1474

2: Edible fruits and nuts

Artocarpus J.R. & G. Forster

Char. Gen.: 101, t. 51, 51a (1776).

MORACEAE

x = 14

Major species and synonyms
— Artocarpus altilis (Parkinson) Fosberg — see separate article.
— Artocarpus chaplasha Roxb., Fl. Ind. 3:525 (1832).
— Artocarpus heterophyllus Lamk — see separate article.
— Artocarpus integer (Thunb.) Merr. — see separate article.
— Artocarpus nitidus Trécul, Ann. Sci. Nat. Bot. 3(8): 119 (1847), with following subspecies (J. Arn. Arb. 41: 121—130, 1960): ssp. borneensis (Merr.) Jarrett, synonym: Artocarpus borneensis Merr. (1922); ssp. griffithii (King) Jarrett, synonyms: Artocarpus eberhardtii Gagnep. (1926), Artocarpus griffithii (King) Merr. (1939); ssp. humilis (Becc.) Jarrett, synonym: Artocarpus humilis Becc. (1902); ssp. lingnanensis (Merr.) Jarrett, synonyms: Artocarpus parva Gagnep. (1926), Artocarpus sampor Gagnep. (1926), Artocarpus lingnanensis Merr. (1931); ssp. nitidus.
— Artocarpus odoratissimus Blanco — see separate article.
— Artocarpus rigidus Blume, Bijdr.: 482 (1825), with the following subspecies (J. Arn. Arb. 40: 150—155, 1959): ssp. asperulus (Gagnep.) Jarrett, synonyms: Artocarpus calophylla Kurz (1876), Artocarpus asperula Gagnep. (1926); ssp. rigidus, synonyms: Artocarpus cuspidatus Griffith (1854), Artocarpus kertau Zoll. ex Miq. (1854), Artocarpus varians Miq. (1861).
— Artocarpus sericicarpus Jarrett, J. Arn. Arb. 40: 350 (1959).
Brief information on other species primarily used for their edible fruit is listed in the chapter on Minor edible fruits and nuts.

General: breadfruit trees (En). Best known are breadfruit (Artocarpus altilis), chempedak (Artocarpus integer) and jackfruit (Artocarpus heterophyllus).
— Artocarpus chaplasha: Burma: toung-peing-nai.
— Artocarpus nitidus ssp. griffithii: shiny tampang (En). Indonesia, Malaysia: tampang. Indo-China: troi.
— Artocarpus nitidus ssp. humilis: Indonesia: betoh, tampang (Kalimantan). Malaysia: selangking (Sarawak).
— Artocarpus nitidus ssp. lingnanensis: Cambodia: sampor. Thailand: ma hat. Vietnam: cây chay.
— Artocarpus nitidus ssp. nitidus: Philippines: kubi.
— Artocarpus rigidus ssp. asperulus: Burma: taung peing. Cambodia: knor prey, dom knol prei. Thailand: kanun pan. Vietnam: cây mit nai, cây da xóp.
— Artocarpus rigidus ssp. rigidus: monkey jack, tempunai (En). Indonesia: pussar (Java), mandalika, kujan (Kalimantan), tempunit, purian (Sumatra). Malaysia: perian, jelatoh, gias, tempunai (Peninsular Malaysia).
— Artocarpus sericicarpus: Malaysia: terap bulu, pedalai (Iban). Philippines: gumihan.

The genus Artocarpus is native to South and South-East Asia, New Guinea and the southern Pacific and comprises about 50 species. Artocarpus altilis and Artocarpus heterophyllus are cultivated throughout the tropics.
Artocarpus chaplasha is native to north-eastern India, lower Burma and the Andaman and Nicobar Islands. In India it is often cultivated. Artocarpus nitidus ssp. borneensis is restricted to Borneo; ssp. griffithii occurs in southern China, Indo-China, Thailand, Peninsular Malaysia, Borneo and Sumatra; ssp. humilis is restricted to Borneo; ssp. lingnanensis occurs in Thailand, Indo-China and southern China and is cultivated, especially in Vietnam; ssp. nitidus is restricted to the central and northern Philippine islands. Artocarpus rigidus ssp. asperulus occurs in Burma, Thailand and Indo-China; ssp. rigidus in Burma, Peninsular Malaysia, Sumatra, Borneo and Java; it is cultivated, especially in Malaysia and Indonesia. Artocarpus sericicarpus occurs in the Philippines, Borneo, Sulawesi and the Moluccas.

All species listed have edible aggregate fruits (syncarps). In some the flesh can be eaten fresh or made into confection, in others it is the pulp around the seed which is eaten.
Most Artocarpus species yield fairly good timber and a dozen species are used in traditional medicine in South-East Asia.
Artocarpus chaplasha is an important timber tree especially in India; its leaves are also used for elephant fodder.
Artocarpus nitidus: especially the fruits of ssp. griffithii (yellow or orange-pink with bright pink flesh) and ssp. lingnanensis (red with pink flesh) are eaten; ssp. lingnanensis is also cultivated for its bark and roots, which are added to betel.
Artocarpus rigidus: both subspecies are cultivated for their edible fruit, the edible portion being the orange, sweet, waxy pulp (perianth) surrounding the seeds. They have a pleasing taste but are apt to give the eater a sore mouth. The timber is used for construction and furniture. The latex is mixed with wax for batik work and is also used as an ointment in veterinary medicine.
Artocarpus sericicarpus bears sweet, aromatic, juicy perianths around the seeds and the seeds themselves are eaten roasted. Probably the bark can be used to make bark cloth.

None of the minor Artocarpus species is traded beyond the local market level. In Sarawak Artocarpus sericicarpus is the one found most frequently in the markets.

Composition of perianth flesh based on dry weight: protein content low, usually between 3.5—7%, fat content very low (less than 2% in most species), ash content 2—4% for all species, fibre content more variable, from 2% in Artocarpus anisophyllus Miq. to 22% in Artocarpus kemando Miq. and 25% in Artocarpus dadah Miq., carbohydrate content from 60% (Artocarpus dadah) to 85—89% (Artocarpus odoratissimus and Artocarpus integer) and 92% (Artocarpus anisophyllus).
Seed composition, also on dry weight basis: protein content 10—13%, fat content low (the highest level ca. 2%), ash content 1.5—4%, fibre content from 5% in Artocarpus integer to 17% in Artocarpus anisophyllus, carbohydrate content from 40—60% (Artocarpus odoratissimus, Artocarpus anisophyllus) to 75—80% (Artocarpus integer).
The wood of Artocarpus chaplasha is brown to dark-brown, moderately hard and strong, with a volumetric mass of 560 kg/m³.

Evergreen or deciduous small to large monoecious trees, all parts containing white latex. Leaves spirally arranged or alternate and distichous, simple and entire to pinnatifid or pinnate, coriaceous, glabrous to pubescent. Inflorescences unisexual, capitate, head cylindric to globose, solitary or paired in leaf axils or rami- or cauliflorous; numerous flowers densely packed together, embedded in the receptacle, the perianths enclosing a single ovary or stamen, usually mixed with abundant stalked interfloral bracts; male head with perianths tubular and bilobed or perforate above, to 2—4-partite, stamens short- to long-exserted; female head with perianths tubular, thin-walled below and enclosing ovary, thick-walled above with a narrow lumen containing the style,; perianths persisting and partially or completely fused with one another to form syncarp; ovary unilocular, style apical to lateral, simple or bifid. Aggregate fruit (syncarp) formed by the enlargement of the entire female head. Seeds large, without endosperm; germination hypogeal.
— A. chaplasha: deciduous tree, 40 m tall, bark grey to brown, peeling off in flakes. Stipules ovate-lanceolate, up to 2.5 cm long, yellow pubescent. Leaves obovate, elliptic to oblong, 14—23 cm x 9—14 cm, margin entire; juvenile leaves pinnatifid; main veins prominent beneath. Inflorescences, axillary, solitary; male head ellipsoid to subglobose, 1.5—3 cm x 1—3 cm; female head with pubescent peltate bracts mostly shed and simple styles exserted to 0.5 mm. Syncarp subglobose, up to 6(—10) cm diameter, yellow, covered by closely set, indurated, cylindric, obtuse, appressed-hispid processes, ca. 1.5 mm x 2 mm; wall ca. 5 mm thick; peduncle 5—8 cm long. 'Seeds' (botanically pericarps) numerous, ellipsoid, 7 mm x 4 mm.
— A. nitidus: evergreen tree, up to 35 m tall, sometimes with buttresses, bark red-brown. Leaves elliptic to obovate-oblong, 3.5—23 cm x 1.5—9 cm, margin entire; main veins prominent beneath. Inflorescences axillary, solitary or paired; male head oblong, obovoid or clavate, 3.5—12 mm x 2.5—7 mm; female head with the styles exserted to 0.5 mm through a covering of peltate bracts. Syncarp subglobose, 1.5—6 cm diameter, surface smooth, with scattered persistent bracts; wall 1—5 mm thick; fruiting perianths 1—12; peduncle 1.5—4(—20) mm long. Pericarp (including the seed) subglobose to ovoid, 8—10 mm x 7—8 mm.
— A. rigidus: evergreen tree, up to 35 m tall, buttressed, bark grey, peeling off in flakes. Stipules ovate-lanceolate, 0.5—3 cm long. Leaves elliptic to oblong-elliptic, 9—32 cm x 5—15 cm, margin entire or shallowly crenate; juvenile leaves pinnatifid; main veins prominent beneath. Inflorescences axillary, solitary; male head (sub)globose, 13—20 mm diameter; female head with pubescent peltate bracts being shed and simple styles exserted to ca. 5 mm. Syncarp globose, up to 7(—13) cm diameter, dull orange, echinate from closely set, rigid, tapering, fluted, acute, hispidulous processes, 7—9 mm long; wall ca. 1 cm thick; fruiting perianths numerous; peduncle 8—25(—40) mm long. Pericarp (including the seed) ellipsoid, 12 mm x 7 mm.
— A. sericicarpus: tree, up to 30—40 m tall. Stipules broadly lanceolate, 6—12 cm long. Leaves elliptic to ovate, 20—70 cm x 10—50 cm, margin entire to slightly crenate; juvenile leaves more or less pinnatifid; main veins prominent beneath. Inflorescences axillary, solitary; male head oblongoid, 3.5—10 cm x 1.5—2 cm; female head subglobose, 4.5 cm x 4 cm, simple style 1 mm exserted. Syncarp cylindrical to ellipsoid, 8.5 cm x 5 cm, yellow-brown, covered with long flexible solid processes, 2—3.5 cm long; wall ca. 2 mm thick; peduncle 10—18 cm long. 'Seeds' (pericarps) numerous, ellipsoid, 10 mm x 6 mm.

Seeds have no dormant period; they germinate immediately and are unable to withstand desiccation. In subgenus Artocarpus, to which belong e.g. Artocarpus altilis, Artocarpus chaplasha, Artocarpus heterophyllus, Artocarpus integer, Artocarpus odoratissimus, and Artocarpus rigidus, the leaves are spirally arranged on ascending stout twigs; the large, deeply dissected leaves of the saplings are replaced by smaller, entire leaves in the mature tree; a few species have compound leaves. In subgenus Pseudojaca Trécul, comprising Artocarpus nitidus, the leaves are alternate and distichous, forming more or less applanate sprays of foliage on slender twigs and the leaves are often simple. In Malaysia new leaves are seen mainly around the months January—February and July—August. Artocarpus rigidus is more or less deciduous. Most other species are evergreen, but when grown in a more seasonal climate they become semi-deciduous, losing most leaves — and fruit — during a cool or dry season.
In Sarawak in primary forest an annual trunk diameter increment of 0.4—3 mm was observed, in secondary forests 1.6—4.75 mm/year in the first four years to 5—9 mm/year for the next 20 years.
Flowering and fruiting is quite variable per year and per region. In Sarawak, during 7 years of observation, Artocarpus kemando did not reproduce at all, Artocarpus anisophyllus reproduced in one year only, Artocarpus odoratissimus in 3 years and Artocarpus integer in 4 years. Fruiting occurred mainly in the last 3 months of the year, following flowering in the second half of the dry season which lasts from April to November in Sarawak.
The male flower heads are generally near the end of the twigs, the female heads below them on the basal part of the twig. When ripe, male heads are dusted with yellow pollen. After flowering the male heads fall off but the female heads persist and in the course of several months (2—6) gradually enlarge into the fruit. Some species appear to be equipped for insect pollination, the male heads having a sweet scent of honey and sticky pollen (Artocarpus heterophyllus, Artocarpus integer). Others are possibly wind-pollinated — male heads emit no scent but on being tapped release clouds of pollen (Artocarpus altilis, Artocarpus rigidus). Pollination requires further study to show that insects visit female as well as male heads, and to relate pollination intensity to fruit size. Generally only a few of the flowers in a female head set seed; the remainder extend as the head swells, to form the straps or strings between the seeds in the ripe fruit. Fruit size tends to correspond with the size of twigs and leaves, a relationship that does not apply to cauliflorous species.
In subgenus Artocarpus the seed is surrounded by a pulpy perianth and the rest of the fruit is fibrous or indurated. In subgenus Pseudojaca the entire syncarp is soft and fleshy.

The 5 subspecies of Artocarpus nitidus can be distinguished as follows: ssp. borneensis: syncarp not velutinous but covered by readily deciduous clavate hairs; ssp. griffithii: syncarp subglabrous; ssp. humilis: leaves with an acumen to 2.5 cm long, lateral veins often markedly ascending, syncarp velutinous; ssp. lingnanensis: leaves obtuse to shortly and bluntly acuminate, syncarp velutinous; ssp. nitidus: syncarp small, up to 1.5(—3) cm across, velutinous.
Distinctive characters of the 2 subspecies of Artocarpus rigidus: ssp. asperulus: syncarp with processes hispid from spreading hairs ca. 0.5 mm long, leaves with a rough upper surface and a rounded or shallowly cordate base; ssp. rigidus: syncarp with processes hispidulous, leaves usually glabrous and smooth above and cuneate at base.

The species of Artocarpus, except for a few species of subgenus Pseudojaca, are large forest trees and are mainly restricted to evergreen forest in the humid tropical zone or in areas with a relatively mild monsoon climate. They are usually found below an altitude of 1000 m, but some occur up to 1700 m.
Artocarpus chaplasha requires a fairly well-marked monsoon climate, but still needs a rainfall of at least 2000 mm/year.
Artocarpus nitidus occurs in forest or savannah up to 1700 m altitude.
Artocarpus rigidus grows in evergreen forests up to 500(—1000) m, very often near streams.
Artocarpus sericicarpus grows in evergreen forests up to 500(—1000) m.

Germplasm collections of mainly the cultivated Artocarpus species are present in Fiji (Koronivia Research Station, Namsoni), Indonesia (Centre for Research and Development in Biology, Bogor), Papua New Guinea (Lowlands Agricultural Experiment Station, Keravat), the Philippines (Institute of Plant Breeding, Los Baños), Thailand (Plew Horticultural Research Centre, Chantaburi), United States (Subtropical Horticultural Research Unit, USDA, Miami; College of Tropical Agriculture and Human Resources, Hawaii). Nearly all the species are being tested in private and/or public collections in Queensland, Australia.

The minor Artocarpus species play an important role in the natural vegetation in South-East Asia. They figure prominently on species lists of sample plots in forests in different parts of the region, providing food for many animals. Man makes use of the fruits, timbers, fibres and medicinal products derived from the trees. However, lack of knowledge of the minor species precludes firm statements on their prospects. A better insight in growth, development and productivity of the trees and acceptability of the fruits is needed. The use of these minor species as rootstocks to adapt the major species to specific conditions also deserves consideration.

Chandlee, D.K., 1988. A guide to Artocarpus fruits. Rare Fruit Council of Australia Newsletter 53(6): 12—18.
Corner, E.J.H., 1988. Wayside trees of Malaya. 3rd ed. Vol. 2. The Malayan Nature Society, Kuala Lumpur. pp. 511—521.
Jarrett, F.M., 1959, 1960. Studies in Artocarpus and allied genera, I—V. Journal of the Arnold Arboretum of Harvard University 40: 1—38, 113—156, 298—369; 41: 73—141, 320—341.
Primack, R.B., 1985. Comparative studies of fruits in wild and cultivated trees of chempedak (Artocarpus integer) and terap (Artocarpus odoratissimus) in Sarawak, East Malaysia, with additional information on the reproductive biology of the Moraceae in South-East Asia. Malayan Nature Journal 39: 1—39.
Voon Boon Hoe, Sim, P. & Chin Thian Hon, 1988. Sayur-sayuran dan buah-buahan hutan di Sarawak [Wild vegetables and fruits in Sarawak]. Department of Agriculture, Sarawak. 55 pp.

B. Seibert & P.C.M. Jansen

Seibert, B. & Jansen, P.C.M., 1991. Artocarpus J.R. & G. Forster. In: Verheij, E.W.M. and Coronel, R.E. (Editors): Plant Resources of South-East Asia No 2: Edible fruits and nuts. PROSEA Foundation, Bogor, Indonesia. Database record: prota4u.org/prosea