have usually only 4 anthers, perianth greenish or yellowish inside, fruits yellow (once orange in the Philippines) and the testa always variegated. In other features the New Guinea specimens agree with those from elsewhere in the range of the species.

2 In Endocomia the seeds are almost always variegated, except in E. macrocoma subsp. prainii from New Guinea, where the seeds are dull greyish brown, not variegated or only faintly so, possibly because the aril is not laciniate.

Tree 10-40 m. Twigs brown, 3.5—5(—14) mm diameter, with rusty hairs 0.5 mm long, early to late glabrescent, bark coarsely striate, tending to flake, lenticels absent or inconspicuous. Leaves membranous to thinly chartaceous, elliptic-oblong to oblong-lanceolate, 18-38 by 6-11 cm, base subcordate, broadly rounded, or short-cuneate, apex acute- acuminate; upper surface drying brown, glabrous, lower surface early glabrescent; midrib on both surfaces often late glabrescent, above rather flat to moderately raised; nerves 17-25(-30) pairs, above (partly) raised, sometimes late glabrescent, lines of interarching often distinct and regular; venation lax, distinct or faint above; petioles 8-25 by 1.5-3 mm, ± late glabrescent; leaf bud 8-18 by 2-3.5 mm, with dense rusty hairs 0.3-0.5 mm. Inflorescences (with male or mixed male and female (?) flowers) between or behind the leaves, sometimes apical, 3-5 times branched, many-flowered, 8-30 by 5-25 cm, peduncle 1.5-5 cm, with dense rusty hairs 0.5-0.7 mm long, bracts elliptic-oblong, 2-6 mm long, caducous. Flowers usually in rather dense clusters of 5-10, sometimes flowers more dispersed and only 2 or 3 together, all ± in the same stage of development; perianth outside with hairs 0.2-0.5 mm long, lobes inside with dense rusty hairs 0.2-0.3 mm. Male flowers: pedicel slender, 2-3.5 mm; buds obovoid to ± obconical, 1.5-1.8 by 1.2-1.5 mm, apex rounded to subtruncate, base ± tapering, in cross section faintly angular or subcircular, pubescent or glabrescent towards the apex, cleft 3/4-4/5, the lobes (3 or) 4 or 5, oblong, 0.2 mm thick; synandrium depressed-globose, (0.2-)0.3 by 0.5-0.6 mm, anthers 4, sessile, androphore 0.3 by 0.2-0.3 mm. Female flowers not seen. Fruits 6-18 in large panicles up to 30 cm long, ellipsoid-oblong, 4.5-6.5 by 1.8-2.5 cm, apex narrowly rounded, base subacute or often tapering into a 4-10 mm long narrowed part, glabrous, drying blackish, finely granulate, without lenticels, not warted; pericarp 1.5-3 (-4) mm thick; fruiting pedicel 8-16 mm long; aril laciniate to 1/5-1/3; seeds ellipsoid- oblong, 3.5-4.5 cm long, apex beaked for 0.2-4 mm, testa with elongate purplish and brownish dapples.

Field-notes Buttresses sometimes present, short and rounded. Bark usually blackish, brittle, smooth with superficially longitudinal cracks or with paper-thin flakes; inner bark 10-15 cm thick, pale yellow, (reddish) brown, or orange, cambium yellowish to red; sapwood pale, whitish to brown-yellow, soft; exudate slight, pale red from inner sapwood. Flowers yellow with reddish or brown-red indumentum. Fruits green to yellow, aril bright red.

Distribution Malesia: Borneo (Sarawak, Sabah, E Kalimantan).

Habitat & Ecology Primary and logged-over lowland rain forest on flat land and hill slopes or on periodically inundated ground; on leached clays, loam soils, black soil, also on sandstone and limestone; 0-400 m altitude; fl.& fr. throughout the year.

Notes 1 Characterized by the inflorescences in which all the flowers are in about the same stage of development, by the conspicuous rusty indumentum on leaf buds and inflorescences, and by the large fruits.

2Endocomia rufirachis resembles Horsfieldia motleyi, which has similarly pubescent flowers. Endocomia macrocoma subsp. macrocoma (Moluccas) may have a similar, rather conspicuous yellowish red indumentum, but differs in the one-clustered flowers in various stages of development, the shape of the androecium, and the smaller fruits.

3 In spite of a fairly large number of flowering specimens, only male flowers have been found. Some of the collections have male inflorescences with fruits separately attached to the herbarium sheet; both are apparently collected from one single tree. In the related Endocomia macrocoma, the female flowers are usually scattered in between the more numerous male ones in one inflorescence, or can be found on separate female inflorescences of the same tree.

Tree 8-20 m. Twigs pale grey-brown, 1.5-4 mm diameter, with grey hairs 0.1 mm or less, early glabrescent, bark finely irregularly striate, slightly longitudinally cracking, not flaking, lenticels absent. Leaves membranous to thinly chartaceous, elliptic(-oblong), 8-20 by 3.5-8 cm, base cuneate, apex acute-acuminate; upper surface dark olivaceous to dull green, glabrous, lower surface olivaceous to dull pale green, glabrous; midrib above slender, flat; nerves 7-12 pairs, above flat or but little raised, lines of interarching distinct or not; venation lax, generally indistinct on both surfaces; petiole 10-18 by 1.5- 2 mm; leaf bud 6-10 by 0.6-1 mm, with rather sparse greyish hairs 0.1 mm or less. Inflorescences slender and lax, 5-15 by 1.5—8(—10) cm, few-flowered, peduncle 0.5-2.5 cm, glabrous or glabrescent, hairs sparse, greyish or pale brown, less than 0.1 mm, early caducous, bracts not seen. Flowers (male) in lax umbel-like clusters of 2-6 flowers, 5-20 mm spaced along the main branches of the inflorescence, the buds in one cluster usually in different stages of development; perianth outside with sparse hairs less than 0.1 mm, soon glabrescent, lobes inside towards the apex with pale hairs 0.2-0.4 mm long in between the anthers. Male flowers: pedicel slender, 4-6 mm long; buds broadly ellipsoid or broadly ovoid, 1.8-2 by 1.5-1.6 mm, ± 3- (or 4-)angular, apex rounded to subacute, base (broadly) rounded, cleft c. 5/6, lobes 3 or 4, widely spreading or ± recurved, 0.2 mm thick; synandrium globose or depressed globose, 0.2-0.3 by 0.3-0.4 mm, anthers 3 or 4, androphore slender, 0.8-1 by 0.2 mm. Female flowers not seen. Fruits 1-4 in little-branched panicles 12-25 cm long, ellipsoid-obovoid to ellipsoid- oblong, (4.5-)5-7 by 2.5-3.5 cm, apex rounded, base subattenuate or with an up to 7 mm long narrowed part, glabrous, drying bright to dark brown, finely granulate; pericarp ± woody, 5-8 mm thick; fruiting pedicel 25-30 mm; aril laciniate to 1/4-1/3; seeds ellipsoid-oblong, 4 cm long, apex subacute, testa elongately purple-brown variegated.

Field-notes Once recorded with buttresses. Bark greenish, brown-yellow, yellowish green, or black, smooth or scaly; inner bark either reddish, orange-yellow, with clear red sap, smelling; sapwood white, soft or medium hard. Perianth greenish to yellowish, anthers yellow. Fruits green.

Distribution Malesia: Borneo (Sarawak: 4th Div.; Brunei; Sabah: Beaufort Hill).

Habitat & Ecology Primary forest on hillsides, ridges; brown or blackish soil; 0-400 m altitude; fl. Jan., May; fr. Jan., Aug., Oct.

Note Characterized by the minute indumentum, the rather small leaves drying green, the slender and lax inflorescences, the flowers which are in each cluster in different stages of development and greenish when mature, the long-stalked synandrium of only 3 or 4 anthers, and the large fruits drying brown. Specimens may resemble those of the Bornean subspecies of Endocomia macrocoma, but they dry brown and have smaller fruits.

Shrubs or trees, dioecious. Twigs often somewhat compressed, lenticellate, not flaky. Leaves rarely brittle when dry, lower surface pale, not papillose, dots absent; reticulation lax, faint. Inflorescences paniculate, branched from near the base (female more condensed); flowers in loose clusters, all in the same stage of development; basal cataphylls caducous; bracts caducous. Flowers short-pedicellate, bracteole absent. Male flowers: perianth urceolate, membranous or leathery, inside pubescent, yellow; buds ellipsoid, cleft to about 1/2 or less, lobes (2 or) 3 or 4, slightly outcurved (in female ± reflexed); androecium sessile or with short and narrow androphore, synandrium (long-)ellipsoid, with 5-12 linear anthers, dorsally connate to the central column, laterally and at apices free, the connective sometimes slenderly protruding. Female flowers: shorter and wider than male, ± ovoid; ovary broadly ovoid, pubescent, stigma small, sessile, 2-lobed, each lobe entire or 3-6-lobulate. Infructescences paniculate, several-fruited. Fruits ellipsoid (globose in G. canarica, S India), 2-3.5 cm long, hairy or glabrescent; pericarp thick- leathery; aril laciniate to near the base (as in Myristica); seeds not variegated; albumen ruminate, containing a fixed oil, starch absent; cotyledons divaricate, connate at base.

Distribution The genus has 7 species, of which one, G. canarica (King) Warb., in S India, and 6 in S Thailand and Malesia east to E New Guinea; not known from Java, the Lesser Sunda Islands, and Palawan (Philippines). Map 2 (see p. 3).

Tree 15-40 m. Twigs 3-6(-8) mm diameter, rusty woolly-tomentose with hairs 0.5-1 mm long, glabrescent, greyish, bark smooth or finely cracked, densely set with lenticels. Leaves coriaceous, elliptic to lanceolate, 18—42 by 7.5—19 cm, base short-attenuate to rounded or subcordate, apex acute(-acuminate), margin often revolute; upper surface olivaceous, often glossy, lower surface grey-brown, hairs dense, rusty, 0.5 mm long, glabrescent; midrib above flat, 1-2 mm wide, nerves (13-) 15-23 pairs, at 60-70° to the midrib, prominent below, venation indistinct on both surfaces; petiole late glabrescent, 10-20 by 3-5 mm; leaf buds 10-20 by 4-6 mm, with dense brown hairs 0.5-1 mm. Inflorescences paniculate, with hairs 1-2 mm long; in male 6-10 cm long, up to 6 cm wide, many-flowered, in female 2-5 cm long, fewer-flowered; bracts triangular, 2-4 mm long, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.2- 0.3 mm long. Male flowers: pedicel 1-3 mm long; buds ellipsoid-oblong, 4-6 by 2-3 mm, cleft 1/4 to nearly 1/2, lobes (long) triangular, slightly spreading, tube 2.5-3.5 mm long; androecium truncately ellipsoid, (sub)sessile, 1.5-2 by 0.8-1.2 mm; anthers (7-)9 or 10, subsessile, free apices ± erect, 0.2-0.3 mm long. Female flowers (immature): pedicel 1 mm long; buds coriaceous, ovoid, 4-6 mm long, lobes 3; ovary subglobose, densely pubescent, stigma sessile. Fruits 2-8 per infructescence, ellipsoid(-oblong), 2.2- 3.5 by 1.5-2.2 cm, with rusty hairs 0.3(-0.5) mm long; pericarp 3 mm thick; fruiting pedicel 3-5 mm long.

Field-notes A handsome tree, especially when in flower; crown dense or spreading; bole smooth, no buttresses. Bark brown to grey, slightly fissured, finely or thickly flaky, or scaly; slash wood white to yellow. Flowers golden yellow with a brownish tinge, with a spicy odour when crushed.

Distribution MalaysiaSumatra (Aceh, Indragiri, Jambi, Palembang, Bangka, Riau), Peninsular Malaysia (Johore), Singapore, Borneo (Sarawak, Brunei).

Habitat & Ecology Primary and degraded dryland forest, also swamp forest and on hillsides and ridges, on granite, sand and sandy loam soil; up to 250 m altitude; fl. Sept., Oct.; fr. throughout the year.

Tree 5-26 m. Twigs subterete to angular or ± ridged, 3-4(-5.5) mm diameter, hairs minute, early glabrescent, bark chocolate to greyish brown, finely longitudinally cracked with lenticels when older. Leaves chartaceous or subcoriaceous, elliptic-oblong to lanceolate, sometimes ± parallel-sided, (16-)20-29 by 6-9.5 cm, base short-attenuate to broadly rounded, apex (acute-)acuminate, margin flat; upper surface ± brown, often glossy, lower surface greyish to violaceous, with scattered appressed hairs, glabrescent; midrib above slightly sunken (grooved), narrow, 1 mm wide, nerves (11-) 13-18 pairs, at 60-70° to the midrib, below slender, distinct but not much prominent, venation indistinct at both surfaces; petiole 10-20(-25) by 2-2.5 mm; leaf buds 4-8 by 2-3 mm, sometimes with scale-like cataphylls, densely minutely appressed-pubescent. Inflorescences in male (broadly) paniculate, 3.5-4.5 cm long, up to 4 cm wide, many-flowered, with rusty hairs 0.3-0.5 mm long, in female 1.5-2 by 1 cm, fewer flowers but more dense; bracts broadly triangular, 2.5 by 3 mm, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.1-0.2 mm long. Male flowers: pedicel 2-2.5 mm long; buds ellipsoid-oblong, 2.5-2.8 by 1.5-2 mm, cleft 1/3 to nearly 2/3, lobes 3 or 4, triangular, slightly spreading, tube 1-1.8 mm long; androecium truncately ellipsoid-oblong, 1.7 by 0.6-0.7 mm, up to 0.2 mm stiped; anthers 6(-8), subsessile, free apices 0.3 mm long, erect. Female flowers: pedicel 1-2 mm long; buds narrowly obovoid, 3-3.5 by 2 mm, cleft 2/3-3/4, lobes 3 or 4, long-triangular, curved outward or reflexed, tube 0.7-1 mm long; ovary subglobose, minutely pubescent, 1.2 mm diameter, stigma sessile, minutely 2-lobed. Fruits c. 7 per infructescence, ellipsoid, 2-2.2 by 1-1.4 cm, with hairs 0.1 mm, or early glabrescent; pericarp 1.2-1.4 mm thick; fruiting pedicel 3-7 mm long.

Field-notes Bark reddish brown, nearly smooth, very fine scaly; inside hard, pale reddish brown. Flowers yellow. Fruits dark red.

Distribution Malesia: Borneo (Sarawak, Brunei, Sabah; possibly also W Kalimantan: Suzuki 9993).

Habitat & Ecology Primary lowland forest, 0-580 m altitude; fl. July-Sept.; fr. Oct.-Dec.

Tree 3-30(-45) m. Twigs faintly angular, ± ridged or lined when young, 1—2(—3.5) mm diameter, with hairs 0.1 mm, early glabrescent, bark chocolate or grey, later on grey- brown, smooth, lenticellate. Leaves thinly chartaceous to coriaceous, elliptic to lanceolate, 5-17 by 1.5—5.5(—6) cm, in E Malesia up to 27 by 8.5 cm, apex acute(-acuminate), base attenuate, margin conspicuously revolute or not, olivaceous or brown, sometimes glossy above; lower surface with sparse appressed hairs 0.1 mm, glabrescent, pale brown to grey-purplish; midrib above flat or sunken (grooved), to 1 mm wide; nerves 7-11 (-15) pairs, flat or raised below, discolorous or not; venation indistinct; petiole 6-15 (-18) by 1-2.5 mm; leaf buds slender, 5-10 by 1-2 mm, with dense appressed greyish hairs 0.1 mm. Inflorescences paniculate, with rusty hairs 0.2 mm long; in male 2.5-12 cm long, up to 8 cm wide, many-flowered, in female up to 4 by 2 cm, fewer flowered; bracts broadly triangular, 1 by 2 mm, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.1 mm long. Male flowers: pedicel 1.5-4 mm long; buds ellipsoid (-oblong), 2.5-4 by 2-3 mm, cleft 1/3-2/3, lobes 3 or 4, (long-)triangular, slightly spreading, tube 1-2.3 mm long; androecium truncately ellipsoid-oblong, 1-2.5 by 0.8-1 mm, stiped up to 0.3 mm; anthers (6 or) 7—11(—13), subsessile, free apices up to 0.5 mm long, erect or somewhat incurved. Female flowers: pedicel 1.5-3 mm long, buds ovoid or obpyriform, 2-3 by 1.5-2 mm, cleft 1/2-3/4, lobes 3 or 4, long-triangular, strongly spreading or recurved, tube short-urceolate, 0.7-1.5 mm long; ovary subglobose, 1 mm diameter, densely minutely pubescent, stigma minute, subsessile. Fruits up to 13 per infructescence, subglobose to ellipsoid-oblong, 1.8-2.8 by 1.1-1.9 cm, glabrescent (glabrous) or inconspicuously pubescent, hairs 0.1 mm long; pericarp 1 mm thick; fruiting pedicel 4-15 mm long.

Distribution A widespread species ranging from Peninsular Malaysia to the Bismarck Archipelago. Four varieties of Gymnacranthera farquhariana are recognized to accommodate its particularly complex variation. The varieties are very close to each other, and several specimens are not easily placed within a variety.

Tree 3-30 m. Leaves coriaceous; blade elliptic(-oblong), widest at or above the middle, 6—15(—17) by 3-5.5(-6) cm, usually with a conspicuously revolute margin; nerves 7-11 pairs, on the lower leaf surface distinct and discolorous or not, but always clearly raised and to be felt with the finger; petiole 8-18 by 1.5-2 mm. Fruits 1-6 per infructescence, globose to short-ellipsoid; fruiting pedicel 6-15 mm long.

Field-notes Crown dense or spreading; bole smooth, in peat swamps sometimes with buttresses or with a few stilt-roots. Bark dark brown, brittle; slash wood whitish. Flowers yellow. Fruits yellow to orange, very spicy.

Distribution Peninsular Thailand; Malesia: S Sumatra (Lampung), Peninsular Malaysia, Singapore, Borneo.

Habitat & Ecology Primary and degraded forest; mostly in peat swamp forest, occasionally on hillsides; 0-1000 m altitude; fl. mainly Apr-July; fr. Sept.-Mar.

Tree 3-30 m. Leaves chartaceous or coriaceous; blade oblong-lanceolate, widest at or below the middle, 5-13.5 by 1.5-4.5 cm, margin little or conspicuously revolute or not; nerves 6-10 pairs, on lower leaf surface not or hardly raised, usually concolorous and not to be felt with the finger; petiole 7-14 by 1-1.5 mm. Fruits l-3(-5) per infructescence, subglobose, 1.8-2.2 by 1.3-1.8 cm; fruiting pedicel 4-13 mm long.

Field-notes Crown small, narrow, dense or not; bole smooth, no buttresses. Bark brown(-grey), finely fissured, with small scales; wood white to pale brown. Flowers bright yellow. Fruits green turning golden yellow to orange, very spicy.

Distribution Malesia: throughout Sumatra, Peninsular Malaysia, Singapore, and Borneo.

Habitat & Ecology Primary and degraded forest, on dry land (hillsides and ridges) as well as in (periodically) wet places, near streams and rivers, and in kerangas; found on limestone and on sandy soils; 0-1300 m altitude; fl. mainly Mar.-Oct.; fr. July-Feb.

Note The distribution of var. eugeniifolia largely coincides with that of var. farquhariana. Specimens from Borneo here referred to var. eugeniifolia were formerly sometimes determined as Gymnacranthera contracta, a species now accepted in a much more restricted sense. Also specimens intermediate with var. zippeliana occur, especially in Sabah and Sarawak.

Tree 8-14 m. Leaves chartaceous; blade (oblong-)lanceolate, 9-21 by 3-6 cm, margin not revolute; nerves (8—>9—11 pairs, on lower leaf surface distinct, sometimes discolorous, little raised; petiole 8-15 by 1-2 mm. Fruits 1 or 2(-4) per infructescence, subglobose or short-ellipsoid, 1.8-2.3 by 1.5-1.9 cm (when large with marked ridge on the suture); fruiting pedicel 8-15 mm long.

Field-notes Bark smooth, brittle, pale grey-brown, 1 cm thick. Fruits orange; seeds banded brown and black, with but little spicy taste.

Distribution Malesia: Philippines.

Habitat & Ecology Forest on ridges as well as by rivers and lakes; up to 1400 m altitude; fl. mainly Apr-June, Aug.-Oct.; fr. Jan.-July.

Note This variety is restricted to the Philippines. It is very close to var. zippeliana from which it differs only in the fruits; in var. zippeliana the fruits are usually ellipsoid- oblong, not subglobose, and have a shorter fruiting pedicel. FB 21074 (the type of G. acuminata) and Sulit 14603, from Samar, deviate in rather small leaves with glossy upper surface.

Tree 3-30(-45) m. Leaves thickly membranous or chartaceous, sometimes subcoriaceous; blade (oblong-)lanceolate, widest at or above the middle, 7.5-27 by 3.5-8.5 cm, margin not revolute; nerves 8—11(—15) pairs, on lower leaf surface distinct or not, but always clearly raised, either pale yellowish, concolorous (W Malesia), or reddish or purplish brown, discolorous (generally in E Malesia); petiole 7-15 by 1—2.5(—3) mm. Fruits (1 or) 2-13 per infructescence, ellipsoid to oblong, rarely subglobose, (1.5-) 1.8-2.5 by 1.1-1.5 cm; fruiting pedicel 4-8 mm long.

Field-notes Crown dense, narrow; bole smooth, no buttresses. Bark brown or grey, slightly fissured and finely flaky or scaly; inner bark dark brown, 5-14 mm thick, slash wood white to yellow; heartwood yellow to brown, hard. Flowers golden yellow to brown, odourless or faintly sweet; androecium brownish; pollen whitish. Seeds dark brown.

Distribution Malesia: Peninsular Malaysia, S Philippines, and New Guinea (incl. theBismarck Archipelago), not in Java or the Lesser Sunda Islands. The most widespread taxon of Gymnacranthera.

Habitat & Ecology Variable; found in primary or in degraded and secondary forest, mostly on hillsides and ridges, in New Guinea usually on foothills and riverbanks, also near the coast; on sandstone, clay, loam, and granite rock; 0-900 m altitude, in Borneo 400-1200 m; fl. mainly Jan.-June, Aug.-Oct. (Borneo mainly July-No v.); fr. Jan.- June, Aug.-Sept. (Borneo Mar.-May, Oct.-Nov.). Locally abundant, but frequently mentioned as growing in regenerating forest.

Uses In Papua Barat (Bird's Head) the bark, together with lime, is used to prepare the skins of birds.

Notes 1Gymnacranthera farquhariana var. zippeliana is variable in leaf size. In Sulawesi, Moluccas, and New Guinea in particular, specimens with large leaves up to 27 cm long are found. The fruits also are variable in shape and size. Particularly distinctive fruits are found in de Vogel 3789, 3795, 3955, from Bacan I., which have small, almost globose fruits 1.2 cm diameter; the trees are 40-45 m high (other specimens have never been recorded as being taller than 33 m). In all other characters these specimens agree with var. zippeliana, but more material may show them to represent a separate taxon.

2 The fruits of some specimens from New Guinea are ± ellipsoid(-oblong), and are intermediate in shape with those of var. paniculata; they are placed in var. zippeliana because of their short fruiting pedicels.

3 Occasionally specimens are found with particularly brittle leaves when dry, a character not often met with in other Gymnacranthera species.

4 Some collections from New Britain have leaves drying dull with the nerves sunken above, and pale, often purplish-whitish below, but intermediate forms exist with the remainder of var. zippeliana.

Tree 5-35 m. Twigs subterete to angular or ridged, (2-)2.4-4(-5.5) mm diameter, hairs 0.1 mm or less, early glabrescent, bark chocolate, later on brown or grey, finely cracked, with many lenticels. Leaves chartaceous or coriaceous, elliptic to oblong-lanceolate, widest at or below the middle, 14-33 by (5.5-)6-13 cm, base (short-)attenuate to broadly rounded, apex (acute-)acuminate, blade coarsely undulate on drying, margin not revolute; upper surface olivaceous to brown, sometimes glossy, lower surface grey (-purple) to brownish, with scattered appressed minute hairs, glabrescent; midrib above somewhat sunken (grooved) above, 1.5 mm wide, nerves 10-18 pairs, at (35-)40-50 (-55)° to the midrib, yellowish, distinct and (very) prominent below, venation sometimes distinct at the lower surface; petiole 8-20 by 1.5-3 mm; leaf buds 5-10 by 1.5-4 mm, with dense greyish hairs 0.1 mm. Inflorescences paniculate, with (grey-)rusty ± woolly hairs 0.2-0.4 mm long; in male 4-12 cm long, up to 8 cm wide, many-flow- ered, in female 1-4 cm long, 1-3 cm wide, generally fewer flowered; bracts triangular, 3 by 3.5 mm, pubescent, caducous. Flowers grey-rusty pubescent inside and outside, hairs 0.1-0.2 mm long. Male flowers: pedicel 2-3 mm long; buds ellipsoid-oblong, 2.5-4 by 1.5-2 mm, cleft 1/3-nearly 1/2, lobes 3 (or 4), (long-)triangular, only slightly spreading, tube 1.5-2.3 mm long; androecium ± truncately ellipsoid-oblong, stiped to 0.2 mm long, 1.5-2.3 by 1 mm; anthers 6-10, subsessile, often somewhat twisted, free apices 0.3-0.5 mm long, ± erect. Female flowers: pedicel 2 mm long; buds ovoid or pyriform, 2-2.5 by 1.7-2 mm, cleft to c. 3/4, lobes (2 or) 3, strongly spreading or recurved, tube short-urceolate, 0.5-0.7 mm long; ovary subglobose, 1-1.3 mm diameter, densely minutely pubescent; stigma sessile, minutely 2-lobed. Fruits 4-25 per infructescence, ellipsoid-oblong, 1.8-2.4 by 1-1.4 cm, (early) glabrescent or short-pubescent, hairs 0.1 mm; pericarp 1 mm thick; fruiting pedicel 4-10 mm long.

Distribution S Thailand; Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo. Somewhat arbitrarily two varieties can be distinguished, of which one confined to Borneo.

Note The leaves of G. forbesii usually dry coarsely undulate, not flat as in other species.

Twigs (2-)2.5-3(-4) mm diameter. Leaves chartaceous or subcoriaceous; blade elliptic to oblong, 14-28 by (5.5—)6—13 cm; nerves on lower surface moderately prominent, 0.3-0.5 mm wide. Infructescences not or little branched at base, with 4-10 fruits.

Field-notes Crown irregular, dense; bole usually straight, not buttressed. Bark soft, grey to brown, smooth, finely fissured, or thinly flaky; the inner bark pink to red-brown, laminated, sometimes fibrous; slash wood white to pale yellow. Flowers brown-green in buds, bright yellow at anthesis; pollen whitish. Fruits brown-green turning orange.

Distribution S Thailand (Pattani); Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo.

Habitat & Ecology Primary and degraded forest; hillsides and riverbanks, alluvial forest; on sandy and limestone-derived soils; up to 600 m altitude; fl. Feb.-Apr., Aug.- Sept.; fr. May-July (-Aug.), Dec.-Jan.

Twigs (2.5-)3-4(-5.5) mm diameter, lower down 3.5-5.5 mm. Leaves coriaceous; blade elliptic-oblong to oblong-lanceolate, 16-33 by 6-12 cm; nerves on lower surface strongly prominent, conspicuous, 0.5-0.7 mm wide. Infructescences often conspicuously branched from the base, with 8-25 fruits.

Field-notes Bole without buttresses. Bark grey to brown, smooth or sometimes slightly flaky, slash wood white-orange-brown. Flowers bright yellow. Fruits brown green, orange-red when ripe.

Distribution Malesia: most of Borneo (according to Sinclair W Kalimantan, but no specimens seen).

Habitat & Ecology Primary and degraded dryland as well as wet forest, alluvial forest; on sandy(-clay) and loamy soils; up to 1250 m altitude; fl. mainly Apr., Aug- Oct.; fr. Apr., Aug.-Dec.

Note Gymnacranthera forbesii var. crassinervis usually is easily recognized and distinguished from var. forbesii and other species of Gymnacranthera by its stout twigs and leaves and usually strong orange-yellowish lateral nerves, which are very distinctly raised on the lower leaf surface. Gymnacranthera bancana also is a stout species, but leaves and young twigs are always rusty tomentose, whereas the present variety is almost glabrous.

Tree 6-20 m. Twigs terete to slightly angular, 1.5-2.5(-4) mm diameter, lower down 3-4 mm, when young rusty pubescent by woolly hairs, not very appressed, hairs woolly, to 0.5 mm long, glabrescent, greyish to brown with the bark smooth or finely longitudinally cracked, densely set with lenti- cels. Leaves chartaceous or coriaceous, (oblong-)lanceolate, 10-28(-30) by 3-7 (-9) cm, base attenuate, apex acute (-acuminate), margin not or but slightly re volute; upper surface olivaceous to brown and often glossy, lower surface brown to purplish grey, with dense ap- pressed hairs 0.2-0.3 mm long, late glabrescent; midrib above flat or slightly sunken, 1 mm wide, nerves 8-17 pairs, at 45-60° to the midrib, distinct but not prominently raised below, venation forming a coarse network ± indistinct at both surfaces; petiole 7-14 by 1.5-2 (-2.5) mm; the leaf buds 10 by 2 mm, densely appressed-pubescent. Inflorescences (broadly) paniculate, with rusty hairs 0.5 mm long; in male 4.5-7 cm long, up to 6 cm wide, many-flowered, in female (from infructescences) little or much branched, rather few- to many-flowered, 3-5 cm long; bracts broadly triangular, 2.5 mm, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.1-0.2 mm long. Male flowers: pedicel 2.5-4 mm long; buds ellipsoid-oblong, 3.5-4 by 2 mm, cleft to 1/4-1/2, lobes 3 or 4, (long-)triangular, somewhat spreading, tube 2-2.7 mm long; androecium ± truncately ellipsoid, 1.8-2.3 by 1.7 mm, stipe 0.2 mm; anthers 8-10, subsessile, the free apices 0.5 mm long, erect or curved slightly inward. Female flowers not seen. Fruits up to 25 in immature infructescences, 3-6 when mature, ellipsoid, 2.3- 3 by 2-2.2 cm, hairs persistent, rusty, 0.2 mm; pericarp 3-5 mm thick; fruiting pedicel stout, 4-8 mm long.

Field-notes Tree to 20 m, dbh to 25 cm, with red sap. Flowers dark yellow. Unripe fruits brown.

Distribution Malesia: E Central Sulawesi, east of Malili.

Habitat & Ecology Primary and degraded forest on ultrabasic (nickel-containing) soils; 200-500 m altitude; fl. Feb., July; fr. Oct.

Note Endemic to Central Sulawesi; restricted to soils derived from ultrabasic rock. Distinguished from the only other species occurring in Sulawesi, G. farquhariana var. zippeliana, by the more conspicuous indumentum of woolly hairs on the young twigs and the large fruits with thick pericarp.

Tree 10-25 m. Twigs somewhat angular or ± compressed, 2-3.5(-6.5) mm diameter, with appressed hairs 0.2 mm high, glabrescent, the bark brown or grey, smooth or finely cracked, with dense conspicuous lenticels of mixed size, even when young; at the base of each innovation a group of scars occurs (see note) Leaves chartaceous or thinly coriaceous, ovate-elliptic to oblong-lanceolate, widest at or below the middle, 10-25 by 4-9.5 cm, base short-attenuate to broadly rounded, apex acute, margin not revolute; upper surface olivaceous or brown, often glossy, lower surface grey-brown, with distinct, but not very dense appressed rusty hairs, late glabrescent; midrib above flat, narrow, 1 mm wide; nerves 11-18 pairs, at c. 45° to the midrib, distinct but not prominent below, venation on both surfaces a coarse, rather indistinct network; petiole 9-18 by 2-2.5 mm; leaf buds ± stout, 10 by 3 mm, composed of the unexpanded leaf often with in addition several scale-like cataphylls, densely appressed-pubescent. Inflorescences (broadly) paniculate, with rusty hairs 0.3-0.5 mm long; in male 3-8.5 cm long, up to 7 cm wide, many-flowered, in female to 2 by 1.5 cm, few-flowered; bracts broadly triangular, 2.5 by 3 mm, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.1(-0.2) mm long. Male flowers: pedicel 1.5-3.5 mm long; buds ellipsoid- oblong, 3-4(-5) by 2-3(-3.5) mm, cleft 1/3-1/2, lobes 3 or 4, (long-)triangular, at anthesis somewhat recurved, tube 2-2.5 mm long; androecium truncately ellipsoid, 1.5- 2.3 by 0.7-1 mm, stipe to 0.2 mm; anthers 7-10, subsessile, the free apices 0.4-0.5 mm long, ± erect. Female flowers: pedicel 1.5-2.5 mm long; buds narrowly ovoid, 2.7-3 by 2 mm, cleft 1/2-3/4, lobes (2 or) 3, long-triangular, ± recurved, tube 0.5-1.3 mm long; ovary subglobose, 1.2-1.5 mm diameter, densely minutely pubescent, stigma sessile, shallowly 2-lobed. Fruits 4-10 per infructescence, ovoid-ellipsoid, usually with truncate base, 1.8-2.2 by 1.1-1.3 cm, hairs 0.1-0.2 mm long; pericarp 0.7-1.3 mm thick; fruiting pedicel 3-7 mm long.

Field-notes Bole smooth, 10-17 m; no buttresses. Bark smooth, fissured, regularly cracked, or flaky, grey or dark red-brown; inner bark 10-15 mm thick, brown; sapwood 4 cm, whitish streaked with pale red; heartwood light brown to blackish brown. Flowers yellow; anthers (pollen) whitish yellow. Fruits green turning orange(-brown).

Distribution Malesia: Borneo (Sarawak, Brunei, Sabah, Kalimantan, incl. Nunukan).

Habitat & Ecology Primary dryland forest, kerangas, forest on ridges and low hills; on tuff, sand, and sandy loam; up to c. 1300 m altitude; fl. mainly June-Nov.; fr. July-Dec.

Note The specific epithet refers to the numerous small eye-like pale lenticels on the twigs. Gymnacranthera ocellata may also be recognized by the conspicuous and numerous scars of cataphylls at the base of each seasonal shoot, possibly related with a marked seasonal growth.

Shrubs or usually trees, dioecious. Twigs sometimes angular or with two raised lines, lenticellate, not flaky. Leaves sometimes dispersed, brittle when dry, lower surface not pale, papillose only in H. iryaghedhi, dots present or absent; reticulation lax (never forming a close raised network as in Knema). Inflorescences (on older wood in H. sabulosa) paniculate, usually branched several times; flowers numerous, all in about the same stage of development, solitary, or clustered (in H. iryaghedhi the male with flowers sessile in dense flower heads); basal cataphylls caducous; bracts small or large, caducous. Flowers small, mostly short-pedicelled, at base articulated or not, bracteole absent. Male flowers: perianth ± globose or cup-shaped, leathery, inside glabrous, yellowish (never red); buds (depressed-)globose, (transversely) ellipsoid, reniform, or clavate, laterally compressed or not, cleft to variable depths, lobes 2 or 3 (or 4), not spread at anthesis; androecium sessile or with short narrow androphore, glabrous, synandrium variously shaped (cup-shaped, globose, ellipsoid, cylindrical, or trigonous), laterally compressed or not; anthers 2-c. 25, largely or entirely connate into a central column hollowed to different depths at apex; anthers straight, curved, or inflexed into the apical cavity to variable depths. Female flowers: slightly larger than male, globose to ovoid-ellipsoid; ovary glabrous or pubescent, style absent, stigma small, deeply 2-lobed (more-lobed in H. iryaghedhi). Infructescences branched, few- to many-fruited. Fruits globose or ellipsoid, 1-8 cm long, pericarp leathery or somewhat fleshy, with or without lenticel-like tubercles, glabrous or pubescent, perianth sometimes persistent; aril entire or only shal- lowly lobed; seeds rarely globose, not variegated; albumen ruminate, with fatty oil but no starch; cotyledons connate at base.

Distribution More than 100 species, ranging from Sri Lanka through NE India to S China (Kwangsi, Hainan) and through Malesia and theCaroline Islands east to the Solomon Islands and N Australia. The genus is absent in the Lesser Sunda Islands and 8 species occur exclusively outside the Malesian area. Except for a few widely distributed species, e.g., H. amygdalina (extra-Malesian), H. glabra, H. irya, H. laevigata, H. tuberculata, most species have a limited distribution. Map 3(see p. 4).

Distinct centres of species development are New Guinea and Borneo, and to a lesser extent Sumatra and Peninsular Malaysia. Three sections have been recognized here, and they occupy largely exclusive areas. Section Horsfieldia, with only H. iryaghedhi, is confined to Sri Lanka. Section Irya (with c. 40 species) is, except for the widespread H. irya, confined to East Malesia, the Solomons and N Australia. Section Pyrrhosa occurs west of Wallace's Line. In distribution, sections Irya and Pyrrhosa overlap for a narrow area in the Philippines and Sulawesi. They are morphologically segregated mainly by a different number of perianth lobes. Some species with the aberrant number of 3 lobes occur in section Irya: H. angularis, H. olens, and H. sepikensis. In section Pyrrhosa the deviating number of 2 lobes is found in H. longiflora, H. thorelii, and H. amygdalina, partly (these three species are extra-Malesian), and H. crassifolia and H. sterilis. For a further explanation, see De Wilde ('1984; 1985).

Habitat & Ecology Trees of primary rain forest, persisting in degraded forest or sometimes in old secondary growths; sometimes in marshy forest (H. irya); stilt-roots are present in some species. Some species reach or occur in montane areas, and the wide altitudinal range contrasts with those of the other Malesian genera of the Myristicaceae.

According to Sinclair (1958) the bark of species of Peninsular Malaysia is usually reddish brown, smooth or more often striate, or rough with circular or irregular dents, sometimes flaking but mostly not. The flowers are usually waxy yellow, and often sweet scented; those of H. iryaghedhi have a particularly strong smell.

Taxonomy There is a large morphological diversity in the flowers of the genus Horsfieldia. Schematic drawings of the androecia of most species have been depicted here on pages 58-61 as Plate 1, Plate 2, Plate 3 [after W. J. de Wilde Gard. Bull. Sing. 37 2 ('1984’, 1985) 129-137].

Notes to the Keys: Besides a general key to the species (1), based on male flowering specimens, five separate regional keys (2- 6) are given, based on female flowering and fruiting specimens and with emphasis on vegetative characters and partly on distribution.

In species with a laterally compressed androecium (generally in flowers with a 2-lobed perianth) the shape and size of the androecium, as given in the keys and descriptions, always concern the outline as seen laterally.

(based on male flowering specimens)

^ Footnote *) In the species descriptions the number of thecae, twice the number of anthers, is given.

(based on female flowering and fruiting specimens)

(based on female flowering and fruiting specimens)

(based on female flowering and fruiting specimens)