Artocarpus altilis (breadfruit)

Identity

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Preferred Scientific Name

Artocarpus altilis (Parkinson) Fosberg

Preferred Common Name

breadfruit

Other Scientific Names

Artocarpus altilis var. non-seminiferus (Duss) Fournet
Artocarpus altilis var. seminiferus (Duss) Fournet
Artocarpus camansi Blanco
Artocarpus communis J. R. Forst. & G. Forst.
Artocarpus incisus (Thunb.) L. f.
Artocarpus incisus var. non-seminiferus Duss
Artocarpus incisus var. seminiferus Duss
Artocarpus laevis Hassk.
Artocarpus rima Blanco
Sitodium altile Parkinson

International Common Names

English: breadnut (seeded varieties); dugdug
Spanish: albopán; àrbo del pan; árbol de pan; castaña; fruta de pan; pana; pana de pepita; panapén
French: âme veritable; arbre à pain; arbre véritable; châtaignier à grains
Portuguese: arbol-pao-de-Brasil; mapen

Local Common Names

: kapiak
Brazil: arvore-do-Pao-fruta; fruta-pao; fruta-pao-de-caroço; pao-de-massa; pao-de-pobre; rima
Cambodia: khanaôr sâmloo; sakéé
Cuba: castaño del malabar; fruto del mar; guapén; mapén; palo de pan
Dominican Republic: abapen; arbapen; buen pan; celipi; juan pan; pan de fruta
Fiji: kulu; uto; uto mbutho; uto ndina
Germany: Brotfruchtbaum, echter
Guatemala: arbol de mazapan
Haiti: châtaignier des Antilles; jaquier; meat fruit; pti-l’arbre à pain; rimier
Indonesia: kelur (seeded); sukun (seedless); timbul (seeded)
Italy: albero del pane; artocarpo
Kiribati: te mai
Lesser Antilles: breadnut; bwapan; châtaigne; châtaignier; fruit á pain; seeded breadfruit
Malaysia: kelor (seeded); kelur; sukun (seedless); timbul
Micronesia, Federated states of: mei
Netherlands: broodvruchtboom
Nicaragua: fruta de pan
Papua New Guinea: kapiak
Philippines: kamansi (seeded); rimas (seedless)
Puerto Rico: lavapén; mapén; pana forastera
Samoa: 'viu
Solomon Islands: bia; nimbalu
Sri Lanka: erapillakai; rata del
Sweden: broedfrukttraed
Thailand: khanun-sampalaor; sa-ke (seedless)
Tonga: mai; mei
Vanuatu: beta
Venezuela: frutepan; pan del año
Vietnam: saké

EPPO code

ABFAL (Artocarpus altilis)

Summary of Invasiveness

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A. altilis is a fast-growing evergreen tree frequently cultivated and naturalized through the tropics (Ragone, 2011). This species is one of the most common trees planted in agroforestry systems in the Pacific islands, but it is also an important starchy staple crop in Oceania, India, Africa, and the West Indies (Orwa et al., 2009). There are seeded and seedless varieties which have been repeatedly introduced and extensively moved by humans within and outside their native distribution range (Morton, 1987; Parrotta, 1994; Ragone, 2006). A. altilis has escaped from cultivation and became naturalized in disturbed and natural areas. Once naturalized, plants grow outcompeting (mainly by shading) seedlings and native understory species (Abreu and Rodrigues, 2010). A. altilis is included in the Global Compendium of Weeds (Randall, 2012) and is listed as invasive in Costa Rica, Brazil, Puerto Rico and the Virgin Islands (Kairo et al., 2003; Chacon and Saborio, 2012; I3N-Brazil, 2014).

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Plantae
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Dicotyledonae
Order: Urticales
Family: Moraceae
Genus: Artocarpus
Species: Artocarpus altilis

Notes on Taxonomy and Nomenclature

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The Moraceae are monoecious or dioecious trees, shrubs, and rarely herbs comprising about 39 genera and 1125 species distributed mostly in tropical to warm temperate regions (Stevens, 2012; Flora of China Editorial Committee, 2014). Nearly all species within this family contain milky latex and have alternate or opposite leaves and small, unisexual, and minute flowers (Wilmott-Dear and Brummitt, 2007). The genus Artocarpus includes 61 species native to the Indian subcontinent, Southeast Asia, and Australasia (Kochummen, 2000; Stevens, 2012). The generic name Artocarpus comes from the Greek words ‘artos’ (bread) and ‘karpos’ (fruit) and many species and varieties produce fruits which are eaten by humans (Orwa et al., 2009; Ragone, 2011).

Description

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Monoecious tree, up to 30 m tall, evergreen in the humid tropics, semi-deciduous in monsoon climates. Trunk straight, 5-8 m tall, 0.6-1.8 m in diameter, often buttressed; trunk of clonally propagated trees branched low; twigs spreading, very thick, with pronounced leaf and stipule scars and lenticels; buds 10-20 cm long, covered with big conical keeled stipules. Leaves alternate, ovate to elliptical in outline, 20-60(-90) x 20-40(-50) cm, undivided when young, older ones entire or deeply pinnately cut into 5-11-pointed lobes, thick, leathery, dark green and shiny above, pale green and rough below, petiole 3-5 cm long. Inflorescences axillary, peduncles 4-8 cm long; male ones drooping, club-shaped, 15-25 x 3-4 cm, spongy, yellow, flowers minute with single stamen; female ones stiffly upright, globose or cylindrical, 8-10 x 5-7 cm, green, flowers numerous, embedded in receptacle, calyx tubular, ovary 2-celled, style narrow, stigma 2-lobed. Fruit a syncarp formed from the entire inflorescence, cylindrical to globose, 10-30 cm in diameter, rind yellow-green, reticulately marked with 4-6-sided faces, sometimes bearing short spines; a large central core is surrounded by numerous abortive flowers which form a pale yellow juicy pulp, the edible portion of the fruit.

Most cultivated breadfruits are seedless, seeded ones are known as breadnuts. Breadnuts bear fleshy prickles, the edible pulp is largely replaced by the seeds, which are brownish, rounded or flattened, 2.5 cm long. All parts of the tree are rich in white gummy latex.

Other Botanical Information

The seeded breadfruit is considered to be the wild form, although both seeded and seedless forms are cultivated. So-called seedless forms occasionally produce fruit with some seeds. Forms with entire leaves and with both seeds and edible pulp are sometimes classified as a separate species: Artocarpus mariannensis Tr cul. Based on leaf, fruit and seed characteristics, hundreds of breadfruit cultivars have been described and named, particularly in the Pacific islands. Usually the cultivars are poorly defined and only grown in one or two localities, so that it is not easy to compare cultivars.

Plant Type

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Perennial
Seed propagated
Tree
Vegetatively propagated

Distribution

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The exact origin of A. altilis is uncertain, but it is believed to be native to Papua New Guinea, the Moluccas (Indonesia), and the Philippines (Kochummen, 2000; Ragone, 2011). This species has long been an important staple food crop in Polynesia and is widely cultivated throughout the tropics (Zerega et al., 2004; Ragone 2006, 2011). A. altilis is now widely distributed throughout the humid tropics, including in Oceania, Asia, Africa, North, Central and South America and the West Indies (see distribution table for details: Morton, 1987; Orwa et al., 2009; Ragone, 2011; Flora of China Editorial Committee, 2014; USDA-ARS, 2014).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 17 Feb 2021

Continent/Country/Region	Distribution	Origin	Invasive	Planted	Reference	Notes
Africa
Benin	Present			Planted	CABI (Undated b)
Cameroon	Present	Introduced			PROTA (2014) CABI (Undated)
Comoros	Present	Introduced			Ragone (2006)
Côte d'Ivoire	Present			Planted	CABI (Undated b)
Gabon	Present			Planted	CABI (Undated b)
Gambia	Present	Introduced			Orwa et al. (2009)
Ghana	Present	Introduced			PROTA (2014) Badii et al. (2015) CABI (Undated)
Guinea	Present	Introduced			Orwa et al. (2009) CABI (Undated)
Kenya	Present			Planted	CABI (Undated b)
Liberia	Present			Planted	CABI (Undated b)
Madagascar	Present	Introduced			Ragone (2006) CABI (Undated)
Mauritius	Present	Introduced			Ragone (2006) CABI (Undated)
Nigeria	Present			Planted	CABI (Undated b)
Seychelles	Present	Introduced			CABI (Undated)
Sierra Leone	Present			Planted	CABI (Undated b)
Tanzania	Present			Planted	CABI (Undated b)
-Zanzibar Island	Present			Planted	CABI (Undated b)
Togo	Present			Planted	CABI (Undated b)
Uganda	Present			Planted	CABI (Undated b)
Asia
Bangladesh	Present			Planted	CABI (Undated b)
Brunei	Present			Planted	CABI (Undated b)
Cambodia	Present			Planted	CABI (Undated b)
China	Present				CABI (Undated a)	Present based on regional distribution.
-Guangdong	Present			Planted	CABI (Undated b)
-Hainan	Present	Introduced			Flora of China Editorial Committee (2014) CABI (Undated)
India	Present	Introduced			Ragone (2006) CABI (Undated)
-Andhra Pradesh	Present	Introduced			Ragone (2006)
-Goa	Present	Introduced			Ragone (2006)
-Karnataka	Present	Introduced			Ragone (2006)
-Kerala	Present	Introduced			Ragone (2006)
-Lakshadweep	Present	Introduced			Ragone (2006)
Indonesia	Present				CABI (Undated a)	Present based on regional distribution.
-Irian Jaya	Present			Planted	CABI (Undated b)
-Java	Present			Planted	CABI (Undated b)
-Maluku Islands	Present	Native			USDA-ARS (2014) CABI (Undated)
-Sulawesi	Present			Planted	CABI (Undated b)
-Sumatra	Present			Planted	CABI (Undated b)
Jordan	Present			Planted	CABI (Undated b)
Laos	Present			Planted	CABI (Undated b)
Malaysia	Present	Native			Orwa et al. (2009)
-Peninsular Malaysia	Present			Planted	CABI (Undated b)
-Sabah	Present			Planted	CABI (Undated b)
-Sarawak	Present			Planted	CABI (Undated b)
Maldives	Present	Introduced			Ragone (2006)
Philippines	Present	Native			USDA-ARS (2014) CABI (Undated)
Singapore	Present			Planted	CABI (Undated b)
Sri Lanka	Present	Introduced			Ragone (2006) Galanihe et al. (2010) CABI (Undated)
Taiwan	Present	Introduced			Flora of China Editorial Committee (2014) CABI (Undated)
Thailand	Present	Introduced			Ragone (2006) CABI (Undated)
Vietnam	Present	Introduced			Orwa et al. (2009) CABI (Undated)
North America
Anguilla	Present, Widespread	Introduced			Broome et al. (2007)
Antigua and Barbuda	Present, Widespread	Introduced			CABI (Undated b)
Bahamas	Present			Planted	CABI (Undated b)
Barbados	Present, Widespread	Introduced			Broome et al. (2007)
Belize	Present	Introduced	Invasive		Meerman (2008)
British Virgin Islands	Present	Introduced			Acevedo-Rodríguez and Strong (2012)	Cultivated
Costa Rica	Present	Introduced	Invasive		CABI (Undated)	Original citation: Chacon and Saborio (2012)
Cuba	Present	Introduced	Naturalized		Kairo et al. (2003)	Cultivated and naturalized
Dominica	Present, Widespread	Introduced			Broome et al. (2007)
Dominican Republic	Present	Introduced	Naturalized		Kairo et al. (2003) CABI (Undated)	Cultivated and naturalized
El Salvador	Present	Introduced			Davidse et al. (2014)
Grenada	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Guadeloupe	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Guatemala	Present	Introduced			Orwa et al. (2009) CABI (Undated)
Haiti	Present	Introduced	Naturalized		Kairo et al. (2003) CABI (Undated)	Cultivated and naturalized
Honduras	Present	Introduced			Davidse et al. (2014) CABI (Undated)
Jamaica	Present	Introduced	Naturalized		Kairo et al. (2003) CABI (Undated)	Cultivated and naturalized
Martinique	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Mexico	Present			Planted	Orwa et al. (2009)
Montserrat	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Netherlands Antilles	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Nicaragua	Present	Introduced			Davidse et al. (2014) CABI (Undated)
Panama	Present	Introduced			Davidse et al. (2014) CABI (Undated)
Puerto Rico	Present	Introduced	Invasive		Acevedo-Rodríguez and Strong (2012) CABI (Undated)
Saint Kitts and Nevis	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Saint Lucia	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Saint Vincent and the Grenadines	Present, Widespread	Introduced			Broome et al. (2007) CABI (Undated)
Trinidad and Tobago	Present	Introduced			Orwa et al. (2009) CABI (Undated)
U.S. Virgin Islands	Present	Introduced	Invasive		Acevedo-Rodríguez and Strong (2012)
United States	Present				CABI (Undated a) Lau et al. (2018)	Present based on regional distribution.
-Florida	Present	Introduced			Morton (1987) CABI (Undated)
-Hawaii	Present	Introduced	Naturalized		Ragone (2006) Lau et al. (2018) CABI (Undated)	Cultivated and naturalised
Oceania
American Samoa	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Australia	Present				CABI (Undated a)	Present based on regional distribution.
-Northern Territory	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
-Queensland	Present			Planted	CABI (Undated b)
Christmas Island	Present			Planted	CABI (Undated b)
Cook Islands	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Federated States of Micronesia	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Fiji	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
French Polynesia	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Guam	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Kiribati	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Marshall Islands	Present	Introduced			Orwa et al. (2009) CABI (Undated)
Nauru	Present			Planted	CABI (Undated b)
New Caledonia	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Niue	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Norfolk Island	Present	Introduced	Naturalized		Orwa et al. (2009)	Cultivated and naturalized
Northern Mariana Islands	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Palau	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Papua New Guinea	Present	Native			USDA-ARS (2014) CABI (Undated)
Pitcairn	Present			Planted	CABI (Undated b)
Samoa	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Solomon Islands	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Tokelau	Present			Planted	CABI (Undated b)
Tonga	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Tuvalu	Present			Planted	CABI (Undated b)
Vanuatu	Present	Introduced	Naturalized		Ragone (2006) CABI (Undated)	Cultivated and naturalized
Wallis and Futuna	Present			Planted	CABI (Undated b)
South America
Bolivia	Present	Introduced			Davidse et al. (2014)
Brazil	Present			Planted	CABI (Undated b)
-Alagoas	Present	Introduced			Romaniuc et al. (2014)
-Amazonas	Present	Introduced			Romaniuc et al. (2014)
-Bahia	Present	Introduced	Invasive		I3N Brasil (2014)
-Ceara	Present	Introduced			Romaniuc et al. (2014)
-Espirito Santo	Present	Introduced			Romaniuc et al. (2014)
-Maranhao	Present	Introduced			Romaniuc et al. (2014)
-Minas Gerais	Present	Introduced			Romaniuc et al. (2014)
-Para	Present	Introduced			Romaniuc et al. (2014)
-Paraiba	Present	Introduced			Romaniuc et al. (2014)
-Pernambuco	Present	Introduced			Romaniuc et al. (2014)
-Piaui	Present	Introduced			Romaniuc et al. (2014)
-Rio de Janeiro	Present	Introduced			Romaniuc et al. (2014)
-Rio Grande do Norte	Present	Introduced			Romaniuc et al. (2014)
-Rio Grande do Sul	Present	Introduced			Romaniuc et al. (2014)
-Rondonia	Present	Introduced			Romaniuc et al. (2014)
-Roraima	Present	Introduced			Romaniuc et al. (2014)
-Sergipe	Present	Introduced			Romaniuc et al. (2014)
Colombia	Present	Introduced			CABI (Undated)	Original citation: Idarraga-Piedrahita et al. (2011)
Ecuador	Present	Introduced			Davidse et al. (2014)
French Guiana	Present			Planted	CABI (Undated b)
Guyana	Present			Planted	CABI (Undated b)
Venezuela	Present	Introduced			Hokche et al. (2008) CABI (Undated)

History of Introduction and Spread

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A. altilis was first domesticated in the western Pacific and was spread by humans throughout Oceania and the numerous islands of Melanesia, Micronesia, and Polynesia where it has been an important staple crop and component of traditional Pacific agroforestry systems for more than 3000 years (Zerega et al., 2004; Ragone 2006, 2011).

During the sixteenth and seventeenth centuries, seeded breadfruit trees from the Philippines were introduced by Spaniards to Mexico and Central America (Parrotta, 1994). In the late 1700s seedless breadfruit trees from Tahiti were introduced by British explorers to Jamaica and St Vincent to provide cheap food for their slaves (Morton, 1987; Ragone, 2011). During this period, a Tongan variety was also introduced to Martinique and Cayenne via Mauritius by French explorers (Ragone, 2011). In 1772, the French navigator Sonnerat obtained seeded breadfruit trees in the Philippines and brought them to the French West Indies, and in 1874, seeded breadfruit trees were introduced to Jamaica (Morton, 1987; Little and Skolmen, 2003). These seedless and seeded plants were then spread to other Caribbean islands, and throughout Central and South America (Morton, 1987; Ragone, 2006). In Brazil, Artocarpus species were introduced around the 1860s (Abreu and Rodrigues, 2010). In 1906, the United States Department of Agriculture introduced breadfruit trees from Panama into Florida (Morton, 1987).

Risk of Introduction

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Even when it has been suggested that A. altilis has a limited potential for invasiveness (Ragone, 2006; 2011), seeded varieties have escaped from cultivation and naturalized in tropical areas (Kairo et al., 2003). In addition, this species has been actively planted by humans for centuries, and currently breadfruit trees are common elements of disturbed secondary forests, forest edges, roadsides, and old plantations and gardens throughout the tropics (Little and Skolmen, 2003, Orwa et al., 2009). Consequently, the likelihood of this species escaping from cultivation and invading new habitats remains high.

Habitat

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A. altilis can be found cultivated and naturalized in the warm and very humid lowlands in house gardens, active and old plantations, agroforestry systems, disturbed secondary forests, forest edges, roadsides, and coastal forests (Parrotta, 1994; Little and Skolmen, 2003; Orwa et al., 2009; Ragone, 2011). This species grows best in lowlands at elevations from sea level to 650 m, but it is occasionally found in highlands up to 1550 m (Orwa et al., 2009; Ragone, 2011).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
Terrestrial	Managed	Cultivated / agricultural land	Present, no further details	Natural
Terrestrial	Managed	Cultivated / agricultural land	Present, no further details	Productive/non-natural
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Natural
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Productive/non-natural
Terrestrial	Managed	Disturbed areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Disturbed areas	Present, no further details	Natural
Terrestrial	Managed	Disturbed areas	Present, no further details	Productive/non-natural
Terrestrial	Managed	Rail / roadsides	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Rail / roadsides	Present, no further details	Natural
Terrestrial	Managed	Rail / roadsides	Present, no further details	Productive/non-natural
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Natural
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Productive/non-natural
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Natural
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Productive/non-natural
Littoral		Coastal areas	Present, no further details	Harmful (pest or invasive)
Littoral		Coastal areas	Present, no further details	Natural
Littoral		Coastal areas	Present, no further details	Productive/non-natural

Biology and Ecology

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Genetics

A. altilis is genetically diverse, especially the seeded varieties in the western Pacific (Ragone, 2011). There are diploid (2n = 56) and triploid (2n = 84) forms and numerous varieties have been developed and are perpetuated by vegetative propagation (Morton, 1987; Ragone, 2001; Orwa et al., 2009).

Reproductive Biology

A. altilis trees are monoecious (i.e., male and female flowers occur on the same tree). The flowering pattern appears to depend on the cultivar rather than the climate, at least in the humid tropics. Most cultivars flower throughout the year, but some flower and fruit seasonally. Further from the equator, however, the seasons determine shoot growth and all cultivars flower and fruit more or less simultaneously. The male and female inflorescences are produced in separate leaf axils of the current season's growth. Male inflorescences emerge before the female ones. The proportion of male inflorescences on a tree may vary from 60-80%. Flowers are visited and pollinated by insects, mostly bees, but wind-aided pollination has also been suggested (Parrotta, 1994): if tapped at anthesis, the flowers release clouds of pollen. The male inflorescences reach anthesis 10-15 days after emergence and well before the female inflorescences on the same tree, thus limiting self-pollination. Female flowers are receptive 3 days after the emergence of the female inflorescence from the bracts and open in successive stages, with basal flowers opening first (Orwa et al., 2009; Flora of China, 2014).

The flowers in a female inflorescence are pollinated and fertilized (in seeded cultivars) within a period of 3-6 days. About 75% of the inflorescences set fruit, but this is considerably reduced during the rainy season, strengthening the impression that the seedless fruits also depend on pollination to stimulate parthenocarpic growth. The fruit matures 60-90(-110) days after the inflorescence emerges. In the seeded form there is little pulp and each fruit contains 20-60 edible seeds.

Physiology and Phenology

Seeds germinate about 2 weeks after sowing. The seedlings tend to grow slowly, but they respond well to better growing conditions. Under favorable environmental conditions, trees grow 0.5-1.5 m in height per year and 0.5-1 m in trunk diameter in the first 10-12 years (Ragone, 2011). Trees growing from seeds begin flowering and fruiting in 6 to 10 years while vegetatively propagated trees start flowering and fruiting in 3 to 6 years (Ragone, 2006). Under stress the trees shed most of their leaves, tufts of young leaves persisting at the tip of the twigs.

A. altilis trees produce flowers and fruits throughout the year. Fruits in all stages of development can be found on trees the year around, but there are two or three main fruiting periods (Morton, 1987). In the Pacific region, the main crop typically occurs during the hot, rainy, summer months, followed by a smaller crop 3 or 4 months later (Ragone, 2006).

Longevity

A. altilis is a perennial, long-lived tree. Small branches often die back after fruiting, but new shoots and branches are continually developed throughout the life of the tree (Ragone, 2006).

Associations

In the Pacific islands, A. altilis is often planted in agroforestry systems with yam (Dioscorea spp), taro (Colocasia spp), Tahitian chestnut (Inocarpus fagifer), cassava, noni (Morinda citrifolia), kava (Piper methysticum), bananas, citrus, papaya, coconut, coffee, and cacao (Ragone, 2006). In Papua New Guinea, A. altilis grows naturally in wet lowland forests in association with Alstonia scholaris, Dracontomelon dao, Garuga spp., Octomeles sumatrana, Allophyluscobbe, Pterocymbium spp., and Terminalia catappoides (Parrotta, 1994). In Puerto Rico, cultivated and naturalized A. altilis plants can be found growing in association with native and alien species such as Andira inermis, Genipa americana, Guarea guidonia, Mangifera indica and Tabebuia heterophylla (Parrotta, 1994).

Environmental Requirements

A. altilis prefers to grow in warm and wet lowland areas of the tropics. The normal latitudinal limits are approximately 17°N and S; the maritime climate of small islands allows growth to 20-23°N. This species prefers climates with bimodal rainfall with mean annual precipitation ranging from 1500 to 3000 mm (Ragone, 2006), and relative humidity of 70-90%. A. altilis grows best in deep, fertile (pH 6.1-7.4), and well drained soils. It tolerates saline soils and shaded conditions (20-50% shade). Plants are able to grow in coralline, sandy, sandy loam and sandy clay soils but do not tolerate frost or water-logged soils (Ragone, 2011). Cultivars differ greatly in their tolerance of adverse conditions; there are cultivars that cope well with shallow calcareous soils, brackish water and salt sprays, annual rainfall of only 1500 or 1000 mm, etc.

The tree is occasionally found in the highlands (even up to 1500 m) and at higher latitudes, but yield and fruit quality suffer in cooler conditions and the tree is more at home in the equatorial lowlands (below 600 m). Whereas the trees shed their leaves under dry conditions, it is said that they shed their fruit when the soil is excessively wet; so yield may be depressed on marginal soils.

Climate

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Climate	Status	Description
Af - Tropical rainforest climate	Preferred	> 60mm precipitation per month
Am - Tropical monsoon climate	Preferred	Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer	Tolerated	< 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate	Tolerated	< 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
20	-20	0	1500

Air Temperature

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Parameter	Lower limit	Upper limit
Absolute minimum temperature (ºC)	7	17
Mean annual temperature (ºC)	23	27
Mean maximum temperature of hottest month (ºC)	26	32
Mean minimum temperature of coldest month (ºC)	20	25

Rainfall

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Parameter	Lower limit	Upper limit	Description
Dry season duration	0	3	number of consecutive months with <40 mm rainfall
Mean annual rainfall	1000	5000	mm; lower/upper limits

Rainfall Regime

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Bimodal
Summer
Uniform

Soil Tolerances

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Soil drainage

free

Soil reaction

acid
alkaline
neutral

Soil texture

heavy
light
medium

Special soil tolerances

saline

Means of Movement and Dispersal

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A. altilis spreads sexually by seeds and vegetatively by root cuttings, suckers, or layering (Orwa et al., 2009; Ragone, 2011). Vegetative propagation is a must for seedless varieties. Seed germination occurs immediately after dispersal, but seeds also lose viability quickly and cannot be dried or stored (Ragone, 2011).

Pathway Causes

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Cause	Notes	Long Distance	Local	References
Crop production	Cultivated for fruits and seeds	Yes	Yes	Ragone, 2011
Disturbance	Naturalized in disturbed areas	Yes	Yes	Little and Skolmen, 2003
Escape from confinement or garden escape		Yes	Yes	Orwa et al., 2009
Food		Yes	Yes	Ragone, 2011
Forestry	Widely cultivated in agroforestry	Yes	Yes	Ragone, 2011
Medicinal use	Used in traditional medicine in the Pacific and Caribbean	Yes	Yes	Ragone, 2006
Ornamental purposes	Planted as shade tree in gardens	Yes	Yes	Ragone, 2006

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Debris and waste associated with human activities		Yes	Yes	Ragone, 2011

Impact Summary

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Category	Impact
Economic/livelihood	Positive and negative
Environment (generally)	Positive and negative
Human health	Positive and negative

Environmental Impact

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A. altilis has escaped from cultivation and became naturalized in disturbed and natural areas. Once naturalized, plants grow outcompeting (shading) seedlings and native species in the understory of native forests (Abreu and Rodrigues, 2010). Because this species spreads by seeds and vegetatively by suckers and cuttings, it has the potential to create monospecific stands displacing slow growing native plants and smothering native plant communities (Kairo et al., 2003; Chacon and Saborio, 2012; I3N-Brazil, 2014).

Risk and Impact Factors

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Invasiveness

Proved invasive outside its native range
Has a broad native range
Abundant in its native range
Highly adaptable to different environments
Is a habitat generalist
Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
Pioneering in disturbed areas
Tolerant of shade
Highly mobile locally
Benefits from human association (i.e. it is a human commensal)
Long lived
Fast growing
Has high reproductive potential
Reproduces asexually

Impact outcomes

Damaged ecosystem services
Ecosystem change/ habitat alteration
Modification of successional patterns
Monoculture formation
Reduced native biodiversity
Threat to/ loss of native species

Impact mechanisms

Causes allergic responses
Competition - monopolizing resources
Competition - shading
Pest and disease transmission
Rapid growth
Rooting

Likelihood of entry/control

Highly likely to be transported internationally deliberately

Uses

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A. altilis is often planted for its edible fruits and seeds, but trees are also attractive for ornament and shade (Orwa et al., 2009). Breadfruit is consumed as a starchy staple when mature mostly in the Pacific islands, Africa and the West Indies. Fruits can be eaten raw, boiled, steamed or roasted (Ragone, 2006; Orwa et al., 2009). The fruit may be cooked whole or after cutting it; thin slices are also fried. A kind of biscuit is made by slicing the ripe cooked fruit and drying in the sun or in an oven; thus prepared it can be kept until the next fruiting season. On many Pacific islands breadfruit is preserved in pits or by burying (Samoa). The stored fruit ferments and is converted into a nutritious but disagreeably smelling cheese-like paste, which is made into cakes and baked. Commercial processing is limited to preserving the boiled cut fruit in brine. In the Philippines, the mature seedless fruit is boiled and eaten with sugar and grated coconut or coated with sugar and dehydrated; immature seeded fruit is cooked as a vegetable with coconut milk.

In other areas, such as Mexico and Central America, breadfruit is used principally as forage and fodder for cattle, goats, pigs, and horses. Foliage is also served as forage for cattle during periods of drought in Hawaii (Little and Skolmen, 2003).

The male flower spikes are blended with fibre of paper mulberry (Broussonetia papyrifera) to make elegant loincloths. The smooth, grey bark is fibrous and was once a source of the native cloth 'tapa'. The milky sap is used to caulk canoes, as a glue to catch birds and as a chewing gum. The light yet quite firm, nicely grained wood is used for surf boards, canoes, furniture, carvings and in construction of houses. Wood is resistant to termites and marine worms (Orwa et al., 2009). Trees are also an important source of firewood on islands in the Pacific (Ragone, 2006).

A. altilis is a cultural icon in the Pacific (Ragone, 2009). All parts of the plant (especially the latex, leaves, and the inner bark) are used in traditional medicine to treat headaches, diarrhea, stomachache, and dysentery (Ragone, 2011). In the West Indies, the yellowing leaf is brewed into a tea and taken to reduce high blood pressure. The tea is also thought to control diabetes (Orwa et al., 2009) and relieve asthma. Dried flowers can be burned to repel mosquitoes and other flying insects (Ragone, 2011). Chewed young leaves are said to counteract food poisoning.

In the Pacific islands, A. altilis is also planted as soil improver and to support yam vines in traditional agroforestry systems (Ragone, 2006). Trees are also planted as windbreaks and sometimes as shade trees for coffee (Jensen, 1995).

Uses List

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Animal feed, fodder, forage

Fodder/animal feed
Forage

Environmental

Agroforestry
Amenity
Boundary, barrier or support
Shade and shelter
Soil improvement
Windbreak

Fuels

Fuelwood

General

Ornamental

Human food and beverage

Flour/starch
Fruits
Gum/mucilage
Honey/honey flora
Seeds
Vegetable

Materials

Carved material
Dye/tanning
Fibre
Gum/resin
Miscellaneous materials
Pesticide
Resins
Rubber/latex
Wood/timber

Medicinal, pharmaceutical

Source of medicine/pharmaceutical
Traditional/folklore

Ornamental

Propagation material
Seed trade

Wood Products

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Boats

Containers

Boxes
Crates

Furniture

Sawn or hewn building timbers

For heavy construction
For light construction

Woodware

Wood carvings

Bibliography

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Barrau J, 1976. Breadfruit and its relatives: Artocarpus spp. In: Simmonds NW, ed. Evolution of Crop Plants. London & New York: Longman, 201-202.

Coenen J, Barrau J, 1961. The breadfruit tree in Micronesia. South Pacific Bulletin, 11(4): 31-39, 65-67.

Morton JF, 1987. Fruits of warm climates. Creative Resource Systems Inc., Winterville N.C., 50-58.

Otanes RT, Ruiz TP, 1956. Propagation of rimas root cuttings. Araneta Journal of Agriculture, 3(2): 56-66.

Reeve RM, 1974. Histological structure and commercial dehydration potential of breadfruit. Economic Botany, 28(1): 82-96.

Sinha MM, Sinha SM, 1968. Breadfruit. Science and Culture, 34:271.

References

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Abreu RC, Rodrigues PJF, 2010. Exotic tree Artocarpus heterophyllus (Moraceae) invades the Brazilian Atlantic Rainforest. Rodriguésia-Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, 61:677-688.

Acevedo-Rodríguez P, Strong MT, 2012. Catalogue of the Seed Plants of the West Indies. Smithsonian Contributions to Botany, 98:1192 pp. Washington DC, USA: Smithsonian Institution. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Agounke D, Agricola U, Bokonon-Ganta A, 1989. Rastrococcus invadens Williams (Homoptera, Pseudococcidae), an exotic pest of fruit trees in West Africa. Bulletin du Service de la Protection des Vegetaux, 11:17-29

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database. Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.html

Cambie RC, Ash J, 1994. Fijian medicinal plants. Commonwealth Scientific and Industrial Research Organisation, Australia.

Chacón E, Saborío G, 2012. Red Interamericana de Información de Especies Invasoras, Costa Rica ([English title not available]). San José, Costa Rica: Asociación para la Conservación y el Estudio de la Biodiversidad. http://invasoras.acebio.org

Davidse G, Sousa M, Knapp S, Chiang F, 2014. Saururaceae a Zygophyllaceae. Flora Mesoamericana [ed. by Davidse, G. \Sousa, M. \Knapp, S. \Chiang, F.]., Mexico: Universidad Nacional Autónoma de México.

FAO, 1982. Fruit-bearing forest trees: technical notes. FAO Forestry Paper No. 34. Rome, Italy: Food and Agriculture Organization.

Flora of China Editorial Committee, 2014. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Gardner DE, 1991. Occurrence of breadfruit rust, caused by Uredo artocarpi, in Hawaii. Plant Disease, 75(9):968

Garner RJ, Chaudhri SA, Commonwealth Bureau of Horticulture and Plantation Crops, 1976. The propagation of tropical fruit trees. Horticultural Review, Commonwealth Bureau of Horticulture and Plantation Crops, No. 4, 566 pp.; 48 fig.; many ref.

Gupta RK, 1993. Multipurpose trees for agroforestry and wasteland utilisation. Multipurpose trees for agroforestry and wasteland utilisation., xv + 562 pp.; [18 pp. of ref + refs in text].

Hearne DA, 1975. Trees for Darwin and northern Australia. Canberra, Australia: Australian Government Publishing Service.

Henderson CP, Hancock IR, 1988. A guide to the Useful Plants of Solomon Islands. Solomon Islands Research Department/Ministry of Agriculture and Lands, Honiara.

Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela (New catalogue of the vascular flora of Venezuela). Caracas, Venezuela: Fundación Instituto Botánico de Venezuela, 860 pp.

I3N Brasil, 2014. National database of exotic invasive species (Base de dados nacional de espécies exóticas invasoras). Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

Idárraga-Piedrahita A, Ortiz RDC, Callejas Posada R, Merello M, 2011. Flora of Antioquia. (Flora de Antioquia.) Catálogo de las Plantas Vasculares, vol. 2. Listado de las Plantas Vasculares del Departamento de Antioquia:939 pp.

Janick, J., Paull, R. E., 2008. The encyclopedia of fruit & nuts., The encyclopedia of fruit & nuts:xviii + 954 pp.

Jensen M, 1995. Trees commonly cultivated in Southeast Asia: an illustrated field guide. Bangkok, Thailand: FAO-Regional Office for Asia and the Pacific (RAP), No-38, vii + 229 pp.; 29 ref.

Kairo M, Ali B, Cheesman O, Haysom K, Murphy S, 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International, 132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Kochummen KM, 2000. Artocarpus J. & G. Forster, nom. conserv. In: Tree flora of Sabah and Sarawak [ed. by Soepadmo, E. \Saw, L. G.]. Kuala Lumpur, Malaysia: Sabah Forestry Department, Forest Research Institute Malaysia, and Sarawak Forestry Department, 187-212.

Lepofsky D, 1992. Arboriculture in the Mussau Islands, Bismarck Archipelago. Economic Botany, 46(2):192-211; 21 ref.

Little EL Jr, Skolmen RG, 1989. Common forest trees of Hawaii (native and introduced). Agriculture Handbook Washington, 679. Washington, DC, USA: US Department of Agriculture, Forest Service.

Little Jr EL, Skolmen RG, 2003. Common Forest Trees of Hawaii (Native and Introduced). Manoa, USA: College of Tropical Agriculture and Human Resources, University of Hawaii. http://www.ctahr.hawaii.edu/forestry/trees/little_skolmen.html

Matthews RF, Bates RP, Graham HD, 1986. Utilization of breadfruit in the tropics. Proceedings of the Interamerican Society for Tropical Horticulture, 30:83-94; [34th Annual Meeting, San José, Costa Rica, 28 Jul.-2 Aug., 1986]; 15 ref.

Meerman JC, 2008. Species List of Non-native Flora of Belize. http://biological-diversity.info/invasive_flora.htm

Moore D, Fischer HU, Agounke D, 1988. Biological control of Rastrococcus invadens Williams in Togo. FAO Plant Protection Bulletin, 36(4):169-174

Morton JF, 1987. Breadfruit. In: Fruits of warm climates [ed. by Morton, J. F.]. Miami, Florida, USA: JF Morton, 50-58.

Mridha AU, Basak AB, Uddin MJ, 1990. A record of leaf spot disease of jack fruit trees by Colletotrichum gloeosporioides [Glomerella cingulata] Penz. from Bangladesh. Bangladesh Journal of Forest Science, 19(1/2):59-61; 7 ref.

Orwa C, Mutua A, Kindt R, Jamnadass R, Simons A, 2009. Artocarpus altilis. Agroforestree Database: a tree reference and selection guide version 4. http://www.worldagroforestry.org/af/treedb/

Parrotta J, 1994. Artocarpus altilis (S. Park.) Fosb. Breadfruit, breadnut. SO-ITF-SM-71. New Orleans, LA, USA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. http://www.fs.fed.us/global/iitf/Artocarpusaltilis.pdf

Paull, R. E., Duarte, O., 2012. Tropical fruits, Volume 2., Tropical fruits, Volume 2:ix + 371 pp. http://www.cabi.org/cabebooks/ebook/20123357661

Peters FE, Wills PA, 1956. Dried breadfruit. Nature, 178:1252.

PROTA, 2014. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Purseglove JW, 1968. Tropical crops: dicotyledons 1 & 2. Longmans, London. pp. xiv + viii + 719.

Ragone D, 1989. Our breadfruit collection grows. Pacific Tropical Botanical Garden Bull., 19(3):86-89.

Ragone D, 1990. Conservation and use of breadfruit in the Pacific Islands. In: Harris W, Kapoor P, eds. Contributions to an international Workshop on Ethnobotany. Botany Division, DSIR, Christchurch, New Zealand, 82-85.

Ragone D, 1991. Collection, establishment and evaluation of a germplasm collection of Pacific Island breadfruit. PhD dissertation. University of Hawaii, Honolulu, USA.

Ragone D, 1997. Breadfruit. Artocarpus altilis (Parkinson) Fosberg. Promoting the conservation and use of underutilised and neglected crops. 10. Institute of Plant Genetics and Crop Plant Research, Gatersleben/International Plant Genetic Resources Institute, Rome, Italy.

Ragone D, 2001. Chromosome numbers and pollen stainability of three species of Pacific Island breadfruit (Artocarpus, Moraceae). American Journal of Botany, 88(4):693-696.

Ragone D, 2006. Artocarpus altilis (breadnut), ver. 2.1. Species profiles for Pacific island agroforestry. Permanent Agriculture resources (PAR), Holualoa, Hawaii [ed. by Elevitch, C. R.]. http://www.agroforestry.net/tti/A.altilis-breadfruit.pdf

Ragone D, 2011. Artocarpus altilis (revised). Farm and Forestry Production and Marketing for Breadfruit. Species profiles for Pacific island agroforestry. Permanent Agriculture resources (PAR), Holualoa, Hawaii [ed. by Elevitch, C. R.]. http://www.agroforestry.net/scps/Breadfruit_specialty_crop.pdf

Ragone D, Tukey HB Jr, 1995. Description of Pacific Island breadfruit cultivars. Proceedings of the second international symposium on the taxonomy of cultivated plants, held at Seattle, Washington, USA, on 10 14 August, 1994. Acta-Horticulturae, No. 413, 93-98; 8 ref.

Rajendran R, 1991. Artocarpus altilis (Parkinson) Fosberg. In: Plant resources of South-East Asia. No. 2. Edible Fruits and Nuts (Verheij EWM, Coronel RE, eds.). PROSEA foundation, Bogor, Indonesia, 83-86.

Randall RP, 2012. A Global Compendium of Weeds. Perth, Australia: Department of Agriculture and Food Western Australia, 1124 pp. http://www.cabi.org/isc/FullTextPDF/2013/20133109119.pdf

Razak AR, 1978. Variation in plant response, gall size and form induced by Meloidogyne on some Malaysian crops. Kasetsart Journal, 12(1):43-45

Reeve RM, 1974. Histological structure and commercial dehydration potential of breadfruit. Economic Botany, 28:82-96.

Romaniuc NS, Carauta JPP, Vianna MDM, Pereira RAS, Ribeiro JEL, Machado AFP, Santos A, Pelissari G, Pederneiras LC, 2014. Moraceae in Lista de Espécies da Flora do Brasil (Moraceae in the list of species of the flora of Brazil). Rio de Janeiro, Brazil: Jardim Botânico do Rio de Janeiro. http://reflora.jbrj.gov.br/jabot/floradobrasil/FB85711

Sasuke N, 1953. Breadfruit, yams and taros of Ponape Island. Proceedings of the 7th Pacific Science Association Congress, 6:159-170.

Smith AC, 1981. Flora Vitiensis nova: a new flora of Fiji (spermatophytes only). Volume 2. 1981, 818 pp.; many pl. (8 col.).

Stevens PF, 2012. Angiosperm Phylogeny Website. http://www.mobot.org/MOBOT/research/APweb/

Streets RJ, 1962. Exotic forest trees in the British Commonwealth. Oxford, UK: Clarendon Press.

Suharban M, Philip S, 1987. Fruit rot of bread fruit (Artocarpus incisa L.). South Indian Horticulture, 35(5):397

USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

Verheij EWM, Coronel RE(Editors), 1991. Plant resources of South-East Asia. No. 2. Edible fruits and nuts. Wageningen, Netherlands; Pudoc, 446 pp.

Waterhouse DF, 1991. Possibilities for the biological control of the breadfruit mealybug, Icerya aegyptiaca, on Pacific Atolls. Micronesica, No. 3, Supplement:117-122

Whistler WA, 1992. Tongan Herbal Medicine. Isle Botanica, Hawaii, USA.

Whistler WA, 1996. Samoa Herbal Medicine. O le Siosiomaga Society Inc. of Western Samoa. Isle Botanica, Hawaii, USA.

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Wootton M, Tumaalii F, 1984. Breadfruit production, utilisation and composition - a review. Food Technology in Australia, 36(10):464-465; 19 ref.

Yen DE, 1974. Arboriculture in the subsistence of Santa Cruz, Solomon Islands. Economic Botany, 28(3):247-284; 47 ref.

Zerega NJC, Ragone D, Motley TJ, 2004. Complex origins of breadfruit (Artocarpus altilis, Moraceae): implications for human migrations in Oceania. American Journal of Botany, 91(5):760-766.

Distribution References

Acevedo-Rodríguez P, Strong M T, 2012. Catalogue of the Seed Plants of the West Indies. Washington, DC, USA: Smithsonian Institution. 1192 pp. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Badii K B, Billah M K, Afreh-Nuamah K, Obeng-Ofori D, 2015. Species composition and host range of fruit-infesting flies (Diptera: Tephritidae) in northern Ghana. International Journal of Tropical Insect Science. 35 (3), 137-151. http://journals.cambridge.org/action/displayJournal?jid=JTI

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database., Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.htm

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Davidse G, Sousa M, Knapp S, Chiang F, 2014. (Saururaceae a Zygophyllaceae). In: Flora Mesoamericana, [ed. by Davidse G, Sousa M, Knapp S, Chiang F]. Mexico: Universidad Nacional Autónoma de México.

Flora of China Editorial Committee, 2014. Flora of China., St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Galanihe L D, Jayasundera M U P, Vithana A, Asselaarachchi N, Watson G W, 2010. Occurrence, distribution and control of papaya mealybug, Paracoccus marginatus (Hemiptera: Pseudococcidae), an invasive alien pest in Sri Lanka. Tropical Agricultural Research and Extension. 13 (3), 81-86. http://www.sljol.info/index.php/TARE/article/view/3143/2522

Hokche O, Berry PE, Huber O, 2008. New catalogue of the vascular flora of Venezuela. (Nuevo Catálogo de la Flora Vascular de Venezuela)., Caracas, Venezuela: Fundación Instituto Botánico de Venezuela. 860 pp.

I3N Brasil, 2014. National database of exotic invasive species. (Base de dados nacional de espécies exóticas invasoras)., Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

Kairo M, Ali B, Cheesman O, Haysom K, Murphy S, 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. In: Invasive species threats in the Caribbean region. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International. 132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Lau J W, Sipes B S, Wang K H, Marahatta S P, Ragone D, 2018. First report of Pratylenchus coffeae on breadfruit (Artocarpus altilis) in the United States. Plant Disease. 102 (9), 1861. DOI:10.1094/pdis-02-18-0242-pdn

Meerman JC, 2008. Species List of Non-native Flora of Belize., http://biological-diversity.info/invasive_flora.htm

Morton JF, 1987. Breadfruit. In: Fruits of warm climates, [ed. by Morton JF]. Miami, Florida, USA: JF Morton. 50-58.

Orwa C, Mutua A, Kindt R, Jamnadass R, Simons A, 2009. Artocarpus altilis. In: Agroforestree Database: a tree reference and selection guide version 4, http://www.worldagroforestry.org/af/treedb/

PROTA, 2014. PROTA4U web database., [ed. by Grubben GJH, Denton OA]. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Ragone D, 2006. Artocarpus altilis (breadnut), ver. 2.1. Species profiles for Pacific island agroforestry. In: Permanent Agriculture resources (PAR), [ed. by Elevitch CR]. Holualoa, Hawaii, http://www.agroforestry.net/tti/A.altilis-breadfruit.pdf

Romaniuc NS, Carauta JPP, Vianna MDM, Pereira RAS, Ribeiro JEL, Machado AFP, Santos A, Pelissari G, Pederneiras LC, 2014. Moraceae in the list of species of the flora of Brazil. (Moraceae in Lista de Espécies da Flora do Brasil)., Rio de Janeiro, Brazil: Jardim Botânico do Rio de Janeiro. http://reflora.jbrj.gov.br/jabot/floradobrasil/FB85711

USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Links to Websites

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Website	URL	Comment
Breadfruit Institute	http://ntbg.org/breadfruit/
PROTA: Plant Resources of Tropical Africa	http://www.prota4u.org/
Purdue University Center for New Crops & Plant Products	https://www.hort.purdue.edu/newcrop/default.html
World Agroforestry Centre	http://www.worldagroforestry.org

Contributors

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25/03/14 Updated by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

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Artocarpus altilis (breadfruit)

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