Annona muricata
(soursop)

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Identity

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Preferred Scientific Name

• Annona muricata L.

Preferred Common Name

• soursop

Other Scientific Names

• Annona macrocarpa Barb. Rodr.
• Annona muricata L. f. mirabilis R.E. Fr.
• Annona muricata L. var. borinquensis Morales

International Common Names

• English: graviola; prickly custard apple
• Spanish: anona; guanabana; guanábana; guanábano; sapote agrio
• French: anone; anone muriquee; cacheimantier èpineux; cachiman; cachiman èpineux; corossol; corossol épineux; corossolier; grand corossol

Local Common Names

• Bolivia: sinini
• Brazil: araticum; araticum-de-comer; araticum-do-Grande; araticum-manso; areticum; ata-coraçao-de-rainha; condessa; coraçao-de-rainha; graviola; graviola-do-norte; guanabano; jaca; jaca-de-pobre; jaca-do-pará; jaqueira-mole; pinha
• Cambodia: tiep banla; tiep barang
• China: ci guo fan li zhi
• Cook Islands: katara‘apa; naponapo taratara
• Fiji: sarifa; seremaia
• French Polynesia: korosor; tapotapo papa‘a; tapotapo urupe
• French Polynesia/Marquesas: koroso
• Germany: Annone, Stachel-; Sauerapfelbaum; Sauersackbaum; Sauersak
• Guam: laguana; laguanaba; laguanaha
• Indonesia: nangka seberang; sirsak; zuurzak
• Indonesia/Java: nangka belanda
• Italy: annona moricata; annona spinosa; casimentaria; pomo di canella
• Japan: reishi; togeban
• Laos: khan thalot; khièp thét
• Malaysia: durian belanda; durian blanda; durian maki; durian makkah; seri kaya belanda
• Marshall Islands: jojaab; sauer sharp
• Mexico: cabeza de negro; caduts-at; catuche; catucho; llama de tehuantepec; polvox; tak-ob; xunapill; zopote de viejas
• Micronesia, Federated states of: saasaf; saasap
• Myanmar: duyin-awza
• Nauru: dawatsip
• Netherlands: zuurzak; zuurzakboom
• New Zealand: sasalapa
• Niue: talapo fotofoto
• Northern Mariana Islands: laguana; laguanaha; syasyap
• Palau: sausab
• Papua New Guinea: saua sap
• Philippines: atti; babana; bayubana; guayabano; guiabano; guyabana; gwabana; labanus; llabanos
• Sierra Leone: soursapi
• Thailand: rian-nam; thu-rian-khack; thurian-khaek; thurian-thet
• Vietnam: mang cân xiem; mang cau xiem

EPPO code

• ANUMU (Annona muricata)

Summary of Invasiveness

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A. muricata is a small American tree, perhaps native to Mexico, Central America, and South America, and also occuring in the West Indies and Caribbean. It is considered an invasive species in CenBIO (Acevedo-Rodriguez and Strong, 2012). The species is tolerant of poor soils, is propagated by both seed and cuttings, and has a history of repeated, intentional introductions in places beyond its native range (PIER, 2014). The species received a low risk score of -3 in an assessment prepared for PIER (2014), indicating it is not currently a major threat, but it is included in the Global Compendium of Weeds as an “agricultural weed, cultivation escape, environmental weed, naturalised, weed” (Randall, 2012), and is cited as invasive in some Pacific islands by references listed by PIER (2014). In the Asia Pacific region it is known to be occasionally adventive or sparingly naturalized in parts of French Polynesia (Wagner and Lorence, 2014), and has been reported to be “moderately invasive on Nauru” and cited by PIER (2014) as invasive in parts of Tonga, Hawaii and the Galapagos Islands.

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Annonales
•                         Family: Annonaceae
•                             Genus: Annona
•                                 Species: Annona muricata

Notes on Taxonomy and Nomenclature

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The genus Annona, commonly known as the custard-apple genus, consists of about 125 species with some species widely cultivated for their edible fruits and often becoming naturalized beyond their native range of tropical America and Africa (Wagner et al., 2014a). The genus name Annona could be derived from the Latin ‘anon’, meaning ‘yearly produce’, referring to “the fruit production habits of the various species in this genus” (Orwa et al., 2009), or, according to Britton and Wilson (1924), from ‘Hanon’, “an aboriginal name for the tree in tropical America, probably Santo Domingo”. The species name ‘muricata’ refers to the soft spinules on the fruit’s exterior that have also led to some vernacular names, like ‘prickly custard apple’ (Quattrocchi, 2012). The common name soursop refers to the slightly acidic taste of the ripe fruit.

Vellozo described a species with the name A. muricata in 1827, but that species was soon after renamed A. montana by Macfadyen in his Flora of Jamaica in 1837. 

Description

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Wagner et al. (2014a, b) give the following description: Small tree 7.5-9 m tall, new growth puberulent with reddish brown hairs. Leaves distichous, petiolate, blade narrowly obovate, narrowly elliptic or obovate-elliptic, 6.5-20 cm long, 2.5-6.5 cm wide, base acute to rounded, apex acuminate, shiny and glabrous above, strigillose along costa and secondary veins beneath, with barbate domatia in secondary vein axils, secondary veins 8-12 pairs, petioles thick, 5-8 mm long. Inflorescences cauliflorous or ramiferous, sometimes in fascicles on knobby outgrowths of trunk, or solitary and leaf-opposed, pedicels stout, 15-20 mm long, 2-2.5 mm in diameter, obconical, rusty puberulent, bracteolate; sepals 3, valvate, broadly triangular, 3-4 mm long, 5-6 mm wide, thick, apex acute, puberulent externally, petals thick, fleshy, glabrous, greenish-yellow to yellow, 3 outer petals 30 mm long, 22 mm wide, ovate, base cordate, apex acuminate, 3 inner petals 20-22 mm long, 10-15 mm wide, elliptic, cucullate, base acute to attenuate, apex obtuse; stamens numerous, 4 mm long, 1 mm in diam., clavate with thickened apex; ovary ca. 5 mm long, 8 mm wide, broadly conical, carpels densely velutinous. Fruit a large, fleshy syncarp to 30 cm long, 15 cm long, ovoid to oblong, often somewhat curved, when fresh green without, the surface bearing regularly well-spaced, soft, conical, curved spines 2-3 mm long, the flesh white, juicy and with cotton-like fibers, surrounding the numerous seeds. Seeds light to dark brown, 13-17 mm long, 9-10 mm wide, ellipsoid, compressed, with low marginal ridge; endosperm ruminate. 

This small, evergreen tree may be slender and upright or low branching and bushy; it often becomes straggly and untidy with age. The dark green leaves emit a strong odour when crushed. The flowers, which have a peculiar smell, are hermaphrodite and are often produced singly or in small clusters on old wood. Normal fruits are generally heart-shaped to oval, but if there is poor pollination, unfertilized ovules fail to develop and the resulting fruit assumes distorted irregular shapes and is usually undersized. The dark green skin has many recurved, soft spines 0.5-1.3 cm apart. There is often a constriction like a fault on the side of the fruit, where the skin has not swollen and the spines are much closer together. The fruit stalk is about 3-8 cm long and woody. The ripe pulp, which adheres to the skin but is easily separated into segments (which were the separate ovaries), has an agreeable subacid flavour with a distinct aroma (Janick and Paull, 2008).

Plant Type

Top of page Perennial
Seed propagated
Shrub
Tree
Vegetatively propagated
Woody

Distribution

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A. muricata is of tropical American origin although its exact origin is unknown (PIER, 2014; Wagner et al., 2014a); it is probably native in Central America and in northern South America (Hanelt et al., 2001). It is known to be cultivated in Africa (mainly the warm lowlands of eastern and western Africa), temperate and tropical Asia, Australasia, North America, the south-central Pacific Islands, the Caribbean, and Mesoamerica (USDA-ARS, 2014).

Some differences were found between sources as to where the species is native. It is reported by some authorities as exotic to the Caribbean and West Indies including Puerto Rico (Acevedo-Rodriguez and Strong, 2012; Randall, 2012), but it was listed as native to Puerto Rico by USDA-NRCS (2014) and as native to Caribbean territories by ITIS (2014). Hanelt et al. (2001) reports the species to be “found wild and cultivated from sea level to 1000 m in the Antilles and from south Mexico to Peru and north Argentina”, although the species is reported as exotic in the Lesser Antilles (Acevedo-Rodriguez and Strong, 2012).

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.

History of Introduction and Spread

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A. muricata is of tropical American origin (PIER, 2014; Wagner et al., 2014a), and it is probably native to Central America and northern South America (Hanelt et al., 2001; Acevedo-Rodriguez and Strong, 2012). Archaeological evidence indicates both A. muricata and A. cherimola were present in pre-Hispanic Peru rather than an introduction by Spaniards, as was thought by 17^th-century records (Bonavia et al., 2004). It was one of the first fruit trees carried from America to the Old World Tropics (Morton, 1987). Both A. cherimola and A. muricata were introduced to Asia by way of the Spanish galleon trade at an early date, and A. muricata is now widely cultivated across the Asia Pacific region for its edible fruit (Koesriharti, 1991; PIER, 2014).

Date of introduction of A. muricata to the West Indies and Caribbean region is uncertain, but it was observed growing in Jamaica in the early 16^th century by Sir Hans Sloane, who collected a specimen that is now in the Sloane herbarium of the UK Natural History Museum in London (specimen BM000594141). Morton (1987) states that Oviedo described the fruit as abundant in the West Indies and in northern South America in 1526. The species has been present in Puerto Rico since before the 1880’s, as it was included in both Bello’s (1883) and Britton and Wilson’s (1924) works on Puerto Rico and the Virgin Islands. It has been grown in Florida since probably the 1870s (Morton, 1987).

Risk of Introduction

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Although A. muricata is known to an agricultural weed and cultivation escape in some places (Randall, 2012), it received a low risk score of -3 in a PIER risk assessment (2014). Risk of introduction for A. muricata is therefore currently low, but it has demonstrated several invasive traits that may cause problems in introduced places where the species has become abundant. 

Habitat

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A. muricata, although widely cultivated, occurs in thickets, hillsides, mountain woodlands, and shaded ravines in Puerto Rico (Britton and Wilson, 1924; Liogier and Martorell, 2000) and in humid premontane forests in Colombia (Vascular Plants of Antioquia, 2014). In Bolivia the species occurs in lowland rainforests (Bolivia Checklist, 2014). It is also cultivated in coastal, island, and Amazonian parts of Ecuador, where the species is native (Vascular Plants of Ecuador, 2014). The species has also been reported to occur in disturbed lowland areas (Peru Checklist, 2014). 

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
	Terrestrial – Managed	Cultivated / agricultural land	Present, no further details	Productive/non-natural
		Disturbed areas	Present, no further details	Productive/non-natural
	Terrestrial ‑ Natural / Semi-natural	Natural forests	Present, no further details	Natural
		Natural forests	Present, no further details	Productive/non-natural
Littoral
		Coastal areas	Present, no further details	Natural
		Coastal areas	Present, no further details	Productive/non-natural

Biology and Ecology

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Genetics

1n = 8; 2n = 14 (IPCN Chromosome Reports, 2014).

Reproductive Biology

Trees can be propagated clonally, in particular through various budding and grafting techniques on seedling stocks, as is the practice in parts of America (e.g. Columbia, Venezuela). However, the species is commonly raised from seed (Morton, 1987).

Growth and Development

A. muricata branches freely through the emergence of sylleptic shoots. Extension growth can occur at any time of the year and proceeds fairly steadily; there are no prominent flushes. The emergence of flower buds follows extension growth. The position of the flowers - mainly terminal on short shoots and anywhere along the axis of long shoots - suggests that they are initiated terminally, the meristem being pushed to a lateral position as extension growth of the shoot is resumed. A dry season imposes synchronous shoot growth and flowering, leading to a harvest peak three months later, but the synchronization is gradually lost in the course of the rainy season. Annona species generally require 27-35 days for flower bud development from initiation to anthesis. The soursop produces fruit throughout the year, but peak production in most areas comes during summer and early autumn, sometimes with a secondary peak during early spring. No photoperiod responses have been reported (Janick and Paull, 2008). Natural pollination in soursop is complex and in most cases results in very low fruit set and yields, with wind- and self-pollination being low (1.5%). The flowers are protandrous, the pollen is shed as the outer petals open towards the evening. The inner petals open much later and only very slightly, admitting small insects attracted by the fragrance of the flowers. Presumably these insects effect cross-pollination, though rather inadequately, for few flowers set fruit and many fruits are misshapen as numerous ovules are not fertilized. These nitidulid beetles (Carpophilus and Uroporus spp.) are considered important pollinators, although no significant effect has been observed from their presence in some cases. These beetles breed very fast in the remains of fruit, so it is recommended to retain rotting fruit as an attractant. Some reports have indicated that the presence of three nitidulid beetles per flower can increase fruit set by 25% (Southampton Centre for Underutilized Crops, 2006). Inadequate pollination appears to be the main factor limiting yield and hand pollination is often recommended for commercial production. However, it is feasible only where there is a definite flowering period. Often, very efficient hand pollination can result in significant economic returns from higher fruit set and larger and more symmetrical fruit. Success in hand pollination is sometimes variable, being less successful on very humid overcast days and with young, vigorous trees. About 150 flowers can be pollinated by a skilled labourer in 1 h with a success rate of 80-100% (Paull and Duarte, 2012).

Fruit growth shows the typical sigmoidal curve with maturation occurring in 16-24 weeks. Low humidity <60% RH) and temperature <13°C) near fruit maturity can increase the severity of fruit skin russeting as well as delaying maturation.

Environmental Requirements

A. muricata requires a warm and humid tropical climate, prefers well-drained and loose, fairly rich, deep loamy soil with a pH range of 5-6.5, is intolerant of waterlogged soil, and can be stunted or killed by cold spells or light frost as the tree is shallow-rooted (Koesriharti, 1991; Orwa et al., 2009). Defoliation and an interruption of fruiting occur when the temperature drops to near freezing. However, soursops are capable of growing in a wide range of soil types from sandy to clay loams provided that the soil has good drainage. Although the tree is commonly grown on slightly acid soils with optimum pH at 5-6.5, it also grows on the porous, oolitic limestone of south Florida and the Bahamas. Higher and more consistent yields are obtained on trees grown on well-drained sandy to sandy loam soils. Waterlogging is a major cause of floral abscission and root rot such as bacterial wilt caused by Pseudomonas spp. Soursop cannot tolerate standing water for any length of time but will tolerate dry soil conditions. A dry season enhances leaf fall and synchronizes extension growth and flowering to some extent. Poor pollination is a frequent problem and occurs at high temperature (30°C) and low humidity (30% relative humidity (RH)), even with hand pollination. Lower temperature (25°C) and high humidity (80% RH) greatly improves pollination (Janick and Paull, 2008).

The soft wood of the trees makes them susceptible to wind damage and limb breakage. Wind may also be partially responsible for the penetration of collar rot organisms. Fruit productivity is improved by the provision of windbreaks.

Its altitudinal range is reportedly 0-2000 m; in Bolivia and Peru the species has been reported growing at 0-500 m (Bolivia Checklist, 2014; Peru Checklist, 2014), while in Colombia and Nicaragua, the species is cultivated at elevations of 0-1000 m (Flora of Nicaragua, 2014; Vascular Plants of Antioquia, 2014), and in Panama it reportedly grows at elevations up to 2000 m (Panama Checklist, 2014).

Climate

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Latitude/Altitude Ranges

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Air Temperature

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Rainfall

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Rainfall Regime

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Uniform

Soil Tolerances

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Soil drainage

• free

Soil reaction

• acid
• neutral

Soil texture

• light
• medium

Natural enemies

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Means of Movement and Dispersal

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Natural Dispersal

The species’ seeds are contained within large fruits sometimes weighing up to 4.5 kg (Flora of Pakistan, 2014) or even 5 kg (Brown, 1950), making its dispersal by wind or water unlikely. Its trademark spinules are also unlikely to function as burrs to attach to animals, considering the weight of the fruit. 

Vector Transmission (Biotic)

Fruits are edible and commonly eaten by humans, animals including “horses, cows, hogs, and indeed every description of stock are fond of it” (Macfadyen, 1837), as well as bats (Koesriharti, 1991), all of which may serve to disperse the species’ seeds.

Accidental Introduction

The species is known to have escaped from cultivation and be accidentally introduced to non-native environments (Randall, 2012).

Intentional Introduction

A. muricata was one of the first plants to be introduced from the New World to the Old Tropics as a crop plant and is now widely cultivated across humid tropical regions of the world for its edible fruit.

Pathway Causes

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Pathway Vectors

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Impact Summary

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Economic Impact

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Although known to be an agricultural weed in some places (Randall, 2012), the species would also have a positive impact on local livelihoods by providing large, edible fruits for foraging and feeding to livestock. 

Environmental Impact

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A. muricata has been reported as a cultivation escape and agricultural weed in some places (Randall, 2012) and occasionally adventive or sparingly naturalized in parts of French Polynesia (Wagner and Lorence, 2014), but is not reported to be problematic in most places. It currently has a low risk of negatively impacting non-native environments (PIER, 2014).

Social Impact

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There is some evidence that A. muricata seeds may negatively affect human health; a 2004 report on Guadeloupe’s atypical Parkinson’s Disease epidemiology found an average of 15 mg of annonacin in each A. muricata fruit, and estimated that an adult who consumed one fruit or can of nectar a day would ingest in one year the amount of annonacin found to induce brain lesions in rats that received purified annonacin via intravenous infusion (Lannuzel et al., 2006; Badrie and Schauss, 2010). Further research on this, especially considering its widespread cultivation and consumption, is warranted.

Risk and Impact Factors

Top of page Invasiveness

• Abundant in its native range
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Tolerant of shade
• Has propagules that can remain viable for more than one year
• Reproduces asexually

Impact outcomes

• Negatively impacts agriculture
• Negatively impacts human health

Impact mechanisms

• Produces spines, thorns or burrs

Likelihood of entry/control

• Highly likely to be transported internationally deliberately

Uses

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A. muricata is most commonly known for its edible fruits. Macfadyen’s flora of Jamaica (1837) describes how the species’ fruit could be “eaten plain, or mixed with sugar and water, to which wine and nutmeg are sometimes added. Horses, cows, hogs, and indeed every description of stock are fond of it”. The fruit pulp is used for making sherbets, ice cream, jellies, and other desserts, as well as a drink as in Java, Cuba, and parts of America (Brown, 1950; Koesriharti, 1991; Flora of Pakistan, 2014). In Indonesia the fruit pulp is boiled and mixed with sugar to make sweetcake ('dodol sirsak'), and in the Philippines young fruits are eaten as a vegetable (Koesriharti, 1991). The species is difficult to produce for large-scale, global commercial purposes, as “yields are generally low: one or two dozen fruit per tree per year, each fruit weighing more than 1 kg on average” and “Mature, firm fruit ripen 3—5 days after harvest and can be held only 2—3 days thereafter, even if cooled. Thus, harvested fruit should be dispatched to distant markets without delay. Because of their tender skin, fruit should be handled with great care”; “because of erratic yield and short shelf life, production of soursop is too scattered to supply a substantial processing industry. Presumably this is why soursop growing in South-East Asia has not expanded convincingly in recent times” and “international trade is virtually limited to processed products” (Koesriharti, 1991).

In addition to its use as a food crop, the seeds and many of the plant parts of A. muricata are used in traditional medicine (Hanelt et al., 2001; Orwa et al., 2009; USDA-ARS, 2014). Badrie and Schauss (2010) report some of its uses in traditional Indian medicine as well as in Jamaica, Haiti, Brazil, the Peruvian Amazon for the treatment of kidney problems, fever, nervousness, ulcers and wounds, with antispasmodic, antidysenteric, and parasiticidal activity, its leaves for fever, its bark as tonic, roots as antispasmodic and parasiticidal, its flowers as bechic [relieving coughs], unripe fruit as antiscrobutic; and seeds as insecticidal, astringent, and as a fish-poison. The plant is used in Ayurvedic medicine as a bitter, tonic, abortifacient, febrifuge, for scorpion stings, high blood pressure, and as a respiratory stimulant. It has become popular as an alternative remedy for cancer, and while there is so far no evidence to support this from clinical trials, some recent laboratory research suggests that extracts can act against cancer cells (Pieme et al., 2014; Yang et al., 2015). Unfortunately, the fruit and seeds also contain annonacin, which has been demonstrated to be toxic in vitro and in vivo to dopaminergic and other neurons, and consumption of the fruit has been associated with an increased risk of developing atypical parkinsonism in humans (Lannuzel et al., 2006; Badrie and Schauss, 2010).

A. muricata has also been used in agriculture as an intercrop species. Koesriharti, (1991) reports that “being a small and early-bearing tree, the soursop may be planted as an intercrop between larger fruit trees such as mango, avocado, and santol. When the main crop requires the space, the soursop trees are grubbed out”.

The seeds of the fruit possess insecticidal properties and have been used for this purpose (Hanelt et al., 2001; Flora of Pakistan, 2014)

Uses List

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Environmental

• Agroforestry

Human food and beverage

• Beverage base
• Fruits
• Vegetable

Materials

• Oils

Medicinal, pharmaceutical

• Traditional/folklore

Prevention and Control

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This species is not considered a high-risk species and therefore little data was found for methods of prevention and control.

Bibliography

Top of page Sunarjono H, 1987. Fruit growing [Indonesian]. Sinar Baru. Bandung, Indonesia,191-195.

Lam PF, Zaipun MZ, 1986. Respiration rates, ethylene productions and chemical compositions of different maturity of soursop (Annona muricata L.) at various temperatures. MARDI Research Bulletin (Malaysia), 14(3): 231-235.

Nakasone HY, 1972. Production feasibility for soursop. Hawaii Farm Science, 21(1): 10-11.

Rismunandar, 1983. Development of fruit crops [Indonesian]. Sinar Baru. Bandung, Indonesia, 147-151.

van der Pijl L, 1953. On the flower biology of some plants from Java with general remarks on fly-traps. Annales Bogoriensis, 1: 77-99.

Wijaya, 1988. Plasma nutfah sirsak [Soursop germplasm]. Warta Penelitian dan Pengembangan Pertanian (Departemen Pertanian Republik Indonesia), 10(6): 1-2.

References

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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

Badrie N, Schauss AG, 2010. Soursop (Annona muricata L.): Composition, Nutritional Value, Medicinal Uses, and Toxicology. In: Bioactive foods in promoting health [ed. by Watson, R. R. \Preedy, V. R.]. Amsterdam, Netherlands: Academic Press, 621-643.

Barbeau G, 1994. Inventory of tropical fruit trees in Central America and the West Indies. Fruits (Paris), 49(5/6):383-389, 469-474.

Bello D, 1883. [English title not available]. (Apuntes para la flora de Puerto Rico. Segunda parte. Monoclamídeas.) Anales de la Sociedad Española de Historia Natural, 12:103-130.

Bolivia Checklist, 2014. Catalogue of the Vascular Plants of Bolivia, Tropicos website. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://tropicos.org/NameSearch.aspx?projectid=13

Bonavia D, Ochoa CM, O´scar TS, Palomino RC, 2004. Archaeological evidence of cherimoya (Annona cherimola Mill.) and Guanabana (Annona muricata L.) in ancient Peru. Economic Botany, 58(4):509-522.

Britton NL, Wilson P, 1924. Scientific Survey of Porto Rico and the Virgin islands, Volume V, Botany of Porto Rico and the Virgin Islands. New York Academy of Sciences, New York.

Brown WH, 1950. Useful Plants of the Philippines, Volume 1:590 pp. [Commonwealth of the Philippines Department of Agriculture and Commerce, Technical Bulletin no 10.]

Carbajal D, Casaco A, Arruzazabala L, Gonzalez R, Fuentes V, 1991. Pharmacological screening of plant decoctions commonly used in Cuban folk medicine. Journal of Ethnopharmacology, 33(1-2):21-24; 7 ref.

Chan YK, Subhadrabandhu S, 1992. Breeding and varietal improvement of tropical fruits at MARDI. Frontier in tropical fruit research. Proceedings of international symposium held on 20-24 May 1991, Pattaya City, Thailand. Acta-Horticulturae, No. 321, 138-151; 12 ref.

Charles Darwin Foundation, 2008. Database inventory of introduced plant species in the rural and urban zones of Galapagos. Database inventory of introduced plant species in the rural and urban zones of Galapagos. Galapagos, Ecuador: Charles Darwin Foundation, unpaginated.

Chong KY, Tan HTW, Corlett RT, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. Singapore: Raffles Museum of Biodiversity Research, National University of Singapore, 273 pp. http://lkcnhm.nus.edu.sg/nus/pdf/PUBLICATION/LKCNH%20Museum%20Books/LKCNHM%20Books/flora_of_singapore_tc.pdf

Clark JL, Neill DA, Asanza M, 2006. Floristic checklist of the Mache-Chindul Mountains of northwest Ecuador. Contributions from the United States National Herbarium, 54:180 pp.

Elzebroek, T., Wind, K., 2008. Guide to cultivated plants., Guide to cultivated plants:vii-xi + 516 pp.

Flora Mesoamericana, 2014. Flora Mesoamericana. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/Project/FM

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

Flora of Nicaragua, 2014. Flora of Nicaragua, Tropicos website. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://tropicos.org/NameSearch.aspx?projectid=7

Flora of Pakistan, 2014. Flora of Pakistan/Pakistan Plant Database (PPD). Tropicos website St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.tropicos.org/Project/Pakistan

Funk V, Hollowell T, Berry P, Kelloff C, Alexander SN, 2007. Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions from the United States National Herbarium, 584 pp.

Gbeassor M, Kedjagni AY, Koumaglo K, Souza C de, Agbo K, Aklikokou K, Amegbo KA, 1990. In vitro antimalarial activity of six medicinal plants. Phytotherapy Research, 4(3):115-117; 10 ref.

Granadino CA, Cave RD, 1994. Inventory of arthropods and pathogenic fungi of Annona in four localities of Honduras. Turrialba, 44(3):129-139; 5 ref.

Granadino CA, Cave RD, 1997. Within-tree distribution of seven insect pests of soursop (Annona muricata) in Honduras. CEIBA, 38(2):161-166.

Hanelt P, Buttner R, Mansfeld R, 2001. Mansfeld's Encyclopedia of Agricultural and Horticultural Crops (except Ornamentals). Berlin, Germany: Springer.

IPCN Chromosome Reports, 2014. Index to Plant Chromosome Numbers (IPCN), Tropicos website. St. Louis, Missouri, USA: Missouri Botanical Garden. http://tropicos.org/Project/IPCN

ITIS, 2014. Integrated Taxonomic Information System. http://www.itis.gov

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Contributors

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28/01/2015 Updated by:

Marianne Jennifer Datiles, Department of Botany-Smithsonian NMNH, Washington DC, USA

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

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