Annona squamosa

{{Short description|Species of tree}} {{Use British English|date=December 2023}} {{Use dmy dates|date=July 2025}} {{speciesbox | image = Sugar apple on tree.jpg | image2 = Sugar apple with cross section.jpg | image2_caption = Cross section of the fruit shown on right | status = LC | status_system = IUCN3.1 | status_ref = <ref>{{cite iucn |author=Botanic Gardens Conservation International (BGCI) |author2=IUCN SSC Global Tree Specialist Group |date=2019 |title=''Annona squamosa'' |article-number=e.T146787183A146787185 |doi=10.2305/IUCN.UK.2019-2.RLTS.T146787183A146787185.en |access-date=16 December 2022}}</ref> | genus = Annona | species = squamosa | authority = L.<ref name="NRCS">{{cite web | url = https://plants.usda.gov/java/profile?symbol=ANSQ | title = PLANTS Profile, ''Annona squamosa'' L | access-date = 17 April 2008 | author = Natural Resources Conservation Service (NRCS) | author-link = Natural Resources Conservation Service | website = The PLANTS Database | publisher = United States Department of Agriculture }}</ref> | range_map_caption = Curre F nt range of uncultivated ''A. squamosa''. | synonyms = *''Annona asiatica'' {{small|L.}} *''Annona asiatica'' {{small|Vahl}} *''Annona cinerea'' {{small|Dunal}} *''Annona distincta'' {{small|Raeusch.}} *''Annona forskahlii'' {{small|DC.}} *''Annona glabra'' {{small|Forssk.}} *''Annona squamosa'' {{small|Delile}} *''Annona squamosa'' f. ''parvifolia'' {{small|Kuntze}} *''Xylopia glabra'' {{small|L.}} *''Xylopicron glabrum'' {{small|(L.) Crantz}} |synonyms_ref = <ref name=":1">{{Cite web |title=''Annona squamosa'' L. {{!}} Plants of the World Online {{!}} Kew Science |url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:72319-1 |website=Plants of the World Online |access-date=30 March 2026 |language=en}}</ref> }}

'''''Annona squamosa''''' is a small, well-branched tree or shrub<ref name="aluka">{{Cite web|title=Compilation: Annona squamosa|url=https://plants.jstor.org/compilation/Annona.squamosa|work=Global Plants|publisher=JSTOR |access-date=5 September 2019}}</ref> from the family Annonaceae that bears edible fruits called '''sugar apples''' or '''sweetsops''' or '''custard apples'''.<ref name=":0">{{Cite book |title=The Complete Guide to Edible Wild Plants |publisher=Skyhorse Publishing |others=United States Department of the Army |year=2009 |isbn=978-1-60239-692-0 |location=New York |page=100 |language=en-US |oclc=277203364}}</ref> It tolerates a tropical lowland climate better than its relatives ''Annona reticulata'' and ''Annona cherimola''<ref name="aft">{{cite web |url=http://www.worldagroforestry.org/Sea/Products/AFDbases/AF/asp/SpeciesInfo.asp?SpID=214 |archive-url=https://web.archive.org/web/20110526114153/http://www.worldagroforestry.org/Sea/Products/AFDbases/AF/asp/SpeciesInfo.asp?SpID=214 |archive-date=26 May 2011 |title=Current name: ''Annona squamosa'' |access-date=17 April 2008 |website=AgroForestryTree Database |publisher=International Center For Research In Agroforestry }}</ref> (whose fruits often share the same name)<ref name="GRIN">{{GRIN | access-date = 17 April 2008}}</ref> helping make it the most widely cultivated of these species.<ref name="purdue">{{cite web | url = http://www.hort.purdue.edu/newcrop/morton/sugar_apple.html | title = Sugar Apple ''Annona squamosa'' | access-date = 17 April 2008 | last = Morton | first = Julia | author-link = Julia Morton | year = 1987 | page = 69 | website = Fruits of warm climates | publisher = Department of Horticulture & Landscape Architecture, Purdue University | archive-url= https://web.archive.org/web/20080405173228/http://www.hort.purdue.edu/newcrop/morton/sugar_apple.html| archive-date= 5 April 2008 | url-status= live}}</ref>

''Annona squamosa'' is semi-(or late) deciduous,<ref name="FNA">{{cite book | last =Kral | first =Robert |editor=Flora of North America Editorial Committee | chapter =''Annona squamosa'' Linnaeus, Sp. Pl. 1: 537. 1753 | title =Flora of North America North of Mexico | volume =3 | chapter-url =http://beta.floranorthamerica.org/Annona_squamosa | access-date =5 September 2019}}</ref> and {{convert|3 to 8|m|ft|abbr=off|frac=2}} tall,<ref name="aluka" /><ref name="FNA" /> similar to soursop (''Annona muricata'').<ref name="PIER">{{cite web | url = http://www.hear.org/pier/species/annona_squamosa.htm | title = Annona squamosa | access-date = 17 April 2008 |website=Pacific Island Ecosystems at Risk (PIER) | date = 5 January 2008 | archive-url= https://web.archive.org/web/20080512181628/http://www.hear.org/pier/species/annona_squamosa.htm| archive-date= 12 May 2008 | url-status= live}}</ref> It is a native of tropical climate in the Americas, and Spanish traders aboard the Manila galleons docking in the Philippines brought it to Asia.<ref name="Morton">{{cite web|last=Morton|first=Julia|author-link=Julia Morton|title=Annona squamosa|url=http://www.hort.purdue.edu/newcrop/morton/sugar_apple.html|work=Fruits of warm climates|access-date=6 March 2013|page=69|year=1987}}</ref> [[File:Custard Apple - Flora Sinensis 1656 (2950756).jpg|thumb|237px|Michał Boym's drawing of, probably, the sugar-apple, in his ''Flora Sinensis'' (1655)]]

The fruit is spherical-conical, {{convert|5|-|10|cm|in|abbr=off|frac=4}} in diameter and {{convert|6|-|10|cm|in|abbr=on|frac=4}} long, and weighing {{convert|100|-|240|g|oz|abbr=off}}, 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 {{convert|13|to|16|mm|in|adj=mid|frac=8|-long}} 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.<ref name=Morton/><ref name=AFTDB>{{cite web|work=AgroForestryTree Database|title=''Annona squamosa''|url=http://www.worldagroforestrycentre.org/SEA/Products/AFDbases/AF/asp/SpeciesInfo.asp?SpID=214|access-date=16 September 2013|archive-url=https://web.archive.org/web/20070314161010/http://www.worldagroforestrycentre.org/SEA/Products/AFDbases/AF/asp/SpeciesInfo.asp?SpID=214|archive-date=14 March 2007}}</ref> The seeds can be ground for use as an insecticide, although this has not been approved by the US EPA or EU authorities.<ref name=":0" /> 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.

=== Stems and leaves === thumb|''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).<ref name="aft" />

Thin, simple, alternate leaves<ref name="PIER" /> occur singly,<ref name="aft" /> {{convert|5 to 17|cm|in|frac=4|abbr=off}} long and {{convert|2 to 6|cm|in|frac=8|abbr=on}} wide;<ref name="FNA" /><ref name="aft" /> rounded at the base and pointed at the tip (oblong-lanceolate).<ref name="FNA" /> They are pale green on both surfaces and mostly hairless<ref name="aft" /> with slight hairs on the underside when young.<ref name="aluka" /> The sides sometimes are slightly unequal and the leaf edges are without teeth, inconspicuously hairy when young.<ref name="aft" /><ref name="PIER" />

The leaf stalks are {{convert|0.4 to 2.2|cm|in|abbr=on|frac=8}} long,<ref name="FNA" /> green, and sparsely pubescent.<ref name="aft" />

=== Flowers === Solitary or in short lateral clusters of 2–4 about {{convert|2.5|cm|in|abbr=on|frac=4}} long,<ref name="FNA" /> greenish-yellow flowers on a hairy, slender<ref name="aft" /> {{convert|2|cm|in|abbr=on|frac=4}} long stalk.<ref name="FNA" /> Three green outer petals, purplish at the base, oblong, {{convert|1.6 to 2.5|cm|in|abbr=on|frac=8}} long, and {{convert|0.6 to 0.75|cm|in|abbr=on|frac=16}} wide, three inner petals reduced to minute scales or absent.<ref name="aluka" /><ref name="FNA" /> Very numerous stamens; crowded, white, less than {{convert|1.6|cm|in|abbr=on|frac=8}} long; ovary light green. Styles white, crowded on the raised axis. Each pistil forms a separate tubercle (small rounded wartlike protuberance), mostly {{convert|1.3 to 1.9|cm|in|abbr=on|frac=4}} long and {{convert|0.6 to 1.3|cm|in|abbr=on|frac=8}} wide which matures into the aggregate fruit.<ref name="aft" />

Flowering occurs in spring-early summer<ref name="FNA" /> and flowers are pollinated by nitidulid beetles.<ref>McGregor, S.E. [http://gears.tucson.ars.ag.gov/book/ Insect Pollination Of Cultivated Crop Plants] {{Webarchive|url=https://web.archive.org/web/20230326235745/http://gears.tucson.ars.ag.gov/book/ |date=26 March 2023 }} USDA, 1976</ref> Its pollen is shed as permanent tetrads.<ref>Walker JW (1971) Pollen Morphology, Phytogeography, and Phylogeny of the Annonaceae. Contributions from the Gray Herbarium of Harvard University, 202: 1-130.</ref>

=== Fruits and reproduction === Fruits ripen 3 to 4 months after flowering.<ref name="Grant2021">{{cite web |last1=Grant |first1=Amy |title=What Is Sugar Apple Fruit: Can You Grow Sugar Apples |url=https://www.gardeningknowhow.com/ornamental/trees/annona/sugar-apple-trees.htm |website=Gardening Know How |access-date=4 August 2023 |date=2021}}</ref> Aggregate and soft fruits form from the numerous and loosely united pistils of a flower<ref name="aft" /> which become enlarged<ref name="FNA" /> and mature into fruits which are distinct from fruits of other species of genus<ref name="aft" /> (and more like a giant raspberry).

The round or heart-shaped<ref name="aft" /> greenish yellow, ripened aggregate fruit is pendulous<ref name="FNA" /> on a thickened stalk; {{convert|5 to 10|cm|in|frac=8|abbr=on}}<ref name="aft" /><ref name="aluka" /> in diameter<ref name="FNA" /><ref name="PIER" /> with many round protuberances<ref name="aft" /> and covered with a powdery bloom. Fruits are formed of loosely cohering or almost free carpels (the ripened pistels).<ref name="aluka" />

The pulp is white tinged yellow,<ref name="aluka" /> edible and sweetly aromatic. Each carpel containing an oblong, shiny and smooth,<ref name="aft" /> dark brown<ref name="aluka" /> to black, {{convert|1.3 to 1.6|cm|in|abbr=on|frac=8}} long seed.<ref name="aft" />

==Nutrition and uses== {{nutritional value | name=Sugar-apples, (sweetsop), raw | kJ=393 | protein=2.06 g | fat=0.29 g | carbs=23.64 g | fiber=4.4 g | calcium_mg=24 | iron_mg=0.6 | magnesium_mg=21 | phosphorus_mg=32 | potassium_mg=247 | sodium_mg=9 | zinc_mg=0.1 | manganese_mg=0.42 | vitC_mg=36.3 | thiamin_mg=0.11 | riboflavin_mg=0.113 | niacin_mg=0.883 | pantothenic_mg=0.226 | vitB6_mg=0.2 | folate_ug=14 | source_usda = 1 | note=[https://web.archive.org/web/20160309144328/https://ndb.nal.usda.gov/ndb/search/list?qlookup=09321&format=Full Link to USDA Database entry] }} Sugar-apple is high in energy, an excellent source of vitamin C and manganese, a good source of vitamin B1 and vitamin B6, and provides vitamin B2, B3 B5, B9, iron, magnesium, phosphorus and potassium in fair quantities.{{citation needed|date=April 2026}}

=== Chemistry === The diterpenoid alkaloid atisine is the most abundant alkaloid in the root. Other constituents of ''Annona squamosa'' include the alkaloids oxophoebine,<ref name="pmid23678817">{{cite journal |author=Dholvitayakhun A, Trachoo N |display-authors=etal |year=2013 |title=Potential applications for ''Annona squamosa'' leaf extract in the treatment and prevention of foodborne bacterial disease |journal=Natural Product Communications |volume=8 |issue=3 |pages=385–388 |doi=10.1177/1934578X1300800327 |pmid=23678817 |doi-access=free}}</ref> reticuline,<ref name="pmid23678817" /> isocorydine,<ref name="pmid21342663">{{cite journal |author=Yadav DK, Singh N |display-authors=etal |year=2011 |title=Anti-ulcer constituents of ''Annona squamosa'' twigs |journal=Fitoterapia |volume=82 |issue=4 |pages=666–675 |doi=10.1016/j.fitote.2011.02.005 |pmid=21342663}}</ref> and methylcorydaldine,<ref name="pmid21342663" /> and the flavonoid quercetin-3-O-glucoside.<ref name="pmid18997283">{{cite journal |author=Panda S, Kar A |year=2007 |title=Antidiabetic and antioxidative effects of ''Annona squamosa'' leaves are possibly mediated through quercetin-3-O-glucoside |journal=BioFactors |volume=31 |issue=3–4 |pages=201–210 |doi=10.1002/biof.5520310307 |pmid=18997283 |s2cid=38336427}}</ref>

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.<ref>{{Cite web |author=Moeschler HF, Pfluger W |display-authors=etal |date=August 1987 |title=Insecticide US 4689232 A |url=https://patents.google.com/patent/US4689232 |access-date=3 December 2014}}</ref> Other acetogenins have been isolated from the seeds,<ref name="pmid22609808">{{cite journal |author=Chen Y, Xu SS |display-authors=etal |year=2012 |title=Anti-tumor activity of ''Annona squamosa'' seeds extract containing annonaceous acetogenin compounds |journal=Journal of Ethnopharmacology |volume=142 |issue=2 |pages=462–466 |doi=10.1016/j.jep.2012.05.019 |pmid=22609808}}</ref> bark,<ref name="pmid2348205">{{cite journal |author=Li XH, Hui YH |display-authors=etal |year=1990 |title=Bullatacin, bullatacinone, and squamone, a new bioactive acetogenin, from the bark of ''Annona squamosa'' |journal=Journal of Natural Products |volume=53 |issue=1 |pages=81–86 |doi=10.1021/np50067a010 |pmid=2348205}}</ref> and leaves.{{citation needed|date=December 2019}}

==Distribution and habitat== ''Annona squamosa'' is native to the tropical Americas, 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;<ref>{{Cite web|url=https://inaturalist.ca/observations/37152727|title = Sweetsop (Annona squamosa)|date = January 2020}}</ref> it was introduced to southern Asia before 1590. It is naturalized as far north as Cuba, south Florida and as far south as Bahia, Brazil, and is an invasive species in some areas.<ref name="aft"/><ref name=purdue/><ref name="PIER"/>

;<nowiki>Native range:</nowiki><ref name=":1"/> :'''Neotropic''' ::'''North America:''' Mexico ::'''Central America''': Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama ::'''South America:''' Colombia

;<nowiki>Naturalised range:</nowiki><ref name=":1" /> ::'''Pacific''': Samoa, Tonga ::'''North America''': United States (Florida) ::'''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 ::'''South America''': Suriname, French Guiana, Guyana, Venezuela, Bolivia, Ecuador, Peru, Argentina, Brazil, Chile, Paraguay, Uruguay<ref name="aft" /> ::'''Afrotropic''': Angola, Namibia, Saudi Arabia, Sudan, Tanzania, Uganda, Yemen ::'''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,<ref name="aft"/> Israel

==Climate and cultivation== {{More citations needed section|date=February 2021}} Like most species of ''Annona'', it requires a tropical or subtropical climate with summer temperatures from {{convert|25|°C|°F}} to {{convert|41|°C|°F}}, and mean winter temperatures above {{convert|15|°C|°F}}. It is sensitive to cold and frost, being defoliated below {{convert|10|C|F}} and killed by temperatures of a couple of degrees below freezing. It is only moderately drought-tolerant, requiring at least {{Convert|700|mm}} of annual rainfall, and does not produce fruit well during droughts.

It will grow from sea level to an altitude of {{convert|2000|m|ft|abbr=off}} 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.<ref name=purdue/><ref name=AFTDB/>

==Ecology== It is a host plant for larvae of the butterfly ''Graphium agamemnon'' (tailed jay). In the Philippines, the fruit is commonly eaten by the Philippine fruit bat (''kabag'' or ''kabog''), which spreads the seeds from island to island.

==Uses== 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.<ref name=" doi10.1016/j.hermed.2016.10.003">{{cite journal| author=Dholvitayakhun A, Trachoo N| title=Using scanning and transmission electron microscopy to investigate the antibacterial mechanism of action of the medicinal plant ''Annona squamosa'' Linn | journal=Journal of Herbal Medicine | year= 2016| volume= 7| pages= 31–36|doi =10.1016/j.hermed.2016.10.003|display-authors=etal| url=https://zenodo.org/record/884833 }}</ref> In traditional Indian medicine, the leaves are also crushed for use as a poultice, and applied to wounds.<ref name=" doi10.1016/j.hermed.2016.10.003" /> In Mexico, the leaves are rubbed on floors and put in hens' nests, to repel lice.<ref name=purdue/> Extracts are used in ethnomedicine.<ref>{{cite journal |last1=Bisht |first1=Manohar S. |last2=Mahajan |first2=Shruti |last3=Chakraborty |first3=Abhisek |last4=Sharma |first4=Vineet K. |title=A high-quality genome assembly of Annona squamosa (custard apple) provides functional insights into an emerging fruit crop |journal=DNA Research |date=June 2025 |volume=32 |issue=3 |doi=10.1093/dnares/dsaf007|pmc=12116420 }}</ref> In Haiti, the fruit is known as ''cachiman'' and is used to simply make juice.<ref>{{cite web |title=Cachiman (Annona reticulata L.) |url=http://caribfruits.cirad.fr/fruits_tropicaux/cachiman |website=Carib Fruits |access-date=8 November 2020}}</ref>

==Gallery== <gallery> File:Annona squamosa fruit from Myanmar.jpg|Annona squamosa fruit from Myanmar File:Pineapple shijia.jpg|Sugar apple (right), with Taiwanese "pineapple ''shijia''" (atemoya) (left) File:Sugar Apple (Annona squamosa) interior.jpg|The sugar apple readily breaks open when ripe. File:Sugar Apple deconstructed.jpg|A deconstruction of a sugar apple shows a lobe of fruit and pulpy segments with seeds. File:Sugar Apple pulp.jpg|A sugar apple ready to eat File:Sugar-apple (Annona squamosa) seeds.jpg|Sugar apple (''Annona squamosa'') seeds File:Red Sugar Apple.jpg|Red sugar apples from Myanmar File:Sugar-apples 5, Taitung County, Dec 06.JPG|Sugar apples in Taitung, Taiwan File:Ata Sugar-apple Pinha Fruta do conde.JPG|A sugar apple in Goiânia, Brazil File:Sugar Apple Tree.jpg|Sugar apple tree in Philippines File:Aata by Mayeenul Islam.jpg|Two sugar apples in Bangladesh File:Sarifa.jpg|A sugar apple in tree in Terai of Nepal File:Squa9.jpg|A Sugar apple in a table </gallery>

==References== {{reflist|2}}

==External links== {{Commons category|Annona squamosa}} {{Wikispecies}} {{EB1911 poster|Sweet-sop}} * {{ITIS |id=18100 |taxon=''Annona squamosa'' L. |access-date=17 March 2008}} *{{cite web | url = http://bio.fiu.edu/trees/sp_pages/Annona_squamosa.html | title = Photographs of trees ''Annona squamosa'' | access-date = 17 April 2008 | author = David Lee | author-link = Florida International University | website = The Miami Tree Puzzle | publisher = Florida International University | archive-url = https://web.archive.org/web/20080407072812/http://bio.fiu.edu/trees/sp_pages/Annona_squamosa.html | archive-date = 7 April 2008 }} * [https://web.archive.org/web/20140517120301/http://libproject.hkbu.edu.hk/was40/detail?lang=en&channelid=1288&searchword=herb_id%3DD01306 ''Annona squamosa'' L.]—Medicinal Plant Images Database (School of Chinese Medicine, Hong Kong Baptist University) {{in lang|zh-hant|en}} * [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200008509 Flora of North America: ''Annona squamosa''] * {{GRIN | ''Annona squamosa'' | 3503}} * [https://web.archive.org/web/20060927224602/http://www.ciat.cgiar.org/ipgri/fruits_from_americas/frutales/Ficha%20Annona%20squamosa.htm Fruits from Americas: ''Annona squamosa''] * [http://www.hear.org/pier/species/annona_squamosa.htm Pacific Island Ecosystems at Risk: ''Annona squamosa''] * [http://balconygardenweb.com/how-to-grow-sugar-apple-growing-sugar-apple-care/ Growing Sugar Apple ''Annona squamosa'']

squamosa Category:Crops originating from South America Category:Crops originating from the Americas Category:Edible fruits Category:Flora of Central America Category:Flora of Colombia Category:Flora of Mexico Category:Trees of Guatemala Category:Trees of the Caribbean Category:Least concern flora of Central America Category:Least concern flora of Mexico Category:Least concern flora of South America Category:Plants described in 1753 Category:Botanical taxa named by Carl Linnaeus