Revisionary monograph
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Revisionary monograph
Flora Malesiana. Series I - Seed Plants, Volume 14. Myristicaceae
expand article infoWillem de Wilde
† NHN, Leiden, Netherlands
Open Access

Abstract

Flora Malesiana. Series I, Volume 14 (2000) iv + 1-634, by W.J.J.O. de Wilde (edited by P. F. Stevens), published by the Nationaal Herbarium Nederland, Universiteit Leiden branch, The Netherlands, under the auspices of Foundation Flora Malesiana.

ISBN 90-71236-47-1

Contains the taxonomic revision of one family, Myristicaceae, for Malesia, i.e. the area covering the countries Indonesia, Malaysia, Brunei Darussalam, Singapore, the Philippines, and Papua New Guinea.

W.J.J.O. de Wilde, Myristicaceae, pp. 1-622.

A pantropical family of trees, in Malesia represented by six genera: Endocomia (4 species), Gymnacranthera (6), Horsfieldia (97), Knema (75, only one species in New Guinea), Myristica (152, of which the majority endemic to New Guinea), and Paramyristica (1, Papua New Guinea). Altogether there are 335 species of the family in the Malesian area. Some species are of economic importance, for instance Myristica fragrans, nutmeg.

The general part consists of 28 pages and also includes paragraphs on vegetative anatomy by P. Baas & J. Koster, on palynology by R.W.J.M. van der Ham, and on phytochemistry and chemotaxonomy by R. Hegnauer.

Myristicaceae are dioecious. In addition to the general keys, mainly based on male specimens, also regional keys are given for the larger genera Horsfieldia, Knema, and Myristica, based on female (fruiting) specimens.

For each species full references, synonymy, keys to infraspecific taxa, diagnostic descriptions, field-notes, distribution, and annotations regarding relationships or differences with resembling species are presented. Genera and species are arranged alpha-betically.

This treatment is illustrated with 94 line drawings (many full-page), 6 maps, and 4 pages with colour photographs* (inserted after p. 8).

Index to scientific plant names of taxa treated in this volume (accepted names and synonyms) on pp. 623-632.

Lists of revised families in Flora Malesiana on pp. 633-634.

^ Footnote *) The grant of the Dr. Hendrik Muller's Vaderlandsch Fonds, The Hague, for the reproduction and inclusion of the colour photographs, is gratefully acknowledged.

MYRISTICACEAE

(W. J. J.O. de Wilde, Leiden, The Netherlands) 1

Myristicaceae R.Br. - Prodr. (1810) 339, 'Myristiceae'

Myristicaceae R.Br. - Warb. Monographie der Myristicaceae Nova Acta Acad. Caes. Leop.-Carol. 68 (1897) 1-680

Myristicaceae R.Br. - J. Sinclair Gard. Bull. Sing. 16 (1958) 205-470, pl. I-XIV

Myristicaceae R.Br. - Whitmore Tree Fl. Malaya 1 (1972) 315-345

Myristicaceae R.Br. - W. J. de Wilde Blumea 30 (1984) 173-196

Myristicaceae R.Br. - 39 (1994) 341-350

Myristicaceae R.Br. - Beitr. Biol. Pflanzen 66 ('1991', 1992) 95-125

Myristicaceae R.Br. - Soepadmo & Saw (eds.) Tree Fl. Sabah & Sarawak 3 (2000) 335-473

Myristicaceae R.Br. - Kühn & Kubitzki Kubitzki et al. Fam. & Genera Vascular Plants 2 (1993) 457-467, (with extensive literature references).

Myristica Gronov.

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.

DISTRIBUTION

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.

HABITAT

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').

ECOLOGY

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

TAXONOMY

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

VEGETATIVE MORPHOLOGY

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.

LEGENDS OF THE COLOUR PHOTOGRAPHS

Photographs 1-9:

Photo 1.

Gymnacranthera bancana (Miq.) J. SinclairDe Wilde & Duyfjes 20635 — Sumatra

Photo 2.

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquharianaDe Wilde & Duyfjes 20542 — Sumatra

Photo 3.

Gymnacranthera forbesii (King) Warb. var. forbesiiDe Wilde & Duyfjes 20648 — Sumatra

Photo 4.

Myristica fatua Houtt. subsp. fatuaDe Wilde & Duyfjes 21921 — Hortus Bogoriensis (origin Ambon)

Photo 5.

Knema glauca (Blume) Petermann var. glaucaDe Wilde & Duyfjes 21923 — Hortus Bogoriensis (origin Ambon)

Photo 6.

Knema kinabaluensis J. SinclairDe Wilde & Duyfjes SAN 141918— Sabah

Photo 7.

Knema sumatrana (Blume) W.J. de WildeDe Wilde & Duyfjes 19860 — Sumatra

Photo 8.

Horsfieldia wallichii (Hook. f. & Thomson) Warb. — Bark slash De Wilde & Duyfjes 20151 — Sumatra

Photo 9.

Myristica iners Blume — Stilt-roots Geesink & Santisuk 5353 — Peninsular Thailand

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

REPRODUCTIVE MORPHOLOGY

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.

Fig. 1.

Schematic representation of male inflorescences in Myristicaceae. — a: single type, of Endocomia rufirachis — b: plural type, distally branched, of Endocomia macrocoma subsp. longipes — c: ditto of Gymnacranthera forbesii var. forbesii — d-g: plural type, generally distally branched, of Horsfieldia, d: H. polyspherula, e: H. clavata, f: H. spic at a, g: H. iryaghedhi (the latter with a strongly aberrant inflorescence within the genus Horsfieldia, the position of the first basal ramifications of the single-type partial inflorescences is often not clear) — h-k: a choice of forms of single-type inflorescences in Knema, h: K. pseudolaurina, i: K. furfur ace a, y. K. celebica, k: K. tridactyla — l-o: a choice of forms of single-type inflorescences as found in Myristica: sect. Myristica: 1: M. iners, m: M. schleinitzii, n: M. fragrans; sect. Fatua: o: M. fatua — p: inflorescence of Paramyristica sepicana; note that here the single-type inflorescences, which are similar in Myristica, are distichously grouped into a short-shoot ending in a vegetative bud.

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).

Fig. 2.

Schematic drawings of the androecium of Endocomia (a), Gymnacranthera (b), Horsfieldia (c), Knema (d), Myristica (e), and Paramyristica (f).

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

FOSSILS

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

CHROMOSOMES

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

USES

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

VEGETATIVE ANATOMY

(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

PALYNOLOGY

(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

PHYTOCHEMISTRY AND CHEMOTAXONOMY

(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 + C1’ ... C9’). According to Gottlieb and Yoshida (1989) lignans and neolignans should be defined biogenetically not purely on chemical arguments. They consider lignans as C8-C8’-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 C9' 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

VERNACULAR NAMES

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

KEY TO THE GENERA

(based on male flowering specimens)

1a Inflorescence a sessile or to 3(-5) mm peduncled, short, tubercle- or worm-like protuberance, usually woody, with scars of fallen pedicels and bracts. Bracteole present 2
b Inflorescence branched, panicle-like, short- or long-peduncled, the distal parts of the branches woody and with scars, or not. Bracteole present or absent 3
2a Androecium a stalked disc with the anthers sessile, contiguous or largely free, stellately attached by their bases. Bracteole mostly small, at the base of the perianth or lower, to median on the pedicel Knema
b Androecium a stalked elongate column with the anthers completely fused dorsally, apex of the column often a sterile protuberance, or ± flat, very rarely shallowly hollowed (M. markgraviana, M. hooglandii). Bracteole embracing the perianth, at or near the apex of the pedicel Myristica
3a Bracteole present, at or towards the apex of the pedicel. [Inflorescences with scar-covered perennial distal parts present or not.] Myristica
b Bracteole absent 4
4a Synandrium variable, with the central column solid or excavated, usually considerably broader than the androphore. Perianth inside glabrous, lobes not reflexed at anthesis 5
b Synandrium elongate or ± globose, central column at apex not excavated, narrow, about as wide as or narrower than the androphore. Perianth inside hairy or papillary hairy, lobes erect or reflexed at anthesis. [Inflorescences panicle-like, usually branched from near the base and with some basal cataphyll scars present.] 6
5a Male perianth small, less than 4 mm long (6-7 mm in H. superba); androphore much shorter than synandrium, glabrous. Inflorescence with peduncle with basal cataphyll scars, not branched from base; apical vegetative bud absent Horsfieldia
b Male perianth 5-6 mm long; androphore nearly as long as synandrium, with minute hairs at base. Inflorescence a brachyblast composed of partial inflorescences of the Myristica sect. Myristica-type (see also p. 360), ending in a vegetative bud Paramyristica
6a Synandrium elongate, anthers free in the apical part. Perianth lobes erect at anthesis Gymnacranthera
b Synandrium short, (depressed) globose, anthers completely sessile, without apically free part. Perianth lobes spreading or reflexed at anthesis Endocomia

KEY TO THE GENERA

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

1a Aril divided into segments to or almost to the base 2
b Aril entire or laciniate up to halfway or less 4
2a Inflorescences or infructescences paniculate, with scars of basal cataphylls, without apical vegetative bud. Bark of twigs smooth or very finely striate. Fruits 1.5-3 cm long. [Bracteole absent. Leaves not conspicuously brittle when dry.] Gymnacranthera
b Inflorescences either 1) simple or furcate, short, (sub)sessile, or panicle-like with peduncle, without basal cataphyll scars and without terminal bud, or 2) compound, consisting of those of the foregoing type distichously arranged on short-shoots, with apical vegetative bud. Bark of twigs usually striate, longitudinally cracked, or flaking. Fruits 1.5-8 cm long 3
3a Inflorescence a compound synflorescence, with apical vegetative bud. Bracteole (female flowers not seen) absent. Fruits 5 cm long, conspicuously pubescent. Crowded linear basal cataphyll scars usually present on innovations. [Leaves brittle when dry.] Paramyristica
b Inflorescences without apical vegetative bud, rarely present (M carrii, M. hooglandii, M. markgraviana). Bracteole scar on fruiting pedicel present. Fruits of variable size, pubescent or glabrescent. Crowded basal cataphyll scars on twigs usually not present. [Leaves brittle when dry.] Myristica
4a Aril at apex convoluted or shallowly laciniate. Seed not variegated, not pointed at one end 5
b Aril coarsely incised for about the upper 1/3, or ± entire in the Philippines. Seeds usually variegated, often bluntly pointed at one end. [Bracteole absent. Stigma narrowly 2- or few-lobed. Inflorescence paniculate. Monoecious. Leaves brittle when dry, not whitish below.] Endocomia
5a Bracteole absent. Inflorescence paniculate. Stigma minutely 2-lobed (or few-lobed in H. iryaghedhi, introduced). Leaves brittle when dry, lower surface usually not whitish (papillose and whitish below only in H. iryaghedhi). Horsfieldia
b Bracteole present. Inflorescences short, wart- or worm-like, simple or forked, scar- covered, (sub)sessile. Stigma few- to many-lobed. Leaves generally not brittle when dry; lower surface usually ± whitish [venation on upper surface finely reticulate and usually clearly visible, unlike most other genera.] Knema
Map 1.

Distribution of the genus Endocomia W.J. de Wilde

Map 2.

Distribution of the genus Gymnacranthera (A. DC.) Warb.

Map 3.

Distribution of the genus Horsfieldia Willd. I = sect. Horsfieldia; II = sect. Irya, east of Wallace's Line (broken line); III = sect. Pyrrhosa, west of Wallace's Line. Distribution of H. irya is indicated by a dotted line, that of H. crassifolia by a line of asterisks. Crosses indicate the approximate areas of species with a number of perianth lobes deviating from that of section areas II and III.

Map 4.

Distribution of the genus Knema Lour.

Map 5.

Distribution of the genus Myristica Gronov. The inlet indicates the eastern part of the total distribution area, with Pacific islands and Fiji

Map 6.

Distribution of the genus Paramyristica W.J. de Wilde.

ENDOCOMIA

Endocomia W. J. de Wilde - Blumea 30 (1984) 179

Endocomia W. J. de Wilde - Beitr. Biol. Pflanzen 66 (‘1991’, 1992) 95

Endocomia W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 338

Endocomia macrocoma (Miq.) W.J. de Wilde.

Horsfieldia sect. Pyrrhosa sub sect. Papillosae Warb. - Warb. Mon. Myrist. (1897) 265

Type: Based on Horsfieldia papillosa Warb., H. prainii (King) Warb., H. canarioides (King) Warb.

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.
See: Fig. 3, Fig. 4.

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.

KEY TO THE SPECIES

1a Flowers in one cluster or semi-cluster all in about the same stage of development. Synandrium depressed-globose, slightly broader than long. Androphore short, about as long as or shorter than the synandrium. Anthers 4-6. Fruits 4.5-7 cm long. 2
b Flowers in one cluster usually in different stages of development. Synandrium globose or short-ellipsoid, about as broad as long or longer than broad. Androphore about as long as or longer than the synandrium or, in Java, sometimes shorter than the synandrium. Anthers mostly 3-6. Fruits of variable sizes 3
2a All male perianths on a plant (3- or) 4-lobed. Anthers 4-6. Leaf buds, twig apex and inflorescences with grey-brown hairs 0.1-0.2 mm long, sometimes glabrescent. Nerves above flat or but little raised. Pericarp 2-10 mm thick. E. canarioides
b Male perianths more or less evenly mixed 4- and 5-lobed. Anthers 4. Leaf buds, twig apex and inflorescences with rusty hairs 0.5 mm long, sometimes late glabrescent. Nerves raised above. Pericarp 1.5-3(-4) mm thick E. rufirachis
3a Leaves 8-20 cm long, drying greenish; nerves 7-12 pairs. Inflorescences weak and slender, rather poorly flowered. Fruits 4.5-7 cm long, pericarp 5-8 mm thick, drying brown E. virella
b Leaves larger, 15-35 cm long, usually drying dark brown; nerves 11-24 pairs. Inflorescences variable. Fruits up to 4.5(-5.5) cm long, pericarp less than 5 mm thick, drying blackish E. macrocoma

Endocomia canarioides (King) W. J. de Wilde

Endocomia canarioides (King) W.J. de Wilde - Blumea 30 (1984) 190, f. 3e-h.

Horsfieldia canarioides (King) Warb. - Mon. Myrist. (1897) 294, t. 21

Horsfieldia canarioides (King) Warb. - Gamble Mat. Fl. Malay Penins. 5 (1913) 208

Horsfieldia canarioides (King) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 55

Myristica canarioides King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 304, pl. 134

Horsfieldia macrocoma (Miq.) Warb. var. canarioides King J. Sinclair - Gard. Bull. Sing. 16 (1958) 389, f. 55

Lectotype: King's coll 10064, Peninsular Malaysia.

Myristica racemosa King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 328, pl. 173

Horsfieldia racemosa (King) Warb. - Mon. Myrist. (1897) 347

Horsfieldia racemosa (King) Warb. - Gamble Mat. Fl. Malay Penins. 5 (1912) 222

Horsfieldia racemosa (King) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 60

Type: Curtis 934, Peninsular Malaysia.

?Embelia ridleyi King & Gamble - Mat. Fl. Malay Penins. 4 (1905) 112

?Embelia ridleyi King & Gamble - J. Sinclair Gard. Bull. Sing. 15 (1956) 31

Type: Ridley 6324, Peninsular Malaysia.

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.
See: Fig. 4e-h.

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.

Endocomia macrocoma (Miq.) W. J. de Wilde

Endocomia macrocoma (Miq.) W. J. de Wilde - Blumea 30 (1984) 182

Endocomia macrocoma (Miq.) W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 339

Myristica macrocoma Miq. - Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 207

Myristica macrocoma Miq. - 2 (1865) 49, p.p. (excl. specimens from Sulawesi = Horsfieldia irya).

Horsfieldia macrocoma (Miq.) Warb. - Mon. Myrist. (1897) 299, t. 21

Horsfieldia macrocoma (Miq.) Warb. - J. Sinclair Gard. Bull. Sing. 16 (1958) 392

Horsfieldia macrocoma (Miq.) Warb. - 23 (1975) 75, p.p.

Lectotype (here designated): Teijsmann 5553, Moluccas, Halmahera.

For more references and synonyms see the subspecies.

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.

KEY TO THE SUBSPECIES

1a Synandrium (0.3-)0.4-0.5 mm long, with (3 or) 4-6 anthers, androphore 0.3-0.6 mm long. Stigma broadly 2-lipped and minutely lobulate 2
b Synandrium 0.2-0.3 mm long, with 2 or 3 anthers, androphore 0.7-1 mm. Stigma narrowly 2-lipped. [Perianth inside greenish to pale yellowish. Fruits 3-4.5 cm long.] subsp. longipes
2a Plants either early glabrescent or leaf buds, inflorescences and flowers with greyish or pale brown hairs 0.1-0.2 mm long or less. Perianth inside greenish to yellowish (continental Southeast Asia, W Malesia, Philippines) or red (New Guinea). Fruits 2.5-4 cm long subsp. prainii
b Leaf buds, inflorescences and flowers with ± conspicuous rusty hairs 0.2-0.5 mm long. Perianth inside greenish yellow. Fruits 1.7-3.5 cm long subsp. macrocoma

Endocomia macrocoma subsp. macrocoma

Endocomia macrocoma (Miq.) W. J. de Wilde subsp. macrocoma - W.J. de Wilde Blumea 30 (1984) 184, f. 2i; 3d

Endocomia macrocoma (Miq.) W. J. de Wilde subsp. macrocoma - 41 (1996) 375

Horsfieldia macrocoma (Miq.) Warb. var. macrocoma - J. Sinclair Gard. Bull. Sing. 16 (1958) 393

Myristica nesophila Miq. - Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 206, p.p.

For the syntype de Vriese s.n., (L) Bacan I. not the lectotype = Horsfieldia.

Horsfieldia leptocarpa Warb. - Mon. Myrist. (1897) 346, t. 21 (excl. Forster s.n., Sulawesi = Horsfieldia irya).

Horsfieldia leptosperma nomen, in obs. sub Horsfieldia olivaeformis Warb. - Warb. Mon. Myrist. (1897) 352

Type: de Vriese s.n., fr., Sulawesi or Burn.

Gymnacranthera ibutii Holthuis - Blumea 5 (1942) 183, f. 4

Type: Lam 2976, Talaud I.

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.
See: Fig. 3i, Fig. 4d.

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.

Endocomia macrocoma subsp. longipes W. J. de Wilde

Endocomia macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde - Blumea 30 (1984) 185, f. 2f-h

Endocomia macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 339

Type: de Vogel 888, SE Kalimantan.

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.
See: Fig. 3f-h.

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.

Fig. 3.

Inflorescences and flowers of Endocomia.- E. rufirachis (J. Sinclair) W. J. de Wilde, a. Flowering twig; flowers all in about the same stage of development and rachis of inflorescence at base without cataphylls; b. male flower in mature bud stage; c. ditto, at anthesis, note papillose-hairy inner surface of perianth lobes; d. section of male perianth showing stalked androecium; e. longitudinal section of androecium, schematic, androphore and central column drawn solid black. — E. macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde, f. Inflorescence with female and male flowers, all in different stages of development; g. male flower at anthesis; note reflexed perianth lobes and slender androphore; h. female flower at anthesis. — E. macrocoma subsp. macrocoma. i. Female flower in section [a-e: BNB For. Dept. 1716; f-h: SAN 66753, i: Atasrip 103]. — Scale bar for a, f = 2 cm; for b-e, g-i = 0.8 mm.

Endocomia macrocoma subsp. prainii (King) W. J. de Wilde

Endocomia macrocoma (Miq.) W.J. de Wilde subsp. prainii King W. J. de Wilde - Blumea 30 (1984) 187, f. 3b, c.

Myristica prainii King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 299, pl. 126

Horsfieldia prainii (King) Warb. - Mon. Myrist. (1897) 292, t. 21

Type: King's coll 417, King's coll 431; Carter s.n., Andaman.

Horsfieldia papillosa Warb. - Mon. Myrist. (1897) 291, t. 21

Myristica papillosa (Warb.) Boerl. - Handl. 3 (1900) 85

Type: male specimen, cultivated in Bogor Botanical Garden, origin unknown.

Horsfieldia merrillii Warb. - Perkins Fragm. Fl. Philipp. (1904) 49

Horsfieldia merrillii Warb. - Merr. Philipp. J. Sci., Bot. 2 (1907) 274

Horsfieldia merrillii Warb. - Enum. Philipp. Flow. PL 2 (1923) 182

Type: Merrill 2233, Mindoro, Merrill 2370, Mindoro.

Horsfieldia oblongata Merr. - Philipp. J. Sci. Bot. 13 (1918) 286

Horsfieldia oblongata Merr. - Enum. Philipp. Flow. pl. 2 (1923) 182

Horsfieldia oblongata Merr. - Markgr. Bot. Jahrb. Syst. 67 (1935) 148, (for the New Guinean specimens only).

Type: Ramos Philip, pi 1393, Luzon.

Horsfieldia trifida A.C.Sm. - J. Arnold Arbor. 22 (1941) 60

Type: Brass & Versteegh 14017, Irian Jaya.

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.
See: Fig. 4b, c.

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.

Fig. 4.

Infructescences, fruits, and seeds of Endocomia. — E. rufirachis (J. Sinclair) W. J. de Wilde. Infructescence. — E. macrocoma (Miq.) W.J. de Wilde subsp. prainii (King) W.J. de Wilde. a. Infructescence; note complete arils and pointed seeds; c. ditto; note arils faintly laciniated, incompletely covering the seeds pointed at apex. — E. macrocoma subsp. macrocoma. d. Infructescence. — E. canarioides (King) W.J. de Wilde, e. Fruit in spirit; f. ditto, opened; note deeply laciniated aril; g & h. seed; note variegated testa (a: Kostermans 9579; b: Edaño PNH 7281; c: Kostermans & Soegeng 478; d: Atasrip 103; e-g: de Wilde/Duyfjes 18877 (spirit); h: King's coll. 10064 ). — Scale bar for all = 2 cm.

Endocomia rufirachis (J. Sinclair) W.J. de Wilde

Endocomia rufirachis (J. Sinclair) W. J. de Wilde - Blumea 30 (1984) 192, f. 2 a-e, 3a

Endocomia rufirachis (J. Sinclair) W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 340

Horsfieldia macrocoma (Miq.) Warb. var. rufirachis J. Sinclair - Gard. Bull. Sing. 16 (1958) 393

Type: Wood SANA 4770, Sabah.

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.
See: Fig. 3a-e, Fig. 4a.

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.

Endocomia virella W. J. de Wilde

Endocomia virella W. J. de Wilde - Blumea 30 (1984) 194

Endocomia virella W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 342

Type: Sadau SAN 49546, Sabah.

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.

GYMNACRANTHERA

Gymnacranthera (A. DC.) Warb. - Ber. Pharm. Ges. 2 (1892) 227

Gymnacranthera (A. DC.) Warb. - Ber. Deutsch. Bot. Ges. 13 (1895) 82

Gymnacranthera (A. DC.) Warb. - Mon. Myrist. (1897) 131

Gymnacranthera (A. DC.) Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 222

Gymnacranthera (A. DC.) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 61

Gymnacranthera (A. DC.) Warb. - J. Sinclair Gard. Bull. Sing. 16 (1958) 434

Gymnacranthera (A. DC.) Warb. - 17 (1958) 96

Gymnacranthera (A. DC.) Warb. - R.T.A. Schouten Blumea 31 (1986) 451

Gymnacranthera (A. DC.) Warb. - W. J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 343

Myristica sect. Gymnacranthera A. DC. - Ann. Sc. Nat. 4 4 (1855) 31

Myristica sect. Gymnacranthera A. DC. - Prodr. 14 1 (1856) 200

Myristica sect. Gymnacranthera A. DC. - Miq. Fl. Ind. Bat. 1 2 (1858) 63

Myristica sect. Gymnacranthera A. DC. - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 304

Gymnacranthera paniculata (A. DC.) Warb.

Myristica paniculata A. DC.

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.
See: Fig. 5-7, Fig. 6, Fig. 7.

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).

KEY TO THE SPECIES

1a Male perianth 4-6 mm long; androecium shorter than the perianth tube. Young twigs and inflorescences with woolly hairs l(-2) mm long; lower leaf surface with persistent woolly hairs 0.5 mm long. Leaves 18-42 cm long, petiole 3-5 mm thick. Fruits 2.5-3.5 cm long, with hairs 0.3-0.5 mm. — Peninsular Malaysia, Sumatra, Borneo G. bancana
b Male perianth 2-5 mm; androecium as long as the perianth tube. Young twigs and inflorescences glabrescent, or with hairs less than 0.5 mm; lower leaf surface glabrescent or with appressed hairs, indumentum less than 0.5 mm high. Leaf size variable, petiole 1-3 mm diameter. Fruits 2-3 cm long, pubescent or glabrescent. 2
2a Young twigs with ± woolly hairs to 0.5 mm long. Leaves densely pubescent below. Fruits 2.3-3 by 2-2.2 cm, indumentum conspicuous, pericarp 3-5 mm thick. — Central Sulawesi G. maliliensis
b Young twigs glabrescent or variously pubescent by appressed hairs, less than 0.2 mm long. Leaves below glabrescent or with ± scattered hairs. Fruits generally smaller, glabrescent or pubescent, pericarp up to 2 mm thick 3
3a Twigs, including the apical part, conspicuously densely set with lenticels. Lower leaf surface with brownish hairs. Midrib flat above; nerves at c. 45° to the midrib in the middle of the leaf. Fruits ellipsoid-ovate with truncate base, 2 cm long, short-pubescent. — Borneo G. ocellata
b Twigs towards the apex without or with but a few lenticels. Lower leaf surface glabrescent or with scattered, inconspicuous, greyish or pale brown hairs. Nerves at more than 45° to the midrib. Fruits globose to ellipsoid-oblong, base not truncate, 1.8-2.8 cm long, glabrescent or pubescent 4
4a Twigs at apex 1-2 mm, about 10 cm lower down 2-3.5 mm diameter. Leaves 5-17 (-27) by 1.5—5.5(—8.5) cm; midrib flat or sunken above. — W & E Malesia (in E Malesia not rarely twigs thicker and leaves larger) G. farquhariana
b Twigs at apex (2-)2.5-4 mm, about 10 cm lower down (3-)3.5-5.5 mm diameter. Leaves larger, 14-33 by (5.5—)6—13 cm; midrib usually sunken 5
5a Lateral nerves on lower leaf surface distinct but little prominent, at 60-70° to the midrib in the middle of the leaf; blade drying flat. Anthers 6(-8), straight. — Borneo G. contracta
b Lateral nerves on lower leaf surface very dinstinct and prominent, at (35-)40-50 (-55)° to the midrib in the middle of the leaf; blade generally drying irregularly undulate. Anthers 6-10, sometimes twisted. — S Thailand, W Malesia. G. forbesii

Gymnacranthera bancana (Miq.) J. Sinclair

Gymnacranthera bancana (Miq.) J. Sinclair - Gard. Bull. Sing. 16 (1958) 436, f. 53, pl. XIII A

Gymnacranthera bancana (Miq.) J. Sinclair - 17 (1958) 99

Gymnacranthera bancana (Miq.) J. Sinclair - R.T.A. Schouten Blumea 31 (1986) 463, f. 3a-f. 4

Gymnacranthera bancana (Miq.) J. Sinclair - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 345

Myristica bancana Miq. - Fl. Ind. Bat. Suppl. (1861) 383

Myristica bancana Miq. - Warb. Mon. Myrist. (1897) 518

Type: Teijsmann 3279, Sumatra, Bangka.

Myristica murtonii Hook f. - Fl. Brit. India 5 (1886) 105

Myristica murtonii Hook f. - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 297, pl. 124 ter.

Gymnacranthera murtonii (Hook, f.) Warb. - Mon. Myrist. (1897) 357, t. 20

Gymnacranthera murtonii (Hook, f.) Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 223

Gymnacranthera murtonii (Hook, f.) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 61

Type: Murton 13, Singapore.

Myristica ferruginea Wall ex King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 298, pl. 125

Wallich Cat. n. 6803, (lecto) Singapore.

Myristica amplifolia Warb. - Mon. Myrist. (1897) 517

Type: Anonymous n. 16, 'Medang Simpai', Palembang, Sumatra.

Gymnacranthera murtonii (Hook, f.) Warb. var. borneensis Warb. - Mon. Myrist. (1897) 359

Myristica murtonii Hook. f. var. borneensis Warb. Boerl. - Handl. 3 (1900) 88, nom. alt.

Gymnacranthera bancana (Miq.) J. Sinclair var. borneensis Warb. J. Sinclair - Gard. Bull. Sing. 16 (1958) 439

Gymnacranthera bancana (Miq.) J. Sinclair var. borneensis Warb. J. Sinclair - 17 (1958) 100

Syntypes:Beccari 1211, Kuching, Sarawak.Beccari 3977, Kuching, Sarawak.

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.
See: Fig. 5a-f, Photo 1.

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.

Fig. 5.

Gymnacranthera bancana (Miq.) J. Sinclair, a. Habit of leafy twig; b. male inflorescence; c. male flower; d. ditto, opened, showing porportionally small androecium; e. fruits; f. fruit, opened, showing thin pericarp and deeply laciniated aril of seed. — G. ocellata R.T.A. Schouten. g. Female flower, opened, showing pubescent ovary with obliquely 2-lipped sessile stigma, each lip shallowly lobulate; h. fruits, note truncate bases [a: Sinclair SF 39502, b-d: Sinclair SF 40045; e, f: Sinclair & Kadim 10436; g: Endert 4835; h: S 39255 Ilias Paie]. — Scale bar for a, b, e, h = 2 cm; for c, d, g = 1.65 mm; for f = 1 cm

Gymnacranthera contracta Warb.

Gymnacranthera contracta Warb. - Mon. Myrist. (1897) 360, t. 20 (excl. Motley 1284 = Gymnacranthera f orb esii)

Gymnacranthera contracta Warb. - J. Sinclair Gard. Bull. Sing. 16 (1958) 439, f. 54 p.p.

Gymnacranthera contracta Warb. - 17 (1958) 100, p.p.

Gymnacranthera contracta Warb. - R.T.A. Schouten Blumea 31 (1986) 471, f. 4

Gymnacranthera contracta Warb. - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 347

Myristica contracta (Warb.) Boerl. - Handl. 3 (1900) 88

Type: Beccari 321, (male, female), Sarawak.

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.

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb.

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. - Mon. Myrist. (1897) 365, t. 20

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. - R.T. A. Schouten Blumea 31 (1986) 476, f. 7

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. - W. J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 348

Myristica farquhariana Wall ex Hook. f. & Thomson - Fl. Ind. (1855) 161, p.p.

Lectotype: Wallich Cat. no 6795, (K) Singapore.

For more references and synonyms see the varieties.

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.

KEY TO THE VARIETIES

1a Leaves oblong-lanceolate, small, 5-13.5 by 1.5-4.5 cm; nerves below not or hardly raised, i.e. usually they cannot be felt with the finger. Fruits short-ellipsoid to globose. — Sumatra, Peninsular Malaysia, Borneo var. eugeniifolia
b Leaves small or large, 6-27 by 2-8.5 cm; nerves below usually clearly visible and contrasting, usually distinctly raised and to be felt with the finger. Fruits various 2 2a. Leaves coriaceous, elliptic(-oblong), 6-15(-17) by 3-5.5(-6) cm, usually with conspicuously revolute edge. [Fruits globose to short-ellipsoid.] — Sumatra, Peninsular Malaysia, Borneo var. farquhariana
b Leaves membranous, chartaceous, or sometimes coriaceous, (elliptic-oblong to) lanceolate, 8-27 by 2-8.5 cm, margin flat, rarely revolute 3
3a Fruits globose to short-ellipsoid, fruiting pedicel 8-15 mm long. — Philippines. var. paniculata
b Fruits ellipsoid to oblong, rarely subglobose (Moluccas), fruiting pedicel 4-8 mm long. — W Malesia (nerves on lower surface of blade usually pale yellowish), E Malesia (nerves usually reddish brown and contrasting), rare in the Philippines var. zippeliana

Gymnacranthera farquhariana var. farquhariana

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquhariana - R.T.A. Schouten Blumea 31 (1986) All

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquhariana - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 349

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. - Mon. Myrist. (1897) 365, t. 20 p.p.

Myristica farquhariana Wall ex Hook. f. & Thomson - Fl. Ind. (1855) 161, p.p.

Myristica farquhariana Wall ex Hook. f. & Thomson - A. DC. Prodr. 14 1 (1856) 200, p.p.

Myristica farquhariana Wall ex Hook. f. & Thomson - Miq. Fl. Ind. Bat. 1 2 (1858) 63, p.p.

Myristica farquhariana Wall ex Hook. f. & Thomson - Hook, f. Fl. Brit. India 5 (1886) 108, p.p.

Myristica farquhariana Wall ex Hook. f. & Thomson - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 305, pl. 138

Myristica griffithii Hook f. - Fl. Brit. India 5 (1886) 109, (excl. Maingay 1306A & B = var. eugeniifolia)

Myristica griffithii Hook f. - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 304, pl. 135 (excl. Curtis 2406, 2458 = Horsfieldia penangiana).

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. griffithii Hook, f. Warb. - Mon. Myrist. (1897) 368

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. griffithii Hook, f. Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 226, p.p.

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. griffithii Hook, f. Warb. - Ridl. Fl. Malay Penins. 3 (1924) 62

Gymnacranthera eugeniifolia (A.DC.) J. Sinclair var. griffithii Hook, f. J. Sinclair - Gard. Bull. Sing. 16 (1958) 447, f. 57

Gymnacranthera eugeniifolia (A.DC.) J. Sinclair var. griffithii Hook, f. J. Sinclair - 17 (1958) 113

Lectotype: Griffith 4356, (K) Malacca.

Myristica farquhariana Wall ex Hook. f. & Thomson var. major King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 306, pl. 136, f. 4

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. major King Gamble - Mat. Fl. Malay Penins. 5 23 (1912) 226

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. major King Gamble - Ridl. Fl. Malay Penins. 3 (1924) 62

Lectotype: Griffith 4355, (K) Malacca.

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.
See: Photo 3.

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.

Gymnacranthera farquhariana var. eugeniifolia (A. DC.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. eugeniifolia A.DC. R.T.A. Schouten - Blumea 31 (1986) 480

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. eugeniifolia A.DC. R.T.A. Schouten - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 349

Myristica eugeniifolia A. DC. - Ann. Sc. Nat. 4 4 (1855) 29

Myristica eugeniifolia A. DC. - Prodr. 14 1 (1856) 190

Myristica eugeniifolia A. DC. - Miq. Fl. Ind. Bat. 1 2 (1858) 58

Myristica eugeniifolia A. DC. - Hook, f. Fl. Brit. India 5 (1886) 113

Gymnacranthera eugeniifolia (A.DC.) J. Sinclair - Gard. Bull. Sing. 16 (1958) 444, p.p.

Gymnacranthera eugeniifolia (A. DC.) J. Sinclair var. eugeniifolia J. Sinclair - Gard. Bull. Sing. 16 (1958) 444, f. 56, pl. XIV, p.p.

Gymnacranthera eugeniifolia (A. DC.) J. Sinclair var. eugeniifolia J. Sinclair - 17 (1958) 112, p.p.

Type: Gaudichaud 116, Penang.

Myristica farquhariana auct. non Hook. f. & Thomson, p.p.: Hook, f. - FL Brit. India 5 (1886) 108

Myristica farquhariana auct. non Hook. f. & Thomson, p.p.: Hook, f. - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 305

Gymnacranthera farquhariana auct. non (Hook. f. & Thomson) Warb., p.p.: Warb. - Mon. Myrist. (1897) 365

Gymnacranthera farquhariana auct. non (Hook. f. & Thomson) Warb., p.p.: Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 225

Gymnacranthera farquhariana auct. non (Hook. f. & Thomson) Warb., p.p.: Warb. - Ridl. Fl. Malay Penins. 3 (1924) 62

Myristica grijfithii auct. non Hook, f.: Hook, f. - Fl. Brit. India 5 (1886) 109, (as for syntype Maingay 1306 only).

Gymnacranthera apiculata Warb. - Mon. Myrist. (1897) 359, t. 20

Myristica apiculata (Warb.) Boerl. - Handl. 3 (1900) 88

Type: Beccari 2246, Borneo, Sarawak.

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.

Gymnacranthera farquhariana var. paniculata (A.DC.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. paniculata A.DC. R.T.A. Schouten - Blumea 31 (1986) 481

Myristica paniculata A. DC. - Ann. Sc. Nat. 4 4 (1855) 31

Myristica paniculata A. DC. - Prodr. 14 1 (1856) 200

Myristica paniculata A. DC. - Miq. Fl. Ind. Bat. 1 2 (1858) 63

Myristica paniculata A. DC. - Fern.-Vill. Nov. App. (1880) 177

Myristica paniculata A. DC. - Vidal Phan. Cuming. (1885) 139

Myristica paniculata A. DC. - Rev. pl. Vase. Filip. (1886) 221

Gymnacranthera paniculata (A. DC.) Warb. - Mon. Myrist. (1897) 370, t. 20

Gymnacranthera paniculata (A. DC.) Warb. - Merr. Philipp. J. Sci. Suppl. 1 (1906) 55

Gymnacranthera paniculata (A. DC.) Warb. - Enum. Philipp. Flow. PL 2 (1923) 181

Gymnacranthera paniculata (A. DC.) Warb. - Elmer Leafl. Philipp. Bot. 3 (1911) 1059

Gymnacranthera paniculata (A. DC.) Warb. - J. Sinclair Gard. Bull. Sing. 17 (1958) 104

Gymnacranthera paniculata (A. DC.) Warb. wax. paniculata - J. Sinclair Gard. Bull. Sing. 17 (1958) 104, f. 2

Type: Cuming 901, Luzon.

Myristica farquhariana auct. non Hook. f. & Thomson, p.p.: Hook, f. - Fl. Brit. India 5 (1886) 108, (for the Philippine specimen).

Gymnacranthera laxa Elmer - Leafl. Philipp. Bot. 8 (1915) 2772

Type: Elmer 13715, Philippines, Mindanao.

Gymnacranthera acuminata Merr. - Philipp. J. Sci. Bot. 12 (1917) 265

Gymnacranthera acuminata Merr. - Enum. Philipp. Flow. pl. 2 (1923) 181

Type: Cenabre & Cortes FB 21074, Philippines, Samar.

Gymnacranthera macrobotrys Merr. - Philipp. J. Sci. Bot. 13 (1918) 284

Gymnacranthera macrobotrys Merr. - Enum. Philipp. Flow. PL 2 (1923) 181

Type: Ramos BS 1171, Philippines, Leyte.

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.

Gymnacranthera farquhariana var. zippeliana (Miq.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. zippeliana Miq. R.T.A. Schouten - Blumea 31 (1986) 482

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. zippeliana Miq. R.T.A. Schouten - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 349

Myristica zippeliana Miq. - Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 50

Myristica zippeliana Miq. - Scheff. Ann. Jard. Bot. Buitenzorg 1 (1876) 45

Gymnacranthera zippeliana (Miq.) Warb. - Mon. Myrist. (1897) 373

Gymnacranthera paniculata (A. DC.) Warb. var. zippeliana Miq. J. Sinclair - Gard. Bull. Sing. 17 (1958) 108, f. 3

Type: Zippelius s.n., Irian Jaya, Bird's Head.

Gymnacranthera suluensis Warb. - Mon. Myrist. (1897) 373

Gymnacranthera suluensis Warb. - Elmer Leafl. Philipp. Bot. 3 (1911) 1058

Gymnacranthera suluensis Warb. - Merr. Enum. Philipp. Flow. PL 2 (1923) 181

Syntypes: Vidal 3546, Philippines, Sulu I., Basilan, Vidal 3561, Philippines, Sulu I., Basilan.

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.

Gymnacranthera forbesii (King) Warb.

Gymnacranthera forbesii (King) Warb. - Mon. Myrist. (1897) 363, t. 20

Gymnacranthera forbesii (King) Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 224

Gymnacranthera forbesii (King) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 61

Gymnacranthera forbesii (King) Warb. - J. Sinclair Gard. Bull. Sing. 16 (1958) 441, f. 55, pl. XIII B

Gymnacranthera forbesii (King) Warb. - 17 (1958) 101, f. 1A, C

Gymnacranthera forbesii (King) Warb. - R.T.A. Schouten Blumea 31 (1986) 472, 474

Gymnacranthera forbesii (King) Warb. - W. J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 350

Myristica forbesii King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 306, t. 137

Lectotype: Forbes 2976, (female fl.) (K), Sumatra, Benkulu .

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.
See: Photo 2.

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.

KEY TO THE VARIETIES

1a Leaves chartaceous to coriaceous; nerves on lower surface moderately prominent, 0.3-0.5 mm wide. Infructescences usually not branched at the base, few-fruited. — Peninsular Malaysia, Sumatra, Borneo var. forbesii
b Leaves (very) coriaceous; nerves on lower surface more prominent, 0.5-0.7 mm wide. Infructescences conspicuously branched from the base, many-fruited. — Borneo var. crassinervis

Gymnacranthera forbesii var. forbesii

Gymnacranthera forbesii (King) Warb. war. forbesii - J. Sinclair Gard. Bull. Sing. 17 (1958) 101, f. 1A,C p.p.

Gymnacranthera forbesii (King) Warb. war. forbesii - R.T.A. Schouten Blumea 31 (1986) 474

Gymnacranthera forbesii (King) Warb. war. forbesii - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 350

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.

Gymnacranthera forbesii var. crassinervis (Warb.) J. Sinclair

Gymnacranthera forbesii (King) Warb. var. crassinervis Warb. J. Sinclair - Gard. Bull. Sing. 17 (1958) 102, f. IB

Gymnacranthera forbesii (King) Warb. var. crassinervis Warb. J. Sinclair - R.T.A. Schouten Blumea 31 (1986) 475

Gymnacranthera forbesii (King) Warb. var. crassinervis Warb. J. Sinclair - W. J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 350

Gymnacranthera crassinervis Warb. - Mon. Myrist. (1897) 362, t. 20

Myristica crassinervis (Warb.) Boerl. - Handl. 3 (1900) 88

Lectotype: Beccari 1119, (K) Sarawak.

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.

Gymnacranthera maliliensis R.T.A. Schouten

Gymnacranthera maliliensis R.T.A. Schouten - Blumea 31 (1986) 467, f. 5

Type: van Balgooy 3960, eastern Central Sulawesi.

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.
See: Fig. 6, Fig. 7.

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.

Fig. 6.

Schematic drawing, in longitudinal and transverse section, of the androecium of Gymnacranthera maliliensis R.T. A. Schouten; note the minute appendix of the connectivum; the broken line indicates the depth of the incision between the anthers. — Scale bar = 1 mm.

Gymnacranthera ocellata R.T. A. Schouten

Gymnacranthera ocellata R.T.A. Schouten - Blumea 31 (1986) 469, 3g, h

Gymnacranthera ocellata R.T.A. Schouten - W.J. de Wilde Tree Fl. Sabah & Sarawak 3 (2000) 351

Type: Saikeh SAN 72177, Sabah.

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.
See: Fig. 5g, h.

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.

Fig. 7.

Gymnacranthera maliliensis R.T.A. Schouten. a. Habit of leafy twig with immature fruits; b. twig with male inflorescence; c. male flower; d. ditto, opened, showing androecium; e. infructescence; f. fruit, opened, showing thick pericarp and deeply laciniated aril of seed [a: van Balgooy 3884; b-d: van Balgooy 3960; e, f: bb Cel./V 255]. — Scale bar for a, b, e = 2 cm; for c, d = 1.65 mm; for f = 1 cm.

HORSFIELDIA

Horsfieldia Willd. - Sp. PL 4 (1806) 872, [non Blume = Harmsiopanax Warb. (Araliaceae)]

Horsfieldia Willd. - Pers. Symb. 2 (1807) 635

Horsfieldia Willd. - Warb. Mon. Myrist. (1897) 130,262

Horsfieldia Willd. - J. Sinclair Gard. Bull. Sing. 16 (1958) 368

Horsfieldia Willd. - 27 (1974) 133-141

Horsfieldia Willd. - 28 (1975) 1-181

Horsfieldia Willd. - W. J. de Wilde Gard. Bull. Sing. 37, 2 ('1984', 1985) 115-179

Horsfieldia Willd. - 38 1 (1985) 55-144

Horsfieldia Willd. - 38 2 ('1985’, 1986) 185-225

Horsfieldia Willd. - 39 1 (1986) 1-65

Horsfieldia Willd. - Blumea 32 (1987) 459-472

Horsfieldia Willd. - 41 (1996) 375-381

Horsfieldia Willd. - Tree Fl. Sabah & Sarawak 3 (2000) 352

Pyrrhosa Endl. - Gen. Pl. (1839) 830, nom. illeg.

Horsfieldia iryaghedhi (Gaertn.) Warb.

Horsfieldia odorata Willd.

Myristica sect. Pyrrhosa Blume - Rumphia 1 (1837) 190-192, t. 62-64

Horsfieldia glabra (Blume) Warb.

Lectotype species: Myristica glabra Blume

Subsequent authors treated the genus Horsfieldia as defined at present partly under Myristica sect. Pyrrhosa as well as under several other sections of Myristica, e.g., sections Caloneura p.p., Eumyristica p.p., Horsfieldia, Irya (see Sinclair, 1958: 368), and under the here accepted and discussed sections.

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.
See: Fig. 8, Fig. 9, Fig. 10, Fig. 11, Fig. 12, Fig. 13, Fig. 14, Fig. 15, Fig. 16, Fig. 17, Fig. 18, Fig. 19, Fig. 20, Fig. 21, Fig. 22, Fig. 23, Fig. 24, Fig. 25, Fig. 26, Fig. 27, Fig. 28, Fig. 29, Fig. 30, Fig. 31, Fig. 32, Fig. 33.

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.

(1) GENERAL KEY TO THE SPECIES

(based on male flowering specimens)

1a Leaves papillose beneath. Male flowers sessile, packed into dense subglobose capi- tula; buds ± obconical, mutually appressed, angular. Perianth 3-lobed, cleft 1/5-2/3. Androecium stalked, anthers * 3-5. — Sri Lanka, elsewhere cultivated H. iryaghedhi
b Leaves not papillose beneath. Male flowers subsessile or usually pedicelled, mutually free or at least not densely clustered; buds variable, not or only somewhat angular. Perianth densely clustered before anthesis in H. sylvestris from E Malesia. 2
2a Leaves in fertile twigs distichous, membranous, usually with whitish marks of irregular shape and size. Perianths globose, 2-lobed, l-1.5(-2) mm diam. Androecium not or hardly laterally compressed; anthers 6-10, for the larger part connate, forming a shallow or deep saucer-shaped column, free apices 0.3 mm, ± incurved; androphore distinct, tapering. — Sri Lanka to Solomon Is. (throughout Malesia), generally coastal-riverine H. irya
b Leaves in fertile twigs distichous (alternate), or dispersed (spirally), or mixed in the same specimen. Leaf of variable consistency, usually without whitish marks. Perianths 2-4-lobed, variable in shape and size. Androecium various; anthers few to many, connate or mutually for some distance free, column variable; if column deeply cup- or saucer-shaped, then anthers at apex free for at least halfway, or deeply inflexed into the cup; androphore various 3
3a Perianth 2-lobed, or sometimes a few flowers in one inflorescence 3- or 4-lobed. — Species mainly from E of Wallace's Line, or the following from W Malesia: H. cras- sifolia (Sumatra, Peninsular Malaysia, Singapore, Borneo), H. penangiana, p.p. (Sumatra), H. sterilis (Borneo), H. sucosa subsp. bifissa (Borneo) 4
b Perianth 3- or 4-lobed, or perianth with both 3 and 4 lobes present; sometimes the odd flower in an inflorescence with 2-lobed perianth. — Species from W Malesia, and the following from New Guinea: H. angularis, H. olens, H. sepikensis. 43
4a Androecium not laterally compressed (in cross section ± circular), not longer than wide. — W Malesia 5
b Androecium laterally compressed or not; if not or only slightly so then the androecium (including androphore) longer than wide; androecium not or little laterally compressed in odd 3- or 4-lobed flowers. — E Malesia 8
5a Leaves coriaceous, lower surface with inconspicuous, subpersistent, dense indumentum beneath (in Borneo the hairs often deciduous) [dark dots and dashes present.] — Swamp forest on peat or sand H. crassifolia
b Leaves ± membranous, glabrous or early glabrescent beneath. — Not from peat swamp forest 6
6a Twigs 2 mm diam.; bark not pale on drying. Leaves 7-12 cm long, with dots beneath. [Lateral nerves flat and inconspicuous above.] H. penangiana
b Twigs 2-5(-10) mm diam.; bark pale on drying, often contrasting with the blackish dried petioles. Leaves 14 cm long or more, without dots beneath 7
7a Androecium depressed-globose, largely consisting of anthers; apical hollow flat and shallow H. sucosa subsp. bifissa
b Androecium broadly obovoid, consisting of a large sterile basal part at apex with 3 or 4 small anthers (i.e., c. 6 thecae) H. sterilis
8a Inflorescences spike-like, unbranched or short-branched, lateral branches to 5 (-10) mm long. Inflorescences, flowers and petioles blackish on drying, usually contrasting with the paler grey-brown twigs. Anthers inflexed. — Moluccas H. spicata
b Inflorescences branched, the side branches at least 5 mm long. Inflorescences, flowers, and petioles brown, not contrasting with the colour of the twigs 9
9a Androecium cup- or saucer-shaped, moderately laterally compressed; anthers at one or both sides of the androecium distally distinctly incurved into the central hollow. — E Malesia: Philippines, Sulawesi, Moluccas, NW New Guinea. 10
b Androecium laterally flattened or not, the column either 1) solid, 2) broadly but shallowly hollowed only up to c. 1/3, or 3) narrowly slit-like channelled to variable depths; anthers straight or only slightly incurved, never inflexed into the central hollow. — E Malesia: Moluccas, whole of New Guinea 17
10a Leaves with dots beneath (lens!). Twigs angular or ridged H. inflexa
b Leaves without dots. Twigs terete, angular, or winged 11
11a Perianth together with the pedicel ± pear-shaped, i.e., pedicel tapering. Petiole comparatively long, 1-2.6 cm. Twigs terete, not angular. [Leaf blade (6-)8-22(-25) cm.] H. moluccana
b Perianth (in lateral view) short-cuneate, rounded, or subtruncate at base; pedicel ± abruptly passing into the perianth, not tapered. Petioles comparatively shorter. Twigs terete or angular 12
12a Perianth cleft about halfway. Anthers connate; androphore short or absent. 13
b Perianth cleft to ± 2/3 or more. Anthers mutually free at least in the incurved or inflexed part 14
13a Anthers 18-25, inflexed at both sides into thin-walled laterally compressed androecium cup. Perianth 2.5-4 mm wide. Leaves membranous, matt on drying. — Moluccas H. parviflora
b Anthers 11-12, inflexed at only one side of the androecium cup; cup thick- and firm-walled. Perianth 2-2.2 mm wide. Leaves chartaceous, ± glossy above. — Philippines (Luzon) H. obscurinervia
14a Anthers free only in the inflexed distal parts, the basal parts connate into a cup- shaped column. Androphore minute, only c. 1/10 of the androecium length. 15
b Anthers free for at least 2/3. Androphore longer, c. 1/3 of the androecium. [Twigs generally angular or ridged. Perianth 2-3 mm wide; pedicel glabrous. Anthers all inflexed into the centre of the androecium.] — Moluccas H. smithii
15a Twigs angular or winged. Perianth 4 mm wide, glabrous. — Philippines H. ardisiifolia
b Twigs terete. Perianth 2.5-3 mm wide 16
16a Pedicel pubescent, shorter than the perianth. Inflorescences densely finely pubescent. Anthers inflexed at both sides of the laterally compressed androecium. — Moluccas (Talaud I.) H. talaudensis
b Pedicel glabrous, longer than the perianth. Inflorescences with sparse hairs less than 0.1 mm long. Anthers inflexed at one side only of the androecium. — Philippines (Samar) H. samarensis
17a Flower buds± angular, arranged into dense semi-globose clusters. Androecium much longer than wide, not or slightly laterally compressed. Leaves lanceolate(-linear), usually ± parallel-sided H. sylvestris
b Buds not angular, not densely clustered 18
18a Twigs angular, ridged, or winged, at the apex as well as lower down 19
b Twigs not winged, i.e., terete or only somewhat angular or lined at apex. 21
19a Perianth 2-, 3-, or 4-lobed, subspherical, hardly or not laterally compressed, ± glossy, not collapsing on drying. — Papua Barat (Bird's Head) H. angularis
b Male perianth predominantly 2-lobed, little or much laterally compressed, matt on drying, slightly or strongly collapsing on drying 20
20a Leaves thinly coriaceous. Pedicel about as long as or longer than the perianth. Buds cleft almost to the base. Hairs of inflorescences and pedicel 0.2-0.3 mm. Anthers 10-14; the column hollow for c. 1/4 H. iriana
b Leaves membranous. Pedicel shorter than the perianth. Buds cleft to 2/3-3/4. Inflorescences and pedicel almost glabrous, hairs 0.1 mm or less. Anthers (12-) 14-18; column solid or almost so H. aruana
21a Inflorescences 25-35 cm long. Buds ± pear-shaped. Androecium longer than broad; androphore 0.5 mm long or more, about half as long as the anthers or longer 22
b Inflorescences 20 cm long or less. Buds of variable shapes. Androecium longer or shorter than broad; androphore short or long 23
22a Buds glabrous (?), 4 by 2 mm. Anthers 10, androphore nearly as long as the anthers. Inflorescences to 25 cm long, glabrescent H. ampla
b Buds pubescent, 3 by 3 mm. Anthers 7, androphore about half as long as the anthers. Inflorescences 25-35 cm long, pubescent H. ampliformis
23a Inflorescences delicate, 2-5(-8) cm long, 1 or 2 (or 3) times branched. — New Guinea, including Aru Is 24
b Inflorescences stouter, 5-20 cm long, not or 1-4 times branched (inflorescences of H. sinclairii and H. basifissa from New Guinea sometimes small). — Whole of E Malesia, including Sulawesi; not in the Philippines 30
24a Buds pubescent or late glabrescent, distinctly or only somewhat longer than broad. Anther-bearing part of the androecium much shorter than the elongate club-shaped androphore 25
b Buds glabrous or early glabrescent, about as long as or shorter than broad. Androphore much shorter than the anthers 29
25a Buds together with tapering pedicel long pear-shaped. Androphore glabrous. 26
b Buds globose or ellipsoid, distinctly marked-off from the slender pedicel. Androphore glabrous or pubescent 27
26a Pedicel and bud together 10-12 mm long H. crux-melitensis
b Pedicel and bud together 5 mm long H. clavata
27a Buds subglobose or ellipsoid in outline, lobes 0.5-0.8 mm thick. Twig apex, leaf bud, and inflorescences with hairs 0.1-0.2 mm 28
b Buds subglobose in outline, lobes 1-1.5 mm thick. Hairs 0.3(-0.4) mm. [Androphore glabrous. Pedicel 3 mm long.] H. urceolata
28a Androphore (at least in the lower half) densely pubescent. Pedicel 3-3.5 mm long. Buds cleft c. 1/8 only H. squamulosa
b Androphore glabrous or with a few scattered hairs. Pedicel (3-)4-6.5 mm long. Buds cleft (1/6—)l/4 H. coryandra
29a Buds ± laterally compressed, ± obtriangular in lateral view, 1.8-3 mm wide, usually ± collapsing on drying. Androphore 0.2-0.5 mm long, shorter than the anthers H. subtilis
b Buds subglobose, 1-2 mm wide, not or but slightly compressed, wrinkled on drying but not collapsing. Androphore 0.4-0.5 mm long, about half the length of the anthers H. schlechteri
30a Bud and pedicel together ± pear-shaped; apex broadly rounded in lateral view, the lower (1/4-) 1/3-1/2 tapered and gradually passing into the ± tapered pedicel (these characters not always clear in certain specimens of H. tuberculata). Buds glabrous, pubescent, or glabrescent 31
b Buds in lateral view circular, ovate, obovate, elliptic, transversely elliptic, or reni- form; at base short-attenuate, rounded, or truncate, not tapered; pedicel ± slender. Buds hairy (at least at base) or in H. basifissa, H. psilantha, and H. sinclairii glabrous or glabrescent 35
31a Buds glabrous. — A variable species H. tuberculata
b Buds minutely pubescent, or in H. corrugata (at 1200-1900 m altitude) early glabrescent 32
32a Leaves ± lanceolate, 5-16 cm long. Buds cleft only c. 1/6. — Sulawesi H. lancifolia
b Leaves elliptic to lanceolate, 12-30 cm long. Buds cleft about halfway 33
33a Pedicel 1.5-2 mm long; buds 2.3 mm long, thinly pubescent. Anthers 6. — Moluccas; at low altitude H. decalvata
b Pedicel generally longer; buds 2.5-3.5 mm long. [Vegetatively and sometimes in fruit much resembling H. laevigata.] — New Guinea; at 450-2000 m 34
34a Pedicel stoutish, 2-4 mm long. Buds ± membranous, glabrescent, with or without a few scattered blackish warts. Anthers 8-12 H. corrugata
b Pedicel stout or slender, 2-5 mm long. Buds ± fleshy, pubescent, blackish warts absent. Anthers 5-10 H. pachycarpa
35a Inflorescences pubescent to nearly glabrous. Leaf bud and twig at apex with rusty or greyish hairs, 0.1-0.4(-0.5) mm long. Leaves ± glabrescent, or with scattered stellate hairs beneath when younger 36
b Inflorescences generally thick-woolly tomentose. Leaf bud and twig at apex with conspicuous, coarse, rusty hairs, (0.3-)0.5-1.5 mm. Leaves with (sub)persistent indumentum, at least on and near the midrib beneath 40
36a Buds ± glabrous, subglobose, 2-3 mm diam., cleft to the base, not collapsing on drying H. basifissa
b Buds glabrous or hairy, size and shape variable, cleft c. 1/2 to near the base; collapsing on drying or not 37
37a Buds (almost) wholly with ± persistent indumentum, hairs may be very minute and scattered. [Leaves olivaceous-brown, without a reddish tinge. Hairs of leaf bud 0.1-0.2 mm, usually greyish.] —Moluccas, New Guinea, incl. New Britain. 38
b Buds (and pedicel) glabrous or glabrescent, at least the upper 4/5. — E Papua New Guinea 39
38a Buds 1.2-1.9 mm diam. — A variable species H. pilifera
b Buds 2-3.3 mm diam. — A variable species H. laevigata
39a Buds (2-)2.5-3.5(-4) mm diam., cleft 1/2-2/3. Leaves 20-40 cm long, olivaceous or brown, not reddish tinged. — Papua New Guinea (Bagabag I., Long I., New Britain, New Ireland) H. psilantha
b Buds (1-) 1.5-2 mm diam., cleft c. 1/2. Leaves 6-20 cm long, generally with a reddish tinge on drying, especially the midrib and nerves. [Hairs of leaf bud to 0.1 mm long, greyish brown. Buds larger and pedicel pubescent in deviating specimens.] — E Papua New Guinea H. sinclairii
40a Buds largely pubescent, towards the base thick-walled and coriaceous, the remainder collapsing on drying; both male and female buds opening at apex by small pore-like slit less than 1 mm long. Androecium subellipsoid, mainly consisting of the column with 2 minute anthers at the apex, just below the pore. Leaves coriaceous, ± bullate; with harsh hairs leaving rough thickened bases. H. pulverulenta
b Buds glabrous or pubescent, membranous or chartaceous, ± not collapsing on drying, cleft at least c. 1/3. Androecium mainly consisting of 10-16 sessile anthers. Hairs not harsh, not leaving rough thickened bases 41
41a Buds pubescent, cleft 3/4-5/6. Anthers 10-14. Leaves membranous or chartaceous H. leptantha
b Buds glabrous, except at the very base; cleft c. 1/2 or less. Leaves generally membranous 42
42a Buds subglobose. Anthers 12-16. Leaves oblong(-lanceolate), at apex (acute-)acuminate, not caudate (always?) H. hellwigii
b Buds obovoid or ellipsoid. Anthers 10 (-12). Leaves oblong-lanceolate, at apex caudate H. ralunensis
43a Leaves in fertile shoots dispersed, i.e. in 3 or more rows along the twigs. Leaf bud proportionally short and broad (about 4 times longer than broad, or less). 44
b Leaves in fertile shoots distichous (in rare cases a few on the same plant in 3 rows). Leaf bud generally more slender 52
44a Leaves ± clustered towards the end of the twigs. Leaf bud and inflorescences with hairs 0.5-1 mm long. Bark of older twigs often blackish, flaking. — Borneo (Sarawak, Brunei); sandy soils H. sabulosa
b Leaves clustered or not. Leaf bud and inflorescences with hairs up to 0.2 mm long. Bark of older twigs not or slightly flaking 45
45a Leaves with dots beneath (lens!) 46
b Leaves without dots 47
46a Bark of twigs (grey-)brown, not contrasting with the dark colour of the petioles; older bark not flaking. Leaves in 2 or 3 rows. — W, C & S Sumatra, Java H. glabra
b Bark pale, grey or yellowish brown, rather contrasting with the petioles; older bark more or less flaking. Leaves in 3-5 rows. — Sumatra (N Aceh); at c. 1300 m. H. atjehensis
47a Bark of twigs brown, not contrasting with the colour of the petioles 48
b Bark of twigs pale, greyish or straw, contrasting with the blackish brown colour of the petioles 50
48a Twigs ridged or short-winged. — Sumatra H. hirtiflora
b Twigs terete, neither ridged nor winged 49
49a Leaf pubescent beneath. — Sumatra, Peninsular Malaysia H. superba
b Leaf glabrous beneath. — Borneo H. fragillima
50a Pedicel articulated. Androecium with depressed apex, apical hollow broad, either shallow or rather deep, reaching up to nearly halfway the column; androphore largely hidden by the anthers. Leaves mosly distichous, sometimes tristichous. — Sumatra, Peninsular Malaysia, H. sucosa subsp. sucosa
b Pedicel mostly not articulated. Androecium not or but slightly depressed, the apical hollow narrow and inconspicuous. Leaves generally in 3-5 rows 51
51a Androphore ± absent. Leaves blackish brown. — Borneo. H. pallidicaula
b Androphore 0.3-0.4 mm long. Leaves bright brown. — Peninsular Thailand, Sumatra, Peninsular Malaysia H. sparsa
52a Buds ± obconical-obovoid, very leathery, cleft 1/6-1/5 (-1/4) only. Androecium turbinate, anthers 3. — Sumatra H. triandra
b Buds of variable shapes, cleft l/5(—1/4) or more. Androecium various, anthers 4 or more (male flowers not known in H. perangusta) 53
53a Buds ellipsoid or obovoid, 3-8 mm long, cleft 1/5—1/4(—1/3). Leaves ± parchmentlike, matt above with finely wrinkled surface (in H. sessilifolia male flowers not known and leaves not so distinctly matt) 54
b Buds either 1) ellipsoid or short pear-shaped, 3.5 mm long or less (though sometimes rather large in H. endertii, H. flocculosa, H. majuscula, and H. wallichii), or 2) (depressed) globose; cleft (1 /4—) 1/3—1/2 or more. Texture of leaves variable including coriaceous, never parchment-like and not typically matt above 58
54a Twigs 5-7(-10) mm diam. Leaves 25-70 cm long, indumentum persistent beneath 57
b Twigs 3-5 mm diam. Leaves up to 30 cm long, (largely) glabrous beneath, or with some hairs persistent on the midrib. [Male perianth 3-5 mm long.] 55
55a Apical leaf bud with hairs 0.1-0.3 mm. Leaf midrib broad, margin flat 56
b Leaf bud with hairs 1 mm long. Leaf midrib above line-shaped, margin (dry) rolled-in H. perangusta
56a Leaves to 32 cm long, dark olivaceous. Twigs (yellowish) brown, coarsely striate and tending to crack longitudinally. Inflorescences (almost) glabrous. Pedicel not articulated. Anthers 12-20 H. tristis
b Leaves up to 24 cm long, fulvous-brown. Twigs brown, finely striate, not cracking. Inflorescences pubescent. Pedicel articulated. Anthers 10-12 H. fulva
57a Buds 7-8 mm long, perianth and pedicel glabrous. Petiole 6-15 mm long. — Sumatra, Peninsular Malaysia H. superba
b Male buds not known; female perianth and pedicel pubescent. Petiole almost absent. — Borneo (Sarawak) H. sessilifolia
58a Leaves with (sub)persistent indumentum beneath; in H. gracilis and H. wallichii sometimes hairs vestigial on and near midrib and nerves 59
b Leaves glabrous or early glabrescent beneath; in some species often with vestigial hairs on the lower midrib 69
59a Leaves not scabrous above 60
b Leaves scabrous above H. grandis
60a Leaves with dots and/or dashes beneath, obscured by hairs or not (lens!). 61
b Leaves without dots or dashes beneath 63
61a Buds short pear-shaped, 2-2.5 mm long; pedicel indistinct, 0.3-1 mm long. Leaves thinly pubescent beneath, often ± glabrescent H. wallichii
b Buds subglobose, 1-1.5 mm diam., pedicel slender, 0.5-1.5 mm long. Leaves with conspicuous indumentum beneath 62
62a Twigs 2-3 mm diam.; leaf blades 7-15 cm long, [lateral nerves 5-9 per side]. — Borneo (Sarawak); sandy soils H. paucinervis
b Twigs 5-8(-14) mm diam.; leaf blades (18-)24-36 cm long. — Sumatra, Peninsular Malaysia H. pulcherrima
63a Buds broadly ellipsoid or obovoid, 2-3 mm long; androecium longer than broad. Pedicel (1.5-)3-4 mm long. Twigs and lower leaf surface with hairs 1.5-2 mm long H. flocculosa
b Buds (depressed) globose, 2.5 mm diameter or less; androecium as broad as or broader than long. Pedicel 3 mm long or less. Twigs and lower leaf surface with hairs to 1.5 mm long 64
64a Buds 1 mm diam., with persistent indumentum or late glabrescent H. motleyi
b Buds 1-2.5 mm diam., glabrous or glabrescent 65
65a Pedicel not or indistinctly articulated (this character not quite clear in H. gracilis and H. rufo-lanata) 66
b Pedicel articulated H. reticulata
66a Upper leaf surface dark brown on drying, with venation ± indistinct. — S Peninsular Thailand, Peninsular Malaysia H. tomentosa
b Upper leaf surface olivaceous on drying, venation distinct or not. — Borneo 67
67a Twigs 3.5—7(—13) mm diam. Leaves 10- 45 cm long; venation distinct above; lateral nerves 11 pairs or more. Buds 1.5-2.3 mm diam.; anthers 8 or more. 68
b Twigs 3 mm diam. Leaves 12-21 cm long; venation indistinct above; lateral nerves 14-17 pairs; [lower leaf surface with sparse subper si stent hairs, vestigial mainly on nerves and midrib]. Buds 0.6-1 mm diam.; anthers 16-18. H. gracilis
68a Leaves 18-45 cm long; lateral nerves 18-25 pairs, sunken above. Buds 1.5-2 mm diam.; anthers 8-10. — Borneo; lowland H. splendida
b Leaves smaller, 10-23 cm long; lateral nerves 11-16 pairs, raised above. Buds 2- 2.3 mm diam.; anthers c. 15. — Borneo (Sarawak, Sabah); montane, 900-1400 m H. rufo-lanata
69a Twigs at apex pale, greyish or straw, contrasting with the dark brown petiole 70
b Twigs at apex (dark) brown, not contrasting with the petiole 75
70a Androecium angular in cross section. Anthers free for the upper half or more. — Borneo; heath forest on sand or peat soil H. oligocarpa
b Androecium (sub)circular in cross section. Anthers largely connate 71
71a Leaves chartaceous, bright brown. [Pedicel not articulated. Androecium with small apical hollow.] — Borneo; mostly in heath forest on sandy soils. 12. H. carnosa b. Leaves membranous, (blackish) brown. — Usually growing in forest on richer soils, including sand 72
72a Pedicel articulated. Androecium strongly depressed-globose; apical hollow broad with flattish bottom, to nearly halfway into the androecium. — Sumatra, Peninsular Malaysia H. sucosa subsp. sucosa
b Pedicel not articulated. Androecium ellipsoid to slightly depressed-globose; with the apical hollow small and narrow (male flowers not known in H. discolor). — Peninsular Malaysia, Borneo 73
73a Buds (sub)globose, 1.5-2(-2.2) mm long, androecium subglobose or short-ellipsoid, 1 mm long. Leaves distichous or dispersed. — Borneo; up to 700 m. 74 b. Buds ellipsoid, 2-2.4 mm long, androecium ellipsoid, 1.8-2 mm long. Leaves distichous. — Peninsular Malaysia; at c. 1300 m H. elongata
74a Fruits 4(-6) cm long or less H. pallidicaula
b Fruits 5 cm long or more H. discolor
75a Twigs ridged or nearly winged, also in the older wood 76
b Twigs not ridged; sometimes twigs faintly ridged, lined, or angular in the apical part only 79
76a Buds cleft nearly to the base. — New Guinea 77
b Buds cleft 1/2-2/3. — West Malesia 78
77a Buds slightly broader than long, short-pubescent in the lower half. Androecium slightly broader than long; anthers erect, not incurved H. angularis
b Buds subglobose to broadly ellipsoid, glabrous. Androecium longer than broad, ± obovoid, the anthers with apex free and incurved, those of one side of the androecium clasping the others H. olens
78a Buds 2.5 mm diam., pubescent. — N Sumatra H. hirtiflora
b Buds 1-1.5 mm diam., glabrous. — S Peninsular Thailand, Sumatra, Peninsular Malaysia, Borneo H. brachiata
79a Buds short pear-shaped, 2(-2.5) mm long, subsessile with the pedicel much shorter than the perianth, 0.3-1 mm long, thickish. Leaves coriaceous, pubescent or glabrescent, with dots and dashes beneath (lens!); lateral nerves flat or sunken above. Twigs hollow H. wallichii
b Buds variable in shape and size, the pedicel proportionally longer and more slender (buds obovoid with pedicel short in H. glabra var. oviflora). Leaves variable, nerves raised or sunken, dots present or absent. Twigs solid or faintly hollow 80
80a Inflorescences stout, the rachis towards the base 5-8 mm diam. Androecium about as broad as long, triquetrous in cross section, [anthers entirely connate]. — W Borneo H. pachyrachis
b Inflorescences large or small, the rachis towards base 4(-4.5) mm thick or less. Androecium triquetrous or circular in cross section. — Whole of W Malesia. 81
81a Androecium 3- or 4-angular in cross section. Anthers ± erect, free for about halfway or more. Buds 1.5(—2) mm diameter or less. Pedicel articulated. Leaves with the lateral nerves raised above; dots absent (H. ridleyana with leaves small, nerves sunken, male buds 1 mm diam.). — Most of W Malesia, not in Sulawesi, rare in the Philippines 82
b Androecium in cross section circular, ellipsoid, or subtriangular with rounded angles. Anthers ± curved, almost entirely connate, free apices c. 1/3 or less. Buds (1.3—)1.5 mm diameter or more. Pedicel articulated or not. Leaves with the lateral nerves raised, level, or sunken above; dots present or absent (lens!) 89
82a Leaves 5-16 cm long; midrib and lateral nerves level or sunken above 83
b Leaves small or large, 5-28 cm long; midrib and lateral nerves raised above. 84
83a Twigs and inflorescences rather glabrescent. Leaf apex acute or acute-acuminate; nerves faint above. Anthers 4-6. — Peninsular Malaysia, Borneo H. ridleyana
b Twigs late glabrescent, inflorescences with persistent indumentum. Leaf apex blunt; nerves flat or but little raised above, clearly visible. Anthers 9 or 10. — Borneo (Sarawak) H. obtusa
84a Leaves early glabrescent beneath, also on the midrib; leaf apex long acute-acuminate. Twigs rather smooth, lower down cracking longitudinally. — Borneo (Brunei) H. disticha
b Leaves early glabrescent beneath, but midrib sometimes late glabrescent; leaf apex acute-acuminate. Twigs striate, lower down coarsely striate or finely cracking 85
85a Twigs l-2(-4) mm diam. Leaves 7-18 cm long, thinly membranous to subcharta- ceous; petiole slender, l-1.5(-2) mm diam. Inflorescences delicate, up to 9 cm long; buds 1 mm diam. 86
b Twigs 1—5(—8) mm diam. Leaves of variable sizes, chartaceous or coriaceous; the petiole 1.5—4(—8) mm diam. Inflorescences up to 15(-20) cm long; buds 1-2 mm diam. 87
86a Leaf bud with hairs (0. l-)0.2 mm long; twigs at apex and leaves glabrous; inflorescences with sparse stellate hairs 0.2 mm long, glabrescent. Leaves drying to a greyish tinge. Male buds short pear-shaped, tapering into the pedicel H. tenuifolia
b Leaf bud, apical part of twig, petiole, midrib beneath and inflorescences with woolly stellate-dendroid hairs (0.2-)0.5 mm long; leaves olivaceous on drying. Male buds globose or depressed-globose H. macilenta
87a Twigs and leaves stout, the midrib broad above, at the transition to the petiole at least 3 mm wide. Inflorescences 10-20 cm long. — Borneo; forests on poor soil, including sand and peat H. laticostata
b Twigs and leaves less robust; midrib above towards the insertion of the petiole less than 3 mm wide. Inflorescences up to 15-20 cm long. — On poor or rich soil 88
88a Leaves 16-28 cm long, leaf base ± rounded or short-attenuate; nerves 16-19 pairs, very prominent above. — Borneo (Sarawak) H. nervosa
b Leaves 7-28 cm long, base short- to long-attenuate; nerves 6-16 pairs, raised to variable degrees above. [On drying, colour of leaves above and beneath usually much contrasting, generally more so than in the related species.] —Variable, with 3 varieties (based on fruit size). Sumatra, Peninsular Malaysia, whole of Borneo, Philippines (Mindanao, Palawan) H. polyspherula
89a Leaf bud, apical part of twig, and inflorescences with hairs 0.2 mm long or more (hairs 0.1-0.4 mm long in H. punctata) 90
b Leaf bud, apical part of twig, and inflorescences with hairs (0.2-)0.1 mm long or less 97
90a Buds ellipsoid, 2.5-3.5 mm long; androecium longer than broad. [Leaves thick and brittle, apex bluntish; lower surface usually with conspicuous pale golden hair scars (lens!).] H. endertii
b Buds (sub)globose; androecium not longer than broad 91
91a Buds 2.5-3 mm diam. (or 1.5 mm in Hallier 624 from W Borneo, see the notes), cleft c. 4/5. — Sumatra H. valida
b Buds (1-)1.2-2.5 mm diam., cleft l/3-2/3(-3/4). — Peninsular Malaysia, Borneo 92
92a Leaves with dots beneath (lens!) 93
b Leaves without dots beneath. [Pedicel not articulated.] 94
93a Pedicel articulated. Lateral nerves flat or sunken above. Lower leaf surface cinnamon or chocolate, contrasting with upper surface. — Borneo H. borneensis
b Pedicel not articulated. Lateral nerves largely raised above. Upper and lower leaf surface not much contrasting. — Peninsular Malaysia H. punctata
94a Buds 2-2.5 mm diam.; androecium sessile, broadly saucer-shaped. Leaves 20-45 cm long. — Lowland forest H. fragillima
b Buds (1-) 1.4-2.2 mm diam.; androecium (depressed-)globose, with the apical hollow small, concealed by the apices of the anthers. Leaves 4-35 cm long. — Montane forest at 800-2000 m 95
95a Androecium with slender androphore 0.3-0.8 mm long, not hidden by the anthers. Leaves membranous, 9-18 cm long, dark brown, [apex acute-acuminate] H. androphora
b Androecium (sub)sessile, androphore absent or up to 0.5 mm, largely hidden by the anthers 96
96a Leaves chartaceous or membranous, to c. 35 cm long, olivaceous-brown; apex acute-acuminate. Inflorescences to 20 cm long. — Borneo (Mt Kinabalu) H. amplomontana
b Leaves coriaceous, 4-14 cm long, blackish; apex obtuse to subacute. Inflorescences 4-16 cm long H. montana
97a Leaves with dots beneath 104
b Leaves without dots beneath (dots should not be confused with smaller, blackish points) 98
98a Buds ± ellipsoid; androecium ± obovoid, the apical part of the anthers deeply inflexed into the apical hollow. — New Guinea H. sepikensis
b Buds and androecium of variable shapes; the anthers ± straight or curved, at apex not inflexed. — W and E Malesia, not in New Guinea 99
99a Perianth coriaceous; lobes thick, towards the base (0.3-)0.4-l mm thick. Androecium ellipsoid-obovoid, longer than broad 102
b Perianth thinner, lobes at base 0.2-0.3 mm thick. Androecium subglobose, broadly ellipsoid, or obovoid, not or but little longer than broad. [Androphore narrow, only 0.2 mm long. Leaves membranous to chartaceous.] 100
100a Pedicel (1—)1.5—2 mm long, about as long as the perianth. — Peninsular Malaysia, Borneo 101
b Pedicel shorter than the perianth, 0.5 mm long. — Sulawesi, Philippines H. costulata
101a Buds cleft c. 1/2 H. subalpina
b Buds cleft 2/3-4/5 H. obscura
102a Bark of twigs not flaking. Leaves ± membranous 103
b Bark of twigs flaking or not. Leaves coriaceous. [Pedicel not articulated. Anthers (3-)4-8; androphore rather broad, tapering, (0.1-)0.2-0.3 mm long.] — Borneo (Sarawak, Sabah); kerangas, montane forest; 800-1550 m. H. xanthina
103a Pedicel articulated. Anthers 7-9; androphore rather broad and tapering, 0.2-0.5 mm long. — Sumatra, Peninsular Malaysia; 0-1000 m H. majuscula
b Pedicel not articulated. Anthers 5 or 6; androphore narrow, 0.1-0.2 mm long, hidden by the anthers. — C Sulawesi; 100-450 m H. coriacea
104a Twigs 2.5-3(-4) mm diam. Leaves (8-)12 cm long or more. Male buds (subglobose, 1.5-4.2 mm diam 105
b Twigs 1.5-2 mm. Leaves 5-12 cm long. Male buds ± ellipsoid or globose, 1.2-1.8 mm long H. penangiana
105a Buds cleft 3/4-4/5; anthers 7-11 106
b Buds cleft 1/3-2/3; anthers 9-20. — Sumatra, Java 107
106a Anthers 7-9. Dry fruits 4-5 cm long, pericarp 10-20 mm thick. — N Sumatra, Peninsular Malaysia, Borneo H. punctatifolia
b Anthers c. 11. Dry fruits 2 cm long, pericarp 1.5 mm thick. — Peninsular Malaysia H. punctata
107a Buds 3-4.2 mm diam.; anthers 15-20. — N & C Sumatra H. macrothyrsa
b Buds 1.5-2.5 mm diam.; anthers 9-15. — Variable, with 3 varieties. Java, W, C & S Sumatra H. glabra

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

(2) REGIONAL KEY TO THE SPECIES — PENINSULAR MALAYSIA, SINGAPORE

(based on female flowering and fruiting specimens)

1a Flower buds pubescent at base. [Ovary pubescent. Fruits pubescent, ellipsoid, 2.5-4 cm long. Flowers in dense clusters and strongly fragrant in male specimens.] — Originating from Sri Lanka, cultivated in Penang, Singapore and elswhere H. iryaghedhi
b Buds glabrous or early glabrescent 2
2a Perianth 2-lobed. Ovary glabrous 3
b Perianth 3- (or 4-)lobed. Ovary glabrous or pubescent 5
3a Twigs ridged or lined. Leaves membranous, often with irregular whitish blotches. Fruits and seeds globose. — Plant usually riverine, in coastal areas. H. irya
b Twigs not lined. Leaves without pale blotches. Fruits and seeds ellipsoid 4
4a Leaves membranous, glabrous beneath, dots absent. — Gardens' Jungle, Singapore (originating from E Malesia) H. parviflora
b Leaves coriaceous, beneath pubescent and with dots (lens!) — Kerangas, peat swamp forest H. crassifolia
5a Leaves with dots beneath (dots not to be confused with smaller dark punctation) 20
b Leaves without dots beneath 6
6a Leaves with persistent indumentum beneath 7
b Leaves glabrous or glabrescent beneath 11
7a Ovary pubescent. Fruits pubescent or at least with vestigial indumentum near the base; perianth not persistent H. tomentosa
b Ovary glabrous or with some incidental minute hairs. Fruits glabrous, perianth (at least at first) persistent 8
8a Hairs on lower (and upper) leaf surface harsh, with hardened hair bases, in older leaves rendering the surface scabrous. Fruits 1-1.4 cm long. H. grandis
b Leaves not scabrous. Fruits 2 cm long or more 9
9a Twigs 3-5 mm diam. Leaf bud and twig apex with hairs 0.2-0.3 mm long. Leaf blade 13-21 cm long, [matt above on drying. Fruits 2.2-2.4(-3) cm long] H. fulva
b Twigs 5-10 mm diam. Leaf bud and twig apex with hairs 0.5 mm long or more. Leaf blade 20-40(-70) cm long 10
10a Leaf bud, twig apex and lower leaf surface with rather stiff, rust-coloured hairs 0.5-1 mm long. Fruits 3.8-5.5 cm long H. superba
b Leaf bud, twig apex and lower leaf surface with yellow-brown or pale brown woolly hairs 1-2 mm long. Fruits 3 cm long H. flocculosa
11a Twigs pale brown or straw, contrasting with the blackish petiole. Leaves either distichous or in 3-5 rows along the twigs 12
b Twigs brown, not contrasting with the petiole. Leaves distichous 14
12a Leaves distichous or in 3 rows 13
b Leaves in 3-5 rows along the twigs. [Pedicel not articulated. Fruits 3-5.5 cm long, perianth not persistent.] H. sparsa
13a Leaves distichous. Pedicel not articulated. Male perianth ± ellipsoid, 2-2.4 mm long. Fruits not known H. elongata
b Leaves distichous or in 3 rows. Pedicel articulated (this character best seen in male flowers). Male perianth globose, smaller. Fruits 2.5-3.5 cm long, perianth persistent H. sucosa subsp. sucosa
14a Leaf upper surface matt on drying caused by fine wrinkles; nerves flat or sunken. Fruits usually with persistent perianth H. fulva
b Leaves above not particularly matt, not finely wrinkled; nerves flat or raised. Perianth not persistent under the fruits 15
15a Lateral nerves flat or but faintly raised above 16
b Lateral nerves distinctly raised above. [Pedicel articulated; this character best seen in male flowers.] 18
16a Leaf bud with hairs 1 mm long. [Midrib narrow, line-shaped, sunken above; blade margin rolled-in.] H. perangusta
b Leaf bud with hairs much shorter 17
17a Leaf bud with hairs 0.1 mm long. Twigs 2.5-5 mm diam. Leaves 15-27 cm long. Pedicel not articulated H. subalpina subsp. subalpina
b Leaf bud with hairs 0.2-0.4 mm long. Twigs 1.5-3.5 mm diam. Leaves 5-15 cm long. Pedicel articulated H. ridleyana
18a Leaf bud with hairs 0.1 mm long. Midrib early glabrescent beneath. Fruits 4.5-6.5 cm long, with thick pericarp H. majuscula
b Leaf bud with hairs 0.1 mm long or usually much longer. Midrib often late glabrescent beneath. Fruits 2-4 cm long, pericarp 2-5(-7) mm thick 19
19a Twigs ± angular, lined or low-ridged. [Fruits 2-3(-4) cm long.] H. brachiata
b Twigs terete, neither lined nor ridged 20
20a Twigs 2-5 mm diam. Leaves usually chartaceous. Fruits 1.9-3.5 cm long H. polyspherula
b Twigs 1-3 mm diam. Leaves membranous. Fruits 2.3-2.4 cm long H. macilenta
21a Leaves usually with persistent indumentum beneath 22
b Leaves glabrous or glabrescent beneath 23
22a Leaves ± pubescent, sometimes late glabrescent, with dots and dashes beneath. Ovary glabrous. Fruits 4-6 cm long, glabrous, perianth usually persistent H. wallichii
b Leaves always pubescent beneath, with dots, not with dashes. Ovary pubescent. Fruits 1.6-1.8 cm long, shaggy-hairy, perianth not persistent H. pulcherrima
23a Twigs 1.5-2 mm diam. Fruits 1.1-2 cm long H. penangiana
b Twigs somewhat stouter, 2.5-5 mm diam. Fruits 2 cm long or more 24
24a Leaves coriaceous, apex blunt or subacute. Fruits 2-2.3 cm long, pericarp thin. — Montane species of C Peninsular Malaysia H. punctata
b Leaves membranous, apex acute-acuminate. Fruits 4.5-8 cm long, pericarp 10-20 mm thick. — Widespread in W Malesia; forests up to c. 1100 m H. punctatifolia

(3) REGIONAL KEY TO THE SPECIES — SUMATRA, JAVA

(based on female flowering and fruiting specimens)

1a Flower buds pubescent at base. [Ovary pubescent. Fruits ellipsoid, 2.5 — 4 cm long, pubescent. In male the flowers in dense clusters, strongly fragrant.] — Cultivated, originating from Sri Lanka H. iryaghedhi
b Buds glabrous (pubescent at base in H. hirtiflora and H. triandra) 2
2a Leaves membranous, often with irregular whitish blotches. Fruits globose, 1.5-2 cm diam., glabrous; pericarp 1-2 mm thick; seeds globose. [Perianth 2-lobed; ovary glabrous.] — Riverine or marshy, usually near the coast H. irya
b Leaves of different consistency, usually not blotched. Fruits and seeds ellipsoid. — Plant not coastal 3
3a Perianth 2-lobed. Leaves with dots beneath (dots not to be confused with smaller punctation of different origin, lens!) 4
b Perianth 3- (or 4-)lobed. Leaves with or without dots beneath 5
4a Leaves coriaceous, 10-20(-28) cm long, finely pubescent beneath. Twigs 2-6 mm diam. Fruits 1.5-2.2 cm long, with persistent perianth. — Peat swamp or padang forest H. crassifolia
b Leaves membranous or thinly chartaceous, 5-12 cm long, glabrous beneath. Twigs 1.5-2 mm diam. Fruits 1-2 cm long; perianth not persistent. — Mixed forest. H. penangiana subsp. penangiana
5a Ovary and fruits (at least at base) pubescent. Leaves with persistent indumentum beneath 6
b Ovary and fruits glabrous (fruits almost glabrous in H. triandra). Leaves glabrous or pubescent beneath 7
6a Twigs 2-5 mm diam. Leaves 9-27 cm long, lower surface without dots. Fruits with hairs 0.5 mm long or less. — Thailand, Peninsular Malaysia; specimens from Sumatra not seen H. tomentosa
b Twigs 5-8 mm diam. Leaves 20-36 cm long, with dots beneath (lens!). Fruits with hairs 2 mm long H. pulcherrima
7a Leaves with dots beneath (lens!). Lateral nerves generally flat or sunken above 21
b Leaves without dots beneath. Nerves either raised or flat to sunken above. 8
8a Leaves with persistent indumentum beneath. Fruits with persistent perianth. 9
b Leaves glabrous or glabrescent beneath. Fruits with persistent perianth or not 11
9a Hairs harsh, older leaves scabrous beneath. [Fruits 1-1.4 cm long.] H. grandis
b Hairs softer; older leaves not scabrous beneath 10
10a Twigs 5-8 mm diam.; leaves 20-40(-70) cm long. Leaf bud with hairs 0.5-1 mm. Fruits 3.8-5.5 cm long H. superba
b Twigs 3-5 mm diam.; leaves 13-21 cm long. Leaf bud with hairs 0.2-0.3 mm long. Fruits 2-3 cm long H. fulva
11a Lateral nerves flat or sunken or but faintly raised above. Colour of lower leaf surface generally greyish brown, not much contrasting with upper surface 12
b Lateral nerves distinctly raised above. Colour of the lower leaf surface bright brown or chocolate, contrasting with the upper surface 16
12a Twigs 1.5-3 mm diam. Leaf bud, twig apex, and young inflorescences with woolly hairs 0.3-0.7 mm. Leaves 5-9 cm long H. triandra
b Twigs (2-)3-10 mm diam. Leaf bud, twig apex, and inflorescences 0.1-0.3 mm long. Leaves more than 10 cm long 13
13a Leaves distichous, matt on drying caused by minutely wrinkled upper surface. Bark of twigs straw or brown 14
b Leaves distichous or in 3-5 rows; not particularly matt, upper surface not finely wrinkled. Bark of twigs pale, grey-brown or straw, contrasting with the blackish petiole 15
14a Leaves elliptic-oblong to oblong, olivaceous-brown above. Stem grey-brown, not much contrasting with petiole. Fruits brown on drying, 2.2-3 cm long, perianth persistent H. fulva
b Leaves elliptic-oblong to lanceolate, dark olivaceous above. Stem pale, ± yellowish brown, rather contrasting with the petiole. Fruits blackish on drying, 1.5 cm long, perianth not persistent H. tristis
15a Leaves distichous or in 3 rows. Fruits 2.5-3.5 cm long, with the perianth persistent. Pedicel articulated (best seen in male flowers). H. sucosa subsp. sucosa
b Leaves in 3-5 rows. Fruits 3-5.5 cm long, perianth not persistent. Pedicel not articulated H. sparsa
16a Perianth 4-lobed. Pedicel not articulated. Fruits 8-9 cm long. [Flowers known only from the remnants persistent under the young fruits.] H. valida
b Perianth generally 3-lobed. Pedicel articulated. Fruits up to 6.5 cm long. 17
17a Twigs ± angular, with lines or ridges 18
b Twigs terete or faintly angular, neither lined nor ridged 19
18a Buds pubescent (known only in male flowers). Fruits 5-6 cm long. — N Sumatra H. hirtiflora
b Buds glabrous. Fruits 2-4 cm long H. brachiata
19a Twigs 1-3 mm diam. Leaves membranous, midrib beneath late glabrescent. Fruits 2.3-2.4 cm long H. macilenta
b Twigs 2-5 mm diam. Leaves chartaceous. Fruits 1.9-6.5 cm long 20
20a Leaf bud and inflorescences with hairs 0.1-0.2 mm long. Midrib beneath early glabrescent. Fruits 4.5-6.5 cm long H. majuscula
b Leaf bud and inflorescences with hairs 0.1-0.6 mm long. Midrib often late glabrescent. Fruits 1.9-3.5 cm long. — Variable; 3 varieties based on fruit size H. polyspherula
21a Leaves usually with persistent indumentum beneath (sometimes glabrescent), and with both dots and dashes (lens!). Twigs conspicuously hollow. Fruits 4-7 cm long, the perianth generally persistent H. wallichii
b Leaves glabrous or glabrescent beneath, with dots, not with dashes. Twigs not conspicuously hollow. Fruits variable 22
22a Bark of twigs pale, greyish to straw-coloured, contrasting with blackish petiole. Leaves dispersed in 3-5 rows. [Female flowers and fruits not known.] — Sumatra (N Aceh); at c. 1300 m H. atjehensis
b Bark of twigs brown, not contrasting with petiole. Leaves distichous (or in 3 rows in H. glabra, p.p.) 23
23a Fruits (4.5-)5-8 cm long, pericarp 10-20 mm thick H. punctatifolia
b Fruits 1-2.5 cm long, pericarp much thinner 24
24a Twigs 1.5-2 mm diam. Leaves 5-12 cm long. Fruits 1.1-2 cm long H. penangiana
b Twigs 2.5-4(-6) mm diam. Leaves (8-) 12 cm long or more. Fruits 1.8-2.5 cm long 25
25a Leaves distichous. — C & N Sumatra H. macrothyrsa
b Leaves distichous or in 3 rows. — S Sumatra, Mentawai Is., north to Simeulue I., Java H. glabra

(4) REGIONAL KEY TO THE SPECIES — BORNEO

(based on female flowering and fruiting specimens)

1a Leaves membranous, usually with irregularly shaped whitish blotches. Fruits globose, 1.5-2 cm diam., glabrous; pericarp 1-2 mm thick; seeds globose. [Perianth 2-lobed; ovary glabrous.] — Plants usually growing not too far from the coast. H. irya
b Leaves variable, usually not white-blotched. Fruits subglobose or ellipsoid; seeds ellipsoid. — Plants coastal or not 2
2a Perianth 2-lobed 3
b Perianth predominantly 3- (or 4-)lobed 5
3a Leaves coriaceous, densely short-pubescent and with dots beneath (lens!). Twigs grey-brown, not contrasting with petioles H. crassifolia
b Leaves membranous, glabrous and without dots beneath. Twigs greyish or straw- coloured, contrasting with blackish petioles 4
4a Inflorescences ± spike-like, 5-10 cm long. Perianth persistent under the fruits (always?). — Borneo (SE Sabah) H. sterilis
b Inflorescences branched, 1-2 cm long. Perianth not persistent under the fruits. — Borneo (Sarawak, Sabah, E, C & S Kalimantan). H. sucosa subsp. bifissa
5a Leaves in 3 or more rows along the twigs 6
b Leaves distichous 8
6a Leaves ± clustered at the apex of the twigs. Petiole proportionally long and slender, 25-50 mm long. — Northern Borneo; sandy soils H. sabulosa
b Leaves not clustered. Petiole proportionally shorter 7
7a Leaves 10-30 cm long. Twigs pale, grey or straw-coloured, contrasting with the blackish petioles. Buds 2.5-3 mm long. Fruits 1.5-6 cm long H. pallidicaula
b Leaves 20-45 cm. Twigs brown, not contrasting with the petioles. Buds 4-5 mm long. Fruits 6-8 cm long H. fragillima
8a Twigs lined or ridged H. brachiata
b Twigs terete or faintly angular, neither distinctly lined nor ridged 9
9a Leaves with (sub)persistent indumentum beneath (sometimes largely glabrescent in H. wallichii) 10
b Leaves glabrous or early glabrescent beneath (midrib sometimes late glabrescent) 18
10a Plant stout; leaves 50 cm long, petiole 3 mm long only. — Borneo (lowland Sarawak) H. sessilifolia
b Plants variable in habit; leaves large, but petiole proportionally much longer.11 11a. Older leaves with scabrous hair scars above and beneath. Fruits 1-1.4 cm long. H. grandis
b Leaves not scabrous. Fruits larger 12
12a Leaves with dots and/or dashes beneath (lens!) H. wallichii
b Leaves with or without dots beneath, never with dashes 13
13a Flower buds with persistent indumentum; fruits without persistent perianth H. motleyi
b Buds glabrous or early glabrescent; fruits with persistent perianth or not. 14
14a Ovary pubescent; fruits sometimes pubescent only towards the base 15
b Ovary and fruits glabrous 16
15a Twigs 3.5-5 mm diam. Leaves 10-23 cm long; nerves 11-16 pairs. Fruits largely glabrescent; perianth not persistent H. rufo-lanata
b Twigs 4-7 mm diam. Leaves 18-45 cm long, nerves 18-25 pairs. Fruits pubescent, with persistent perianth H. splendida
16a Twigs 3-6 mm diam. Leaves 18-35 cm long; upper surface sometimes bullate; [nerves 17-20 pairs.] Fruits 2.3-2.7 cm long H. reticulata
b Twigs 1.5-3 mm diam. Leaves often smaller, not bullate. Fruits 1-1.5 cm long 17 17a. Leaves membranous, beneath without dots; nerves 14-17 pairs. H. gracilis
b Leaves thinly chartaceous, beneath with dots; nerves 5—9(—11) pairs H. paucinervis
18a Leaves with dots or with dots and dashes beneath (lens!) (dots not to be confused with smaller, irregularly spaced points, which are usually present) 19
b Leaves without dots beneath (enlarged hair scars sometimes present) 22
19a Leaf bud, twig apex, and young inflorescences with hairs 0.2 mm long or more 20
b Leaf bud, twig apex, and young inflorescences with hairs 0.1 mm long or less 21 20a. Twigs hollow. Leaves often with persistent indumentum beneath. Pedicel not articulated. Fruits with persistent perianth H. wallichii
b Twigs (almost) solid. Leaves glabrescent beneath. Pedicel articulated. Fruits without persistent perianth H. borneensis
21a Twigs 1.5-2 mm diam. Leaves 5-12(-17) cm long; nerves 8-11 pairs. Fruits 1.1-2 cm long, pericarp thin H. penangiana
b Twigs 2.5-4 mm diam. Leaves 9-21 cm long, nerves 11-16 pairs. Fruits 4.5-8 cm long, with thick pericarp H. punctatifolia
22a Twigs pale, grey-brown or yellowish, contrasting with the dark brown petioles 23
b Twigs brown on drying, ± not contrasting with the petioles 27
23a Leaves membranous, (blackish) brown above, somewhat paler beneath. Perianth persistent under the fruits. — Mixed forest H. pallidicaula
b Leaves usually chartaceous, bright brown or olivaceous above. Perianth not persistent under the fruits 24
24a Twigs 2-3 mm diam. Leaves 7-16 cm long, bright brown or chocolate beneath, contrasting with the grey-olivaceous upper surface. Fruits 1.8-2.7 cm long. — Kerangas, peat forest H. oligocarpa
b Twigs 3-10 mm diam. Leaves 13-35 cm long, the lower surface not conspicuously contrasting with upper surface 25
25a Fruits 5 cm long or more. Leaves not conspicuously matt above. 21. H. discolor b. Fruits 2 cm long or less. Leaves matt, caused by finely wrinkled upper surface. 26
26a Leaves (elliptic-)oblong. Fruits 1.6-2 cm long. — Heath forest, peat swamp forest H. carnosa
b Leaves elliptic-oblong to lanceolate. Fruits 1.5 cm long. — Mixed forest H. tristis
27a Leaf bud and immature inflorescences with hairs 0.1 mm long or less. Lateral nerves flat or sunken, or but little raised above 28
b Leaf bud and immature inflorescences with hairs 0.1 mm long or more; lateral nerves above raised or not; if hairs only 0.1 mm long, then the lateral nerves above distinctly raised, at least in the lower half 30
28a Species from lowland limestone, up to c. 700 m. Leaves membranous. Fruits not known. — Borneo (NE Kalimantan) H. obscura
b Montane species; 800-1800 m. Leaves membranous or coriaceous. Fruits 5 cm long or less. — Borneo (Sarawak, Sabah) 29
29a Leaves without distinct large hair scars beneath (lens!). Fruits 3-5 cm long. H. subalpina subsp. kinabaluensis
b Leaves with (usually) distinct yellowish enlarged hair scars beneath. Fruits 3.5-5 cm long H. xanthina
30a Pedicel not articulated (best seen in male flowers). Fruits with perianth persistent or not 31
b Pedicel articulated. Perianth not persistent under the fruits 36
31a Leaves 15-45 cm long. Fruits 6 cm long or more; perianth ± persistent. — Lowland or montane forest 32
b Leaves 5-20 cm long. Fruits 2-4 cm long; perianth not persistent. — Montane forest at 800-2000 m 34
32a Nerves 11-22 pairs. — Borneo (Sabah: Mt Kinabalu); 1000-1500 m H. amplomontana
b Nerves 20-30 pairs. — Forests up to c. 1000 m 33
33a Female flowers and fruits not known. — Hallier 624, Mt Damoes, W Kalimantan; probably an undescribed species close to H. valida, see there. aff. H. valida
b Perianth 4-5 mm long. Fruits 6-8 cm long, pericarp 10-20 mm thick H. fragillima
34a Leaves membranous, apex acute-acuminate. Fruits 2.4-3 cm long H. androphora
b Leaves chartaceous or coriaceous, apex rounded to (sub)acute, not acute-acuminate 35
35a Leaves chartaceous or coriaceous, without large hair scars beneath (lens!). Perianth 2 mm long. Fruits 2-2.7 cm long H. montana
b Leaves usually strongly coriaceous, usually with large hair scars beneath. Perianth 2.5-3 mm long. Fruits 3-4 cm long H. endertii
36a Male inflorescences very stout, the rachis towards the base 5-8 mm diam. [Female inflorescences, flowers, and fruits not known.] — Borneo (W Kalimantan) H. pachyrachis
b Male inflorescences less stout 37
37a Midrib on upper leaf surface towards the transition to the petiole 3 mm broad or more H. laticostata
b Midrib at base narrower 38
38a Leaves 16-28 cm long, base rounded or short-attenuate; nerves 16-19 pairs. — Borneo (Sarawak) H. nervosa
b Leaves 5-28 cm long, base rounded, short-, or long-attenuate; nerves 5-15(-20) pairs 39
39a Lateral nerves sunken, flattish, or but slightly raised above 40
b Lateral nerves raised above 41
40a Leaf apex rounded. Fruits not seen H. obtusa
b Leaf apex acute-acuminate. Fruits 1.5-2 cm long H. ridleyana
41a Leaf bud and young inflorescences with hairs 0.1-0.2 mm long. Leaves on drying dull, greyish brown, colour of upper and lower surface not much contrasting. Fruits 1.7-2 cm long H. tenuifolia
b Leaf bud and young inflorescences with hairs 0.2 mm long or more; if hairs 0.1 mm long, then the olivaceous to dark-brown upper leaf surface much contrasting with the cinnamon colour beneath 42
42a Twigs 1-3 mm diam. Leaves membranous, 10—18(—27) cm long. Fruits 2.3-2.4 cm long H. macilenta
b Twigs 2-5 mm diam. Leaves chartaceous; leaves and fruits of variable sizes. 43
43a Twigs early glabrescent; bark ± longitudinally cracking. Leaf apex long acute- acuminate. Fruits 2.8-3.2 cm long; pericarp hard-woody, 8-10 mm thick. — Borneo (Brunei) H. disticha
b Twigs late glabrescent; bark striate, not cracking. Leaf apex acute-acuminate, the acumen not conspicuously long. Fruits 1.9-6 cm long. — Variable, with 3 varieties (based on fruit size). Whole of Borneo H. polyspherula

(5) REGIONAL KEY TO THE SPECIES — PHILIPPINES, SULAWESI, MOLUCCAS

(based on female flowering and fruiting specimens)

1a Leaves membranous, usually irregularly whitish blotched. Perianth 2-lobed; ovary glabrous. Fruits globose, 1.5-2 cm diam., glabrous; pericarp 1-2 mm thick; seeds globose. — Riverine or marshy, mostly near the coast H. irya
b Leaves variable, usually not whitish blotched. Fruits subglobose or ellipsoid; seeds ellipsoid. — Coastal or not 2
2a Perianth 3-lobed. [Ovary and fruits glabrous.] 3
b Perianth 2-lobed 5
3a Leaf bud and inflorescences with hairs 0.2-0.6 mm long. Pedicel articulated. [Fruits 2(-2.5) cm long.] — Philippines (Mindanao, Palawan) H. polyspherula var. polyspherula
b Leaf bud and inflorescences with hairs 0.1-0.2 mm long. Pedicel not articulated 4
4a Fruits 3.5-7 cm long; (dry) pericarp (4—)8—15 mm thick. Leaves ± membranous, olivaceous-brown, midrib glabrous above; leaves sometimes with whitish blotches as in H. irya. — Philippines, Sulawesi H. costulata
b Fruits 4 cm long, (dry) pericarp 3.5-8 mm thick. Leaves membranous to thinly coriaceous, brown; midrib towards the base pubescent above in younger leaves. — C Sulawesi H. coriacea
5a Twigs 4-14(-20) mm diam. Leaves 20-45 cm long, petiole 2-7 mm long. Leaf bud and inflorescences with hairs 0.3-1 (-1.5) mm long. Buds 3.5-5 mm long, glabrous; ovary glabrous. Fruits 3.5-5.5 cm long, glabrous H. sylvestris
b Twigs more slender. Leaves smaller, petiole comparatively longer. Leaf bud and inflorescences with hairs up to 0.2 mm long. Buds 3(-3.5) mm long or less. Fruits up to 3 cm long (in H. lancifolia to 3.5 cm long) 6
6a Leaves ± chartaceous, oblong-lanceolate to lanceolate. Buds cleft c. 1/4; ovary pubescent. Fruits often ± pear-shaped, 2.5-3.5 cm long, early glabrescent; (dry) pericarp 4-8 mm thick. — Sulawesi H. lancifolia
b Leaves of different consistency, generally broader, oblong to oblong-lanceolate. Buds cleft c. 1/3 or more. Fruits 1-3 cm long 7
7a Ovary and fruits pubescent; hairs on the fruits may be inconspicuous and only remaining at the very base near the insertion of the pedicel (lens!); pericarp thick or thin 8
b Ovary and fruits glabrous; pericarp 1-2 mm thick. [Female flowers and fruits not known in H. aruana and H. samarensis.] 11
8a Flower buds 2.5-3 mm long, cleft 1/3-1/2. Fruits (1.6-) 1.8-3 cm long; pericarp 2-3 mm thick H. laevigata
b Female flowers not known. Fruits smaller, pericarp thinner 9
9a Fruits 1.5-1.6 cm long, short-ellipsoid. [Male buds transversely ellipsoid, cleft 2/3-4/5. Leaves 8-30 cm long, membranous or chartaceous; nerves flat, inconspicuous. Twigs terete, not ridged.] — Moluccas (Talaud Is.), possibly Sulawesi. H. talaudensis
b Fruits smaller, 1.1-1.3 cm long 10
10a Fruits subglobose. Twigs ± flattened, usually lined or low-ridged. Leaves 12-25 cm long, membranous; nerves flat, inconspicuous. Male buds ± pear-shaped, cleft to c. 2/3. — Moluccas H. decalvata
b Fruits short-ellipsoid. Twigs terete, not lined. Leaves 5-14 cm long, chartaceous, nerves inconspicuous on both surfaces. Male buds ± obtriangular, cleft about halfway. — Philippines (Luzon) H. obscurinervia
11a Twigs angular or ridged. [Species distinctive only in male flowering specimens.] 12
b Twigs (sub)terete or sometimes faintly angular, or shallowly lined 13
12a SW New Guinea, possibly Aru and Tanimbar Is. [Female flowers and fruits not known.] H. aruana
b Moluccas (Seram, Banda, Dammar I., possibly Ternate) H. smithii
c Philippines. [Ovary glabrous or almost so.] H. ardisiifolia
13a Bark of twigs pale, grey-brown, contrasting with the blackish petioles. [Fruits blackish on drying, 1.5-2 cm long.] — Moluccas H. spicata
b Twigs brown, in colour not contrasting with the petioles 14
14a Fruits ± globose to subellipsoid, 0.9-1.2 cm long (to 2 cm in New Guinea); blackish on drying. — Aru Is., New Guinea H. subtilis var. subtilis
b Fruits 1.1-1.6 cm long (fruits not known in H. samarensis) 15
15a Philippines (Samar) H. samarensis
b Moluccas, Sulawesi 16
16a Fruits ellipsoid, 1.5 cm long; blackish on drying. — Moluccas (Morotai, Obi Is.) H. moluccana var. moluccana
b Fruits subglobose or ellipsoid, l.l-1.6(-2) cm long, brown on drying. — Sulawesi (Kabaena Is.), Moluccas (Seram) H. parviflora

(6) REGIONAL KEY TO THE SPECIES — NEW GUINEA

(based on female flowering and fruiting specimens)

1a Leaves membranous, often with irregular whitish blotches. Perianth 2-lobed; ovary glabrous. Fruits glabrous, globose, 1.5-2 cm diam.; pericarp 1-2 mm thick; seeds globose. — Riverine or marshy, usually not far from the coast H. irya
b Leaves of different consistency, generally without whitish blotches. Fruits glabrous or pubescent, globose or ellipsoid; if globose either only 1 cm diam. (H. subtilis), or the pericarp more than 2 mm thick, at least at one side; seeds mostly ellipsoid. — Coastal or not 2
2a Twigs angled or ridged. — Aru Is., New Guinea 3
b Twigs terete, sometimes lined in-between the bases of petioles but neither angled nor ridged. — New Guinea to Solomon Is 8
3a Leaves with dots beneath (lens!). Perianth 2-lobed. — New Guinea (Bird's Head to W Sepik Prov.) H. inflexa
b Leaves without dots beneath 4
4a Perianth 3- (or 4-)lobed 5
b Perianth 2-lobed 6
5a Ovary glabrous (?). Fruits 10-16 mm long, glabrous. Leaves chartaceous, 7-14 cm long; petiole comparatively long and slender, 11-20 mm long. — SW & S New Guinea (Digul, Western Prov.) H. olens
b Ovary pubescent. Fruits 17-20 mm long, pubescent at base. Leaves membranous or thinly chartaceous, 10-27 cm long; petiole 7-15 mm. — Papua Barat (Bird's Head) H. angularis
6a Flower buds depressed-globose, lobes nearly 1 mm thick; ovary pubescent. Fruits 17-20 mm long, pubescent. Leaves membranous or thinly chartaceous. — Papua Barat (Bird's Head) H. angularis
b Female flowers and fruits not known 7
7a Leaves membranous. — SW New Guinea; possibly Aru and Tanimbar Is H. aruana
b Leaves thinly coriaceous. — SW New Guinea (a species close to H. aruana). H. iriana
8a Perianth 3- (or 4-)lobed, [cleft almost to the base.] — Papua New Guinea (East Sepik Prov.) H. sepikensis
b Perianth 2-lobed 9
9a Ovary and fruits glabrous 10
b Ovary and fruits pubescent. [Hairs on fruits either distinct or small and inconspicuous and usually only to be seen at the base of the fruits near the insertion of the stalk (lens!). Fruits ellipsoid, sometimes globose in H. sinclairii.] 16
10a Leaf bud, twig apex, and inflorescences with hairs 0.3-1.5 mm long. Leaves 17-45 cm long, often ± parallel-sided; nerves 30-40 pairs. [Fruits ellipsoid, 3.4-5.5 cm long, glabrous.] — Moluccas and W & C New Guinea H. sylvestris
b Leaf bud, twig apex, and inflorescences with hairs 0.2 mm long or less; hairs in H. moluccana and H. tuberculata 0.1-0.3 mm long. Leaves generally smaller; nerves fewer 11
11a Fruits globose or subglobose, [not beaked and without pseudostalk], 1.4 cm diam. or less 12
b Fruits ellipsoid, 1.3 cm long or more 13
12a Fruits brown on drying; pericarp 1.5-3 mm thick. Buds cleft nearly to the base. — Northern parts of Papua Barat and Papua New Guinea H. basifissa
b Fruits blackish on drying; pericarp 1 mm thick. Buds cleft c. 1/3. — Aru Is., whole of New Guinea H. subtilis var. subtilis
13a Fruits to 2 cm long, blackish on drying; apex pointed/beaked or not, base without or with long or short pseudostalk 14
b Fruits 1.3-3.7 cm long, (dark) brown on drying; apex rounded, base mostly without pseudostalk 15
14a Pseudostalk of fruit (1.5-)2-6 mm long. — Papua Barat (Jayapura), Papua New Guinea (W Sepik Prov.) H. schlechteri
b Pseudostalk absent or up to 3 mm long.—Whole of New Guinea H. subtilis
15a Buds 2 mm long, cleft 1/2-4/5. Fruits 1.3-2.8 cm long, pericarp 1-2 mm thick. — Moluccas, W New Guinea H. moluccana
b Buds 2-3 mm long, cleft 1/2-2/3. Fruits 1.5-3.7 cm long, pericarp 1-8 mm thick. — Papua New Guinea (Milne Bay Prov., Bismarck Archipelago, Papuan Islands) H. tuberculata
16a Leaf bud, twig apex, and inflorescences with hairs 0.2 mm long or less (0.1-0.3 mm long in H. psilantha) 17
b Leaf bud, twig apex, and inflorescences with hairs 0.5-1 mm long (0.2-0.5 mm long in H. ampliformis; indumentum not known in H. ampla) 27
17a Hairs 0.1-0.3 mm long. Infructescences and female inflorescences large, much branched, 10-16 cm long. Fruits 1.7-2.2 cm long, pericarp 1-2 mm thick. — Papua New Guinea (Bismarck Archipelago, Bagabag I., Long I.). H. psilantha
b Hairs 0.1-0.2 mm long, or less. Infructescences and female inflorescences 10 cm long or less. Fruits variable 18
18a Fruits 1.6 cm long or less; pericarp 1-3 mm thick. Buds pubescent 19
b Fruits 1.5 cm long or more; pericarp 2 mm thick or more; if fruit 1.5 cm long, then almost globose and buds glabrous 24
19a Fruits at apex rounded, not apiculate; pseudostalk absent H. pilifera
b Fruits apiculate; pseudostalk to 5 mm long 20
20a Leaves broadly obovate to oblong, 12-20 by 5-11 cm. Pedicel widening to above and gradually passing into the bud. Fruits (including 1 mm long pseudostalk and 2 mm long apiculum) 1.4 by 0.8-0.9 cm, fruiting pedicel 9-14 mm long, distinctly tapering H. crux-melitensis
b Leaves elliptic to lanceolate, 4.5-20 by 0.7-6 cm. Pedicel clearly marked off from the bud. Pseudostalk of fruits 1.5-5 mm; fruiting pedicel up to 10 mm long, not or but little tapering 21
21a Pseudostalk of fruits 5 mm long. [Fruits excluding pseudostalk but including 2-3 mm long apiculum 1.6-1.7 by 0.9-1 cm.] H. squamulosa
b Pseudostalk of fruits 1.5-3 mm long 22
22a Leaf bud, twig apex, and inflorescences with hairs 0.3 mm. Tertiary venation of leaves below coarse and distinct. [Female flowers not known; fruits not known with certainty; male buds subglobose, with thick perianth.]. H. urceolata
b Leaf bud, twig apex, and inflorescences with hairs 0.1-0.2 mm. Tertiary venation generally less distinct. [Male perianth different.] 23
23a Fruits excluding the 1.5-2.5 mm long pseudostalk, but including the 0.5-2 mm long apiculum, 1.2-1.5 by 0.8-1 cm. Male buds clearly marked off from the pedicel H. coryandra
b Fruits excluding the 1.5-2 mm long pseudostalk, but including the 2 mm long apiculum, 1.3 by 1 cm long. Male pedicel broadening to above and gradually passing into the bud H. clavata
24a Buds 2-2.4 mm long, glabrous. Leaves 6-14 cm long. [Fruits (sub)globose, short- ellipsoid, or obovoid, 1.5-2.5 by 1.5-2 cm; pericarp 4-6 mm thick.] — E Papua New Guinea H. sinclairii
b Buds 2.5 mm long or more, pubescent or glabrescent. Leaves 10 cm long or more 25
25a Fruits 1.6-3 cm long, usually with coarse pale wart-like lenticels; (dry) pericarp 2-6 mm thick (sometimes much resembling small-fruited H. pachycarpa). — Moluccas, whole of New Guinea including Bismarck Archipelago; 0-1000 m. H. laevigata
b Fruits (3-)3.5-7.5 cm long; pericarp (4-)5 mm thick or more. — New Guinea; (450-) 1000-2000 m 26
26a Buds pubescent. Fruits 3-4.5 cm long; pericarp 4-10 mm thick H. pachycarpa
b Buds glabrescent. Fruits 6-7.5 cm long; pericarp 10-20 mm thick H. corrugata
27a Leaf bud and inflorescences with hairs 0.2-0.5 mm long. Fruits not known. [Leaf bud, female flowers, and fruits not known in H. ampla.] 28
b Leaf bud and inflorescences with hairs 0.5—1(—1.5) mm long. Fruits usually conspicuously pubescent 29
28a Inflorescences glabrescent. — Papua New Guinea (Sepik Prov.). H. ampla
b Inflorescences pubescent. — Papua New Guinea (Sepik and Morobe Prov.) H. ampliformis
29a Leaves coriaceous, beneath with harsh hairs, when shed leaving thickened scars. Buds 4 mm long, opening with narrow pore-like slit. [Fruits 3-5 cm long, pericarp 4-7 mm thick.] — New Guinea H. pulverulenta
b Leaves membranous or chartaceous. Buds cleft 1/4-1/2 30
30a Flowers [only the male known] entirely pubescent. [Fruits 2-2.4 cm long, pericarp 4-7 mm thick.] — New Guinea H. leptantha
b Flowers largely glabrescent 31
31a Leaves generally oblong-lanceolate, at apex caudate. Buds 4 mm long. Fruits 2.5-3 cm long. — Papua New Guinea (New Britain) H. ralunensis
b Leaves oblong(-lanceolate), at apex not caudate (always?). Buds 3 mm long. Fruits 1.2-2.8 cm long. — Most of Papua New Guinea (incl. New Britain and New Ireland) H. hellwigii

Three sections can be recognized and are supposedly of unequal taxonomic weight but with significant different ranges of distribution (see above): 1) sect. Horsfieldia, containing one single species, the type species of the genus, rather deviating from all other species, 2) sect. Irya, containing most species with predominantly a 2-lobed perianth, and 3) sect. Pyrrhosa, most of its species with predominantly a 3- or 4-lobed perianth. The descriptions of the three sections have been given here separately and are not included in the treatment of the species, which are all listed alphabetically. [For an extensive discussion of the subdivision of Horsfieldia into the three sections, see W.J. de Wilde ('1984', 1985) 125-136]

Plate 1.

Legends to Plates 1-3:

Semi-schematic drawings of the androecia of most species of Horsfieldia, except H. ampla[1], H. discolor[21], H. disticha[22], H. perangusta[59], and H. sessilifolia[75], of which no male flowers are available.

Between square brackets the number has been given of the alphabetically arranged species of the present revision.

Lateral view (left), longitudinal section (right), apical view (top); white: anthers; black: sterile tissue (i.e., androphore and central column).

Magnification for 3-11, 15-19, 21-27, 29- 40, 45- 48, 50, 52, 74, 75, 81 = x 5; Magnification for 1, 2, 12-14, 20, 28, 41-44, 49, 51, 53-73, 76-80, 82-92a, b = x 10.

1: H. iryaghedhi[36] — 2: H. irya[35] — 3: H. spicata[79] — 4: H. inflexa[33] — 5: H. moluccana var. moluccana[44a] — 6: H. parviflora[56] — 7: H. ob- scurinervia[49] — 8: H. ardisiifolia[6] -9: H. talaudensis[88] -10: H. samarensis[72] -11: H smithii[77] -12: H olens[51] — 13: H sepikensis[74] — 14: H. syl- vestris[87] -15: H. coryandra[16] -16: H. urceolata[94] -17: H. crux-meliten- sis[19] -18: H. clavata[13] -19: H. squamulosa[81] — 20: H. ampliformis[2] — 21: H. angularis[5] — 22: H. iriana[34] — 23: H. aruana[7] — 24: H. subtilis var. subtilis[84a] — 25: H. schlechteri[73] — 26: H. basifissa[9] — 27: H. sinclairii[76] -28: H. psilantha[62] — 29: H. laevigata var. laevigata[37a] — 30: H. pilifera[60] — 31: H. lancifolia[38].

Plate 2.

32: H. decalvata[20] — 33: H. tuberculata[93] — 34: H. corrugata[15] — 35: H. pachycarpa[53] — 36: H. pulverulenta[64] — 37: H. leptantha[40] — 38: H. hellwigii var. hellwigii[31a] — 39: H. ralunensis[67] -40: H. sabulosa[71] -47; H. atjehensis[8] — 42: H. sucosa subsp. sucosa[85a] — 43: H. pallidicaula var. pallidicaula[55a] — 44: H. sparsa[78] — 45: H. triandra[91] — 46: H. tristis[92] — 47: H. fulva[27] — 48: H superba[86] — 49: H grandis[30] — 50: H wallichii[96] — 51: H. pulcherrima[63] — 52: H. flocculosa[25] — 53: H. motleyi[46] — 54: H. tomentosa[90] — 55: H. cf gracilis[29] — 56: H. paucinervis[57] — 57: H. splendida[80] — 58: H. rufo-lanata[70] — 59: H reticulata[68] — 60: H. crassifolia[18] — 61: H carnosa[12].

Plate 3.

62: H. sterilis[82] — 63: H hirtiflora[32] — 64: H. brachiata[11] — 65: H pachyrachis[54] — 66: H. ridleyana[69] — 67: H. obtusa[50] — 68: H. tenuifolia[89] — 69: H. macilenta[41] — 70: H. laticostata[39] — 71: H. nervosa[47] -72: H. polyspherula var. polyspherula[61a] — 73: H. oligocarpa[52] — 74: H. endertii[24] — 75: H valida[95] — 76: H. borneensis[10] — 77: Hfragillima[26] — 78: H androphora[4] — 79: H. amplomontana[3] — 80: H. montana[45] — 81: H. elongata[23] — 82: H. punctata[65] — 83: H. costulata[17] — 84: H. subalpina subsp. sub- alpina [83a] — 85: H. obscura[48] — 86: H. xanthina subsp. xanthina[97a] — 87: H. majuscula[43] — 88: H. coriacea[14] — 89: H. penangiana[58] — 90: H. punc- tatifolia[66] — 91: H. macrothyrsa[42] — 92a: H. glabra var. glabra[28a] — 92b: H. glabra var. javanica[28b].

Section Horsfieldia

Myristica Gronov. sect. Horsfieldia A. DC. - Prodr. 14 1 (1856) 200

Myristica Gronov. sect. Horsfieldia A. DC. - Miq. Fl. Ind. Bat. 1 2 (1858) 63

Myristica Gronov. sect. Irya auct. non Hook. f. & Thomson: Benth. & Hook, f. - Gen. PL 3 (1880) 137, for Horsfieldia only.

Myristica Gronov. sect. Eumyristica Hook. f. & Thomson sub sect. Horsfieldia (A. DC.) King - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 282

Horsfieldia odorata Willd.

Myristica Gronov. sect. Pyrrhosa Blume - Rumphia 1 (1835) 190, p.p., for Myristica horsfieldii only, not the lectotype species.

Myristica Gronov. sect. Eumyristica Hook. f. & Thomson - Fl. Ind. (1855) 162, p.p., for Myristica horsfieldii only.

Horsfieldia Willd. sect. Orthanthera Warb. - Mon. Myrist. (1897) 268, p.p., for the lectotype species only.

Horsfieldia Willd. sect. Trivalves J. Sinclair sub sect. Orthanthera (Warb.) J. Sinclair - J. Sinclair Gard. Bull. Sing. 16 (1958) 371, p.p., nom inval., provisional name onl

Lectotype species: Horsfieldia iryaghedhi (Gaertn.) Warb.

Phyllotaxis of plagiotropic shoots distichous. Lower leaf surface with alveolar tissue, epidermis papillose, stomatal complex sunken; without larger dark-coloured dots (cork warts). Flowers in male inflorescences sessile, at base not articulated, arranged in many- flowered subglobose dense heads; perianth in buds elongate-obconical, ± angled, cleft 1/2-3/4, 3- or 4-(in female: 2- or 3-)lobed. Androecium elongate, subcylindrical, androphore distinct; column narrow, narrowly hollowed for over halfway; anthers 3-5, erect. Stigma sessile, many-lobulate.

One species, Sri Lanka Map 3 (p. 4).

Section Horsfieldia is monotypic, the species H. iryaghedhi deviating from all other Horsfieldias by some anatomical characters of the leaf, male flowers sessile and arranged in dense heads with a thick receptacle, angular buds, anthers largely connate, but not back to back so that a narrowly hollowed central column is formed; the stigma in the female flowers is many-lobed, not 2-lobed as in the other Horsfieldia species.

Section Irya

Horsfieldia Willd. sect. Irya Hook.f. & Thomson Warb. - Mon. Myrist. (1897) 123, 267, p.p.

Horsfieldia Willd. sect. Irya Hook.f. & Thomson Warb. - WJ. de Wilde Gard. Bull. Sing. 37 2 (‘1984’, 1985) 127

Myristica Gronov. sect. Irya Hook. f. & Thomson - Fl. Ind. (1855) 159

Myristica Gronov. sect. Irya Hook. f. & Thomson - A. DC. Prodr. 14 1 (1856) 202

Myristica Gronov. sect. Irya Hook. f. & Thomson - Miq. Fl. Ind. Bat. 1 2 (1858) 64

Myristica Gronov. sect. Irya Hook. f. & Thomson - Benth. & Hook, f. Gen. PL 3 (1880) 137, p.p., excl. sect. Horsfieldia

Myristica Gronov. sect. Irya Hook. f. & Thomson - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 284, p.p., for the smaller part only.

Horsfieldia Willd. sect. Irya (Hook. f. & Thomson) Warb. sub sect. Euirya Warb. - Warb. Mon. Myrist. (1897) 123, 267, p.p., for the type species only.

Horsfieldia irya (Gaertn.) Warb.

Myristica irya Gaertn.

Myristica Gronov. sect. Pyrrhosa Blume - Rumphia 1 (1837) 190, p.p., for Myristica javanica and a few other species only, excl. lectotype species Myristica glabra (= sect. Pyrrhosa) and Myristica horsfieldii (= Horsfieldia iryaghedhi, sect. Horsfieldia)

Myristica Gronov. sect. Pyrrhosa Blume - A.DC. Prodr. 14 1 (1856) 202, p.p.

Myristica Gronov. sect. Pyrrhosa Blume - Miq. Fl. Ind. Bat. 1 2 (1858) 64, p.p., excl. Myristica glabra (= sect. Pyrrhosa).

Horsfieldia Willd. sect. Pyrrhosa Blume Warb. sub sect. Bivalves Warb. - Warb. Mon. Myrist. (1897) 262, (incl. series Smithii and series Globularia).

Horsfieldia Willd. sect. Bivalves J. Sinclair - Gard. Bull. Sing. 16 (1958) 370, 371, comb. inval., provisional name only.

Phyllotaxis in plagiotropic shoots distichous. Lower leaf surface without alveolar tissue, epidermis not papillose, stomatal complex not sunken; rarely (H. inflexa) with larger dark-coloured dots. Flowers pedicelled, not articulated at base, solitary or in loose clusters; buds rarely globose or obovoid, usually laterally compressed, in outline circular, elliptic, or pear-shaped, not angular (in young stages of H. sylvestris faintly so), cleft to variable depths, perianth mostly predominantly 2-lobed, rarely 3-lobed. Androecium various, often ± zygomorphic, laterally compressed, sometimes anthers at apex bi-laterally incurved, or obconical and ± actinomorphic; androphore distinct or not, central column broad or narrow, little to much hollowed; anthers few to many, (sub)-erect, or at apex incurved into apical hollow of the column. Stigma minutely 2-lobed.

Mainly E Malesia (including the Philippines), and one species, H. irya, distributed over almost the whole area of the genus. Map 3 (p. 4).

Almost all of the 40 species of this section have predominantly 2-lobed perianths, with a more or less zygomorphic androecium, because it is laterally compressed or with the anthers at apex incurved from two sides into an apical hollow of the column.

Aberrant are H. olens and H. sepikensis with 3- or 4-lobed perianths, but with the androecium tending to be zygomorphic. Horsfieldia angularis has 2-4-lobed perianths.

Also aberrant are the species of the group of H. clavata, with a 2-lobed perianth but a club-shaped non-zygomorphic androecium.

A few mutually related species from continental SE Asia, placed in section Pyrrhosa, viz. H. longiflora, H. thorelii and H. amygdalina, have (partly) a 2-lobed perianth, and a zygomorphic androecium, especially H. longiflora. They blur the distinction between sections Irya and Pyrrhosa. Section Irya occurs mainly in East Malesia, with only H. irya extending far beyond the main range of distribution of the section. Within section Irya, eight groups of species can be distinguished, a survey of which is given by De Wilde ('1984', 1985: 128).

Section Pyrrhosa

Horsfieldia Willd. sect. Pyrrhosa Blume Warb. - Mon. Myrist. (1897) 262, p.p.

Horsfieldia Willd. sect. Pyrrhosa Blume Warb. - W.J. de Wilde Gard. Bull. Sing. 37 2 ('1984', 1985) 130

Myristica Gronov. sect. Pyrrhosa Blume - Rumphia 1 (1837) 190, t. 62- 64, p.p. for the smallest part incl. the lectotype t. 64 f. 1A, B

Myristica Gronov. sect. Pyrrhosa Blume - Hook. f. & Thomson Fl. Ind. (1855) 160

Myristica Gronov. sect. Pyrrhosa Blume - A. DC. Prodr. 14 1 (1856) 202, p.p.

Myristica Gronov. sect. Pyrrhosa Blume - Miq. Fl. Ind. Bat. 1 2 (1859) 64, p.p.

Myristica Gronov. sect. Pyrrhosa Blume - Benth. & Hook.f. Gen. pl. 3 (1880) 136

Myristica Gronov. sect. Pyrrhosa Blume - King Ann. Roy. Bot. Gard. Calc. 3 (1891) 282

Horsfieldia Willd. sect. Pyrrhosa Blume Warb. subsect.EupyrrhosaWarb. - Mon. Myrist. (1897) 265, (excl. Horsfieldia macrocoma = Endocomia).

Horsfieldia glabra (Blume) Warb.

Lectotype species: Myristica glabra Blume

Myristica Gronov. sect. Eumyristica Hook.f. & Thomson - Fl. Ind. (1855) 162, p.p., for Myristica superba = Horsfieldia superba (Hook. f. & Thomson) Warb. only.

Myristica Gronov. sect. Caloneura A. DC. - Prodr. 14 1 (1856) 192 p.p., for Myristica superba Hook.f. & Thomson only [= Horsfieldia superba (Hook. f. & Thomson) Warb.].

Myristica Gronov. sect. Irya auct. non Hook. f. & Thomson: King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 284, p.p.

Horsfieldia Willd. sect. Irya (Hook. f. & Thomson) Warb. sub sect. Euirya Warb. - Warb. Mon. Myrist. (1897) 267, p.p., excl. Horsfieldia irya (type species of sect. Irya)

Horsfieldia Willd. sect. Irya Hook.f. & Thomson Warb. sub sect. Trivalves Warb. - Warb. Mon. Myrist. (1897) 267

Horsfieldia Willd. sect. Trivalves J. Sinclair sub sect. Trivalves - J. Sinclair Gard. Bull. Sing. 16 (1958) 370, 371, comb.inval., provisional name only.

Horsfieldia Willd. sect. Orthanthera Warb. - Mon. Myrist. (1897) 268 p.p., for Horsfieldia ralunensis and Horsfieldia sylvestris only, excl. the lectotype species Horsfieldia iryaghedhi.

Phyllotaxis of plagiotropic shoots dispersed or distichous. Lower leaf surface without alveolar tissue, epidermis not papillose, stomatal complex not sunken; with or without larger dark-coloured dots (or cork warts). Flowers with a pedicel (short in H. wallichii), at base with or without articulation, solitary or in loose clusters; buds (depressed) globose, obovoid, or ellipsoid, not laterally compressed, not angular, cleft c. 2/3 or less; perianths predominantly 3- (or 4-)lobed (rarely 2-lobed, H. longiflora, H. sterilis). Androecium (sub)circular or more or less triquetrous in cross section, never laterally compressed, depressed-globose, ellipsoid, or obovoid, usually with a broad central column with an apical hollow of variable shape and depth; androphore usually narrow at base; anthers few to many, ± straight or curved, almost entirely connate or free for about the upper half (H. polyspherula-group). Stigma minutely 2-lobed.

Continental SE Asia, West Malesia (including the Philippines). Map 3 (p. 4).

Section Pyrrhosa contains c. 60 species, mainly with exclusively or predominantly a 3- (or 4-)lobed perianth; species with a 2-lobed perianth are H. longiflora (Vietnam), H. crassifolia (Sumatra, Peninsular Malaysia, Borneo) and H. sterilis (Sabah). For a brief discussion see De Wilde ('1984’, 1985: 132).

Horsfieldia ampla Markgr.

Horsfieldia ampla Markgr. - Bot. Jahrb. Syst. 67 (1935) 148

Horsfieldia ampla Markgr. - W.J. de Wilde Gard. Bull. Sing. 38 1 (1985) 95

Type: Ledermann 9639, (B, lost), Papua New Guinea, Sepik Prov.

Small tree 4-5 m. Twigs terete. Leaves cuneate-obovate, up to 40 by 16 cm, base ± attenuate, apex short-acuminate; nerves 16-18 pairs, straight, sharply raised beneath and connected towards the margin; petiole 1 cm long. Inflorescences on the older wood, to 25 by 10 cm, glabrescent, loosely flowered. Male flowers yellow, clavate, 4 by 2 mm (excluding pedicel?), the perianth 2-lobed, cleft to hardly 1/4. Staminal column thick; thecae to c. 20, the androphore about as long as the anthers or slightly shorter.

Distribution Malesia: NE Papua New Guinea (Sepik Prov., 'Aprilfluss'; mountain slope near camp 18).

Habitat & Ecology Dense, very humid forest, on mountain slope at 200-400 m altitude; male fl. Nov. 1912.

Notes 1 Known only from the type, now destroyed. Keyed out by Markgraf with crux-melitensis, both having clavate flowers. He mentions in the key that the perianth (not the androecium) is largely hollow, and in a note that the species is peculiar amongst the New Guinea Horsfieldias because of its large male flowers, which in other species are smaller and almost always broader than long, and that it is without close relatives.

2 The species is possibly related to or may be identical with H. ampliformis.

Horsfieldia ampliformis W. J. de Wilde

Horsfieldia ampliformis W. J. de Wilde - Gard. Bull. Sing. 38 1 (1985) 95, f. 14

Type: Hoogland & Craven 11085, (male fl.), New Guinea, Sepik Prov.

Tree 5-8 m. Twigs when young narrowly ridged, 4-7(-10) mm diameter, early or late glabrescent, with ± woolly hairs 0.2-0.5 mm long; bark coarsely striate, lenticellate, not flaking. Leaves thickly membranous, (elliptic-)oblong, (18—)25—38 by (6-)7-13 cm, base short to long-attenuate, apex attenuate-acuminate; upper surface drying dark brown, lower surface early or late glabrescent or with rather scattered stellate hairs 0.2-0.5 mm long; dots absent; midrib above ± narrow, flattish; nerves 18-22 pairs, above thin, flat or sunken; lines of interarching beneath not very prominent; venation lax, indistinct on both surfaces; petiole 4-6 by 3-4 mm; leaf bud 25-40 mm long, with hairs 0.2-0.5 mm. Inflorescences below the leaves; in male: many-flowered, 4 to 5 times branched, 25-35 by 20-30 cm, peduncle 4-5 cm; in female: 9-10 by 6-8 cm; all branches with rather loose hairs 0.2-0.5 mm long; bracts (seen only in female) 5 mm long, caducous. Flowers 2-5 together in male and female, with loose hairs (0. l-)0.2-0.3 mm long, in female glabrescent towards apex; perianth 2-lobed; pedicel not articulated. Male flowers: pedicel 2-4 mm long, buds largely hollow, broadly obovoid, laterally ± flattened, 3-3.3 by 3-3.2 mm, apex obtuse to broadly rounded, base shortly tapering, cleft c. 1/3, lobes 0.3 mm thick; androecium small, ± flattened, 2.5 by 1-1.2 mm, apex broadly rounded, synandrium 1.5-1.8 by 1-1.2 mm, narrowly hollowed for 1/5-1/3 at apex (Plate 1: 20); thecae 14, free apices 0.1-0.2 mm; androphore 0.8-1 by 0.5-0.6 mm. Female flowers: pedicel 1-2 mm long, buds broadly ovoid, 3 by 2.6-2.8 mm, cleft c. 2/3; ovary broadly ovoid, 2-2.2 by 1.8-2 mm, densely pubescent with hairs 0.1 mm or less, stigma short, not or hardly lobed, 0.1 by 0.4 mm. Fruits not seen.
See: Fig. 8.

Field-notes Small tree, 8 m high. Flowers medium green, yellow at anthesis.

Distribution Malesia: northern Papua New Guinea (Sepik, Morobe Prov.).

Habitat & Ecology Lower montane rain forest; 1200-1300 m altitude; fl. Apr., Aug.

Notes 1Horsfieldia ampliformis is close to H. ampla, of which no material has been seen. According to the description the latter differs by the more elongate, possibly glabrous perianths, 4 by 2 mm, the androecium with 20 thecae, and the glabrescent inflorescences. The two species have a peculiar long-stalked androecium and larg, male inflorescences. Horsfieldia ampla was collected at only 200-400 m altitude.

2 Known only from a male and a female flowering specimen. The perianths of the female specimen, Craven & Schodde 1463 (Morobe Prov.), are glabrescent in the upper half; it could be H. ampla. Moreover, as the hairs on the leaf buds are slightly shorter than those of the male specimen, this specimen is difficult to distinguish from the variable and widespread H. laevigata.

Horsfieldia amplomontana W.J. de Wilde

Horsfieldia amplomontana W.J. de Wilde - Gard. Bull. Sing. 39 1 (1986) 34

Horsfieldia amplomontana W.J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 361

Type: Clemens 30536, Sabah.

Tree 10-20 m. Twigs 3.5-6(-10) mm diameter, early to rather late glabrescent, hairs 0.3—1(—1.5) mm long; bark coarsely striate, not flaking, lenticels small, not contrasting in colour and inconspicuous. Leaves membranous to chartaceous, elliptic-oblong to oblong-lanceolate, 15-35 by 5-11 cm, base short-attenuate to narrowly rounded, apex acute- acuminate; upper surface glabrescent, except towards the base of the midrib in young leaves, olivaceous to brown, lower surface glabrous (glabrescent), without dots or hair scars, sometimes pale and contrasting with upper surface; midrib slender, ± raised above, nerves 11 or 12 pairs, above flattish or raised, lines of interarching not distinct; venation lax, ± distinct or not; petiole 8-15 by 2.5-3.5 mm, glabrescent; leaf bud 15-22 by 3-4 mm, with dense rusty hairs 0.5—1(—1.5) mm long. Inflorescences behind the leaves, with shaggy rusty hairs 0.5 mm; in male: 3 or 4 times branched, many-flowered, 10-21 by 10-16 cm, peduncle 1.5-4 cm long, in male in loose clusters of 5-10; in female (in fruit): 7-8 cm long; bracts broadly ovate-ellipsoid, 2-7 mm long, finely pubescent, caducous; flowers glabrous, perianth 3- (or 4-)lobed, pedicel sometimes ± articulated (see note 2). Male flowers: pedicel 0.8-1.5(-2) mm long, buds (depressed-)globose, 1.5-2 by 2-2.3 mm, apex and base (broadly) rounded, glabrous, cleft 1/2-2/3, lobes 0.2 mm thick, not or only slightly collapsing on drying; androecium depressed-globose, 0.6-1 by 1.1- 1.8 mm, apex broadly rounded, base rounded or sagged (Plate 3: 79); thecae 20-26, almost completely sessile, 0.8-1.2 mm long, free apices up to 0.1 mm, incurved, concealing a ± 3-radiate apical slit or cavity 0.2-0.5 mm deep; column broad, solid; androphore rather narrow, (0.1-)0.2-0.4 mm long, completely or partly hidden by the anthers. Female flowers (from fruit): 3-lobed, 3 mm long. Fruits 1-3 per infructescence, ellipsoid, apex and base rounded, 7-8 by 4.5-5 cm, glabrous, drying dark brown, finely to coarsely tubercled, pericarp 15 mm thick; fruiting pedicel 3 mm long; perianth persistent.

Field-notes Large tree. Bark grey, fissured; outer bark soft, 5 mm thick; inner bark white, soft, 5 mm; cambium pale; sap wood white; exudate from bark sticky. Flowers golden. Ripe fruits orange.

Distribution Malesia: Borneo (Sabah: Mt Kinabalu).

Habitat & Ecology Primary and degraded forest, ridge forest; on sandstone; 1000- 1500 m altitude; fl. Nov., Dec., Feb.; fr. Nov.

Notes 1 Close to H. montana, both having very similar male flowers, but H. amplomontana differs considerably by its stouter twigs, larger leaves, larger male inflorescences, and very much larger fruits with a thick pericarp; in H. montana the fruit is only 2-2.7 cm long, and the perianth is not persistent.

2 The pedicels are generally not articulated, although some flowers of SAN 18843 seem to have an articulation, but this may be an artefact caused by drying.

Fig. 8.

Horsfieldia ampliformis W. J. de Wilde, a. Twig apex with leaves; b. twig with male inflorescence axillary to fallen leaf; c. mature male flower, perianth opened, showing androecium; d. twig with female inflorescence; e. female flower, opened, showing finely pubescent ovary and minute 2-lobed stigma [a-c: Hoogland & Craven 11085; d, e: Craven & Schodde 1463]. — Scale bar for a, b, d = 2 cm; for c, e = 1.7 mm.

Fig. 9.

Horsfieldia androphora W.J. de Wilde. a. Branch with leafy twig and male inflorescence; b. mature male flower; c. ditto, longitudinally opened, showing androecium; d. androecium, longitudinal section, schematic; e. twig with infructescence, fruits mature, aril complete [a-d: Nooteboom & Chai 01710, type; e: Sinclair, Kadim & Kapis 8977].— Scale bar for a, e = 2 cm; for b-d = 0.85 mm.

Horsfieldia androphora W. J. de Wilde

Horsfieldia androphora W. J. de Wilde - Gard. Bull. Sing. 39 1 (1986) 32, f. 30

Horsfieldia androphora W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 362

Type: Nooteboom & Chai 01710, Sarawak.

Tree 7-20 m. Twigs 2-4(-5) mm diameter, with rusty hairs 0.3-0.6 mm, rather late glabrescent; bark blackish brown, finely striate, not flaking; lenticels small, inconspicuous. Leaves membranous, elliptic to oblong, 9-18 by 3.5-6.5 cm, base attenuate, apex acute-acuminate; upper surface drying dark brown or blackish brown, glabrous, lower surface glabrescent (except midrib); dots absent; midrib above raised, beneath with some vestigial indumentum or late glabrescent; nerves 9-13 pairs, raised above, lines of inter- arching on the lower surface irregular and not very conspicuous; venation lax, distinct or not; petiole 10-12 by 1.5-2 mm, glabrescent; leaf bud 8-12 by 2-3 mm, with rusty hairs 0.3-0.6 mm. Inflorescences with ± dense hairs 0.2-0.6 mm; in male: rather many-flowered, 3 (or 4) times branched, 6-14 by 3.5-9 cm, peduncle 1-2 cm long; in female (from infructescences): 3-4 cm long; bracts densely short-pubescent, ovate-elliptic, ± acute, 3 mm long, caducous; flowers (male) in clusters of 2-6 each, perianth 3-lobed, glabrous, pedicel glabrous or with a few minute hairs 0.1 mm at the very base, not articulated. Male flowers: pedicel 0.5-2 mm, slender; buds globose, 1.4-2(-2.2) mm diameter, cleft 1/3 to nearly 1/2, lobes 0.2 mm thick; synandrium depressed-globose, somewhat flattened or impressed at apex and/or base, in cross section rounded, (0.6-) 0.8-1 by (0.8—)1—1.3 mm, androphore slender, (0.3-)0.4-0.8 mm long (Plate 3: 78); thecae 14-22, almost completely connate, incurved, concealing the apical hollow, 0.2-0.3 mm deep. Female flowers not seen. Fruits 2-5 per infructescence, ellipsoid, 2.4-3 by 1.4-2 cm, apex subacute to rounded, base rounded or shortly narrowed, glabrous, drying dark brown, finely tuberculate, without lenticels, pericarp 1.5-2 mm thick; fruiting pedicel 2 mm long; perianth not persistent.
See: Fig. 9.

Field-notes Bark chocolate to reddish brown, narrowly cracked, longitudinally furrowed, or cut into rectangular blocks; sap watery, more or less colourless (tree in flower), or blood red (tree in fruit). Twigs chocolate, with rusty hairs. Flowers yellow. Fruits smooth, orange, testa whitish grey.

Distribution Malesia: Borneo (Sarawak, Sabah).

Habitat & Ecology Montane forest, mossy forest, wooded sandstone ridges, 800- 1200 m altitude; fl. Mar., Oct.; fr. Mar., June.

Notes 1 Regarding the general morphology of the androecium, the present species belongs to the H. grandis group. It seems closest to H. tomentosa from S Thailand and Peninsular Malaysia, the two having a long-stalked synandrium in common. Horsfieldia tomentosa has generally larger flowers, a pubescent lower leaf surface, and smaller fruits which are pubescent or glabrescent. Horsfieldia androphora grows in mountains; H. tomentosa is restricted to lowlands.

2Horsfieldia androphora keys out beside H. fragillima (also with non-articulated pedicels), but the latter species differs in many characters including general habit and fruit size; its saucer-shaped androecium is quite different.

Horsfieldia angularis W. J. de Wilde

Horsfieldia angularis W. J. de Wilde - Gard. Bull. Sing. 38 1 (1985) 97

Type: BW 5828, (male fl.) New Guinea, Bird's Head.

Tree 15-30 m. Twigs 2-angular, lower down subterete, with two ridges, 3-7(-10) mm diameter, early glabrescent, hairs grey-brown, 0.1 mm or less; bark striate, distinctly coarsely lenticellate, not flaking. Leaves membranous to thinly chartaceous, oblong (-lanceolate), 10-27 by 3-7.5 cm, base attenuate, apex acute-acuminate; upper surface drying pale to dark brown, often finely paler pustulate, lower surface glabrescent, hairs very minute, grey, stellate, less than 0.1 mm; dots absent; midrib slightly raised above; nerves 12-15 pairs, above thin and flattish or slightly sunken, lines of interarching not distinct; venation lax, faint; petiole 7-15 by 2-3 mm; leaf bud 10-15 by 2-2.5 mm, with hairs 0.1 mm. Inflorescences with rather dense hairs 0.2-0.3 mm; in male and female: 2 or 3 times branched, rather few-flowered, 3-4 by 2-2.5 cm, peduncle 0.3-0.6 cm long; bracts not seen, caducous; flowers (male) generally 2-4 together; perianth 2-4-lobed in male, 2(-3)-lobed in female, in the lower half with hairs 0.1 (-0.2) mm long; pedicel pubescent, not articulated. Male flowers: pedicel not tapering, 1-2 mm long; buds in lateral view circular or slightly transversely ellipsoid, not or only slightly laterally compressed, not collapsing on drying, 1.7-2.3 by 2.2-3.2 mm, cleft c. 9/10, lobes (0.2-)0.3 mm thick; androecium slightly laterally flattened (in 3- or 4-lobed flowers about 3- or 4- angular in cross section), above broadly rounded, 1.2-1.5 by 1.5-2.2 mm (Plate 1: 27); thecae 24 to c. 40 (in 4-lobed flowers), ± erect, free parts at apex to 0.1 mm, central column at apex narrowly hollowed for (1/3—) 1/2; androphore absent, the androecium ± broadly attached. Female flowers: pedicel 1-1.5 mm long; buds depressed globose, 2.5 by 3-3.2 mm, cleft c. 3/4; ovary ± depressed globose-ovoid, 1.2 by 1.5 mm, densely short-pubescent, style and stigma minutely 2-lobed, 0.1 by 0.3 mm. Fruits 5-10 per infructescence, short-ellipsoid, 1.7-2 by 1.4-1.7 cm, pubescent at very base, with coarse paler-coloured lenticel-like tubercles; pericarp thick-woody, 3-5 mm thick; fruiting pedicel 3-5 mm long; perianth not persistent.

Field-notes Sometimes buttressed to 1 by 0.5 m; bark sometimes fissured, or peeling off in small scales; with red exudate; sapwood pale brown or white; heartwood not discernible or pinkish. Flowers greenish. Fruits yellow(-brown), sour and edible.

Distribution Malesia: Papua Barat (Bird's Head, subprov. Manokwari).

Habitat & Ecology Primary forest; on clayey soils; locally common on the coastal plain up to 600 m in Kebar Valley; 0-600 m altitude; fl. Feb., Aug.; fr. Feb., Oct.

Note Much related to H. basifissa, of which sterile specimens are difficult to identify since their twigs too are rather ridged. Horsfieldia angularis is distinguished from H. basifissa by 1) the more strongly ridged and somewhat stouter twigs, 2) the more hairy and 2-4-lobed flowers with thicker lobes, 3) the hairy ovary and the thinly pubescent ellipsoid fruits. Both species have thickish, subglobose male buds, which hardly collapse on drying, and which at anthesis are cleft to the base.

Horsfieldia ardisiifolia (A. DC.) Warb.

Horsfieldia ardisiifolia (A. DC.) Warb. - Mon. Myrist. (1897) 274

Horsfieldia ardisiifolia (A. DC.) Warb. - J. Sinclair Gard. Bull. Sing. 28 (1975) 3

Horsfieldia ardisiifolia (A. DC.) Warb. - W. J. de Wilde Gard. Bull. Sing. 38 1 (1985) 72, f. 9

Myristica ardisiifolia A. DC. - Ann. Sc. Nat. Bot. 4 4 (1855) 31, t. 4

Myristica ardisiifolia A. DC. - Prodr. 14 1 (1856) 203, 'ardisiaefolia.

Type: Cuming 1702, Philippines.

Horsfieldia warburgiana Elmer - Leafl. Philipp. Bot. 3 (1911) 1061

Horsfieldia warburgiana Elmer - Merr. Enum. Philipp. Flow. PL 2 (1923) 183

Type: Elmer 12297, Philippines.

Horsfieldia gigantifolia Elmer - Leafl. Philipp. Bot. 9 (1925) 3120, 3129

Horsfieldia gigantifolia Elmer - 10 (1939) 3763, nom. nud.

Tree 5-10 m. Twigs flattened 2-angular, lower down terete with two ridges, 3—6(—13) mm diameter, at first with bright rusty hairs 0.3-0.5(-0.8) mm long, early glabrescent; bark smooth to striate, distinctly lenticellate, not flaking. Leaves membranous, (elliptic-)oblong, 20-40 by 5.5-15 cm, base nearly rounded to attenuate, apex acute-acuminate; upper surface drying olivaceous to blackish brown, finely minutely paler pustulate or not, lower surface early glabrescent except for some indumentum remaining on the midrib, hairs coarse 0.3-0.5 mm; dots absent; midrib fairly broad, flattish above; nerves 18—28 pairs, slender above, flattish, lines of interarching regular and distinct beneath; venation lax, inconspicuous; petiole 13-16 by 3-4.5 mm; leaf bud 10-20 by 3-4 mm, with hairs 0.3-0.8 mm long. Inflorescences thinly with stellate-dendroid hairs 0.3 mm; in male: 3 or 4 times branched, rather many-flowered, broadly pyramidal, 7-16 by 6-14 cm, peduncle 0.5-l(-2) cm long; in female: 4-8 cm long; bracts broadly ovate, pubescent, 3-4 mm long, caducous; flowers (male) solitary or 2-4 together, perianth 2-lobed, glabrous, pedicel sometimes at first with sparse hairs, slender, not articulated. Male flowers: pedicel 1-2(-4) mm; buds transversely ellipsoid or reniform, moderately laterally compressed, drying dull, more or less collapsed on drying or not, 2.5-3 by 4-4.5 mm, below sometimes with a basal sinus, cleft 4/5-5/6, the lobes 0.2(-0.3) mm thick; androecium broadly transversely ellipsoid, slightly laterally flattened, hollow, 1.5 by 3- 3.5 mm (Plate 1: 8); thecae (36-)40-48, connate for about halfway, forming a cup with the anthers from one side deeply inflexed, those from the other side for a large part overarching the former; anthers sometimes slightly sagged at base, hiding the narrow androphore, 0.2-0.3 mm long. Female flowers: pedicel 2(-2.5) mm long; buds subglobose- ovoid, 2.5 mm diameter, cleft c. 1/2; the ovary broadly ovoid-subglobose, 1.5-1.7 mm diameter, glabrous, stigma consisting of 2 minute sessile lobes 0.1-0.2 mm. Fruits 2-6 per infructescence, broadly ellipsoid to subglobose, 20-25 by 17-20 mm, glabrous (or possibly with few minute hairs at base), finely rugulose, without marked tubercles, drying (reddish) brown; pericarp 1.5-2 mm thick; fruiting pedicel 3-6 mm long; perianth not persistent.
See: Fig. 10.

Field-notes Flowers yellow, fragrant. Fruits orange-red.

Distribution Malesia: Philippines (Luzon, Mindoro, Sibuyan, Samar, Leyte).

Habitat & Ecology Lowland forest in moist valleys; 0-400 m altitude; fl. & fr. throughout the year.

Note Horsfieldia ardisiifolia is close to species like H. parviflora and H. smithii, both from the Moluccas, all of which have anthers strongly incurved or inflexed into the androecium cup. Horsfieldia ardisiifolia is distinguished by thick winged or ridged twigs, large leaves, coarse hairs on the leaf buds, male buds 4-4.5 mm wide, and a broad androecium with the anthers deeply incurved and clasping each other.

Horsfieldia aruana (Blume) W. J. de Wilde

Horsfieldia aruana (Blume) W J. de Wilde - Gard. Bull. Sing. 38 1 (1985) 100

Palala aruana Rumph. - Herb. Amb. 7 (1755) t. 24

Myristica aruana Blume - Rumphia 1 (1837) 191

Myristica aruana Blume - J. Sinclair Gard. Bull. Sing. 28 (1975) 112, 118, 119, 122-124, in the synonymy of Horsfieldia spicata.

Horsfieldia novo-guineensis Warb. - Mon. Myrist. (1897) 271, t. 23 nom. nov., p.p., for the lectotype only.

Lectotype: those specimens of Zippelius s.n. at L, annotated by Blume, W New Guinea.

Tree c. 15 m. Twigs 2-angled, becoming subterete with two ridges, 3-5 mm diameter, early glabrescent, hairs c. 0.1 mm long; bark striate, not flaking; lenticels small, inconspicuous. Leaves membranous, elliptic-oblong, 15-29 by 5-9.5 cm, base attenuate, apex acute-acuminate; upper surface drying olivaceous to brown, lower surface early glabrescent; dots absent; midrib slightly raised above; nerves 13-15 pairs, slender, flattish; venation lax, indistinct; petiole 10-15 by 1.5-2.5 mm; leaf bud c. 10 by 1.5 mm, hairs c. 0.1 mm long. Inflorescences among or below the leaves, with sparse hairs c. 0.1 mm or less; in male: 3 or 4 times branched, 5-8 by 4-5 cm, rather many-flowered, peduncle 0.5-1.5 cm long; bracts not seen, caducous; flowers (male) in loose clusters of 2-5 each, perianth 2-lobed, glabrous; pedicel slender, sparsely pubescent, not articulated. Male flowers: pedicel 1-1.5 mm long; buds in lateral view circular to somewhat transversely elliptic, laterally compressed, blackish and collapsing (always?) on drying, 1.5-2 by 2-2.5 mm, cleft 2/3-3/4, lobes c. 0.2 mm thick; androecium much compressed laterally, c. 1.5 by 2 mm, above broadly truncate-rounded (Plate 1: 23); thecae 28-36, distal free parts 0-0.1 mm, column almost completely solid; androphore to 0.1 (-0.2) mm. Female flowers and fruits not seen.

Distribution Malesia: SW Papua Barat; possibly also Moluccas (Aru and Tanimbar Islands, see note).

Habitat & Ecology Not known.

Note Specimens perhaps to be included in H. aruana are Buwalda 4969 from the Aru Is. and bb 24414 from the Tanimbar Is.; the male flowers of both are immature. The synandrium of Buwalda 4969 is cleft to c. 1/10 only; however, in bb 24414 it appears cleft nearly 1/4 or 1/5; the irregular whitish blotches on the leaves are similar to those usually found in H. irya and H. smithii.

Horsfieldia atjehensis W.J. de Wilde

Horsfieldia atjehensis W. J. de Wilde - Gard. Bull. Sing. 38 2 ('1985', 1986) 186

Type: Bangham 882, Sumatra, N Aceh.

Horsfieldia amygdalina auct. non (Wall.) Warb.: Merr. - J. Arnold Arbor. 8 (1934) 61

Tree c. 10 m. Twigs terete, 3.5-5(-8) mm diameter, pale grey to yellowish brown, early glabrescent, with grey-brown hairs less than 0.1 mm; bark coarsely striate and tending to flake; lenticels rather conspicuous towards the apex of the twig. Leaves in 3- 5 rows, thinly chartaceous, obovate-oblong to oblong-lanceolate, 13-25 by 4.5-9 cm, base long-attenuate, apex acute-acuminate; upper surface drying dark brown, lower surface early glabrescent, with scattered dots (lens!); midrib flat above; nerves 10-12 pairs, flat above, lines of interarching indistinct; venation lax, indistinct or invisible on both surfaces; petiole 12-15 by 2.5-3.5 mm; leaf bud 15 by 3.5-4 mm, with dense grey-brown hairs less than 0.1 mm. Inflorescences (female not seen) behind the leaves, glabrescent, or thinly haired, less than 0.1 mm; in male about 3 times branched, rather many-flowered, 7-14 by 4-10 cm, peduncle 1-2 cm long; bracts elliptic-oblong, 2-4 mm, finely pubescent, caducous; flowers in male in loose clusters of 4-8 each, glabrous, perianth 3-lobed, pedicel not articulated. Male flowers (± immature): pedicel slender, 1-1.5 mm long; buds globose, 1.5 mm diameter, cleft c. 1/2, lobes 0.2(—0.3) mm thick; androecium subglobose to short-ellipsoid, 1.2 by 1 mm, apex broadly rounded, in cross section sub- circular (Plate 2: 41); thecae 22, almost completely sessile, free apices 0.1 (-0.2) mm, curved over and more or less into the rather narrow apical cavity 0.3 mm deep; column broad, androphore narrow, 0.2(-0.3) mm long. Female flowers and fruits not seen.

Field-notes Leaves leathery, glabrous. Flower buds green.

Distribution Malesia: Known from only one collection in N Aceh, Sumatra.

Habitat & Ecology Montane forest, possibly on limestone; 1200-1800 m altitude; male fl. (immature) in Jan.

Note Horsfieldia atjehensis is in many respects closely related to and ± intermediate between H. amygdalina (Wall.) Warb. (from continental SE Asia), H. glabra, H. macrothyrsa, and H. sparsa, but is still markedly distinct from these species (De Wilde, I.e.: 188).

Fig. 10.

Horsfieldia ardisiifolia (A.DC.) Warb. a. Leafy twig apex, note ridged twig; b. twig with male inflorescence in axil of fallen leaf; c. mature male flower, lateral view; d. ditto, opened, showing androecium; e. androecium, longitudinal section, schematic; f. mature female flower, lateral view; g. ditto, opened, showing glabrous ovary with minute stigma; h. twig with infructescence with ripe fruits [a: Ramos BS 39770, b: Sulit PNH 6236; c-e: Elmer 12337; f, g: Elmer 17220; h: Conklin PNH 17461]. — Scale bar for a, b, h = 2 cm; for c, d, f, g = 1.7 mm; for e = 0.85 mm.

Horsfieldia basifissa W. J. de Wilde

Horsfieldia basifissa W. J. de Wilde - Gard. Bull. Sing. 38 1 (1985) 109

Type: White NGF10242, New Guinea.

Horsfieldia polyantha auct. non Warb.: J. Sinclair - Gard. Bull. Sing. 28 (1975) 95, p.p.

Tree 10-25 m. Twigs faintly ridged or not, 2-4(-8) mm diameter, with grey-brown hairs 0.1 mm, early glabrescent; bark finely striate, not flaking; lenticels inconspicuous. Leaves membranous or thinly chartaceous, elliptic-oblong to oblong-lanceolate, 10-22 by 3-8 cm, base attenuate, apex acute-acuminate; upper surface drying olivaceous to brown, often with paler markings, sometimes faintly pale pustulate, lower surface glabrescent, hairs 0.1 mm; dots absent; midrib above flattish; nerves 10-15 pairs not particularly contrasting, above thin, flattish or sunken, beneath with lines of interarching neither regular nor prominent; venation lax, rather faint; petiole 5-10 by 1.5-2.5 mm; leaf bud 10 by 1.5 mm, with hairs 0.1 mm. Inflorescences in male 3 (or 4) times branched, many-flowered, 4-10 by 2.5-6 cm, peduncle 0.2-2 cm long; in female: 5 by 3.5 cm; with dense to sparse stellate hairs 0.1-0.2 mm; bracts elliptic-oblong, acute, l-2(-4) mm long, caducous; flowers generally 1-3 together; perianth 2-lobed, glabrescent except at the very base, pedicel with hairs 0.1 mm long, not articulated. Male flowers: pedicel 1.5-3 mm long; buds as seen laterally ± circular, slightly broader than long, slightly laterally compressed, not or but slightly collapsed on drying, 2.2-2.7 by 2.6-3 mm, cleft to the base, lobes 0.1-0.2 mm thick; androecium laterally much flattened, above broadly rounded, 1.5-1.7 by 2 mm (Plate 1: 26); thecae 24-28(-32), erect, at apex free for 0.1 mm long, column at apex narrowly hollowed for 1/3-2/3; androphore to 0.1 mm, broadly attached. Female flowers (immature): pedicel 1.5-2 mm long, buds broadly ovoid, 1.5 by 1.4 mm, cleft nearly to the base; ovary ovoid, 1.1 by 0.6 mm, glabrous, style and stigma minutely 2-lobed. Fruits 1-20 per infructescence, globose or subglobose, 1.1-1.4 cm diameter, glabrous, drying light to dark brown, with or without coarse, paler coloured lenticels or warts; pericarp 1.5-3 mm thick, woody-granular; fruiting pedicel 3-4 mm; perianth not persistent.

Field-notes Slender tree, branches horizontal. Flowers yellow. Fruits green, turning orange.

Distribution Malesia: New Guinea (NE Papua Barat, including Memberamo River area; N Papua New Guinea: Sepik, Madang Prov.).

Habitat & Ecology Primary and degraded forest, marshy forest, locally common; recorded from Pometia-Intsia forest; on clays and marls; 0-200 m altitude; fl. Sept.; fr. Mar., June, Oct.

Note Apart from H. angularis (see the note under that species) H. basifissa is possibly closely related to H. parviflora, both have glabrous fruits. The globose fruits are often very similar to those of H. pilifera or H. sinclairii; in these two species, however, the fruits are always hairy, at least towards the base. Horsfieldia basifissa has much in common with H. laevigata var. novobritannica, which also has the androecium deeply hollowed inside; the latter has a more hairy perianth. The female flowers of var. novobritannica are not known, but its globose fruits are larger than those of H. basifissa and somewhat hairy at the base. Horsfieldia basifissa is characterized by the subglabrous male flowers with a very deeply cleft perianth, glabrous ovary, and glabrous, globose fruits.

Horsfieldia borneensis W. J. de Wilde

Horsfieldia borneensis W. J. de Wilde - Gard. Bull. Sing. 39 1 (1986) 27

Horsfieldia borneensis W. J. de Wilde - Blumea 32 (1987) 468;

Horsfieldia borneensis W. J. de Wilde - Tree Fl. Sabah & Sarawak 3 (2000) 363

Type: Bojang S 14610, Sarawak.

Tree 10-30 m. Twigs subterete, (1.5-)2-4(-10) mm diameter, sometimes blackish, early to rather late glabrescent, hairs rusty, 0.2-0.4 mm; bark finely striate, not distinctly lenticellate, sometimes finely cracking or slightly flaking. Leaves chartaceous to thinly coriaceous, elliptic-oblong to oblong-lanceolate, 7-18 by 2-6 cm, base (short-)attenuate, apex acute to (short-)acuminate; upper surface drying dull olivaceous to (partially) blackish brown, glabrous, lower surface drying pale brown to chocolate, glabrescent, hairs densely branched dendroid 0.3-0.4 mm (especially on midrib); dots present and obvious; midrib slightly raised above, (late) glabrescent; nerves 10-16 pairs, slender above, flat or sunken (slightly raised only close to the midrib) or in thinner-leaved specimens slightly raised, glabrous, lines of interarching ± regularly shaped, not distinct; venation hardly or not visible on both surfaces; petiole 12-25 by 1.5-2.5 mm, sometimes late glabrescent; leaf bud 10-17 by 2-4 mm, with dense hairs 0.3 mm long. Inflorescences behind the leaves, with dense short-woolly rusty hairs up to 0.7 mm long; in male: fairly large, many-flowered, about 4 times branched, (8-) 13-20 by (5—)10—18 cm, peduncle 1.5-3.5 cm long; in female: 8—10(—13) by 4-5 cm, less branched; bracts elliptic to elliptic-oblong, pubescent as the inflorescences, 1.5-5 mm long, caducous; flowers (male) in loose clusters of 2-6, glabrous; perianth 3-lobed, pedicel distinctly articulated. Male flowers: pedicel 1-1.5 mm; buds subglobose to broadly ellipsoid or broadly obovoid, 1.3-1.8 by 1.2-1.7 mm, cleft c. 1/3 (to nearly 1/2), not collapsing on drying, lobes 0.2-0.3 mm thick; androecium broadly ellipsoid to subglobose, 0.7-1.2 by 0.6- 1.3 mm, the apex broadly rounded, slightly impressed in the centre with cavity to c. 1/4, base rounded, in cross section circular (Plate 3: 76); thecae 16-20, almost completely sessile and mutually closely appressed, at apex incurved over the cavity, free apices about none; column broad; androphore narrow, at most 0.1 mm long. Female flowers: pedicel thickish, glabrous, 1-1.5 mm long, distinctly articulated; buds subglobose or broadly ellipsoid, 3-3.5 by 3 mm, glabrous, cleft c. 1/3; ovary subglobose to broadly ovoid-ellipsoid, 2(-2.5) by 2 mm, glabrous, stigma 2-lobed, 0.2 mm high, descending 1.5 mm down the ovary. Fruits 1-7 per infructescence, ovoid, 4-6 by 3-4.5 cm, glabrous, somewhat laterally flattened and slightly flanged, apex and base rounded, drying brown and often with a glaucous tinge, smooth; pericarp 10-15 mm thick; fruiting pedicel stout, 4-6 mm long; perianth not persistent.

Field-notes Bark usually dark brown, reddish, or blackish, rough, deeply fissured, flaking in squares, strips or flakes up to 5 cm wide, up to 1 cm thick (strips with rounded edges, appearing smooth); living bark 5-10 mm thick, red-brown, the sap red; sapwood 10 cm, reddish white to pale red; heartwood red-brown. Fruits bluish green, turning green- yellow to yellow or reddish, pericarp pink inside.

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

Habitat & Ecology Primary lowland dipterocarp forest, swamp forest; on sandy soils, flat clayey soil, sandstone, sandy ridges; 0-200 m altitude; fl. Apr., Aug., Sept.; fr. throughout the year.

Notes 1Horsfieldia borneensis and H. wallichii both have characteristic blackish dots on the lower leaf surface, a dull upper leaf surface with largely sunken nerves, and similar fruits (although the perianth in H. wallichii is persistent). However, H. wallichii, which also occurs in Borneo, is generally stouter and has much larger leaves, often with a persistent indumentum. Above all, it differs in general appearance, shape, and structure of the androecium, and the pedicel which is not articulated.

2Horsfieldia borneensis, with its dotted lower leaf surface, belongs to a group of species including H. wallichii and H. pulcherrima, while the structure of its androecium links it to species such as H. flocculosa, H. grandis, and H. pulcherrima; compare also H. punctatifolia.

Horsfieldia brachiata (King) Warb.

Horsfieldia brachiata (King) Warb. - Mon. Myrist. (1897) 325

Horsfieldia brachiata (King) Warb. - Gamble Mat. Fl. Malay Penins. 5 23 (1912) 218

Horsfieldia brachiata (King) Warb. - Ridl. Fl. Malay Penins. 3 (1924) 59

Horsfieldia brachiata (King) Warb. - W.J. de Wilde Gard. Bull. Sing. 39 1 (1986) 3

Horsfieldia brachiata (King) Warb. - Tree Fl. Sabah & Sarawak 3 (2000) 363

Myristica brachiata King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 311, pl. 144

Horsfieldia subglobosa (Miq.) Warb. var. brachiata King J. Sinclair - Gard. Bull. Sing. 16 (1958) 431, f. 51E.

Horsfieldia brachiata (King) Warb. var. brachiata J. Sinclair - Gard. Bull. Sing. 28 (1975)) 9

Lectotype: Griffith 4351, Peninsular Malaysia.

Tree 10-35 m. Twigs ± angular or subterete, or more or less flattened, distinctly lined or ridged (sometimes lines evident only in part of the material), 2-7(-18) mm diameter, generally early glabrescent, hairs rusty, (0.1-)0.2-0.4 mm long; bark finely to coarsely striate, not flaking; lenticels present but not much contrasting. Leaves membranous, elliptic-oblong to oblong(-lanceolate), 12-26(-30) by 4—9(—11) cm, base cuneate, apex acute-acuminate; upper surface glabrous, drying olivaceous to brown or sometimes blackish, lower surface drying light brown, early glabrescent, midrib sometimes later glabrescent; dots absent; midrib raised above, glabrous; nerves 12-20 pairs, raised above, lines of interarching usually not distinct; venation lax, usually not distinct above; petiole 8-13(-20) by 2-3 mm, glabrescent; leaf bud 8-15 by 3-4 mm, hairs 0.2-0.4 mm long. Inflorescences with sparse to dense dendroid hairs 0.2-0.5 mm, sometimes glabrescent; in male: 3 or 4 times branched, many-flowered, 7-18(-22) by 5—16(—18) cm, peduncle 0.6-1.8 cm long, the flowers in loose clusters of 3-6; in female: many-flowered, 3-8 by 2-6 cm; bracts oblong-lanceolate, acutish, 3-5 mm long, pubescent, caducous; flowers with perianth 3- (or 4-)lobed, glabrous, pedicel pubescent in various degrees, hairs 0.1- 0.2 mm long (in female glabrescent), articulated. Male flowers: pedicel (1—)1.5—2.5 mm long; buds (depressed-)globose to broadly obovoid, in cross section rounded or slightly angular, 1-1.5 by 1.2-1.8 mm, base rounded to short-cuneate, not or but little collapsing on drying, cleft 1/2-2/3, lobes 0.2-0.4(-0.5) mm thick; androecium depressed-globose to obovoid in outline, (0.5-)0.7-l by 0.8-1.2 mm, ± rounded or usually ± triangular in cross section (Plate 3: 64)\ thecae 12-20, anthers 0.5-0.7 mm long, mutually free for about halfway, usually curved towards the centre, column largely hollowed out, at base continuing into the 0.2-0.3 mm long androphore, slightly tapering or not. Female flowers: pedicel 1-1.5 mm long; buds broadly ellipsoid, 2.2-2.5 by 1.8-2 mm, cleft c. 1/3; ovary ovoid, 1-1.4 by 0.8-1.2 mm, glabrous, the stigma 2-lobed, 0.2 by 0.4 mm. Fruits 4-12(-20) per infructescence, broadly ellipsoid, apex narrowly rounded, base (broadly) rounded, 2-2.8(-3) by 1.8-2.2(-2.6) cm (see note 3), glabrous, drying brown to dark brown, neither warted nor lenticellate, pericarp 1.5-4 mm thick; fruiting pedicel 1.5-3 mm long; perianth not persistent.

Field-notes Usually a slender tree with straight bole, once recorded as with buttresses to 50 cm high; bark ± smooth, pale to dark brown, generally with shallow vertical fissures 1 cm apart, sometimes ± laminated, scaly, or cracked; living bark 8-10 mm thick, pinkish to reddish brown, exuding reddish sap; wood whitish to pale brown; no heartwood; twigs with raised lines. Flowers greenish yellow to dark yellow, scented. Fruits yellow(-green) or yellow-orange.

Distribution Peninsular Thailand; Malesia: Sumatra, Peninsular Malaysia (Kedah, Kelantan, Perak, Trengganu, Pahang, Malacca, Johore), Borneo (Sarawak, including one deviating collection, see note 3; Sabah; C, E & NE Kalimantan; Brunei); not found in Singapore and large parts of Kalimantan

Habitat & Ecology Primary and degraded lowland rain forest; often near streams in flatland; marshy, riverside, and peaty forests, forest on alluvial plains, poor forest on soil with stagnant water, but also on hillsides; on alluvial soils, brown and sandy soil (in Tristania forest, Sabah), sandstone, peaty soils, loam soil with lime; 0-400 m altitude; fl. & fr. throughout the year.

Notes 1Horsfieldia brachiata is close to H. polyspherula, and in most cases easily recognized by its weak to strong raised lines on the twigs. Its fruits are rather uniform in shape and size, 20-28 mm long, and thus ± intermediate between those of H. polyspherula var. polyspherula and var. sumatrana (see there). The leaves of H. brachiata are rather like those of H. polyspherula var. sumatrana, namely generally membranous and drying pale, dull olivaceous above and pale cinnamon below. Its flowers are rather uniform, with mature male buds 1.2-1.8 mm in diameter, and the androecium usually ± triquetrous with 6-10 stamens (but see note 2), and they do not differ from those of H. polyspherula s.l. Sterile and flowering collections in which the apical and lower twig parts are not sufficiently represented may be difficult to place. Horsfieldia brachiata generally has stouter inflorescences than H. polyspherula and is quite common in evergreen forests in Peninsular Thailand; H. polyspherula has not been found there.

2 The species usually has 12-16 thecae in the androecium, but material from Peninsular Thailand may have 18 or 20 thecae.

3 S 34908 from Sarawak (Kapit, 5th Div.) is a stout specimen, in bad condition, with female flowers; at L there is a single fruit measuring 40 by 30 mm, with the pericarp ± woody, 5-7 mm thick. Horsfieldia brachiata is not common in Sarawak and this large- fruited specimen probably represents a separate taxon. It was collected in a kerangasmossy forest at c. 800 m altitude, higher than any other specimen of the species.

Horsfieldia carnosa Warb.

Horsfieldia carnosa Warb. - Mon. Myrist. (1897) 348, 619

Horsfieldia carnosa Warb. - Merr. Enum. Born. (1921) 268

Horsfieldia carnosa Warb. - J. Sinclair Gard. Bull. Sing. 28 (1975) 21

Horsfieldia carnosa Warb. - W. J. de Wilde Gard. Bull. Sing. 38 2 ('1985', 1986) 222, f. 26

Horsfieldia carnosa Warb. - Tree Fl. Sabah & Sarawak 3 (2000) 364

Myristica carnosa (Warb.) Boerl. - Handl. 3 (1900) 87

Lectotype: Beccari 1242, (Fl acc. 7625) fr. Sarawak.

Tree 4-10 m. Twigs 3—10(—16) mm diameter, early glabrescent, hairs grey-brown, 0.1 mm; bark coarsely or finely striate, tending to flake, drying somewhat pale, yellow- brown or light grey-brown, generally contrasting with the blackish brown of the petioles; lenticels usually not conspicuous. Leaves chartaceous-coriaceous, rarely ± membranous, (elliptic-)oblong, 13-35 by 5-11 cm, base long- or sometimes short-attenuate, apex acute-acuminate; upper surface drying bright dark brown, finely wrinkled-granulate, glabrous, lower surface early glabrescent (glabrous); dots absent; midrib above flat; nerves 13-18 pairs, flat above, lines of interarching indistinct; venation lax, faint or invisible on both surfaces; petiole 10-16 by 2-4 mm, glabrous or early glabrescent; leaf bud 9-13 by 2-3 mm, hairs grey-brown, 0.1 mm long. Inflorescences with dense to sparse hairs 0.1 mm or less; in male: below the leaves, many-flowered, 3 or 4 times branched, 6-17 by 5-14 cm, peduncle 1-3 cm long; in female: ramiflorous, rather many- flowered, 1-2 cm long; bracts elliptic to oblong, 4-10 mm long, pubescent, caducous; flowers glabrous, in male in loose clusters of 3-9, perianth 3-lobed, pedicel not articulated. Male flowers: pedicel 1-1.5 mm long; buds (sub)globose, 1.9-2.1 by 1.8-2 mm, cleft 1/3-1/2, not collapsing on drying, lobes 0.2 mm thick; androecium (sub)globose, 1-1.2 by 1-1.3 mm, circular in cross section (Plate 2: 61); thecae 18-22, completely sessile, without free apices, incurved, concealing a small apical cavity 0.2-0.4 mm deep; column broad, ± spongy, androphore rather narrow, 0.2-0.5 mm long, completely hidden by the anthers. Female flowers: pedicel 1-1.5 mm long; buds ellipsoid, 3.5 by 2.5 mm, cleft c. 1/3, lobes 0.3-0.4 mm thick; ovary ellipsoid, 2 by 1.5 mm, glabrous, stigma consisting of two broad lips 0.2 mm high. Fruits 2-11 per infructescence, ellipsoid, 1.6-2 by 1.2-1.5 cm, glabrous, drying brown, the surface finely granulated; pericarp 1.5 mm thick; fruiting pedicel 1-2 mm long; perianth not persistent.
See: Fig. 11.

Field-notes Small tree, trunk slender; the bark often flaking or shallowly fissured; inner bark yellow, thin, sap watery, clear, not reddish; sapwood whitish, twigs light brown. Flowers green-yellow, anthers whitish. Fruits (immature) greenish yellow, aril orange.

Distribution Malesia: Borneo (Sarawak, Brunei, Sabah; W Kalimantan: Mt Klam).

Habitat & Ecology Heath forest, wet kerangas, peat swamp forest, Agathis- Casuarina forest; on white sandy soils; 0-100 m altitude; fl. mainly July-Nov.; fr. throughout the year. An extensive note on the ecology is given by Sinclair, l.c.

Note Horsfsieldia carnosa is a well-characterized species, a small tree of kerangas or peat swamp forest, on white sandy soils. It is distantly related to H. glabra, which is distinguished by a less stout habit, dark twigs, bark not tending to flake, smaller and usually membranous leaves, globose male flowers, pedicels ± articulated, globose or ellipsoid androecium with short androphore, and somewhat longer, not densely clustered fruits, 1.8-2.4 cm long.

Fig. 11.

Horsfieldia carnosa Warb. a. Twig with leaf and male inflorescences; b. apical part of leafy twig; c. mature male flower, lateral view; d. ditto, opened, showing androecium; e. androecium, longitudinal section, schematic; f. twig with female inflorescence axillary to leaf scar; g. female flower at anthesis, lateral view; h. ditto, longitudinally opened, showing glabrous ovary with broad 2-lobed stigmas; i. older twig with infructescences, fruits mature, aril complete but torn on drying [a: van Niel 5419; b-e: S 18011; f-h: SAN 63191; i: SAN 17438]. — Scale bar for a, b, f, i = 2 cm; for c-e, g, h = 0.85 mm.

Horsfieldia clavata W. J. de Wilde

Horsfieldia clavata W. J. de Wilde - Gard. Bull. Sing. 38 1 (1985) 92, f. 13d-f.

Type: Hoogland 3663, New Guinea.

Shrub or tree, 3-6 m. Twigs 1.5-3 mm diameter, glabrescent, hairs grey-rusty, 0.1 mm long; bark finely striate, not flaking; lenticels absent or inconspicuous. Leaves membranous, elliptic or oblong, 7-18 by 3-6 cm, base short- to long-attenuate, apex acute- acuminate (in Hoogland 3523 2 cm caudate); upper surface drying olivaceous, lower surface with persistent, scattered, stellate-dendroid scale-like hairs 0.1-0.2 mm long, especially on midrib, the nerves not much contrasting; dots absent; midrib slender above, raised; nerves 10-20 pairs (including some intersecondary nerves), above thin and flat or slightly raised, beneath much raised (not much contrasting in colour), lines of inter- arching regularly looping, distinct; venation lax, rather indistinct; petiole 7-14 by 1-1.5 mm; leaf bud 7-10 by 1-1.5 mm, hairs 0.1 mm. Inflorescences with scale-like hairs 0.1 mm or less, among the leaves, delicate, 1 or 2 (or 3) times branched, lowest branch from near the base; in male 2-3 by 1.5-2 cm, rather many-flowered; in female: 1-2 cm long, 2- or 3-flowered; bracts densely pubescent, 1-1.5 mm long, caducous; flowers solitary or 2 or 3 together; perianth 2-lobed, with stellate-dendroid hairs 0.1 mm; pedicel not articulated. Male flowers: pedicel 2.5-3.5 by 1-1.5 mm, pubescent; buds clavate, with tapering pedicel, together 4-5.5 by 1.5-2.2 mm; perianth rounded above, 1.5-2 by 1.5- 2.2 mm, cleft c. 1/10, lobes 0.2 mm thick, lower down perianth wall 0.5-0.7 mm thick; androecium clavate, 1.5 by 0.7 mm, anthers 3 (or c. 6 thecae), 0.3 mm long, ± stellate, sessile, column not hollowed out; androphore thickish subcylindrical, slightly bullate- striate, glabrous (Plate 1:18). Female flowers: pedicel ± slender, 2 mm long; buds ellipsoid, 1.8(-2) by 1.2 mm, cleft c. 1/4; ovary ovoid, 1 by 0.6 mm, with dense stellate scalelike hairs 0.1 mm or less, style 0.4 mm long, stigma 2-lobed, 0.2 mm long. Fruits 1 (or 2) per infructescence, broadly ellipsoid-ovoid, 1.3 by 1 cm (excluding pseudostalk), base broadly rounded, apex ± acuminate, beak 2 mm, with hairs 0.1 mm long, drying brownish, without lenticels; pericarp 1 mm thick; fruiting pedicel 6-10 mm, the pseudostalk 1.5-2 mm long; perianth not persistent.
See: Fig. 14d-f.

Field-notes Shrub or treelet. Flowers yellow. Fruits orange or red.

Distribution Malesia: Papua New Guinea (Northern Prov.).

Habitat & Ecology Locally common in regrowth in tall lowland forest on welldrained soil; 0-50 m altitude; fl. & fr. Aug.

Note Horsfieldia clavata is related to H. squamulosa and H. crux-melitensis which have a similar clavate androecium. Horsfieldia squamulosa differs in its slender, male pedicels. The pedicel, and hence the whole male flower of H. crux-melitensis is similarly club-shaped as in the present species, but about twice as large; its leaves are also larger and darker, and both male and female flowers have much thickened pedicels.

Horsfieldia coriacea W. J. de Wilde

Horsfieldia coriacea W.J. de Wilde - Gard. Bull. Sing. 39 1 (1986) 50

Type: bb Cel. 111-27, Sulawesi.

Tree 8-25 m. Twigs 2.5-4(-10) mm diameter, early glabrescent, hairs greyish brown, 0.1 mm; bark finely striate, not flaking; lenticels conspicuous or not. Leaves membranous to thinly chartaceous, (elliptic-)oblong, 14-27 by 5-10 cm, base attenuate, apex acute-acuminate; upper surface glabrous, drying olivaceous brown to blackish brown, the midrib glabrous but towards the base in younger leaves finely pubescent, lower surface glabrous; dots absent; midrib moderately raised above; nerves 13-18 pairs, raised above, lines of interarching not distinct; venation lax, barely visible on either surface; petiole 12-16 by 2.5-3.5 mm; leaf bud 12-17 by 2-3 mm, hairs dense, grey-brown to rusty, 0.1 mm long. Inflorescences behind the leaves, with sparse hairs 0.1 mm; in male: 2 or 3 times branched, few-flowered, 4-10 by 3-5 cm, peduncle 1-2 cm long, the flowers in loose clusters of 3-5; in female (from infructescences): 2-5 cm long; bracts not seen, caducous; flowers glabrous, perianth 3- or 4-lobed, pedicel not articulated. Male flowers: pedicel 1.5-2 mm long; buds subglobose to broadly ovoid, 2-2.5 by 2-2.3 mm, apex shortly rounded to subacute, not or only slightly collapsing on drying, cleft 1/2-2/3, lobes 0.4-0.5 mm thick, coriaceous; androecium subellipsoid, 1.5-1.6 by 0.8-0.9 mm, in cross section ± blunt-triangular (Plate 3: 88); thecae 10 or 12, at the base curved, and towards the apex erect or somewhat curved, 1.6 mm long, free apices 0.1-0.2(-0.3) mm, apical cavity narrow, 0.2-0.3(-0.5) mm deep, androphore narrow, 0.1-0.2 mm long, hidden by the anthers. Female flowers not seen. Fruits 1-5 per infructescence, ellipsoid, 4-4.2 by 2.5-3.2 cm, glabrous, drying rust-brown, finely granulate and with at most a few tubercles or lenticels; pericarp rather coriaceous, 3.5-8 mm thick; fruiting pedicel 2-4 mm long; perianth not persistent.

Field-notes Bark and leaves with aromatic scent; branches horizontal; cauliflorous. Flowers yellow, strongly scented; perianth fleshy. Ripe fruits orange.

Distribution Malesia: Endemic in C Sulawesi.

Habitat & Ecology Primary and disturbed forest (with Imperata, Gleichenia, and Melastoma) on ultrabasic soil; 100-700 m altitude; fl. Mar., Nov.; fr. Apr., July.

Notes 1 In most flowers there are a few minute wart-like appendages 0.1 mm high around the insertion of the androecium.

2Horsfieldia coriacea, vegetatively and in fruit, resembles H. costulata, which has a much larger distribution in Sulawesi and the Philippines. However, the latter differs in thinner membranous leaves, drying generally more olivaceous, with the midrib on the upper surface entirely glabrous, the lateral nerves usually forming a greater angle with the midrib, and less conspicuous lenticels on the twigs. The fruits are generally larger with a pericarp 8—10(—15) mm thick. Furthermore, the male flowers are quite different, those of H. costulata being arranged in rather dense clusters of 5-10.

3Horsfieldia coriacea seems closely related to H. majuscula (Sumatra, Peninsular Malaysia) and H. xanthina (Borneo), both also having an elongate androecium, but with a broader and tapered androphore, not hidden by the anthers; in H. majuscula the pedicel is articulated.

Horsfieldia corrugata Foreman

Horsfieldia corrugata Foreman - Contr. Herb. Austral n. 10 (1974) 45, f. 1

Horsfieldia corrugata Foreman - W.J. de Wilde Gard. Bull. Sing. 38 1 (1985) 130, f. 20a-c.

Type: LAE 52461, Papua New Guinea.

Tree 5-12 m. Twigs (3-)4-5(-12) mm diameter, early glabrescent, hairs greyish to rusty, 0.1 mm; bark striate, not flaking, lenticels large, usually not much contrasting in colour. Leaves thinly coriaceous, elliptic-oblong, 12-29(-32) by 4.5-8.5(-10) cm, base attenuate, apex acute-acuminate; upper surface drying dark brown, minutely pustulate or not, lower surface early glabrescent; dots absent; midrib slender to rather broad, flat-tish above; nerves 12-18 pairs, thin and flat above, beneath lines of interarching with irregular loops, distinct or not, venation lax, indistinct; petiole 6-18 by 2-3.5 mm; leaf bud 10-20 by 1.5-3 mm, hairs 0.1 mm. Inflorescences below the leaves, with rusty stellate hairs 0.1 mm long or less; in male: 2 or 3 times branched, rather slender, (4-)6-14 by 2-9 cm, peduncle 1-2.5 cm long; in female: up to 5 cm long, peduncle 1 cm long; bracts pubescent, 1.5-4 mm long, caducous; flowers (in male) solitary or in loose clusters of 2-5, glabrous or glabrescent, hairs scattered, less than 0.1 mm, perianth 2-lobed, pedicel ± tapering, not articulated. Male flowers: buds in lateral view subcircular, 3-3.5 by 3(-4) mm, apex broadly rounded, the lower half ± tapering into the thickish tapering pedicel, (2-)3-4 mm long; perianth cleft 1/2 to nearly 2/3, lobes 0.2-0.3 mm thick, often with a few coarse blackish brown wart-like dots; androecium thickish, not much laterally compressed, above broadly rounded, (1.5-)2-2.2 by 2-2.2(-3) mm (Plate 2: 34); thecae 16-24, erect, 2 mm long, free apical parts 0.1-0.2 mm, column narrowly hollowed for 1/5-1/4, androphore 0.2-0.3 mm long. Female flowers: pedicel 4-5 mm long, minutely pubescent; buds narrowly ovoid, almost glabrous, with a few coarse, dark brown wart-like dots, 4.5 by 3 mm, cleft 1/4-1/3, lobes 0.3-0.4 mm thick, coriaceous; ovary ovoid, somewhat dented or corrugated, 2.5-3 by 2.5 mm, with dense hairs less than 0.1 mm long, style and 2-lobed stigma glabrous, 0.8-1 mm long. Fruits l(-4) per infructescence, ramiflorous, broadly ellipsoid to subglobose, somewhat flattened, 6-7.5 by 4.5-6.5 cm, coarsely flanged and corrugated, drying blackish brown, with scattered, coarse, paler coloured tubercles, glabrescent, at base sometimes a short pseudostalk, apex acutish, pericarp ± woody-corky, 10-20 mm thick; fruiting pedicel 5-10 mm long; perianth not persistent.
See: Fig. 12a-c.

Field-notes Wood very light brown. Flowers yellow or orange. Fruits green, strongly wrinkled or corrugated, and strongly ridged.

Distribution Malesia: Papua New Guinea (Central, Northern, Milne Bay Prov.).

Habitat & Ecology Primary and degraded rain forest of mountainous terrain on slopes and ridges, fagaceous forest; 1200-1900 m altitude; fl. & fr. July to Dec.

Note When in flower, H. corrugata may be difficult to distinguish from, e.g., H. pachycarpa, H. tuberculata, or certain forms of H. laevigata. However, the few coarse and conspicuous blackish brown wart-like dots on the perianth, found in male and female flowers, help to characterize H. corrugata. The large, corrugated and ridged thick-lobed fruits are also distinctive, those of the other species may be similar but not ridged.

Fig. 12.

Horsfieldia corrugata Foreman, a. Longitudinally opened male flower showing androecium; b. ditto, female flower, showing pubescent ovary and narrow 2-lobed style; c. fruit. — H. pachycarpa A.C.Sm. d. Leafy twig with infructescence; e. longitudinally opened male flower showing androecium; f. ditto, female flower with pubescent ovary with short 2-lobed stigma; g. almost mature fruit [a: Carr 14123; b: LAE 60020; c: Carr 14334; d: LAE 62196; e: LAE 51940; f: Clemens 5378; g: NGF 38895]. — Scale bar for c, d, g = 2 cm; for a, b, e, f = 1.7 mm.

Horsfieldia coryandra W.J. de Wilde

Horsfieldia coryandra W.J. de Wilde - Blumea 32 (1987) 464

Horsfieldia squamulosa auct. non WJ. de Wilde: WJ. de Wilde - Gard. Bull. Sing. 38 1 (1985) 93, p.p.

Type: NGF 46892, (male fl.; fr.), Papua New Guinea.

Shrub or treelet, 1.5-6 m Twigs 1.5-2 mm diameter, with rusty hairs to 0.1 mm long, glabrescent; bark finely striate, neither cracking nor flaking; lenticels few and inconspicuous or absent. Leaves membranous, elliptic-oblong to lanceolate, 5-20 by 0.7-5 cm, base (long-)attenuate or acute, apex long-acuminate, acumen often slender, to 3 cm long, or gradually narrowed from slightly above the middle, blade above glabrous, drying dark brown (sometimes slightly olivaceous), beneath glabrescent with scattered hairs 0.1 mm or less remaining on and near the midrib, drying brown or olivaceous brown; dots absent; midrib above slender, flat or raised; nerves 11-19 pairs (usually with some intersecondary nerves not reaching the marginal nerve), above indistinct, flat or sunken, beneath distinct, with lines of interarching usually distinct; venation coarse and distinct; petiole 6-14 by 0.5-1.5 mm, glabrescent; leaf bud 7-12 by 1 (—1.5) mm, with dense rusty hairs 0.1 mm or less. Inflorescences among the leaves, 2 or 3 times branched, peduncle 0.2-1 cm long, with grey-brown hairs 0.1 mm or less, rather few-flowered; in male: 2-3 by 1.5-3 cm, flowers solitary or 2 or 3 together; in female: 1-1.5 cm long, not or little branched, few-flowered; bracts oblong, 1-2 mm long, caducous; flowers 2-lobed, with sparse hairs 0.1 (-0.2) mm or less, glabrescent in the apical part, pedicel not articulated. Male flowers: pedicel slender, (2.5-)4-6.5 mm long; buds slightly compressed elliptic(-obovate) to elliptic-oblong, 2-3 by 1.5-2 mm, cleft 1/6 to nearly 1/4; perianth 0.5-0.7 mm thick, towards the apex of the lobes 0.2-0.3 mm; androecium club-shaped, the apex subacute (to bluntish), 1.5-2.5 by 0.6-0.8 mm; anthers 0.5-0.7(-l) mm long, mutually touching, at apex free for c. 0.3 mm (Plate 1: 75); thecae 8, column not hollowed at apex; androphore glabrous or basally with scattered pale brown hairs less than 0.1 mm, the upper part with somewhat warted or wrinkled surface. Female flowers: pedicel 2.5-3.5 mm; buds ellipsoid or ± fusiform, 2-2.5 by 1.5-2 mm, cleft c. 1/3; perianth (and lobes) 0.2-0.3 mm thick; ovary ovoid, 1.5 by 1.4 mm, densely minutely pubescent, style erect with 2 ± acute lobes together 0.4 mm long. Fruits 1 or 2 per in-fructescence, broadly ellipsoid-ovoid, including the 0.5-2 mm long apiculum 1.2-1.5 by 0.8-1 cm, pseudostalk 1.5-2.5 mm, all with hairs 0.1 mm or less, pericarp 0.5(-l) mm thick, drying blackish, without lenticels; fruiting pedicel (5—)10—12 mm long, not or hardly broadened towards the apex; perianth not persistent.

Field-notes Bark smooth, greenish brown or dark green, underbark red; exudate red; inner bark brown; wood cream turning brown on exposure. Flowers yellow or orange. Fruits (yellow-)green to orange; aril complete (orifice very small and folded away), thin, red.

Distribution Malesia: Papua New Guinea (Milne Bay Prov. incl. Normanby I., Northern Prov., Morobe Prov.).

Habitat & Ecology Understorey shrub or low tree, sometimes gregarious. Lower hill forest; Castanopsis forest on steep slopes, Eucalyptus-dominated forest, ridge forest, on riverbanks; forest on limestone; 200-500 m altitude; fl. & fr. throughout the year.

Note Close to H. squamulosa, with similar, rather ellipsoid perianth, but differing in some small features in flowers and fruits.

Horsfieldia costulata (Miq.) Warb.

Horsfieldia costulata (Miq.) Warb. - Mon. Myrist. (1897) 350

Horsfieldia costulata (Miq.) Warb. - W.J. de Wilde Gard. Bull. Sing. 39 1 (1986) 38

Myristica costulata Miq. - Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 48

Type: de Vriese & Teijsmann s.n., Sulawesi.

Horsfieldia pachythyrsa Warb. - Mon. Myrist. (1897) 618

Horsfieldia pachythyrsa Warb. - Koord. Meded. Lands pl. Tuin 19 (1898) 70, 'crassithyrsa’.

Myristica pachythyrsa (Warb.) Boerl. - Handl. 3 (1900) 86, 87, ' crassithyrsa’.

Horsfieldia minahassae auct. non (Warb.) Koord.: Koord. - Meded. Lands pl. Tuin 19 (1898) 70, p.p., quoad Koorders 18158

Syntypes: Koorders 18156, (male, L lecto) Sulawesi, Koorders 18158, (L) Sulawesi, Koorders 18170, (female, L) Sulawesi.

Horsfieldia confertiflora Merr. - Philipp. J. Sci. Bot. 13 (1918) 285

Type: Ahern's Coll. FB 3183, Philippines.

Horsfieldia megacarpa Merr. - Philipp. J. Sci. Bot. 13 (1918) 286

Type: Ramos BS 16527, Philippines.

Horsfieldia villamilii Elmer ex Merr. - Enum. Philipp. Flow. pl. 2 (1923) 182, nom. nud.

Horsfieldia vulcanica Elmer ex Merr. - Enum. Philipp. Flow. pl. 2 (1923) 182, nom. nud.

Tree 9-30 m. Twigs 2.5-5(-10) mm diameter, early glabrescent, with grey-brown to light rusty hairs, 0. l(-0.2) mm; bark finely to coarsely striate, not flaking; lenticels small, generally inconspicuous. Leaves membranous or subchartaceous, elliptic-oblong to oblong-lanceolate, 15-30 by 5-13 cm, base narrowly rounded to attenuate, apex acute-acuminate; upper surface drying olivaceous to dark brown, sometimes with whitish marks as in H. irya; lower surface early glabrescent; dots absent; midrib above flat or slightly raised, early glabrescent; nerves 14-21 pairs, above thin, flat or raised, lines of interarching generally indistinct; venation lax, faint on both surfaces; petiole 7-14 by 2-4 mm; leaf bud 8-14 by 2-2.5 mm, with dense (grey-)rusty hairs 0. l(-0.2) mm. Inflorescences mostly behind the leaves, with dense or sparse hairs 0.1-0.2 mm; in male: 3 or 4 times branched, many-flowered, 6-14 by 5-13 cm, peduncle 1-3 cm long; in female: 2-6 cm long, shortly branched; bracts broadly triangular to elliptic-oblong, 2-A(-5) mm long, short-pubescent, caducous; flowers in male in clusters of 5-10 each, in female fewer, glabrous, perianth 3- (or 4-)lobed, pedicel not articulated. Male flowers: pedicel slender, 0.4-0.6 (-0.7) mm long; buds (± depressed-)globose, 1.5-1.8 by 1.5-2 mm; cleft c. 1/2, lobes 0.2 mm thick; androecium (depressed-)globose or broadly ovoid 0.5-0.8 by 0.7-1.1 mm, circular in cross section (Plate 3: 83); thecae 14-20, completely sessile (free apices to 0.1 mm), incurved, apical cavity narrow, (0.1-)0.2 mm deep; androphore rather stout, 0.2-0.4 mm long, completely or partly hidden by the anthers. Female flowers: pedicel 0.5-1 mm long; buds subglobose, 2.3-2.5 mm diameter, cleft 1/3-1/2; ovary ovoid, glabrous, 1.2-1.5 mm diameter, stigma minutely 2-lobed, 0.1-0.2 mm. Fruits 1-3 per infructescence, subglobose to broadly ellipsoid or obovoid, 3.5-6 by 3-4 cm, glabrous, finely granulate, drying bright brown to blackish brown; pericarp (4-) 8-10 mm thick; fruiting pedicel 2-4 mm long; perianth not persistent.

Field-notes Tree with or without low buttresses, 30 by 10 cm; bark fissured or with longitudinal grooves, often peeling off, sap first clear, turning red to brown-red; heart-wood reddish. Flowers yellow. Fruits yellow to red, on the larger branches.

Distribution Malesia: Philippines (incl. Palawan), Sulawesi.

Habitat & Ecology Mixed rain forest, primary dipterocarp forest; recorded from alluvial soil and volcanic soil, with Eucalyptus deglupta dominance; 250-1200 m altitude; fl. & fr. throughout the year, but fl. mainly July-Sept.

Horsfieldia crassifolia (Hook, f. & Thomson) Warb.

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - Mon. Myrist. (1897) 323, p.p.

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - J.Sinclair Gard. Bull. Sing. 16 (1958) 386, f. 34, pl. X-A

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - 28 (1975) 23

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - J.A.R. Anderson Gard. Bull. Sing. 20 (1963) 195

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - W. J. de Wilde Gard. Bull. Sing. 38 2 ('1985', 1986) (219) (f. 25)

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. - Tree Fl. Sabah & Sarawak 3 (2000) 366

Myristica crassifolia Hook f. & Thomson - Fl. Ind. (1855) 160

Myristica crassifolia Hook f. & Thomson - A. DC. Prodr. 14 1 (1856) 204

Myristica crassifolia Hook f. & Thomson - Miq. Fl. Ind. Bat. 1 2 (1858) 68

Myristica crassifolia Hook f. & Thomson - Hook, f. Fl. Brit. India 5 (1886) 108

Myristica crassifolia Hook f. & Thomson - King Ann. Roy. Bot. Gard. Calc. (1891) 308, pl. 140

Myristica irya Gaertn. var. crassifolia Miq. ex Hook f. - Fl. Brit. India 5 (1886) 108, pro syn.

Type: Griffith 4350, (see notes by Sinclair, 1975: 25) Peninsular Malaysia

Myristica horsfieldia auct. non Blume: Wall. - Cat. (1832) n. 6806 p.p. (other parts are Horsfieldia polyspherula and H. wallichii).

Myristica subglobosa Miq. - Fl. Ind. Bat. Suppl. 1 (1861) 383, p.p. (other part is Horsfieldia irya).

Myristica paludicola King - Ann. Roy. Bot. Gard. Calc. 3 (1891) 328, pl. 169

Horsfieldia fulva (King) Warb. var. paludicola King Warb. - Mon. Myrist. (1897) 299

Syntypes: King's coll. 4267, Peninsular Malaysia, King's coll. 4706, Peninsular Malaysia, King's coll. 6688, Peninsular Malaysia, Wray 3071, Peninsular Malaysia.

Tree 10-25 m. Twigs 2-6(-8) mm diameter, rather early glabrescent, hairs yellow-brown or rusty, woolly, 0.2-0.5 mm; bark coarsely striate, not flaking; lenticels sparse to dense, distinct or not. Leaves coriaceous, elliptic to oblong, 10-20(-28) by 3.5-7(-10) cm, base rounded to attenuate, apex rounded to subacute or rarely emarginate; upper surface drying dull greenish brown to dark brown, lower surface ± covered with sub-persistent interwoven or spaced stellate scale-like hairs 0.1 mm high (when very young with dendroid emergents), or glabrescent and then showing distinct hair scars, and scattered dots and dashes; midrib above flattish; nerves 11-16 pairs, above thin and flat or sunken; venation faint on both surfaces; petiole 9-20(-30) by 1.5-4.5 mm, leaf bud 7-12 by 2-3 mm with hairs 0.2-0.5 mm. Inflorescences among or behind the leaves, late glabrescent or with persistent, dense, woolly dendroid hairs 0.2-0.5 mm; in male: 3-5 times branched, broadly paniculate, many-flowered, 6-20 by 4-15 cm, peduncle 0.5-2 cm long; in female: 3-14 cm long; bracts elliptic-lanceolate, 2-5(-7) mm long, pubescent, caducous; flowers (male) in loose clusters of 2-7, glabrous; perianth 2-lobed, pedicel slender, not articulated. Male flowers: pedicel (0.3-)l mm long; buds globose or slightly transversely ellipsoid, (0.8-)l-1.3 by 1-1.5 mm, cleft l/3-l/2(-2/3), lobes 0.2-0.3 mm thick; androecium globose or ± transversely ellipsoid, barely laterally compressed, 0.4-0.5 by 0.5-0.8 mm (Plate 2: 60); thecae (6-)8-12, widely spaced, connectives broad (and androecium angular), the anthers free for almost the upper half or more; androphore 0.2(-0.3) mm long, slender. Female flowers: pedicel 1.5-2.5 mm long; buds broadly obovoid, 2-3 by 2-2.5 mm, cleft 1/5-1/3, lobes 0.6-1 mm thick; ovary obovoid, glabrous, 1.5 by 1.2-1.5 mm, stigma of 2 sessile small lobes 0.1-0.2 mm high, running out into a faint ridge at one side of the ovary. Fruits (1—)2—10 per infructescence, ovoid to obovoid, 1.5-2.2 by 1.2-1.8 cm, glabrous, drying dark brown, with at most few lenticel-like tubercles; pericarp 1.5-2 mm thick; fruiting pedicel 2-5(-7) mm long; perianth persistent.
See: Fig. 13.

Field-notes A few stilt-roots or low buttresses occasionally recorded; bark greyish, fissured, flaking in small rectangular scales. Flowers yellow, strongly scented.

Distribution S Thailand; Malesia: Sumatra (including Indragiri, Riau, Bangka, Belitung), Peninsular Malaysia (Perak, Trengganu, Selangor, Negri Sembilan, Malacca, Johore), Singapore, whole of Borneo.

Habitat & Ecology Mostly in marshy forest, freshwater and peat-swamp forest; on sandy soils, 0-200 m altitude; fl. & fr. throughout the year.

Notes 1 The lower leaf surfaces of Borneo material of H. crassifolia are earlier glabrescent as compared to those in Sumatra and Peninsular Malaysia.

2Horsfieldia crassifolia may be confused with H. fulva, a species also with more or less coriaceous leaves and a perianth persistent on the fruit, but with a 3-lobed perianth. Sterile specimens of H. crassifolia may be recognized by the coriaceous leaves, which have usually persistent scale-like hairs on the lower surface, and sparse to rather dense irregularly shaped dark dots and dashes. Sinclair (1975: 26) remarked that the species can easily be recognized from a distance by the rusty or cinnamon-brown colour of the lower leaf surface. The species is very constant in habit, characters, and habitat.

Fig. 13.

Horsfieldia crassifolia (Hook. f. & Thomson) Warb. a. Branch with leafy twig and infructescence with mature fruits; note persistent perianth and completely closed aril; b. twig with male inflorescence; c. mature male flower, lateral view; d. ditto, longitudinally opened, showing androecium; e. older twig with female inflorescence axillary to leaf scar; f. opened mature female flower showing glabrous ovary and 2-lobed stigma; note that the flower is considerably larger than the male flower; g. part of lower leaf surface with persistent indumentum and irregularly shaped dark-coloured dots [a: Mondi 51; b-d: S 9226; e-g: SAN 27183]. — Scale bar for a, b, e = 2 cm; for c, d, f = 0.85 mm; for g = 0.4 mm.

Horsfieldia crux-melitensis Markgr.

Horsfieldia crux-melitensis Markgr. - Bot. Jahrb. Syst. 67 (1935) 148, 'crux melitensis'

Horsfieldia crux-melitensis Markgr. - J. Sinclair Gard. Bull. Sing. 28 (1975) 26, ''cruxmilitensis'

Horsfieldia crux-melitensis Markgr. - W. J. de Wilde Gard. Bull. Sing. 38 1 (1985) 90, f. 13a-c

Horsfieldia crux-melitensis Markgr. - Blumea 32 (1987) 459

Type: Schlechter 19246, Papua New Guinea.

Shrub or treelet 2-7 m.