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Datasheet

Annona squamosa (sugar apple)

Summary

  • Last modified
  • 22 November 2017
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Annona squamosa
  • Preferred Common Name
  • sugar apple
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • A. squamosa is a shrub or small tree of American origin. The species is widely grown as a commercial fruit tree both within its native range and in tropical regions around the world, and although less common th...

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Pictures

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PictureTitleCaptionCopyright
One-year-old seedling planted in home garden.
TitleSeedling
CaptionOne-year-old seedling planted in home garden.
Copyright©Rafael T. Cadiz
One-year-old seedling planted in home garden.
SeedlingOne-year-old seedling planted in home garden.©Rafael T. Cadiz
Annona squamosa (sugar apple); 1. flowering branch. 2. branch with fruit (pseudocarp). 3. halved fruit
TitleLine artwork
CaptionAnnona squamosa (sugar apple); 1. flowering branch. 2. branch with fruit (pseudocarp). 3. halved fruit
Copyright©PROSEA Foundation
Annona squamosa (sugar apple); 1. flowering branch. 2. branch with fruit (pseudocarp). 3. halved fruit
Line artworkAnnona squamosa (sugar apple); 1. flowering branch. 2. branch with fruit (pseudocarp). 3. halved fruit©PROSEA Foundation

Identity

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Preferred Scientific Name

  • Annona squamosa L.

Preferred Common Name

  • sugar apple

Other Scientific Names

  • Annona asiatica L.
  • Annona cinerea Dunal
  • Guanabanus squamosus M. Gomez
  • Xylopia frutescens Sieb. ex Presl
  • Xylopia glabra L.

International Common Names

  • English: atemoya; sugarapple; sugar-apple; sweetsop; sweetsop tree
  • Spanish: anon; anona (blanca); anona blanca; chirimoyo
  • French: annone ecailleuse; attiee; attier pomme cannelle; Corossolier a fruits ecailleux; fruta del conde; fruta-do-conde; grand corossolier; pomme cannelle; pommier canelle; pommier cannelle
  • Chinese: fan li zhi
  • Portuguese: ata

Local Common Names

  • Australia: custard apple; sweet apple; sweet sop
  • Bolivia: chirimoya
  • Brazil: ata; fruta do conde; pinha do conde
  • Cambodia: tiep baay; tiep srôk
  • Cook Islands: katara apa Maori; katara‘apa Maori; naponapo; tapotapo; tapotapo Maori
  • Cuba: anón sin semilla; hanón
  • Dominican Republic: cachimán; cachiman canelle; cachiment canelle; candongo
  • Fiji: apeli
  • French Polynesia: noni haoe; nonihaoé; tapo tapo; tapotapo
  • French Polynesia/Marquesas: manini puteketeke
  • Germany: Rahm- Annone; Rahmapfel; Schuppenannone; Sussack; Zimtapfel; Zuckerapfel
  • Guam: atis
  • Haiti: candón; pomme de canelle; pommier de canelle
  • India: aatoa; amritaphala; ata; athichakku; luna; seethapalam; seethaphala; sharifa; shariffa; sitaphal
  • Indonesia/Java: sirkaja
  • Indonesia/Moluccas: atis
  • Indonesia/Sumatra: sarikaja
  • Italy: pomo canella
  • Japan: banreishi
  • Laos: khièb
  • Malaysia: buah nona; nona sri kaya; sri kaya
  • Mexico: saramuya
  • Middle East: achta
  • Nauru: dawatsip
  • Netherlands: Kaneel-appel
  • Niue: talapo
  • Pakistan: sharifa; sitaphal
  • Palau: ngel ra ngebard
  • Philippines: ates; atis; atit; atti; yates
  • Taiwan: sek-khia; sek-kia
  • Thailand: lanang; makkhiap; noina
  • Tonga: apele papalangi
  • Tuvalu: nameana
  • Vietnam: mang câù ta; na

EPPO code

  • ANUSQ (Annona squamosa)

Summary of Invasiveness

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A. squamosa is a shrub or small tree of American origin. The species is widely grown as a commercial fruit tree both within its native range and in tropical regions around the world, and although less common than other Annona spp., production areas are increasing. The species is listed as “cultivation escape, naturalised, weed” in the Global Compendium of Weeds (Randall, 2012); it is known to escape from cultivation, often naturalising, and even becoming invasive in places such as French Polynesia and Nauru in the Pacific, and Mayotte in the Indian Ocean (PIER, 2015). The species is known to be a weed in Jamaica and Cambodia (Holm et al., 1979). It is likely to become invasive in other countries, and may be particularly problematic on tropical 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 squamosa

Notes on Taxonomy and Nomenclature

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The genus Annona consists of some 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., 2015). 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 Annona squamosa is commonly known as ‘sugar apple’ or ‘sweetsop’ in English, but it is also sometimes known as ‘custard-apple’, especially in South Asia, though this common name generally refers to Annona reticulata and so can only cause confusion.

Description

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A. squamosa, a small deciduous tree about 3-5(-6) m tall, produces its first branches near the base of the trunk. The branches are irregularly spreading and the young growth is densely pubescent (Leon, 1917; Carangal et al., 1961). The leaves are alternate, ovate-oblong or elliptic-oblong, thin, sparsely downy, dark green above, 8-15 cm long and 2-5 cm wide. When young, leaves are pubescent and give a peculiar smell when crushed (Troup et al., 1975). The petiole is about 1.0-1.5 cm long (Coronel, 1983). The small, pendulous flowers occur singly or in pairs in the leaf axils of young shoots or opposite the leaves. The pedicel is 1.5-2.5 cm long and hairy. The three sepals are short, deciduous, densely or thinly pubescent, and 0.2-0.3 cm long. The 6 petals are biseriate. The three outer petals are lanceolate, thick, fleshy, trigonous, finely pubescent, yellowish-green on the outside, yellowish-white inside, 2.0-2.5 cm long and 0.5-1.0 cm wide. The three inner petals alternate with the outer ones and are minute, sometimes absent, ovate and never more than 0.1 cm long. The stamens are numerous, yellowish-white in many rows on the glabrous, raised receptacle (torus), 0.12-0.15 cm long and crowded in a whorl around the gynoecium. The pistils are also numerous, dark violet, finely pubescent, and are found above the stamens. The stigmas are sessile, stuck together, and deciduous. The stamens and pistils form a cone-shaped structure at the centre of the flower (Coronel, 1983). The fruit is a syncarp developed from the fusion of numerous ovaries. It is irregularly heart-shaped, about 5-20 cm in diameter. The exterior is marked by polygonal tubercles which correspond to the fused carpels from which the fruit is formed. The ripe fruit is light yellowish-green or purple and the exterior readily separates along the lines between the tubercles. The flesh is white, soft and juicy with a mild, agreeable flavour. The numerous seeds are obovoid or elliptic, dark brown or black, shiny, slightly compressed, 1.0-1.5 cm long and 0.5-0.8 cm wide, and each is enclosed in the edible pulp.

Plant Type

Top of page Broadleaved
Perennial
Seed propagated
Shrub
Tree
Woody

Distribution

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A. squamosa is indigenous to tropical South America and the West Indies according to Wester (1912); however, Pinto et al. (2005) states that the sugar apple originated in lowland Central America where it is indigenous, and from there it was distributed to Mexico and throughout tropical America. In this datasheet, the more restrictive native range of mainland Central America is adopted, as it is likely that it was spread in pre-history. In the lowlands of Mexico it is found in a naturalized or wild state, and it is grown from Central America southwards to northern South America, extending to northeastern Brazil where it is one of the most popular fruits. It is a common species in the West Indies and has been present since at least 1689 (UK Natural History Museum, 2015).

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

CambodiaPresentIntroducedWorld Agroforestry Centre, 2010
ChinaPresentIntroducedWorld Agroforestry Centre, 2010; PIER, 2015
-FujianPresentIntroducedFlora of China Editorial Committee, 2015
-GuangdongPresentIntroducedFlora of China Editorial Committee, 2015
-GuangxiPresentIntroducedFlora of China Editorial Committee, 2015
-HainanPresentIntroducedFlora of China Editorial Committee, 2015
-Hong KongPresentIntroducedPIER, 2015
-YunnanPresentIntroducedFlora of China Editorial Committee, 2015
-ZhejiangPresentIntroducedFlora of North America Editorial Committee, 2015
IndiaPresentIntroducedWorld Agroforestry Centre, 2010
IndonesiaPresentIntroducedWorld Agroforestry Centre, 2010
IsraelPresentMorton, 1987Lowlands of Palestine
LaosPresentIntroducedWorld Agroforestry Centre, 2010
MalaysiaPresentCABI, 2005
MaldivesPresentIntroduced Not invasive PIER, 2009
PakistanPresentIntroducedFlora of Pakistan, 2015
PhilippinesPresentIntroducedWorld Agroforestry Centre, 2010; Pelser et al., 2014; PIER, 2015
TaiwanPresentIntroducedFlora of Taiwan Editorial Committee, 2014; Flora of China Editorial Committee, 2015
ThailandPresentIntroducedWorld Agroforestry Centre, 2010
VietnamPresentIntroducedWorld Agroforestry Centre, 2010

Africa

BurundiPresentIntroducedMissouri Botanical Garden, 2009
EgyptPresentMorton, 1987
GabonPresentIntroducedMissouri Botanical Garden, 2009
MadagascarPresentIntroducedMissouri Botanical Garden, 2009
MayottePresentIntroduced Invasive Barthelat, 2005
RéunionPresentIntroduced Not invasive PIER, 2009
SeychellesPresentIntroduced Not invasive PIER, 2009
SudanPresentIntroducedWorld Agroforestry Centre, 2010
TanzaniaPresentIntroducedMissouri Botanical Garden, 2009
-ZanzibarPresentIntroducedWorld Agroforestry Centre, 2010
UgandaPresentIntroducedWorld Agroforestry Centre, 2010

North America

BermudaPresentMorton, 1987
MexicoPresentIntroducedHanelt et al., 2001; Missouri Botanical Garden, 2009; Flora Mesoamericana, 2014
USAPresentIntroducedMissouri Botanical Garden, 2009
-CaliforniaPresentIntroducedMissouri Botanical Garden, 2009
-FloridaPresentIntroducedMissouri Botanical Garden, 2009; USDA-NRCS, 2015
-HawaiiPresentMorton, 1987; PIER, 2015

Central America and Caribbean

Antigua and BarbudaPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
ArubaPresentIntroducedAcevedo-Rodriguez and Strong, 2012
BahamasPresentIntroducedBritton and Millspaugh, 1920; Missouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
BarbadosPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
BelizePresentIntroducedMissouri Botanical Garden, 2009; Randall, 2012Naturalized
British Virgin IslandsPresentIntroducedAcevedo-Rodriguez and Strong, 2012
Cayman IslandsPresentIntroducedAcevedo-Rodriguez and Strong, 2012
Costa RicaPresentIntroducedMissouri Botanical Garden, 2009; Randall, 2012
CubaPresentIntroducedMissouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
CuraçaoPresentIntroducedAcevedo-Rodriguez and Strong, 2012
DominicaPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
Dominican RepublicPresentIntroducedMissouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
El SalvadorPresentNativeMissouri Botanical Garden, 2009; Flora Mesoamericana, 2014
GrenadaPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
GuadeloupePresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
GuatemalaPresentNativeMissouri Botanical Garden, 2009; Flora Mesoamericana, 2014
HaitiPresentIntroducedMissouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
HondurasPresentNativeMissouri Botanical Garden, 2009
JamaicaPresentIntroducedMacFadyen, 1837; Missouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
MartiniquePresentIntroducedMissouri Botanical Garden, 2009
MontserratPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
Netherlands AntillesPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
NicaraguaPresentIntroducedMissouri Botanical Garden, 2009; Flora of Nicaragua, 2015Not common, cultivated in lowland parts of Pacific region
PanamaPresentNativeMissouri Botanical Garden, 2009; Flora Mesoamericana, 2014
Puerto RicoPresentIntroducedLiogier and Martorell, 2000; Missouri Botanical Garden, 2009; USDA-NRCS, 2015'in thickets, on roadsides, and in valleys, in the southern districts, Puerto Rico'
Saint Kitts and NevisPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
Saint LuciaPresentNativeWorld Agroforestry Centre, 2010
Saint Vincent and the GrenadinesPresentIntroducedWorld Agroforestry Centre, 2010; Acevedo-Rodriguez and Strong, 2012
Trinidad and TobagoPresentIntroducedMissouri Botanical Garden, 2009; Acevedo-Rodriguez and Strong, 2012
Turks and Caicos IslandsPresentIntroducedMissouri Botanical Garden, 2009
United States Virgin IslandsPresentIntroducedMissouri Botanical Garden, 2009

South America

ArgentinaPresentNativeWorld Agroforestry Centre, 2010
BoliviaPresentIntroducedMissouri Botanical Garden, 2009; Bolivia Checklist, 2015
BrazilPresentIntroducedMissouri Botanical Garden, 2009
ChilePresentNativeWorld Agroforestry Centre, 2010
ColombiaPresentNativeMissouri Botanical Garden, 2009; Vascular Plants of Antioquia, 2015
EcuadorPresentIntroducedMissouri Botanical Garden, 2009; Vascular Plants of Ecuador, 2015
French GuianaPresentNativeWorld Agroforestry Centre, 2010
GuyanaPresentNativeWorld Agroforestry Centre, 2010
ParaguayPresentIntroducedMissouri Botanical Garden, 2009; Paraguay Checklist, 2015Cordillera
PeruPresentIntroducedMissouri Botanical Garden, 2009
SurinamePresentNativeWorld Agroforestry Centre, 2010
UruguayPresentNativeWorld Agroforestry Centre, 2010
VenezuelaPresentNativeWorld Agroforestry Centre, 2010

Europe

CyprusPresentIntroducedWorld Agroforestry Centre, 2010
GreecePresentIntroducedWorld Agroforestry Centre, 2010
MaltaPresentIntroducedWorld Agroforestry Centre, 2010

Oceania

AustraliaPresentIntroducedWorld Agroforestry Centre, 2010
-Australian Northern TerritoryPresentIntroducedRandall, 2012Naturalized; weed
-New South WalesPresentIntroducedRandall, 2012Naturalized
-QueenslandPresentMorton, 1987
Caroline IslandsPresentWagner et al., 2015
Cook IslandsPresentIntroduced Not invasive PIER, 2009
FijiPresentIntroducedWorld Agroforestry Centre, 2010; PIER, 2015
French PolynesiaPresentIntroduced Invasive PIER, 2015; Wagner and Lorence, 2015
GuamPresentIntroduced Not invasive PIER, 2009
KiribatiPresentIntroduced Not invasive PIER, 2009
Marshall IslandsPresentIntroduced Not invasive PIER, 2009; Wagner et al., 2015
Micronesia, Federated states ofPresentWagner et al., 2015
NauruPresentIntroduced Invasive PIER, 2015
New CaledoniaPresentIntroduced Not invasive PIER, 2009
New ZealandPresentIntroducedWorld Agroforestry Centre, 2010
NiuePresentIntroduced Not invasive PIER, 2009
Northern Mariana IslandsPresentIntroduced Not invasive PIER, 2009; Wagner et al., 2015
PalauPresentIntroduced Not invasive PIER, 2009; Wagner et al., 2015
Papua New GuineaPresentIntroducedWorld Agroforestry Centre, 2010
Pitcairn IslandPresentIntroduced Not invasive PIER, 2009
SamoaPresentIntroducedWorld Agroforestry Centre, 2010
Solomon IslandsPresentIntroduced Not invasive PIER, 2009
TongaPresentIntroduced Not invasive PIER, 2009

History of Introduction and Spread

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Date of introduction of A. squamosa to the West Indies is unknown, but it was present by the time of Sir Hans Sloane’s 1687-1689 voyage to Jamaica (UK Natural History Museum, 2015), during which he collected specimens now present in the British Museum (British Museum specimen BM000594147). The species was also included in Macfadyen's (1837) work on Jamaica, and was present in Puerto Rico by 1883, as it was included in Bello’s (1883) work on the island. It was also listed in Britton’s (1918) flora of Bermuda and Britton and Millspaugh’s (1920) work on the Bahamas.

A. squamosa may have been first introduced into Brazil via Bahia state in 1626 by Conde de Miranda, which explains the species’ vernacular name in Brazil, ‘fruta do conde’. The species was probably brought to the Old World tropics by the Spaniards to the Philippines and by the Portuguese to India by 1590, and from there cultivation eventually spread to Indonesia, China, Australia, Polynesia, and Hawaii (Morton, 1987; Hanelt et al, 2001). In India there is a very large, diverse population, and its commercial importance is so great that some botanists have considered it to be a native fruit of that country. Some of the arguments used by those who favour an Asiatic origin include the occurrence of common names for it in Sanskrit; the existence of large, apparently wild populations in several parts of India; and the presence of carvings and wall-paintings, maybe representing the fruit, in the ruins of ancient temples. However, according to Pinto et al. (2005), this is a secondary centre of diversity, created during the last 500 years.

The purple-fruited form originated in India and was introduced to the Philippines from India in 1911 (Merrill, 1912; Juliano, 1935; Galang, 1955). It has a prominent place in the Philippines, Thailand and India for the production of fruit. In Cuba, the sugar apple ranks with mango as one of the favourite fruits and it is common on other islands of the West Indies. In the USA, it is commercially planted in the subtropical region of Florida (Noonan, 1953) but has never been grown to fruiting size in California (Popenoe, 1952).

Risk of Introduction

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Risk of introduction for this species is low but not insignificant. The species received a low risk score of -2 in a risk assessment prepared for Hawaii (PIER, 2015), but the species is known to have escaped from cultivation in Australia and Costa Rica (Morton, 1987; Randall, 2012), and is known to be invasive to non-native habitats including French Polynesia, Nauru, and Mayotte (PIER, 2015). As a commercial fruit, it is likely to be further introduced.

Habitat

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A. squamosa is a lowland tropical or marginally subtropical species and native to the warmest and driest places in Central America, growing between latitudes 23°N and S, but also yields well in humid regions and is frequently reported in cultivation in semi-arid climates, such as northeastern Brazil (Pinto et al., 2005). In North America the species is reportedly found in “dryish, sandy substrates and dry hammocks” (Flora of North America Editorial Committee, 2015) and in Puerto Rico, “in thickets, on roadsides, and in valleys, in the southern districts” (Liogier and Martorell, 2000); on Saint John, US Virgin Island, this species is naturalized and found along roadsides and secondary forests (Acevedo-Rodríguez, 1996). In the Bahamas it was reportedly growing in scrublands (Britton and Millspaugh, 1920) and similarly, in dry regions of North Queensland, Australia, it is found wild in pastures, forests, and along roadsides (Morton, 1987). The species occurs in moist tropical forests of Colombia (Vascular Plants of Antioquia, 2015), and in coastal regions of Ecuador (Vascular Plants of Ecuador, 2015).

In India, wild A. squamosa predominantly inhabits hillocks, gravelly soils and waste land. It can be common on dry areas at lower elevations. The species may be the most adaptable of any Annona spp., doing well on poor, sandy, or limestone soil as long as it is well drained; it also tolerates more dryness and wind than other Annona spp.

Habitat List

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CategoryHabitatPresenceStatus
Littoral
Coastal areas Present, no further details Harmful (pest or invasive)
Coastal areas Present, no further details Natural
Coastal areas Present, no further details Productive/non-natural
Terrestrial-managed
Cultivated / agricultural land Principal habitat Productive/non-natural
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Managed forests, plantations and orchards Principal habitat Productive/non-natural
Rail / roadsides Present, no further details Natural
Urban / peri-urban areas Secondary/tolerated habitat Productive/non-natural
Terrestrial-natural/semi-natural
Arid regions Secondary/tolerated habitat Natural
Arid regions Secondary/tolerated habitat Productive/non-natural
Natural forests Present, no further details Natural
Natural grasslands Present, no further details Natural
Riverbanks Present, no further details Natural
Rocky areas / lava flows Secondary/tolerated habitat Natural
Scrub / shrublands Principal habitat Natural

Biology and Ecology

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Genetics

There are some recognized cultivars of sugar apple, most of these being in India, their names giving an idea of their origin as introductions: ‘Mammoth’, ‘Barbados’, ‘British Guinea’, ‘Balondegar’, ‘Red Sitaphal’, and ‘Sindhan’, the last being local to Gujarat, and a dwarf cultivar, ‘Lal Sitiphal’ (Pinto et al., 2005). Three forms of A. squamosa exist in the Philippines, a green-fruited seeded form grown in most parts of the country, a purple-fruited seeded form reportedly introduced from India (Galang, 1955) and a seedless green-fruited form whose origin is unknown. It is the green-fruited seeded form from which selection of superior strains is advised (Coronel, 1983); however, to date there has been very little selection of superior seedlings.

Sugar apple (A. squamosa) and cherimoya (A. reticulata) were hybridized by PJ Webster in 1907 in Florida, USA, producing a new fruit called atemoya. Crossing also occurred naturally in the field in Australia in 1850 and again in Palestine in 1930, and the hybrid is in commercial production in Australia (where it is confusingly called custard apple), as well as the USA, Israel, South Africa, the Philippines and numerous parts of Central and South America. This hybrid is preferred because there appears to be no pollination difficulties (Pinto et al., 2005).

Reproductive Biology

A. squamosa can regenerate rapidly by seed or asexual propagation. The seeds stay viable for some, and 90-95% germination occurs 20-30 days after sowing if the seeds are freshly harvested (Miraflores, 1915; Paguirigan, 1951; Galang, 1955). A. squamosa can also be propagated by stem cuttings using healthy branches of matured wood during the dormant period (Noonan, 1953; Ochse et al., 1961). Wester (1912) observed that the fruitfulness of A. squamosa in the Philippines is due to the presence of certain species of coleopterous insects which abound there and pollinate the flowers.

A. squamosa begins to bear fruit 2-4 years after planting, but the yield per tree is usually very low, on average 40-75 fruits per year (Cabbab and Soliven, 1938). Fruits are considered mature when the skin between the segments has turned greenish-yellow to creamy-yellow. Fruits left to ripen on the tree are eaten by birds and bats, and the fruit has a tendency to burst open on the tree if left to overmature. The fruits mature at irregular intervals over a period of about 3 months and are harvested as they mature. Fully mature fruits ripen 2-5 days after harvest and they should be eaten as soon as they ripen because they are perishable (Gonzalez, 1934; Ochse et al., 1961; Viñas, 1972).

Physiology and Phenology

A. squamosa grows in the open or in partly shaded areas. When young, it requires shaded areas and at a later stage full overhead light for vigorous growth. It can withstand drought as it has deciduous growth characteristics during dry periods.

In India, fruit set takes place after the commencement of the rainy season (Thakur and Singh, 1965, 1967). Those fruits that are set during the summer will generally dry up from high temperatures and low relative humidity. Fruit set during summer may be increased by irrigating the trees and providing shade. Fruit growth of green-fruited and red-fruited A. squamosa is rapid from August to mid-September. Afterwards it gradually slows down in the red-fruited form. The growth rate of the green-fruited form, however, still increases into October, after which it slows down and ceases by mid-November (Thakur and Singh, 1965).

In Florida, the fruiting season begins in mid summer (wet season) with irregular ripening lasting 3 months. In the Philippines, fruiting occurs during the beginning of the rainy season (summer) and in India, fruiting also occurs in the wet season (August to mid September) and can occur from October to November (the end of the wet season) (Pinto et al., 2005). In Mexico, flowering occurs at the end of the dry season (March to May) with fruiting at the end of the wet season (September to November). In Brazil, flowering occurs at the end of the dry season (March to May) and fruiting in the wet season (December to January) (Pinto et al., 2005).

Environmental Requirements

A. squamosa thrives well in warm and humid tropical and subtropical lowlands below 1000-1200 m altitude (Paguirigan, 1951; Galang, 1955; Viñas, 1972), and is relatively drought-tolerant but does not fruit well in high rainfall regimes. While it is less tender than some of its relatives, it cannot cope with freezing temperatures or prolonged periods of cold weather (Noonan, 1953). It can, however, withstand drought well and can be grown successfully in areas with a prolonged dry season (Verkataratanam and Satyanaranaswamy, 1956; Cantillang, 1976). Its deciduous growth habit contributes to its drought resistance because it does not have leaves during the majority of months in the dry season. Dry weather during the flowering season seems to be preferable, but fruits are frequently set at the beginning of the rainy season (Noonan, 1953; Ochse et al., 1961).

A. squamosa is not demanding in its soil requirements, perhaps being the most adaptable of the Annona species, as it can tolerate wind and dry climates as well as infertile, sandy, and limestone soil so long as it is well-drained (Staples and Herbst, 2005; Paraguay Checklist, 2015). It cannot tolerate waterlogging for any length of time. A. squamosa grows well on sands, sandy or silty loams, and clays. However, consistently higher yields appear to be obtained from trees grown on sandy loams (Coronel, 1983; Verheij and Coronel, 1991). The root system is relatively shallow, so that it does not require very deep soil. Like most other fruit crops, it prefers slightly acidic soil conditions with a pH of 5.5-6.5 (Coronel, 1983) and can tolerate mild salinity (Pinto et al., 2005).

Elevation range for the species is generally low. In North America it has been reported growing between 0-50 m (Flora of North America Editorial Committee, 2015) and 0-100 m in Panama (Panama Checklist, 2014), while in the lowland rainforests of Bolivia it grows between 0-500 m (Bolivia Checklist, 2015), and along coastal regions of Ecuador between 0-500 m (Vascular Plants of Ecuador, 2015). Similarly, in Nicaragua the species is cultivated in lowland parts of the Pacific region between 30-145 m (Flora of Nicaragua, 2015). In Colombia the species has been reported at slightly higher altitudes of 500-1000 m in moist tropical forest (Vascular Plants of Antioquia, 2015).

 

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
BW - Desert climate Tolerated < 430mm annual precipitation

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
16 18 0 1200

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) >18
Mean annual temperature (ºC) 29 32
Mean maximum temperature of hottest month (ºC) 32 35
Mean minimum temperature of coldest month (ºC) 24 28

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration25number of consecutive months with <40 mm rainfall
Mean annual rainfall7504500mm; lower/upper limits

Rainfall Regime

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Uniform

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • acid

Soil texture

  • light
  • medium

Special soil tolerances

  • infertile
  • saline
  • shallow

Notes on Natural Enemies

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A. squamosa is attacked by a number of pests that feed on its roots, stems, leaves and fruits (Gabriel, 1975). The moth borer Annonaepestis bengalella is the most destructive pest in the Philippines (de Leon, 1917; Estalilla, 1921; Galang, 1955), and larvae eat and tunnel into the interior of the fruits. The surface of attacked fruits shows frasses which are held together by fine silk. The eggs are laid singly in the sutures and stems of fruits (rarely on leaves) and hatch after 4-5 days. Once hatched the larvae immediately bore into the fruits, frequently making new tunnels. It is at the time that they reach their third molt that the most serious damage is done. The fruit may almost fail to develop and in severe cases fall off the tree. The larvae make cocoons near the skin where they pupate. The moths emerge 12 days after pupation.
 
The root grub (Anomala sp.) is another pest of A. squamosa, attacking the roots and in advanced stages causing sudden wilting at the base of the plant (Viñas, 1972; Gabriel, 1975). Mealybugs are also common pests of A. squamosa, and the grey mealybug (Ferrisia virgata) and cottony cushion mealybug (Planococcus lilacinus) suck the sap of young leaves and fruits, causing leaves to turn yellow and wither (Coronel, 1983). Coffee carpenter moth (Zeuzera coffeae) larvae damage the stem by boring into the heart of the wood where they feed and grow (Viñas, 1972; Gabriel, 1975). Inflorescence rot, pink disease and rhizoctonia thread blight may also infect A. squamosa.

Means of Movement and Dispersal

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Vector Transmission (Biotic)
 
Fruits left to ripen on the tree are eaten by birds and bats. In the Philippines, the fruit is commonly eaten by the Philippine fruit bat (kabag or kabog) which then spreads the seeds from island to island.
 
Intentional Introduction

A. squamosa has been widely introduced as a commericial, if underutilized, fruit tree species.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Crop production Yes Yes Pinto et al., 2005
Digestion and excretion Yes Pinto et al., 2005
Escape from confinement or garden escape Yes Pinto et al., 2005
Horticulture Yes Yes Pinto et al., 2005

Impact Summary

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CategoryImpact
Cultural/amenity Positive
Economic/livelihood Positive
Environment (generally) Negative
Human health Positive

Economic Impact

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For a full review of the production and uses of A. squamosa, see Pinto et al. (2005). Although sugar apple production data are scarce, the information collected shows that the potential for expanding the sugar apple market is high in many countries. This species is grown commercially in the West Indies and Dominican Republic, the USA (Florida), the Middle East, India, Malaysia and Thailand. Although it is still considered a backyard fruit used mainly for domestic consumption in the Philippines, this country's production is considered one of the largest in the world. In Brazil, sugar apple production is concentrated in Alagoas and São Paulo states. Other areas of production are in Mexico, Egypt and India, where most fruits come to market from semi-wild forests of the Deccan Plateau where sugar apple has gone wild.

Risk and Impact Factors

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Uses

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The fruit of A. squamosa is usually eaten fresh. It contains 50-61% edible matter, and is a source of carbohydrates, vitamins and proteins. The fruit is utilized commercially as a flavouring for ice cream and can also be made into sherbet. The pulp, after removing the seeds, can be passed through a strainer or homogenized to make a delicious and refreshing drink.

The leaves, bark, roots, seeds and fruit of A. squamosa have various important medicinal uses. The green fruit and seed have effective vermicidal and insecticidal properties and are used as astringents in diarrhoea and dysentery. The seeds contain 45% of a yellow, non-dying oil which is an irritant poison for lice. Crushed leaves are applied as an effective cure for ulcers and malignant sores. A poultice from fresh leaves is used for dyspepsia and when mixed with oil is used for diseases of the scalp. Crushed fresh leaves are applied to the nasal area in cases of fainting spells. A decoction of roots is used as a drastic purgative (Coronel, 1983). The astringent bark, leaves, unripe fruit and seed can be used as a source of the alkaloid anonaine (Troup, 1975).

A. squamosa is usually grown as a backyard fruit tree and as a component of agroforestry systems. Its fruit is a source of food and the flowers are used for apiculture. In some instances, A. squamosa is planted in parks or plazas as a shade and ornamental tree due to its attractive fruit colour (Coronel, 1983).

Uses List

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Animal feed, fodder, forage

  • Invertebrate food

Fuels

  • Fuelwood

Genetic importance

  • Progenitor of

Human food and beverage

  • Beverage base
  • Food additive
  • Fruits

Materials

  • Pesticide

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Similarities to Other Species/Conditions

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Four other Annona spp. are also invasive in the Pacific, Annona cherimola, Annona glabra, Annona muricata and Annona reticulata (PIER, 2009), though these are readily distinguishable by the fruit.

Prevention and Control

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No specific information is available for the control of A. squamosa. However, control methods for other Annona spp. may also be suitable for A. squamosa.

For example, Land Protection (2006) made the following recommendations on chemical control of A. glabra in Queensland, Australia: Triclopyr or glyphosate can both be used for stem injection and cut stump treatments, triclopyr or fluroxypyr as a basal bark treatment, and imazapyr or glyphosate for foliar applications. Stem injectionis recommended for aquatic areas as it minimises herbicide run-off and off-target impacts, and can be carried out with an axe or drill, downwards, holes 5 cm apart, so it is not so suited to larger trees due to the number of cuts/holes required and it is also difficult to control multi-stemmed trees where each separate stem requires treatment. The cut stump method is more suitable for use on large trees and multi-stemmed plants. Basal bark treatment involves spraying or painting herbicide from ground level up to 50 cm which is a rapid method of control in areas with large monocultures but should not be used in aquatic situations for both environmental and effectiveness reasons. Foliar application of herbicides is useful for dense monocultures of young plants up to 1 m tall where there is no risk of damaging native vegetation.

Gaps in Knowledge/Research Needs

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Areas of recommended research include the environmental impact of this species in places where it has become weedy or invasive, in order to better determine the risk of its introduction. Considering its long history of cultivation around the world, however, at present the species does not appear to be a priority invasive species.

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Contributors

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28/1/2015 Updated by:

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

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

21/07/2009 Updated by:

Nick Pasiecznik, Consultant, France

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