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Annona squamosa
Annona squamosa
Annona squamosa
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Annona squamosa
Cross section of the fruit shown on right
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Magnoliids
Order: Magnoliales
Family: Annonaceae
Genus: Annona
Species:
A. squamosa
Binomial name
Annona squamosa
Synonyms

Annona asiatica L.[3]
Annona cinerea Dunal
Guanabanus squamosus (L.)M.Gómez[4] Xylopia glabra L.[5]
Annona forskahlii DC.[6]

Annona squamosa is a small, well-branched tree or shrub[7] from the family Annonaceae that bears edible fruits called sugar apples or sweetsops or custard apples.[8] It tolerates a tropical lowland climate better than its relatives Annona reticulata and Annona cherimola[6] (whose fruits often share the same name)[3] helping make it the most widely cultivated of these species.[9]

Annona squamosa is semi-(or late) deciduous,[10] and 3 to 8 metres (10 to 26 feet) tall[7][10], similar to soursop (Annona muricata).[11] It is native of tropical climate in the Americas and West Indies, and Spanish traders aboard the Manila galleons docking in the Philippines brought it to Asia.[12]

Michał Boym's drawing of, probably, the sugar-apple, in his Flora Sinensis (1655)

The fruit is spherical-conical, 5–10 centimetres (2–4 inches) in diameter and 6–10 cm (2+1⁄4–4 in) long, and weighing 100–240 grams (3.5–8.5 ounces), with a thick rind composed of knobby segments. The colour is typically pale green through blue-green, with a deep pink blush in certain varieties, and typically has a bloom. It is unique among Annona fruits in being segmented; the segments tend to separate when ripe, exposing the innards.

The flesh is fragrant and sweet, creamy white through light yellow, and resembles and tastes like custard. The seeds are coated with the flesh, It is found adhering to 13-to-16-millimetre-long (1⁄2 to 5⁄8 in) seeds forming individual segments arranged in a single layer around a conical core. It is soft, slightly grainy, and slippery. The hard, shiny seeds may number 20–40 or more per fruit and have a brown to black coat, although varieties exist that are almost seedless.[12][13] The seeds can be ground for use as an insecticide, although this has not been approved by the US EPA or EU authorities.[8] The stems run through the centre of the fruit connecting it to the outside. The skin is shaped like a Reuleaux triangle coloured green and rough in texture. Due to the soft flesh and structure of the sugar apple it is very fragile to pressure when ripe.

New varieties are also being developed in Taiwan and Hong Kong. The atemoya or "pineapple sugar-apple", a hybrid between the sugar-apple and the cherimoya, is popular in Taiwan, although it was first developed in the United States in 1908. The fruit is similar in sweetness to the sugar-apple, but has a very different taste. As its name suggests, it tastes like pineapple.

Description

[edit]
Flower
Seedling
Branches

The fruit of A. squamosa (sugar-apple) has sweet whitish pulp, and is popular in tropical markets.[10]

Stems and leaves

[edit]
A. squamosa leaves

Branches with light brown bark and visible leaf scars; inner bark light yellow and slightly bitter; twigs become brown with light brown dots (lenticels – small, oval, rounded spots upon the stem or branch of a plant, from which the underlying tissues may protrude or roots may issue).[6]

Thin, simple, alternate leaves[11] occur singly,[6] 5 to 17 centimetres (2 to 6+3⁄4 inches) long and 2 to 6 cm (3⁄4 to 2+3⁄8 in) wide;[10][6] rounded at the base and pointed at the tip (oblong-lanceolate).[10] They are pale green on both surfaces and mostly hairless[6] with slight hairs on the underside when young.[7] The sides sometimes are slightly unequal and the leaf edges are without teeth, inconspicuously hairy when young.[6][11]

The leaf stalks are 0.4 to 2.2 cm (1⁄8 to 7⁄8 in) long,[10] green, and sparsely pubescent.[6]

Flowers

[edit]

Solitary or in short lateral clusters of 2–4 about 2.5 cm (1 in) long,[10] greenish-yellow flowers on a hairy, slender[6] 2 cm (3⁄4 in) long stalk.[10] Three green outer petals, purplish at the base, oblong, 1.6 to 2.5 cm (5⁄8 to 1 in) long, and 0.6 to 0.75 cm (1⁄4 to 5⁄16 in) wide, three inner petals reduced to minute scales or absent.[7][10] Very numerous stamens; crowded, white, less than 1.6 cm (5⁄8 in) long; ovary light green. Styles white, crowded on the raised axis. Each pistil forms a separate tubercle (small rounded wartlike protuberance), mostly 1.3 to 1.9 cm (1⁄2 to 3⁄4 in) long and 0.6 to 1.3 cm (1⁄4 to 1⁄2 in) wide which matures into the aggregate fruit.[6]

Flowering occurs in spring-early summer[10] and flowers are pollinated by nitidulid beetles.[14] Its pollen is shed as permanent tetrads.[15]

Fruits and reproduction

[edit]

Fruits ripen 3 to 4 months after flowering.[16]

Aggregate and soft fruits form from the numerous and loosely united pistils of a flower[6] which become enlarged[10] and mature into fruits which are distinct from fruits of other species of genus[6] (and more like a giant raspberry instead).

The round or heart-shaped[6] greenish yellow, ripened aggregate fruit is pendulous[10] on a thickened stalk; 5 to 10 cm (2 to 3+7⁄8 in)[6][7] in diameter[10][11] with many round protuberances[6] and covered with a powdery bloom. Fruits are formed of loosely cohering or almost free carpels (the ripened pistels).[7]

The pulp is white tinged yellow,[7] edible and sweetly aromatic. Each carpel containing an oblong, shiny and smooth,[6] dark brown[7] to black, 1.3 to 1.6 cm (1⁄2 to 5⁄8 in) long seed.[6]

Nutrition and uses

[edit]
Sugar-apples, (sweetsop), raw
Nutritional value per 100 g (3.5 oz)
Energy393 kJ (94 kcal)
23.64 g
Dietary fiber4.4 g
0.29 g
2.06 g
Vitamins and minerals
VitaminsQuantity
%DV†
Thiamine (B1)
9%
0.11 mg
Riboflavin (B2)
9%
0.113 mg
Niacin (B3)
6%
0.883 mg
Pantothenic acid (B5)
5%
0.226 mg
Vitamin B6
12%
0.2 mg
Folate (B9)
4%
14 μg
Vitamin C
40%
36.3 mg
MineralsQuantity
%DV†
Calcium
2%
24 mg
Iron
3%
0.6 mg
Magnesium
5%
21 mg
Manganese
18%
0.42 mg
Phosphorus
3%
32 mg
Potassium
8%
247 mg
Sodium
0%
9 mg
Zinc
1%
0.1 mg
Link to USDA Database entry
†Percentages estimated using US recommendations for adults,[17] except for potassium, which is estimated based on expert recommendation from the National Academies.[18]

Sugar-apple is high in energy, an excellent source of vitamin C and manganese, a good source of thiamine and vitamin B6, and provides vitamin B2, B3 B5, B9, iron, magnesium, phosphorus and potassium in fair quantities.[19]

Chemistry

[edit]

The diterpenoid alkaloid atisine is the most abundant alkaloid in the root. Other constituents of Annona squamosa include the alkaloids oxophoebine,[20] reticuline,[20] isocorydine,[21] and methylcorydaldine,[21] and the flavonoid quercetin-3-O-glucoside.[22]

Bayer AG has patented the extraction process and molecular identity of the annonaceous acetogenin annonin, as well as its use as a biopesticide, although this use has not been approved by US or EU authorities.[23] Other acetogenins have been isolated from the seeds,[24] bark,[25] and leaves.[citation needed]

Distribution and habitat

[edit]

Annona squamosa is native to the tropical Americas and West Indies, but the exact origin is unknown. It is now the most widely cultivated of all the species of Annona, being grown for its fruit throughout the tropics and warmer subtropics, such as India, Indonesia, Thailand, Taiwan, and China as far north as Suzhou;[26] it was introduced to southern Asia before 1590. It is naturalized as far north as Cuba, South Florida, United States and as far south as Bahia, Brazil, and is an invasive species in some areas.[6][9][11]

Native
Neotropic
Caribbean: Antigua and Barbuda, Bahamas, Barbados, Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe, Haiti, Jamaica, Martinique, Montserrat, Netherlands Antilles, Puerto Rico, St Kitts and Nevis, St Lucia, St Vincent and the Grenadines, Suriname, Trinidad and Tobago, Virgin Islands.
Central America: Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama
Northern South America: Suriname, French Guiana, Guyana, Venezuela
Western South America: Bolivia, Colombia, Ecuador, Peru
Southern South America: Argentina, Brazil, Chile, Paraguay, Uruguay[6]
Naturalised
Pacific: Samoa, Tonga
North America: Mexico, Belize
Afrotropic: Angola, Namibia, Sudan, Tanzania, Uganda, Zanzibar, Kenya
Australasia: Australia, Fiji, New Zealand, Papua New Guinea, Solomon Islands
Indomalaya: Bangladesh, Cambodia, China, India, Indonesia, Laos, Malaysia, Nepal, Pakistan, Philippines, Sri Lanka, Taiwan, Thailand, Myanmar, Vietnam
Palearctic: Cyprus, Greece, Lebanon, Malta,[6] Israel

Climate and cultivation

[edit]

Like most species of Annona, it requires a tropical or subtropical climate with summer temperatures from 25 °C (77 °F) to 41 °C (106 °F), and mean winter temperatures above 15 °C (59 °F). It is sensitive to cold and frost, being defoliated below 10 °C (50 °F) and killed by temperatures of a couple of degrees below freezing. It is only moderately drought-tolerant, requiring at least 700 millimetres (28 in) of annual rainfall, and does not produce fruit well during droughts.

It will grow from sea level to an altitude of 2,000 metres (6,600 feet) and thrives in hot dry climates, differing in its tolerance of lowland tropics from many of the other fruit bearers in the Annona family.

It is quite a prolific bearer, and it produces fruit within as little as two to three years. A five-year-old tree can produce as many as 50 sugar apples. Poor fruit production has been reported in Florida because there are few natural pollinators (honeybees have a difficult time penetrating the tightly closed female flowers); however, hand pollination with a natural fibre brush is effective in increasing yield. Natural pollinators include beetles (coleoptera) of the families Nitidulidae, Staphylinidae, Chrysomelidae, Curculionidae and Scarabaeidae.[9][13]

Ecology

[edit]

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.

It is a host plant for larvae of the butterfly Graphium agamemnon (tailed jay).

Uses

[edit]

In traditional Indian, Thai, and Native American medicines, the leaves are boiled down with water, possibly mixed with other specific botanicals, and used in a decoction to treat dysentery and urinary tract infection.[27] In traditional Indian medicine, the leaves are also crushed for use as a poultice, and applied to wounds.[27] In Mexico, the leaves are rubbed on floors and put in hens' nests, to repel lice.[9]. Extracts are used in ethnomedicine.[28] In Haiti, the fruit is known as cachiman and is used to simply make juice.[29]

[edit]
  • Annona squamosa fruit from Myanmar
    Annona squamosa fruit from Myanmar
  • Sugar apple (right), with Taiwanese "pineapple shijia" (atemoya) (left)
    Sugar apple (right), with Taiwanese "pineapple shijia" (atemoya) (left)
  • The sugar apple readily breaks open when ripe.
    The sugar apple readily breaks open when ripe.
  • A deconstruction of a sugar apple shows a lobe of fruit and pulpy segments with seeds.
    A deconstruction of a sugar apple shows a lobe of fruit and pulpy segments with seeds.
  • A sugar apple ready to eat
    A sugar apple ready to eat
  • Sugar apple (Annona squamosa) seeds
    Sugar apple (Annona squamosa) seeds
  • Red sugar apples from Myanmar
    Red sugar apples from Myanmar
  • Sugar apples in Taitung, Taiwan
    Sugar apples in Taitung, Taiwan
  • A sugar apple in Goiânia, Brazil
    A sugar apple in Goiânia, Brazil
  • Sugar apple tree in Philippines
    Sugar apple tree in Philippines
  • Two sugar apples in Bangladesh
    Two sugar apples in Bangladesh
  • A sugar apple in tree in Terai of Nepal
    A sugar apple in tree in Terai of Nepal

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Annona squamosa

View on Grokipedia
from Grokipedia
Annona squamosa, commonly known as the sugar apple, sweetsop, or , is a small in the family native to the tropical regions of the , including the and Central and . It typically grows to a height of 3-8 meters with a broad, open crown and lanceolate leaves that are pale green and . The produces fragrant, greenish-yellow flowers and distinctive aggregate fruits that are heart-shaped, round, or conical, measuring 5-10 cm in diameter, with a knobby, scaly rind and creamy white, custard-like pulp containing numerous black seeds; the fruit is sweet and edible, containing 50-80% pulp rich in carbohydrates, (35-42 mg/100 g). Widely cultivated in tropical and subtropical areas worldwide, including parts of , , and , A. squamosa thrives in hot, dry climates with well-drained sandy soils, tolerating temperatures up to 41°C and altitudes from 0-2000 m, though it is sensitive to and requires at least 700 mm annual rainfall. The begins fruiting at 3-4 years of age and is propagated mainly by , with fruiting seasons varying by region—such as July to August in —and yields varying based on cultivars like the green-fruited 'Thai Lessard' or purplish-red 'Kampong '. It is valued primarily for its fruit, consumed fresh or in desserts like and milkshakes, providing nutritional benefits including 18-26 g carbohydrates per 100 g. Beyond its culinary role, A. squamosa has medicinal applications, with leaves, bark, roots, and seeds used traditionally to treat diarrhea, dysentery, and as vermifuges, while the seeds contain insecticidal compounds; the wood serves for fuel and light construction, and the tree provides shade and ornamental value in landscapes. Despite its benefits, the tree can become invasive in some non-native regions, such as parts of the West Indies, and requires careful management in cultivation to control pests and diseases.

Taxonomy and nomenclature

Etymology

The genus name is a Latinization of indigenous or terms like anon or hanon used by Hispaniolan peoples for fruits in this group; Linnaeus associated it with the Latin word annōna, meaning "annual produce" or "provisions." The specific squamosa comes from the Latin squāmōsus, meaning "scaly" or "covered in scales," alluding to the knobby, scale-like segments on the immature . Common names for Annona squamosa reflect its sensory qualities and regional adaptations, with "sugar apple" or "sweetsop" emphasizing the sweet, granular pulp, while "" highlights the creamy, custard-like texture—though the latter is more precisely applied to A. reticulata in some contexts. In the , it is known as atis or ata, and in as pinha or fruta-do-conde. The species was first formally described by in his in 1753, under the binomial Annona squamosa, based on specimens from tropical America.

Classification and synonyms

Annona squamosa belongs to the kingdom Plantae, phylum Tracheophyta, class Magnoliopsida, order , family , genus , and species squamosa. Within the family, A. squamosa exemplifies key characteristics such as an apocarpous composed of numerous free carpels and syncarpous fruits formed by the fusion of these carpels into an aggregate structure. Accepted synonyms for A. squamosa include Annona asiatica L., Annona cinerea Dunal, and Annona distincta Raeusch., reflecting historical taxonomic variations in classifications. Recent genomic studies, including a high-quality chromosome-level genome assembly published in 2025, have confirmed the phylogenetic position of A. squamosa within Annonaceae, revealing it as most closely related to Annona cherimola (cherimoya), with the two species diverging approximately 4.2 million years ago and forming a sister subclade with Annona montana.

Botanical description

Habit and stems

Annona squamosa is a deciduous or semi-deciduous small tree or shrub that typically reaches a height of 3 to 8 meters, forming a low, spreading canopy with an open, rounded or irregular outline. The plant exhibits a moderate growth rate, with upright to spreading branches that contribute to its bushy habit, and in cultivation, it can have a lifespan of up to 20-30 years. The stems feature light brown to grayish bark that is slightly furrowed and thin, providing a smooth texture on mature branches. Young branches are pubescent, covered in hairs that eventually become glabrous as they mature, while the overall branching pattern is irregular and often low on the cylindrical trunk. This supports an adaptive wood composition that enhances , allowing the plant to withstand periods of water stress in tropical environments. However, A. squamosa is highly susceptible to damage, with temperatures below -2°C causing severe injury or death to young trees and disrupting growth in established ones. Leaves are alternately attached along the stems, contributing to the plant's overall architecture without altering its primary .

Leaves

The leaves of Annona squamosa are simple, alternate, and distichous (two-ranked), often appearing clustered at the tips of branches. They exhibit an oblong-lanceolate to elliptic shape, typically measuring 5–17 in length and 2–6 in width, with an acute to obtuse apex and cuneate to rounded base, petiole 0.4–1 long. The adaxial surface is pale green and glabrous upon maturity, though young leaves are slightly pubescent; the abaxial surface is paler, bearing a prominent midrib and secondary veins that are raised beneath and impressed above. When crushed, the leaves release a distinctive aromatic scent attributable to their content, which includes compounds such as β-caryophyllene and germacrene D. Annona squamosa displays a habit, shedding its leaves during prolonged dry seasons to enhance , with new flushes emerging at the start of wet periods to support renewed growth.

Flowers

The flowers of Annona squamosa are bisexual, pendulous, and typically greenish-yellow, measuring 2–3 cm in length. They emerge solitary or in small clusters of 2–4 from short peduncles (1–2 cm long) in the axils of year-old shoots or new growth. Flowering occurs primarily in spring to early summer, coinciding with periods of new vegetative flushing in subtropical and tropical regions. Structurally, the flowers lack prominent sepals beyond three small, inconspicuous, triangular green ones (1.5–2 mm long) that are persistent but minimally visible. The features three outer petals that are oblong, 1.6–2.5 cm long, 6–7.5 mm wide, thick, fleshy, green with purple at the base, and keeled inside; the inner petals are reduced to minute scales or absent. The androecium consists of numerous (typically 20–30), arranged in a dense, central mass around a conical receptacle, with each stamen featuring a short filament and oblong anther (about 1 mm long). The includes 20–40 free, oblong carpels (1.5–2 mm long), each with a single and a capitate stigma. These flowers exhibit protogyny, with the stigmas becoming receptive in the female phase prior to anther dehiscence in the male phase, facilitating cross-pollination. The blooms produce a mildly fruity scent, which serves to attract specific nitidulid beetles as pollinators. This odor, combined with the pendulous orientation and enclosed floral chamber formed by the , creates an environment suited to beetle activity within the flower.

Fruits

The of Annona squamosa is an aggregate type known as a syncarp, developed from the fusion of numerous carpels derived from a single flower. It typically exhibits a spherical to heart-shaped form, with dimensions of 5–10 cm in and a weight ranging from 100–240 g. The exterior surface is characterized by 20–40 knobby, scale-like carpels that create a distinctive bumpy or tuberculate appearance, separated by shallow grooves. The skin is thin and pale green when unripe, transitioning to yellowish with a slight powdery bloom upon maturation. The interior features creamy white, custard-like pulp that is soft, slightly grainy, and aromatic, surrounding individual seeds within each carpel. Following , the fruit begins to form and develops over 3–4 months, with maturation occurring irregularly across the tree's . Due to this uneven ripening on the plant, fruits are optimally harvested while still green and allowed to ripen off-tree, a process that usually takes 2–5 days. Each fruit contains numerous hard, shiny black seeds, typically 1–2 cm long and ovoid-ellipsoid in shape, with one seed per carpel. These seeds are poisonous if consumed raw, primarily due to annonacin, a potent neurotoxic acetogenin that can cause irritation and toxicity.

Reproduction

Pollination

Annona squamosa is primarily pollinated by nitidulid beetles, such as Carpophilus dimidiatus and Carpophilus hemipterus, which enter flowers in the female phase, remain trapped overnight, and transfer pollen during anther dehiscence the following day. These beetles feed on floral tissues and mate within the flowers, facilitating pollination before exiting in the male phase. The inner petals form a specialized chamber that traps the beetles, enhancing their role as effective pollinators. Pollination by bees or wind is ineffective due to the flowers' structure and the beetles' exclusive behavior. The species exhibits protogynous dichogamy, with the stigma receptive in the female phase before anthers dehisce in the male phase, promoting through temporal separation. However, A. squamosa is self-compatible, allowing set from self-pollen if transferred, though cross-pollination often yields higher seed viability. In introduced ranges such as and , low densities of native nitidulid beetles result in poor natural set, typically ranging from 1% to 8% without intervention. This scarcity leads to deformed or sparse development, limiting commercial yields. Hand , involving manual transfer of from male-phase to female-phase flowers using a or similar tool, achieves set rates exceeding 80% and substantially boosts yields in pollinator-poor areas. This technique is particularly vital in regions like , where natural set can approach zero, and in Indian orchards, where it enhances quality and quantity despite labor demands.

Seed dispersal and propagation

The seeds of Annona squamosa are primarily dispersed through zoochory, where animals consume the sweet, pulp of the ripe and excrete the intact seeds away from the parent plant. Birds and fruit bats, such as the Philippine flying fox (Acerodon jubatus), play key roles in this , particularly in tropical regions where the fruit ripens on the and attracts these dispersers. Mammals like rats and squirrels also contribute by feeding on fallen fruits and scattering seeds. In flood-prone habitats, hydrochory occurs as buoyant seeds are carried by water currents, aiding long-distance dispersal. Each fruit typically contains 20-40 hard, brown to black , which exhibit short viability, often lasting only a few weeks to two months under natural conditions before potential declines significantly. Fresh achieve 50-95% rates under optimal moist, warm conditions (25-30°C), typically emerging in 3-4 weeks after testa rupture and protrusion. , such as soaking in water for 3 days or mechanical nicking, enhances by overcoming the impermeable coat. Human propagation of A. squamosa commonly begins with direct seed sowing in well-drained media, yielding variable offspring due to cross-pollination. For superior cultivars, vegetative methods like air-layering, , inarching, or onto rootstocks (e.g., A. squamosa or other species) ensure true-to-type plants and dwarfing for easier management. Cuttings are less reliable but can succeed with rooting hormones. Vegetative propagation offers advantages over seeding, including faster time to fruiting (1-2 years versus 3-4 years from ), uniform crop characteristics, and improved resistance to soil-borne diseases through selected rootstocks.

Distribution and habitat

Native range

Annona squamosa, commonly known as the sugar apple or sweetsop, is native to the Neotropical region, encompassing from southward to , northern including countries such as and , and the islands of the . This distribution spans diverse lowland ecosystems where the species likely originated and evolved, with its exact center of origin remaining somewhat uncertain due to extensive pre-Columbian human influence. Within its native habitats, A. squamosa thrives in lowland tropical moist forests, open savannas, and disturbed sites such as hillocks, gravelly slopes, and waste lands, generally at elevations ranging from to 1,500 meters. The plant shows a strong preference for well-drained, sandy or gravelly soils in warm, subtropical to tropical climates with seasonal dry periods, allowing it to tolerate both moist lowlands and drier upland margins. These ecological niches support its habit and opportunistic growth in areas altered by natural disturbances or early human activity. Historical evidence from archaeological contexts reveals that in the domesticated A. squamosa, with independent centers of cultivation emerging in and northwestern . Botanical remains and linguistic records from pre-Columbian sites in the and mainland regions indicate its integration into early agricultural systems, where it was valued for its edible fruit and propagated in managed landscapes. The holds a of Least Concern on the , owing to its broad native distribution across approximately 3.5 million square kilometers and a stable overall population trend. Nonetheless, localized declines occur in fragmented habitats due to and , underscoring the need for monitoring in vulnerable subpopulations.

Introduced ranges

Annona squamosa was introduced to , particularly the , prior to 1590 by Spanish traders via the Manila galleons, marking one of the earliest trans-Pacific dispersals of the from its native tropical American range. By the early 17th century, explorers had facilitated its spread to southern , where it quickly became established in tropical lowlands. Introduction to occurred through European colonial trade routes in the 1600s, primarily to West African regions, leading to its cultivation and in suitable tropical climates. Today, the is , cultivated or naturalized in over 50 countries across , , , and the due to its valued fruit and adaptability. Major cultivation occurs in , the world's leading producer with an annual output of approximately 514,000 metric tons from 56,300 hectares (as of 2023-24), particularly in states like , , , and . Other key regions include , where it supports significant commercial orchards; , with concentrated production in and ; and , focused on subtropical areas for export markets. The species has naturalized in subtropical zones such as , where it grows wild in the Keys, and , though it remains primarily planted rather than widespread. In some introduced areas, particularly Pacific islands like , , and , Annona squamosa exhibits weedy tendencies, forming dense thickets that compete with native vegetation for resources and space. This invasiveness is generally limited, and the species is often managed through practices to mitigate ecological impacts while preserving its economic value. Recent phylogeographic analyses, including a 2024 high-quality assembly, reveal that populations in introduced ranges show genetic with wild relatives, enhancing adaptive traits like and contributing to local diversity.

Cultivation

Climate and soil requirements

Annona squamosa, commonly known as sugar apple or sweetsop, thrives in tropical climates with daytime temperatures ranging from 25°C to 41°C and minimum temperatures not falling below 15°C, as these conditions support optimal growth and fruiting. The is highly frost-sensitive, with young trees suffering damage at 0°C to -1°C and mature trees tolerating brief exposures to -2°C but dying at lower temperatures; it is best suited to USDA hardiness zones 10 to 11. Annual rainfall of 700 to 2,000 mm is ideal, with adequate moisture during the and fruit development to prevent reduced yields, though the exhibits moderate and benefits from dry periods preceding flowering to promote bloom initiation. In arid regions, supplemental is essential to maintain without waterlogging. The species prefers well-drained sandy or alluvial soils with a range of 5.5 to 7.5, tolerating a broader spectrum from slightly acidic to mildly alkaline conditions but performing poorly in heavy clay or waterlogged environments that lead to . It is intolerant of flooding for more than 7 to 10 days, emphasizing the need for good drainage to ensure healthy . For fruit production, full sun exposure is required, providing at least 6 to 8 hours of direct sunlight daily, though juvenile plants show some tolerance to partial shade during establishment.

Planting and care

Annona squamosa trees are typically planted at spacings of 4 to 6 meters between trees to accommodate their canopy spread of up to 5 meters, allowing for adequate light penetration and air circulation in commercial or home orchards. Planting holes should be dug to dimensions of approximately 50 x 50 x 50 cm, filled with a mixture of topsoil and 10-20 kg of organic manure such as farmyard manure to improve soil fertility and drainage before inserting the seedling. In suitable tropical climates with well-drained soils, trees establish roots quickly and begin bearing fruit within 3-4 years. Routine care involves annual to maintain health and productivity, focusing on the removal of dead wood, suckers, and crossing branches to enhance and reduce risk while training the to an open-center shape for better light distribution. is best performed in early spring or post-harvest, limiting cuts to about one-third of the previous year's growth to avoid stressing the . Fertilization supports vigorous growth and fruiting, with balanced NPK fertilizers such as 10-10-10 or similar formulations applied at rates of 200-300 g per annually for young trees, increasing to 1-2 kg for mature specimens divided into 2-4 applications per year. Micronutrients like and are supplemented via foliar sprays 2-4 times yearly to address deficiencies common in sandy soils, while a 5-8 cm layer of organic around the base helps retain and suppress weeds. Harvesting occurs 2-3 times per year in tropical regions after the reaches 3 years of age, with mature trees yielding 10-50 kg of depending on and management, typically picked when the skin turns yellowish-green to avoid damage. Post-harvest, fruits ripen optimally at 20-25°C over 2-5 days, after which they can be stored in for up to 4 days to extend .

Varieties and cultivars

Annona squamosa cultivars vary in fruit characteristics, yield, and adaptability to local conditions. In , 'Thai Lessard' is a recommended selection with green-peeled fruits weighing 227–454 g and excellent eating quality, while 'Kampong Mauve' features reddish-purple peel and fruits of 136–398 g with good to very good quality. In , the 'Balanagar' variety from stands out for its high yield under rain-fed, rocky soils, thick rind, high pulp content, few seeds, inherent sweetness, and prolonged shelf life without irrigation or fertilizers. In June 2025, a geographical indication (GI) tag application was filed for Balanagar (application no. 1569), which remains pending as of November 2025. Breeding programs have emphasized selections for and reduced seediness, such as Indian varieties including 'Balanagar', 'Arka Sahan', and 'APK (Ca)1', evaluated for performance in rainfed vertisols with yields varying by but favoring adaptable, high-pulp types. Hybrids like (A. squamosa × A. cherimola) enhance desirable traits over pure A. squamosa. The 'Gefner' Atemoya , suited to , produces large conical to ovate fruits up to 908 g with creamy, sweet pulp, high yields of 75–200 fruits per tree, and no need for . 'African Pride', an Australian Atemoya selection, is an early, consistent bearer of medium to large heart-shaped fruits (500–800 g) with low seed counts and self-fertility, though prone to skin blackening from diseases like purple spot. Atemoya hybrids are widely cultivated in , particularly in , for their superior flavor, larger sizes, and export potential to markets like . Cultivar and hybrid selection prioritizes size (up to 500 g or more in hybrids), sweetness (15–20° ), and , as seen in evaluations of genotypes showing soluble solids in this range alongside high pulp recovery.

Pests and diseases

Annona squamosa is susceptible to several pests that can significantly impact quality and vigor. Mealybugs, particularly Planococcus citri and Pseudococcus spp., infest stems, leaves, and , excreting honeydew that promotes growth, leading to reduced and aesthetic damage. Fruit borers, including the annona seed borer Bephratelloides cubensis and the yellow moth Conogethes punctiferalis, bore into developing and , causing internal decay, premature fruit drop, and up to 20-80% crop loss in severe cases, especially during late summer. Fungal diseases pose major threats, particularly in humid environments. Anthracnose, caused by Colletotrichum gloeosporioides, affects flowers, leaves, and fruits, manifesting as dark lesions, blossom blight, and post-harvest rot, with reported crop losses ranging from 10-100%. Root rot, primarily from Phytophthora spp. such as P. capsici and P. nicotianae in poorly drained soils, results in wilting, yellowing leaves, stunted growth, and root decay, often leading to tree decline or death. Plant-parasitic nematodes, notably root-knot nematodes (Meloidogyne spp.), attack roots, forming that disrupt water and nutrient uptake, causing and reduced vigor in infested trees. Management of these biotic threats relies on (IPM) strategies combining cultural, biological, and targeted chemical approaches. Cultural practices include pruning for improved air circulation to minimize humidity-related diseases and to break nematode cycles, while avoiding overwatering prevents . Biological controls, such as releasing predatory like ladybugs for mealybugs or parasitic wasps for borers, enhance natural enemy populations. Chemical interventions are limited to essentials like for soft-bodied pests such as mealybugs, and copper-based fungicides (e.g., copper hydroxide) for anthracnose, applied sparingly to preserve beneficial organisms; resistant rootstocks are recommended for nematode-prone areas.

Ecology

Interactions with pollinators and dispersers

Annona squamosa exhibits a specialized reliant primarily on nitidulid beetles (Coleoptera: Nitidulidae), such as species in the genera Carpophilus and Colopterus, which are attracted to the flowers' yeasty and feed on floral tissues during their nocturnal visits. These beetles enter the flowers in the female phase, become trapped overnight within the sturdy petals that form a protective chamber, and effect by transferring during the subsequent male phase when anthers dehisce. In its native range, secondary may occur through , which visit flowers frequently but contribute less effectively due to their behavior limiting transfer, while play a minor role in some related species but are not primary for A. squamosa. Seed dispersal in A. squamosa is facilitated by frugivorous animals that consume the sweet, aril-covered seeds and deposit them away from the parent plant, promoting wider distribution. In its native American range, bats such as Artibeus jamaicensis and birds like (Pipridae) serve as key dispersers, ingesting the pulp and excreting intact seeds during flight or perching. In introduced Asian regions, including rats ( spp.) and primates such as macaques contribute to dispersal by carrying and dropping seeds after feeding on the ripe . These interactions represent mutualistic relationships where A. squamosa provides resources in exchange for reproductive services. Flowers offer nitidulid beetles shelter from predators and adverse weather within the pollination chamber, along with protein-rich tissues for feeding and breeding, enabling the beetles to survive and reproduce effectively. Similarly, the nutritious, fleshy arils of the reward dispersers with energy, while the hard-coated pass through their digestive systems unharmed, facilitating transport to suitable sites. In cultivated monocultures, these mutualisms can be disrupted by broad-spectrum pesticides, which reduce nitidulid beetle populations and lead to lower fruit set and yields, often necessitating to compensate for diminished natural activity.

Role in ecosystems

Annona squamosa contributes to in tropical ecosystems by serving as a larval host plant for the butterfly Graphium agamemnon (tailed jay), providing essential foliage for caterpillar development and supporting lepidopteran populations in native and naturalized ranges. Its flowers attract nitidulid beetles as primary , offering and resources that sustain these and promote their role in broader networks. These interactions enhance local insect diversity, particularly in systems where the tree is integrated. The species forms mutualistic associations with arbuscular mycorrhizal fungi (AMF), which colonize its roots to improve nutrient absorption, especially , thereby bolstering plant vigor and indirectly supporting microbial communities in nutrient-poor tropical . Although its is relatively shallow, it helps stabilize surface in sloped tropical terrains, contributing to mitigation when planted in mixed agroecosystems. In introduced regions such as parts of the , , , and , A. squamosa exhibits invasive potential, establishing dense thickets that alter light availability and suppress native through competitive shading and . As an indicator for tropical disturbances, A. squamosa thrives in degraded or human-modified landscapes, reflecting stress from land-use changes. It also plays a role in within modified habitats.

Nutritional composition

Macronutrients and calories

The fruit of Annona squamosa, commonly known as sugar apple or sweetsop, provides approximately 94 kcal of per 100 g of portion, with the majority derived from carbohydrates. This caloric content positions it as a moderate- fruit suitable for snacks or dietary inclusion in tropical regions. The macronutrient profile is dominated by carbohydrates at 23.6 g per 100 g, comprising sugars—primarily , , and —and 4.4 g of , which supports digestive health. Protein contributes a modest 2.1 g, while is minimal at 0.29 g, making the fruit low in overall. ranges from 68% to 75%, which accounts for its juicy texture and relatively low nutrient density compared to drier fruits. Nutritional content can vary by and maturity. In comparison to other tropical fruits such as bananas or mangos, A. squamosa offers a similar overall caloric and carbohydrate profile but stands out with higher levels of readily accessible sugars, providing quick energy without excessive fat or protein. This composition aligns with its role as a fresh, hydrating fruit in diets where rapid carbohydrate replenishment is beneficial.

Vitamins and minerals

The pulp of Annona squamosa fruit is notably rich in vitamin C, containing 36.3 mg per 100 g, which provides approximately 60% of the recommended daily value and plays a key role in supporting immune function by acting as an antioxidant and aiding collagen synthesis. It also supplies vitamin B6 at 0.2 mg per 100 g (12% DV), essential for protein metabolism, neurotransmitter synthesis, and red blood cell formation. Key minerals in the fruit include at 0.42 mg per 100 g (18% DV), which contributes to bone health and activity; at 0.11 mg per 100 g (12% DV), supporting formation and iron utilization; and at 247 mg per 100 g (5% DV), helping maintain balance and nerve function. The fruit is low in sodium, with levels typically under 10 mg per 100 g, making it suitable for sodium-restricted diets. The ascorbic acid () in A. squamosa enhances non-heme iron absorption in the body, where iron content is about 0.6 mg per 100 g, thereby improving overall of this mineral for preventing . Nutrient concentrations, particularly vitamins, tend to be higher in fully ripe fruits due to continued during maturation.
NutrientAmount per 100 g% Daily Value
36.3 mg60%
0.2 mg12%
0.42 mg18%
0.11 mg12%
247 mg5%
Iron0.6 mg3%
Sodium<10 mg<1%

Phytochemicals and bioactive compounds

Annona squamosa is rich in secondary metabolites, particularly alkaloids and concentrated in the seeds, which contribute to its bioactive profile. , a prominent annonaceous and , is found in the seeds, where it acts by depleting ATP through inhibition of mitochondrial complex I. Other , such as squamocin and annosquatin, are also present in the seeds and exhibit cytotoxic effects against tumor cell lines by disrupting cellular energy production. Flavonoids and phenolic compounds dominate the phytochemical content of the fruit pulp, with quercetin and rutin serving as key antioxidants at concentrations of 50-100 mg/100 g. These compounds scavenge free radicals and modulate , supporting responses through inhibition of pro-inflammatory pathways like . Total phenolic content in the pulp reaches approximately 405 mg equivalents per 100 g, underscoring the fruit's potential as a source of these bioactives. Essential oils extracted from the leaves of Annona squamosa contain monoterpenes such as , comprising about 7.7% of the oil composition, alongside and other volatiles that demonstrate activity against bacterial and fungal pathogens. These oils inhibit microbial growth by disrupting cell membranes and function. Regarding toxicity, the seeds pose risks when consumed raw due to elevated levels, potentially leading to neurotoxic effects like atypical , whereas the pulp contains negligible amounts of these compounds, rendering it safe in moderation as per recent assessments.

Uses

Culinary applications

The fruit of Annona squamosa, commonly known as sugar apple or , is primarily consumed fresh by breaking open the knobby, green-to-yellow skin and scooping out the creamy white pulp with a , while discarding the large, inedible black seeds. The pulp's sweet, custard-like texture and flavor profile, reminiscent of a blend of , , and , make it a favored fresh treat in tropical regions. In culinary preparations, the pulp is versatile and incorporated into various desserts and beverages, including ice creams, sherbets, smoothies, milkshakes, and baked goods such as cakes. It is also pureed for use in fruit cups, salads, juices, and other beverages, enhancing their sweetness and nutritional value with its high vitamin C content. Regionally, in India, the pulp features prominently in sitaphal milkshakes blended with milk and sugar for a refreshing drink. In Thailand, it appears in fresh salads combining the fruit with herbs and dressings for a sweet-savory contrast. Brazilian cuisine utilizes the pulp in sorbet-like frozen desserts, capitalizing on its cooling properties in warm climates. For processing, the ripe is often converted into purees or dried into , which serves as a base for beverages, jams, and jellies to extend usability and incorporate into year-round recipes. Unripe fruits, being more , can be cooked into chutneys with spices for a tangy in South Asian dishes. Ripe A. squamosa fruits are highly perishable, lasting only 1-2 days at ambient (25-30°C) due to rapid respiration and softening. Under refrigerated storage at 15-20°C with (low oxygen, 10% CO₂), they maintain quality for up to 7-12 days, avoiding chilling injury that occurs below 10°C. For commercial export, fruits are harvested at the mature green stage and treated to delay , allowing transport over longer distances while preserving firmness.

Medicinal and therapeutic uses

In , leaf decoctions of Annona squamosa have been used to treat and diarrhea, particularly in regions like and tropical America where the plant is native. Bark preparations are employed in Ayurvedic practices for managing , leveraging the plant's purported antihypertensive properties. In folk medicine, the fruit is valued for its cooling effects to alleviate fevers and related inflammatory conditions. Scientific research has explored the therapeutic potential of A. squamosa, with from seeds and bark showing promising anti-cancer activity in studies, including cytotoxicity against breast () and colon lines. A 2024 review highlights the role of these compounds in targeting liver, colon, and cancers through mechanisms like mitochondrial inhibition. , such as rutin and quercetin-3-O-glucoside in leaf extracts, demonstrate antidiabetic effects in animal models; for instance, oral administration reduced blood glucose by 18-30% in streptozotocin-induced diabetic rats and high-fat-fed mice during glucose tolerance tests. Recent studies (as of 2024-2025) have also identified potential anti-arthritic and larvicidal applications from leaf extracts, supporting expanded therapeutic uses. Seed extracts exhibit properties, inhibiting bacterial growth including in agar diffusion assays, attributed to polyphenols and other bioactive compounds with minimum inhibitory concentrations around 50 mg/mL. However, overconsumption of the fruit or seeds poses risks of due to annonaceous like squamocin, which inhibit mitochondrial complex I and may lead to atypical Parkinson's-like symptoms, including tremors and , as observed in chronic exposure cases in tropical regions. Moderate consumption of the pulp is considered safe based on nutritional profiles, and clinical trials on therapeutic uses remain limited; A. squamosa extracts are not approved by the FDA for medical treatment.

Other economic and cultural uses

Annona squamosa plays a significant role in tropical economies through its production and trade, with as the leading producer at approximately 449,000 metric tons annually as of 2021 data from the National Horticulture Board. In 2023, international trade in the fruit, primarily from , generated over $40 million in export value, with volumes reaching 26,660 metric tons, underscoring its growing commercial importance. supporting systems where the tree is intercropped with crops like to enhance and in tropical regions. Industrially, the seeds yield an oil rich in unsaturated fatty acids, utilized in the manufacture of soaps and for its emollient and moisturizing properties. , known for its durability, is employed in crafting tool handles and pegs, providing a sustainable material for local applications. Additionally, bark extracts exhibit insecticidal properties, particularly against larvae, offering a natural alternative for in eco-friendly formulations. Culturally, Annona squamosa holds symbolic value and practical uses in various traditions; in , the , known as Ramphal, is offered during the Hindu festival of Ram Navami to honor Lord Rama's birth. Local harvest celebrations, such as festivals in producing regions, highlight community gatherings and traditional processing methods. The is also widely planted as an ornamental in tropical gardens for its attractive foliage and unique clusters, enhancing landscape aesthetics. In terms of , cultivation and processing of Annona squamosa bolster rural livelihoods in tropical areas, particularly for indigenous , where initiatives in enabled earnings of up to Rs. 80,000 annually through fair pricing and value-added products like pulp, compared to previous minimal incomes below Rs. 5,000. These efforts, often aligned with sustainable harvesting practices, promote economic empowerment and environmental conservation in contexts.

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