The fifty species of Artocarpus are mostly large trees of the tropical everwet zone, or they are from areas that have a relatively mild monsoon climate; that is with a short dry season. They are usually found below 1000 metres above sea level in the rainforests, though several species may occur up to 1600 m (5000 feet). In Borneo, the area where we particularly studied them, the average height of the forest canopy is about 45 m (150 feet), and Artocarpus trees in these forests reach about that height. They range from 20 m to 65 m height. The majority of the species are evergreen, though there are a few that are deciduous, for example Artocarpus dadah. Under cultivation, however, most trees are semi-deciduous, that is they will lose leaves if there is cool or dry weather. They will also drop their fruit at these times. The somewhat erratic rainfall in North Queensland, as well as the mild winter (Mean Minimum for July at El Arish is 14.7°C.) result in a matching erratic growth pattern in most Artocarpus species.
Although one species of Artocarpus, the breadnut (A. communis), grows abundantly in the swamp forests of New Guinea, they are found mainly on level or sloped ground in various soils - heavier soils in tropical areas but some lighter soils as well.
In most species the leaves are arranged spirally and the twigs are very stout. There is one species, A. anisophyllus (Tawak) - which has pinnate leaves, but most species have simple leaves on the mature tree. Like many rainforest tree genera though, Artocarpus sapling or juvenile leaves can be very large and dissected in appearance. A. elasticus (Mendi) can have these leaves triply pinnatifid (i.e. lobed like Philodendron selloum) and up to 200 cm (6½ feet) long. Later, the leaves become smaller and entire in shape, with no lobes.
Except in the two cauliflorous species (which bear fruits on the trunk and larger branches), the Jakfruit and Chempedak, the size of the leaves and twigs correspond to the size of the fruit. Large-fruited trees usually have stout twigs and large leaves, while small-fruited trees have thin twigs and small leaves.
The fruits are born with the male inflorescence among the foliage. As male inflorescences drop to the ground and rot, small flies breed in them. These flies pollinate the female inflorescence, thus establishing a brief symbiotic relationship. The fruit structure is called a syncarp - an aggregate fruit something like a strawberry. The syncarp is an advanced evolutionary fruit structure in the flowering plants. In it, the perianths (various floral organs) are united and reduced in number, and the many carpels form a single 'fruit.' Jarrett considers that the Artocarpus represent "a highly specialized evolutionary end-point" in the development of the syncarp.
The fruits are primarily adapted for distribution by arboreal mammals such as monkeys, squirrels and civet-cats. Ridley1 suggests that the huge fruits of jakfruit and chempedak are eaten by forest ungulates such as wild cattle, pigs, and elephants, which can easily reach them.
Looking at a map of South East Asia, notice that the island of Borneo is centrally located. Although the different species occur over different areas, most species occur in Borneo because of overlapping distribution. This is where we did most of our research and collecting. Figure 1 illustrates the area where most of the fruits grow naturally. The Sunda Shelf, the largest Continental Shelf in the world, is the area from which rise the great islands of Borneo, Sumatra and Java, as well as a portion of mainland Asia, including the Malay Peninsula, lower Thailand, lower Burma, and southern Vietnam. Separate from the Sunda Shelf are the Philippines (except Palawan) and eastern Indonesia and New Guinea. During the Ice Ages the Shelf was exposed above the sea, and plants 'migrated' across the great shelf through the vast forests which covered it.
The wide diversity of the fruits of the Artocarpus, with their different colours, textures, and flavours, offers the tropical gardener or farmer the opportunity to grow a group of beautiful exotic trees which will provide his family with ample crops of delicious dessert and starchy food over a long season. The fact that all these fruits are, at some time, available in markets near their natural habitats, indicates that they can also provide income to growers in many countries.
Trees of the Artocarpus generally fruit reliably and quite heavily, even, for example, in Asia when durians (Durio spp.) and other seasonal fruits fail, Artocarpus are still available. The size of fruit varies depending on which part of the tree it is borne. All Artocarpus should be picked from the tree and not allowed to fall to the ground.
Virtually all the members of the genus are fast growing trees which don't require a lot of care. They do better in rich soils, however, and they need plenty of water, especially if there is a fruit crop on; they will require irrigation. Most species like semi-shade when they are young and they should be protected from cold snaps in winter. Propagation is by seed exclusively except for seedless breadfruit, which is by root cuttings. The seeds have no dormant period, and so must be planted immediately, or if to be shipped, packed in a suitable damp medium such as sawdust. Jakfruit, chempedak, and marang have now been approach- and cleft-grafted successfully. If sap flow is too great, the scions may have to be allowed to harden, or be 'bled' in advance.
Below are descriptions of the species of Artocarpus which have so far been introduced into Australia and have edible fruits. They are grouped, the members within each group being closely related. Refer to Table 1 for a correlation of the scientific names, many common names, and the distributions. This table can be used as a guide for visitors to or residents of South East Asia.
The breadfruit (A. communis) is known from Polynesia of course, but it is believed to have been introduced there long ago from the island of New Guinea, the Moluccas, and Melanesia. There it is common in the forests.
It can provide abundant and versatile crops which are useful both as starchy vegetables and fruits. The leaves of the breadfruit are dark green and deeply-incised, only slightly so in the breadnut, and 30 to 100 cm in length. The beautiful tree can be propagated by seed, producing the breadnut, the seeds of which resemble jakfruit or chempedak seeds in taste, or more commonly by root or stem cutting or suckers or sucker marcots, producing the seedless varieties, a more desirable form.
The green to yellow, smooth-skinned (seedless), or covered with small hard protuberances (seeded), fruit can be 30 cm in diameter. The flesh of the seedless varieties is white and smooth throughout. It has a mild flavour varying with the degree of ripeness at picking and at eating, and with the variety. The fully ripe fruit is quite sweet.
There are many varieties or clones, a few of which have been introduced to Australia, and new varieties are continuing to arise through back-crossing with seeded forms. Most varieties fruit during January to April in North Queensland, however early varieties do exist, and by planting both early and mid-to-late season varieties a grower can have a continuous supply of the delicious starchy staple for 6 months or more.
Since they are susceptible to cold snaps, they are restricted to areas free of these. They are difficult to grow south of the high rainfall area around Tully (Lat. 18°S.), however they may survive in many warm coastal pockets further south. The breadfruit is not too particular about soils but it does grow best if the soil is enriched and it is well-watered. Maintaining a heavy crop requires about a minimum of 50 mm of water per week.
There are many ways to cook breadfruit; both the South Pacific Commission and the Pacific Tropical Botanical Gardens have excellent booklets with numerous recipes.
Jakfruit (A. heterophyllus) and chempedak, the following fruit, are closely related. The jakfruit is native to the rainforests of southern India and is now cultivated throughout the tropics. It is the largest tree fruit in the world but some of the best varieties are the small finer-fleshed or crisp ones. There is, for example, a small crisp variety only 20 to 25 cm (8-10") long, with a good flavour and a high proportion of edible part. Some fruits have bright orange or pink flesh. With future selection of varieties it should be possible to have jakfruit available all year in many areas.
The jak was the first Artocarpus introduced into Australia and is now well-established. Because it is well-known, only a few points about harvesting will be mentioned.
Maturity of jakfruit can be determined by several characteristics: a dull hollow sound is produced when the fruit is tapped by the fingers; the last leaf on the fruit stem (peduncle) turns yellow; the fruit spines become well developed and move apart from one another; the spines yield to moderate pressure. Finally, an aroma develops. When you are picking jakfruit for distant markets, the spines should be developed and widely-spaced but the fruit should still be firm and not have any aroma. If you are just using it at home all the characteristics should be present in a fully mature fruit.
If wild animals are a problem the fruits should be bagged on the tree. This will also protect the fruit against insect attack. Another problem with jakfruit is too many fruit! The solution? Pick your fruit green and use it as a vegetable, preferably before the seed coat develops. Test for this with a skewer. It can be used as a tender vegetable, with herbs and spices, boiled, fried or roasted. The seeds of mature fruits can be eaten boiled or roasted with other vegetables or separately, but they must be peeled.
The chempedak (A. integer), which is widely cultivated in Malaysia and Indonesia, is like a small cylindrical jakfruit, but the skin is almost smooth with only small protrusions. The tree grows to 20 metres in height in forests to 1200 m (4000 feet). The fruit lacks the annulus or ring at the base of the fruit around the peduncle that the jak has. It has a similar internal appearance to the jak with smaller segments of 2-3 cm across with flesh of a similar texture surrounding each seed. The segments are attached to the central, inedible core. There are two distinct types; the larger one is 30 to 45 cm in length with pale yellow flesh, and the smaller type is 25 cm long with darker yellow flesh. Both have a very rich, sweet, juicy flavour and creamy texture with some fibre. The smaller variety is generally sweeter and creamier. Compared with the jakfruit the chempedak is sweeter and has less acidity. It has a strong aroma. Among the small chempedaks are an orange-fleshed variety, which is a good-flavoured, sweet, creamy fruit about 30 cm long, and a green-fleshed variety.
Chempedak is quite good chilled, and the Malays make an 'ice' out of it. The immature fruits are used as a vegetable, but once again, before the seed coat forms. When boiled in a soup with onions and seasonings it is quite tender and flavoursome. Chempedak fritters with or without the seed are deep-fried in batter and make a tasty dish. Chempedak requires some microclimate establishment during the early years of the tree's life. Apart from that it is easy to grow and will tolerate any soil. It is also moderately tolerant of near-freezing temperatures.
A hybrid, A. heterophyllus X integer, occurs readily between the two closely related species, and it combines the characteristics of the two fruits in one fruit.
The Mendi (A. elasticus), also known by many other names, is in a group closely allied with the following three fruits, the Pedalai, Pudau, and Anjili. Mendi is a very large, distinctive tree in nature, strongly buttressed and up to 65 metres in height. It is common-to-rare, depending on the area, and grows mainly on clay soils. It occurs in Borneo up to 1400 m altitude. It is also occasionally cultivated throughout the Sunda Shelf. The large, beautiful leaves have already been referred to (Introduction). These become ovate-elliptic and sized 60 by 35 cm on the mature tree. They are covered with very short hairs on both sides and have a sandpapery feel. The yellowish-brown, cylindrical-ovate fruit, up to 17 by 10 cm in size, is covered with long fleshy, curly hairs like those on the pedalai, interspersed with short protuberances. The soft, white-fleshed segments are quite sweet, and the seeds may also be eaten roasted. Ripe fruit have a strong aroma. The tree tolerates a short dry season, and is more cold resistant than the pedalai. Individual trees react differently to the vagaries of the North Queensland climate. Growth of the best tree at El Arish, after 3½ years, has been rapid, up to 2m per annum. Irrigation and microclimate establishment is recommended in the early years.
The Pedalai (A. sericicarpus) is locally abundant in the wet forests throughout its reasonably wide distribution and is a large, well-branched, strongly-buttressed tree growing to about 40m in height when mature. It is found on quite steep hillsides, on clay soils, but it also grows well near the coast. Although the digitate young leaves with many lobes, can measure up to 180 by 70 cm, later the leaves of the mature tree are elliptic to ovate and up to 70 by 50 cm.
The globular fruit is about 15 cm in diameter, with an attractive bright orange skin. Some of the small, soft protrusions sprout curly yellow hairs 40 mm long, like a giant rambutan. Pedalai is similar to the marang (A. odoratissimus) inside but has a superior flavour, firmer flesh, and slightly larger segments. The sweet creamy white flesh is easy to eat, and the segments cling to the central core when the skin is peeled off.
There is little aroma to the ripe fruit. The seeds when fried in oil for a short time taste like peanuts, and they can also be boiled or roasted.
Given the fact that the pedalai is a very beautiful brightly-coloured fruit with little aroma, that the flavour is sweet and delicious, and that the handsome buttressed tree is easy to grow in a warm climate, it seems certain that it will become better known around the tropical world. It grows best under light or partial shade for the first three years. The tree is very fast growing, the fastest of the Artocarpus, attaining up to 6 m in 3½ years from seed. It does best on alluvial soils but also grows well on clay, and on basalt soils.
Pedalai resembles breadfruit in having little cold tolerance. Although there are trees surviving as far south as Rockhampton (lat. 23ºS.), many trees are killed by about 6°C. Some seedlings, however, appear to have more cold tolerance than others.
The Pudau (A. kemando) is a large, rare tree throughout the primary and secondary forests of its area. The leaves are small (15 by 8 cm), smooth above, rough below. It occurs in lowland and highland areas. In the latter case it was found on alluvial clay soils at 1000 metres.
It has a small round fruit, 4 cm in diameter, with smooth green skin like chempedak, and there is not a lot to it (8 segments) but it is all edible with little waste. It has a mild flavour without sweetness. The pudau is unusual in that latex from the tree is edible in small quantities. It tastes like coconut milk and is used as a sauce. Although rare, the tree is widespread, and seems to favour swampy ground. It is easy to grow.
Anjili (A. hirsutus) is an evergreen tree to 70 m in height which grows in the wet forests of the Western Ghats Mountains in southern India from sea level to 1200 m (4000'). The juvenile leaves are pinnatifid, to 50 by 35 cm in size, while the mature leaves are 25 by 12 cm, elliptic to ovate.
The yellow to orange fruit, which is up to 8 cm across and 400 g in weight, varies in shape from round to ellipsoid, and is covered with closely set rigid protuberances like the marang, but pointed. The flesh of the ripe fruit is dark yellow or reddish yellow in colour, and of a subacid flavour. There are many of these delicious mealy-textured segments which melt in your mouth. Apart from being eaten out of hand, anjili may be fried. The seeds also are good fried, and taste like peanuts.
So far the tree has proved easy to grow in the Northern Territory, where it has attained 1.8 m in one year. Attempts are also being made to reproduce it by vegetative means.
The Tawak (A. anisophyllus) is closely allied to the following fruit, keledang (A. lanceifolius), and somewhat less related to the following three fruits - marang or tarap, monkey jak, and pingan.
Tawak is an evergreen tree to 45 m height, occurring occasionally throughout the lowland and montane Dipterocarp forests of Borneo, southern Malay Peninsula and southern Sumatra to an altitude of 1200 m (4000'). The beautiful dark green, slightly rough leaves are unique in the Artocarpus in being pinnate in the adult form. In juvenile plants these may reach 150 cm long, and the individual pinnae are lobed.
The golden-brown, round to oblong fruit are about 20 by 13 cm in size, including the densely set, rigid, blunt spines with which it is covered, just like the marang. The many small arils inside, light orange in colour, firmer in texture and less sweet than marang and pedalai, are delicious, and preferred by some to these fruits. The tree is fast growing.
Keledang (A. lanceifolius) is a common tree in the Dipterocarp forests of parts of the Sunda Shelf, occurring on sandy and clay soils to 1400 m (4500') altitude. The evergreen tree has small buttresses and reaches 35 m in height. The elliptic to ovate leaves (to 33 by 17 cm) are pinnate or pinnatifid when juvenile, and slightly rough.
The round fruit is 12 to 15 cm across, orange-brown in colour, and is regularly divided or 'tiled' by small bumps. There are a small number of large, bright orange, fleshy segments, which have a sweet taste well-liked by Europeans. The core is thick.
MARANG or TARAP
Marang (A. odoratissimus) (the Philippine name for the Borneo Tarap) is one of the dominant early trees in secondary forests. It is consequently one of the commonest trees in the northern 2/3 of Borneo now that most of the forest has been logged. It has been introduced to Mindanao and other islands in the past. In Borneo it grows to an altitude of about 1600 m (5000'). The attractive evergreen tree can attain 25 m in height, and has dark green, broadly elliptic to ovate leaves up to 50 by 30 cm in size. Some adult leaves have two lobes while the juvenile leaves are more lobed (pinnatifid), and both are covered with hairs like a 'five o'clock shadow.'
The tarap fruit is round to oblong in shape, large (up to 30 by 20 cm), and covered with hundreds of closely set, blunt, green or brown spines which feel rough. The small (10 mm) seeds are surrounded by the creamy white flesh of the many pulp segments, which cling to the small core when the skin is removed. The flavour is sweet and juicy with a tang to it. It has a fibreless melting texture, and the fruits develop a strong aroma after they ripen. The spines should snap when the fruit is ripe; they bend and exude latex when not ripe. Like all Artocarpus, the marang must be picked as it will not fall. The seeds are good to eat when fried. The fresh fruit can also be used in pies, cakes and to flavour ice cream. Since marang grows in regions with abundant and equally distributed rainfall, it will require irrigation if grown in a monsoon climate. There is a marang tree growing successfully in a favoured location near Yeppoon, central Queensland (lat. 23°S.) which has been exposed to about 4°C. repeatedly, so the tree's tolerance of cold is substantial, and with proper microclimate conditions it should prosper. In rich soil in North Queensland it has attained 3 metres growth per annum.
Monkey Jak (A. rigidus) is a semi-deciduous tree of the wet dipterocarp forests, to an altitude of 900 m (3000'), and of the mixed swamp forests, throughout the Sunda Shelf. It is buttressed and grows to a height of 35 m, and prefers heavy clay or peat soils. Although it is rare in nature it, is cultivated in Malaya and Java. The dark green, shiny elliptic to ovate leaves are smooth above and up to 32 by 15 cm in size. Juvenile leaves are pinnatifid. Monkey jak changes its leaves about twice a year, after dry weather, and flowers are borne on the new twigs.
The round fruit is yellow to dull orange or golden-brown and 11 to 15 cm in diameter, and has a brush-like surface covered with closely set stiff conical spines like marang. The thick skin is white inside with inedible 'rags', and only a few large golden-orange segments of waxy texture cling to the thick core. It has a pleasant sweet-sour taste when fully ripe, but is usually eaten before this stage. It is said to give a rawness to the mouth. The fruit takes six months to develop and ripen on the tree, which would preclude its growth in a cool climate.
Since monkey jak has two subspecies and is so widespread in its distribution, it should be possible to make selections of good varieties, with the best characters, for future propagation.
The Pingan (A. sarawakensis) is a rare tree endemic to the forests of Sarawak. The leaves resemble those of marang or tarap, having 'five o'clock shadow' and being broadly elliptic to ovate and up to 60 cm long, and in being lobed in the juvenile stage. The lobes, however, are more pointed than those of A. odoratissimus. The leaves of pingan are dark green with golden hairs on the petiole. The fruit is 7.5 to 10 cm and round, being covered with orange-brown short stubby projections like marang. Inside there are many small yellow segments which have a good, sweet flavour.
Nothing is known of its specific requirements, however it does grow slowly, tolerates clay soils, and appreciates shade when young.
Dadak2 (A. dadah) and the following four fruits - utu, lakuch, and selanking, and to a lesser extent, kwaimuk, are a closely related group of Artocarpus trees with small, opposite leaves, which do not differ markedly from each other; however the fruits can distinguish them.
This common and variable species, also called tampang, is widespread over the Sunda Shelf. It is a deciduous tree growing in wet forests up to 900 m (3000') altitude. The tree, which can reach 35 m in height, has obovate or elliptic-oblong dark green leaves to 30 by 17 cm in size, the underside of which are reddish gold, with prominent darker veins.
The yellowish-green, velvety-skinned, 5 to 8 cm fruit is almost round, with deep pink or red flesh. Dadak needs selection, and may have promise.
Utu (A. reticulatus) is a beautiful smallish tree (to 30 m) from Sulawesi and the Moluccas, growing up to 600 m (2000') above sea level. The tree has small buttresses, and the small leaves (to 35 by 16 cm) are oblong to elliptic with an entire edge. The main veins and lesser veins are prominent beneath, and the top surface is green and glossy, giving the tree a dense, shiny crown.
Utu fruits look like a small (12-15 cm) golden-brown jakfruit, being irregular or almost round, and covered with small protuberances, or in some varieties, smooth and velvety. The creamy-coloured flesh, however, is "eat-all", with no waste, and the hard-shelled, large seeds can be roasted and eaten. The flavour of the flesh is neither sweet (like most Artocarpus) nor sour, but tangy and very pleasing, reminiscent of yogurt. Can be used in the same way as breadfruit.
The area to which utu is endemic has a mild monsoon climate, and this may help it adapt to Queensland's seasonal regime. Growth is rapid and vigorous.
Lakuch (A. lakoocha) is a deciduous tree, to 20 m in height, of the wet, semi-wet, and wet deciduous forests from northern India to southern China. It grows to an altitude of 1900 m (6000') in areas where there is a distinct dry season. The smooth, plain green leaves are elliptic, oblong or ovate in shape, and up to 37 by 21 cm in size. Juvenile leaves have slight lobes.
The fruit of lakuch is round or irregular in shape and up to 12 cm across. The yellow skin is smooth and velvety. The colour and taste of the flesh is unknown to the author.
As well as being the most wide-ranging species on the Asian continent, lakuch is also the most tolerant of cool temperatures and dry conditions. However it is damaged by frost. It is frequently cultivated for its fruit throughout its range and south as far as Bombay (lat. 19°6 N ), so it would seem well-suited to both tropical and subtropical, but frost-free areas.
Selanking (A. nitidus), an evergreen tree which reaches 35 m tall, is occasional but widespread in wet forests and savannah woodlands of much of South East Asia, growing from sea level to 1600 m (5000') in clay-rich soils. There are many subspecies and varieties. The leaves are small, elliptic or with small lobes, to 23 by 9 cm, and dark green and smooth.
The round or oval fruit, 6 by 4 cm or larger, has smooth, soft orange skin and a sweet-acid orange flesh with few very small seeds. It is cultivated.
(This name is also used to refer to other Artocarpus in China, including the preceding fruit, A. nitidus.)
This small tree, A. hypargyreus, a native of China, reaches 14 m in height, and has an evergreen, dense, rounded canopy. It withstands frost. The small elliptic leaves are up to 17 by 8 cm in size. The round to ovoid fruit is 4 to 5 cm across, and has a yellow, finely-velvet surface. The soft pulp surrounding the dozen seeds is reddish orange to red, and has a melting texture and a subacid flavour. It is usually eaten fresh.
The research for this article was done by the author and Lauren Gartrell. Extensive use was made of Jarrett's monograph. Many thanks are due to members of the following departments for their assistance or use of facilities: Department of Agriculture, Sabah; Department of Agriculture, Sarawak; Department of Forestry, Sarawak. Help was also received from numerous individual Malaysians.
Jarrett, Frances M., 1959-60, Studies in Artocarpus and Allied Genera, Jour. Arnold Arb., XL 1-37, 113-155, 298-368, XLI, 73-140
Browne, F.G., 1955, Forest Trees of Sarawak and Brunei
Anderson, J.A.R., 1980, A Checklist of the Trees of Sarawak, Forest Dept.
Kohua, Na Lima, 1976, Breadfruit, Pacific Tropical Botanical Garden
SPC Community Education Training Centre, 1983. Breadfruit, South Pacific Commission, Fiji
Pantastico, Mendoza, and Hapitan, 1970, Harvesting Handling and Storage of Leading Philippine Fruits, Dept. of AGronomy, University of Philippines
1. Ridley, 1940, The Dispersal of Plants
2. Beware: Dadah, an alternative pronunciation of dadak, also means drugs in Malay.
Other uses: It has not been the objective of this paper to cover these, however they are many, including the extensive use of A. elasticus and its close relatives for removal of bark (should be from relatively young trees) for the making of straps and ritual clothing, and the great value as timber of many of the species which have hard wood.
|Scientific Name||Common Name||Distribution|
|A. anisophyllus Miq.||Tawak, Entawak, Bintawak, Kelidang, Bakil, Puan||Borneo, S. Malay Peninsula, S. Sumatra|
|A. communis Forst.||Breadfruit||South East Asia and South Pacific|
|A. dadah Miq.||Dadak, Tampang, Selanking dadak||Sunda Shelf, lower Thailand|
|A. elasticus Reinw. ex Bl.||Mendi, Tekalong, Benda, Terap Togop, Ahbat, Jerami, Ho, Aw, Ka aw||Sunda Shelf|
|A. heterophyllus Lamarck.||Jakfruit, Nangka||India, South East Asia (cult.)|
|A. hirsutus Lamarck.||Anjili, Hebhalsina, Ran-phunnas||South-west India|
|A. hypargyreus Han. ex Benth.||Kwaimuk||Guandong and Hainan, China|
|A. integer (Thunb.)Merr.||Chempedak, Temedak, Nakan Pudau,||Sunda Shelf, Sulawesi, Irian Jaya|
|A. kemando Miq.||Pudu, Selibut, Kudu||S. Borneo, S. Sumatra, S. Malay Peninsula|
|A. lakoocha Roxb.||Lakuch, Barhal, Oahu, Myauklok, Hat lom, Hat non||E. and N. India, Burma, Thailand, Laos|
|A. lanceifolius Roxb.||Keledang, Klidang, Keliput||Malay Peninsula, Sumatra, N. and E. Borneo|
|A. nitidus Trec.||Selanking, Butong, Empatah, Sinojoh, Ma hat Sampor, Cay Chay||Sunda Shelf, N. Philippines, Bali to Timor|
|A. odoratissimus Blanco||Marang, Terap, Madang||N. Borneo, S. Philippines|
|A. reticulatus Miq.||Utu, Maumbi||Sulawesi, Moluccas|
|A. rigidus Bl||Monkey jak, Pujan, Perian, Tampuni, Pussar, PalaMusoh||Sunda Shelf, S. Indochina|
|A. sarawakensis Jarrett||Pingan||Sarawak|
|A. sericicarpus Jarrett||Pedalai, Belalai, Gumihan, Terap||Borneo, Philippines, Sulawesi,|
DATE: November 1988
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