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Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Cladus: Lamiids
Ordo: Solanales

Familia: Solanaceae
Subfamilia: Solanoideae
Tribus: Physaleae
Subtribus: Physalidinae
Genus: Physalis

Species: P. acutifolia – P. aggregata – P. ampla – P. angulata – P. angustifolia – P. angustior – P. angustiphysa – P. arborescens – P. arenicola – P. campanula – P. campechiana – P. carnosa – P. caudella – P. chenopodiifolia – P. cinerascens – P. cinerea – P. cordata – P. coztomatl – P. crassifolia – P. eggersii – P. fendleri – P. filipendula – P. flava – P. galapagoensis – P. glabra – P. glutinosa – P. gracilis – P. greenei – P. greenmanii – P. grisea – P. halicacabum – P. hastatula – P. hederifolia – P. heterophylla – P. hintonii – P. hirsuta – P. hispida – P. hunzikeriana – P. ignota – P. ingrata – P. integrifolia – P. ixocarpa – P. jaliscensis – P. lagascae – P. lanceolata – P. lassa – P. latecorollata – P. leptophylla – P. lignescens – P. longicaulis – P. longifolia – P. longiloba – P. longipedicellata – P. macrosperma – P. mcvaughii – P. melanocystis – P. michoacanensis – P. microcarpa – P. microphysa – P. mimulus – P. minimaculata – P. minuta – P. missouriensis – P. mollis – P. muelleri – P. muriculata – P. neomexicana – P. nicandroides – P. orizabae – P. parvianthera – P. patula – P. pennellii – P. peruviana – P. philadelphica – P. philippensis – P. porphyrophysa – P. porrecta – P. pringlei – P. pruinosa – P. pubescens – P. pumila – P. purpurea – P. queretaroensis – P. quillabambensis – P. rydbergii – P. sancti-josephi – P. solanacea – P. sordida – P. spathulifolia – P. stapelioides – P. subilsiana – P. subrepens – P. sulphurea – P. tamayoi – P. tehuacanensis – P. texana – P. turbinatoides – P. vestita – P. victoriana – P. virginiana – P. viscosa – P. volubilis – P. walteri

Nothospecies: P. × elliottii
Name

Physalis L., Sp. Pl. 1: 182. (1753) nom. et typ. cons.

Type species: Physalis pubescens L., Sp. Pl. 1: 183. (1753)

Synonyms

Heterotypic
Alicabon Raf.
Alkekengi Tourn. (1763) [[nom. illeg. hom.]] non Alkekengi Mill., Gard. Dict. Abr., ed. 4. [unpaged] (1754)
Boberella E.H.L.Krause
Epetorhiza Steud.
Margaranthus Schltdl.
Pentaphiltrum Rchb.

References

Linnaeus, C. 1753. Species Plantarum. Tomus I: 182. Reference page.
Applequist, W.L. 2012. Report of the Nomenclature Committee for Vascular Plants 64. Taxon 61(5): 1108–1117. DOI: 10.1002/tax.615019 Open access Reference page.
Govaerts, R. et al. 2020. Physalis in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2020 Oct. 31. Reference page.
Physalis – Taxon details on Integrated Taxonomic Information System (ITIS).
Tropicos.org 2020. Physalis. Missouri Botanical Garden. Published online. Accessed: 31 Oct. 2020.
Whitson, M. 2011. (2016) Proposal to conserve the name Physalis (Solanaceae) with a conserved type. Taxon 60(2): 608-609. PDF Reference page.

Vernacular names
беларуская: Фізаліс
English: Ground Cherries
suomi: Lyhtykoisot
日本語: ホオズキ属
македонски: Зрнавец, плускавец
português: camapu, juapoca

Physalis (/ˈfaɪsəlɪs/, /fɪ-/, /faɪˈseɪlɪs/, /-ˈsæ-/, from φυσαλλίς phusallís "bladder"[2]) is a genus of flowering plants in the nightshade family (Solanaceae), which grow in warm temperate and subtropical regions of the world. Most of the species, of which there may be 75 to 90, are indigenous to the Americas. Cultivated species and weedy annuals have been introduced worldwide. A notable feature is the formation of a large, papery husk derived from the calyx, which partly or fully encloses the fruit.[3] Some species bear edible fruit, which is small and yellow to orange, similar in size, shape, and structure to a small tomato (hence the name husk tomatoes). They are also grown as ornamental plants, and used in Chinese medicine.

At least 46 species are endemic to Mexico.[4]

Many Physalis species are called groundcherries.[5] One name for Physalis peruviana is Inca berry; another is Cape gooseberry, though unrelated to gooseberries of the genus Ribes (family Grossulariaceae). Other names used to refer to the fruit are husk cherries, poha berries, and golden berries.[6]

Description

Physalis species are herbaceous plants growing to 0.4 to 3.0 m tall, similar to the common tomato, a plant of the same family, but usually with a stiffer, more upright stem. They can be either annual or perennial. Most require full sun and fairly warm to hot temperatures. Some species are sensitive to frost, but others, such as the Chinese lantern, P. alkekengi, tolerate severe cold when dormant in winter.
Cultivation and uses
Physalis peruviana fruit with calix open

These plants grow in most soil types and do very well in poor soils and in pots. They require moisture until fruiting. Plants are susceptible to many of the common tomato diseases and pests, and other pests such as aphids, whiteflies, spider mites, and the false potato beetle (Leptinotarsa juncta) also attack them. Propagation is by seed. Some species are self-incompatible and require pollen from other plants to bear fruit.

Some but not all Physalis species bear edible fruit. Select species are cultivated for their edible fruit; the typical Physalis fruit is similar to a firm tomato in texture, and like strawberries or pineapple in flavor, with a mild acidity. Some species, such as the Cape gooseberry and tomatillo, have been bred into many cultivars with varying flavors, from tart to sweet to savory. Physalis fruit are rich in cryptoxanthin. The fruit can be used like the tomato. Once extracted from its husk, it can be eaten raw[7] and used in salads. Some varieties are added to desserts, used as flavoring, made into fruit preserves, or dried and used like raisins. They contain pectin and can be used in pie filling. Ground cherries are called poha in the Hawaiian language, and poha jam and preserves are traditional desserts made from Physalis plants grown on the Hawaiian Islands.[8]

The Cape gooseberry is native to the Americas, but is common in many subtropical areas. Its use in South Africa near the Cape of Good Hope inspired its common name. Other species of commercial importance include the tomatillo (P. philadelphica). Some nations, such as Colombia, have a significant economic trade in Physalis fruit. Physalis spp. are widely cultivated in India.

Some species are grown as ornamental plants. For example, the hardy Physalis alkekengi has edible small fruits, but is most popular for its large, bright orange to red husks.

In Chinese medicine, Physalis species are used to treat such conditions as abscesses, coughs, fevers, and sore throat.[9] Smooth groundcherry (P. subglabrata) is classified (erroneously) as a hallucinogenic plant, and its cultivation for other than ornamental purposes is outlawed in the US state of Louisiana under State Act 159 of 2005.

The extinct Dacian language has left few traces, but in De Materia Medica by Pedanius Dioscorides, a plant called Strychnos alikakabos (Στρύχνος άλικακάβος) is discussed, which was called kykolis (or cycolis) by the Dacians. Some have considered this plant to be P. alkekengi, but the name more likely refers to ashwagandha (Withania somnifera).[10]
Diversity
Yellow nightshade groundcherry (Physalis crassifolia)
Physalis peruviana fruits

As of 2005, about 75 to 90 species were placed in the genus.[3]

Species include:[5][11]

Physalis acutifolia (Miers) Sandw. – sharp-leaved groundcherry, Wright groundcherry
Physalis alkekengi L. – Chinese lantern, Japanese lantern, bladder-cherry, winter-cherry, hōzuki (Japanese)
Physalis angulata L. – cut-leaved groundcherry, lance-leaved groundcherry, camapu
Physalis angustifolia Nutt. – coastal groundcherry
Physalis arenicola Kearney – cypress-headed groundcherry
Physalis carpenteri Riddell ex Rydb. – Carpenter's groundcherry
Physalis caudella Standl. – southwestern groundcherry
Physalis chenopodifolia
Physalis cinerascens (Dunal) A.S. Hitchc. – small-flowered groundcherry
Physalis clarionensis
Physalis cordata Mill. – heart-leaved groundcherry
Physalis coztomatl Moc. & Sessé ex Dunal
Physalis crassifolia Benth. – thick-leaved groundcherry, yellow nightshade groundcherry
Physalis foetens Poir. – tropical groundcherry
Physalis grisea (Waterfall) Martínez – strawberry-tomato
Physalis hederifolia A.Gray – ivy-leaved groundcherry
Physalis heterophylla Nees – clammy groundcherry
Physalis hispida (Waterfall) Cronq. – prairie groundcherry
†Physalis infinemundi Wilf et al. 2017 fossil from the Ypresian of Argentina[12]
Physalis ixocarpa
Physalis latiphysa Waterfall – broad-leaved groundcherry
Physalis longifolia Nutt. – common groundcherry, long-leaved groundcherry
Physalis longiloba[4]
Physalis mimulus
Physalis minima L. – pygmy groundcherry, native gooseberry (Australia)
Physalis missouriensis Mackenzie & Bush – Missouri groundcherry
Physalis mollis Nutt. – field groundcherry
Physalis noronhae
Physalis peruviana L. – Cape gooseberry, Peruvian groundcherry, Inca berry, uchuva (Colombia), poha
Physalis philadelphica Lam. (syn. P. ixocarpa) – tomatillo, Mexican groundcherry, jamberry, Mexican tomato, tomate de cáscara, tomate de fresadilla, tomate milpero, tomate verde

Strawberry groundcherry (Physalis pruinosa)

Physalis pruinosa L. – strawberry groundcherry
Physalis pubescens L. – golden strawberry, Chinese lantern
Physalis pumila Nutt. – dwarf groundcherry
Physalis subulata Rydb. – Chihuahuan groundcherry
Physalis tamayoi[4]
Physalis turbinata Medik. – thicket groundcherry
Physalis virginiana Mill. – Virginia groundcherry
Physalis viscosa L. – grape groundcherry, star-haired groundcherry
Physalis walteri Nutt. – Walter's groundcherry

Walter's groundcherry plant with fruit at Honeymoon Island State Park, Florida

Formerly placed here

Deprea orinocensis (Kunth) Raf. (as P. orinocensis Kunth)
Leucophysalis grandiflora (Hook.) Rydb. (as P. grandiflora Hook.)
Quincula lobata (Torr.) Raf. (as P. lobata Torr.)[11]
Salpichroa origanifolia (Lam.) Baill. (as P. origanifolia Lam.)
Withania somnifera (L.) Dunal (as P. somnifera L.)[11]

Fossil record

A 52 million year old fossil fruit of Physalis has been found in Patagonia.[13][14]
References

"Genus: Physalis L." Germplasm Resources Information Network. United States Department of Agriculture. 2009-09-01. Archived from the original on 2010-05-29. Retrieved 2010-04-14.
"Physalis | Definition of physalis in English by Oxford Dictionaries".
Whitson, M.; Manos, P. S. (2005). "Untangling Physalis (Solanaceae) from the physaloids: a two-gene phylogeny of the Physalinae". Systematic Botany. 30 (1): 216–30. doi:10.1600/0363644053661841. JSTOR 25064051. S2CID 86411770.
Vargas, O.; et al. (2001). "Two new species of Physalis (Solanaceae) endemic to Jalisco, Mexico". Brittonia. 53 (4): 505–10. doi:10.1007/bf02809650. S2CID 11564.
"Physalis". Integrated Taxonomic Information System. Retrieved 2011-05-21.
Doctor, Vikram (4 March 2013). "Golden berry: Decoding the acid freshness and wild sweet taste of physalis". The Economic Times. Retrieved 6 Sep 2014.
Angier, Bradford (1974). Field Guide to Edible Wild Plants. Harrisburg, PA: Stackpole Books. p. 90. ISBN 0-8117-0616-8. OCLC 799792.
Gibbons, Euell (1962). Stalking the Wild Asparagus. Chambersburg, Pennsylvania: Alan C. Hood & Company, Inc. p. 104. ISBN 978-0-911469-03-5.
Duke, J. A.; Ayensu, E. S (1985). Reference Publications, Inc. (ed.). Medicinal Plants of China. ISBN 978-0-917256-20-2. Retrieved 2009-05-15.
Berendes, J. (ed.) Arzneimittellehre in fünf Büchern des Pedanios Dioskurides aus Anazarbos. Stuttgart. 1902. 405-08.
"GRIN Species Records of Physalis". Germplasm Resources Information Network. United States Department of Agriculture. Archived from the original on 2008-10-05. Retrieved 2011-05-21.
Switek, Brian. "Paleo Profile: Tomatillo from the End of the World".
bbc.com/news/science-environment-38511034
Wilf, Peter (6 Jan 2017). "Eocene lantern fruits from Gondwanan Patagonia and the early origins of Solanaceae". Science. 355 (6320): 71–75 – via Science.

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