Physalis L.

Arenas, Pastor & Kamienkowski, Nicolás Martín, 2013, Ethnobotany of the genus Physalis L. (Solanaceae) in the South American Gran Chaco, Candollea 68 (2), pp. 251-266 : 252-258

Physalis L. : taxonomy, biogeography, ecology

The genus Physalis comprises about 90 species, most of them native to America, excepting P. alkekengi L. which is native to the Old World ( HUNZIKER, 2001; TOLEDO & BARBOZA, 2005: 69). Physalis is a clearly defined genus, in the tribe Solaneae , and has a distinctive fruit. This is a globose two-carpelate berry, small or large (4-7 / 10-20 mm diameter) with an either juicy or rather dry pericarp. The berry is loosely enclosed by the accrescent bladdery inflated calyx. The seeds are usually numerous (around 50, 100 or 180 per fruit) but this number tends to diminish as the flowering season advances, often to 5-6 seeds ( MARTÍNEZ, 1998: 76; HUNZIKER, 2001: 204).

The species are low annual or perennials herbs and shrubby or arborescent perennials ( MARTÍNEZ, 1998: 73; HUNZIKER, 2001: 202-204). The centre of diversity of Physalis is Mexico with over 70 species, most of which are endemic; two other centres of diversity are United States and Central America, also with endemic species ( MARTÍNEZ, 1998: 72; HUNZIKER, 2001: 207). In South America barely 12 species live, some of them are also endemic ( MARTÍNEZ, 1998: 72; HUNZIKER, 2001: 207).

A few species are cultivated in the temperate and tropical Old World and in Australia, while others (mainly P. peruviana L. and P. angulata L.) are ruderal plants or weeds ( NATIONAL RESEARCH COUNCIL, 1989: 240-251; HUNZIKER, 2001: 207). These plants were introduced into warm areas of the world in post-Columbian times, with the voyages of exploration, discovery and commercial exploitation that began in the 16th century ( HEDRICK, 1919).

On the basis of voucher specimens available from the Gran Chaco and bibliographical data, the genus Physalis is represented by 6 taxa in this region ( TOLEDO & BARBOZA, 2005). In the Gran Chaco, Physalis flowers in spring and summer, and its fructification extends simultaneously until autumn. Occasionally, the fruits are also available in winter, when wild foods are scarce, particularly in the xerophytic parts of the region, where there is a limited amount of fresh food available for both people and animals. The different species of Physalis of the Gran Chaco grow in a diversity of environments and conditions, such as clearings and the edges of forests, xerophytic scrubland, and thus tend to invade crops, roadside verges, natural grasslands, sandy beaches along streams and rivers, land which is prone to flooding and the herbaceous stratum of gallery forests ( FIEBRIG & ROJAS, 1933: 51).

Ethnobotany of Physalis

The ethnobotanical information is presented in two sections: the first contains a basic review of the literature on the species that live in other geographical regions, particularly in America, while the second section brings together specific data on the Gran Chaco and its neighbouring zones of influence. They will thus provide comparative elements to highlight the role of this genus of plants for the people of the Chaco. The bibliographic review does not seek to be exhaustive but aims to provide a prospective vision that allows general comparisons and conclusions.

Physalis : a general review

In works of a general nature that deal with the subject of useful plants, several authors have mentioned the nutritional use of the fruits of various species in different regions, indicating their area of origin, and whether they are consumed raw, or in sauces, compotes, pies, jams or relishes ( HEDRICK, 1919: 431-433; BOIS, 1928: 367-369; BAILEY, 1938: 657-658; KÜNKEL, 1984: 275; FACCIOLA, 1990: 206-207). They also mention the consumption of the leaves of P. angulata in salads ( HEDRICK, 1919: 431; KÜNKEL, 1984: 275).

Herbarium labels usually provide additional data recorded by collectors which are of ethnobotanical interest. Reviews of specimens of the Harvard University Herbarium and the NYBG collections provide information on plants collected in the Neotropics, recording vernacular names, uses and other curiosities. In the case of Physalis , the edible and medicinal uses prevailed ( REIS ALTSCHUL, 1973: 269; REIS & LIPP, 1982: 270).

The Eurasian species P. alkekengi is used as an ornamental plant and is also mentioned as medicinal and edible, although it has been mentioned that its taste is rather unpleasant and it may be toxic when unripe ( BOIS, 1928: 366; BAILEY, 1938: 657; UPHOF, 1959: 277; ROMO, 1996: 67). This plant is now domesticated and sold all around Europe in fruit markets.

In North America, several edible species are mentioned. Their fruits are consumed raw or boiled in various culinary preparations among indigenous groups in the region ( YANOVSKY, 1936: 56; UPHOF, 1959: 277; MOERMAN, 1998: 395-396). MOERMAN (1998: 395, 396; 2010: 180, 181) reports several medicinal qualities attributed to 6 species and the edible use of 10 species. In Mexico various representatives of the genus with edible fruits and medicinal applications among the indigenous peoples and the “mestizo” population are also mentioned ( CABALLERO & al., 1978: 123; CABALLERO & MAPES, 1985: 35, 45-46; WILLIAMS, 1985: 95-127; CASAS & al., 1987: 331; MARTÍNEZ ALFARO & al., 1995: 238-240; GISPERT CRUELLS & RODRÍGUEZ GONZÁLEZ, 1998: 50-51).

In Mexico and in the Andean region, some species of the genus have been cultivated for their edible fruits since ancient times ( HEISER, 1984: 50). The original species of Mesoamerica P. philadelphica Lam. has been cultivated in Mexico and Guatemala from immemorial time, to which archaeological deposits bear witness. Domesticated in Mexico, it was taken to Spain and other parts of the world ( MONTES HERNÁNDEZ & AGUIRRE RIVERA, 1992). The fruits of P. peruviana were in use in Ancient Peru, in pre- Columbian times. This plant spread through the Old World after the Conquest of America as well as to other warm regions of America, extending southwards to the River Plate region, as well as to Africa and Australia ( BOIS, 1928: 367- 368; LEÓN, 1964: 109-110; MATHON, 1981: 103; BALBIN ORDAYA, 1982: 6; HEISER, 1984: 50; MONTES HERNÁNDEZ & AGUIRRE RIVERA, 1992: 115; HURRELL & al., 2010: 236). Other species cultivated, albeit in a more restricted form, are P. grisea (Waterf.) M. Martínez from the eastern United States, well appreciated for its juicy fruits ( MARTÍNEZ, 1998: 72), and P. ixocarpa Hornem. , considered of excellent nutritional quality and grown in orchards in Mexico and Central America ( LEÓN, 1964: 110; MANGELSDORF & al., 1964: 435; MATHON, 1981: 96; KATZ, 1990: 265).

There are numerous reports on around ten South American species, as well as others which have spread throughout America. For many of them, edible and medicinal uses are reported as well as several other applications ( UPHOF, 1959: 277). In several countries ( Bolivia, Chile, Peru, Colombia, Nicaragua and Venezuela) mention is made of species of Physalis with edible fruits while other parts of the plant are used as medicines ( SOUKUP, 1970: 261; GARCÍA BARRIGA, 1975: 81-84; SECAB, 1983: 186; CÁRDENAS, 1989: 133; MÖSBACH, 1991: 104; COE & ANDERSON, 1996: 105). Information on medicinal and food uses are also reported in the Amazonia ( Brazil, Ecuador and Colombia) ( GLENBOSKI, 1983: 56; VICKERS & PLOWMAN, 1984: 31; ALBERT & MILLIKEN, 2009: 56, 180). Similar records can be found in other regions in Brazil ( CORRÊA, 1926: 408- 409; DI STASI & al., 1989: 45-46; SOUZA & al., 2003: 100). Similarly, there are references concerning the edible and medicinal use of various species by different human groups in Bolivia ( GIRAULT, 1987: 382; CÁRDENAS, 1989: 13; MOSTACEDO & USLAR, 1999: 22; BOURDY, 2002: 147-149). The nutritional and medicinal use of P. viscosa L. in eastern Paraguay is also widespread ( MICHALOWSKI, 1955: 12; GATTI, 1985: 58, 59; PAVETTI & al., 1985: c.33; PIN & al., 2009: 152). Similar data are repeated in the north of Argentina, particularly in the provinces bordering on the Gran Chaco, where P. viscosa is mentioned as a food and medicine, with limited mentions of other representatives of the genus ( HIERONYMUS, 1929: 203- 204; RAGONESE & MARTÍNEZ CROVETTO, 1947: 204; PARODI, 1886: 34; SCHULZ, 1963: 61-62; MARTÍNEZ CROVETTO, 1981: 99). Physalis pruinosa L. is a weed in the north of Chile and was used in past decades as a medicine (MÖBASCH, 1991: 104).

There is a body of information on the phytochemical components and the pharmacological activity of various species, which in some way would appear to validate the medicinal uses assigned to them. Several authors provide references on this subject, and also include other references to provide greater depth in the matter ( DI STASI & al., 1989; SCHULTES & RAFFAUF, 1990: 436; HUNZIKER, 2001: 207, 208; PIN & al., 2009: 152). Given the characteristics of its clearly ethnobotanical approach, this paper will offer no further information on this phytopharmaceutical point.

Vernacular names

There are lists of vernacular names assigned to species of Physalis in works of a floristic nature, or in works on the subject of useful plants, However, they have compiled names that in many cases give no specification as to where they are applied or by whom, nor to which species they refer ( CORRÊA, 1926: 408-409). This frequently happens in the literature of the Southern Cone of America, although the names are usually restricted to P. viscosa , the most common species in the region (BONDENBENDER, 1941: 18; MARZOCCA, 1957: 338; XIFREDA, 1992: 44; HURRELL & al., 2010: 236). MARTÍNEZ (1998) condenses a long, very detailed list of names with indications as to the places where they are applied, giving special importance to those used in Mexico and Central America. Similar care has been taken to present the vernacular nomenclature of P. peruviana by the compilers of the work that deals with the lost crops of the Incas ( NATIONAL RESEARCH COUNCIL, 1989: 249- 250). Indigenous names from the Gran Chaco are given in Appendix 2, which includes data from published works on ethnic groups that were not studied by the authors.

In order to give the Criollo names or those given by the rural population greater precision we organized the vernacular nomenclature according to these human and regional groups. These generic vernacular names are also applied to various species of Physalis with no distinction. Among the Spanishspeaking population in Argentina the plants receive different names: “uvilla”, “uvilla del campo”, “uvilla camambú”, “pocote”, “pocote de víbora”, “pocote de perro” and “meloncillo” ( HIERONYMUS, 1929: 203; BODENBENDER, 1941: 18; ACUÑA, 1945: 23; SCHULZ, 1976: 42; SCARPA, 2000: 265; BIURRUN & al., 2007: 126; TORRES & al., 2007: 163, 166, 180). In Paraguay, where the population is bilingual in Spanish and Guarani, the name generally applied to the different species is the Guarani term “kamambu” (FIEBRIG- GERTZ, 1923:119; CADOGAN, 1957: 31). This designation is also applied in the Argentine regions bordering on Paraguay (such as the provinces of Misiones, Corrientes, Formosa, Chaco and Salta) as well as in the east of Bolivia ( FRANZÉ, 1925: 14; RAGONESE & MARTÍNEZ CROVETTO, 1947: 204; SCHULZ, 1963: 61-62; BOURDY, 2002: 147-149). BERTONI (1980: 22, 43) offers it as a generic word for Physalis and gives in addition the phonetic variant “kamapú, ” possibly more linked to the Tupi language, since it is the word that is habitually reported for Brazil ( TASTEVIN, 1923: 16; CORRÊA, 1926: 408; GONZÁLEZ TORRES, 1981: 226; DI STASI & al., 1989: 45; DA MATTA, 2003: 87). “Kamambu” means “blister, bladder formed by elevation of the epidermis, bubble”; by extension, or similarity, it is applied to those inflated fruits such as the species of Physalis and those of some Sapindaceae ( GATTI & al., 1947: 37; GUASCH, 1981: 569; GATTI, 1985: 58-59). It should once again be stressed that the Tupi Guarani-speaking peoples are refined observers of nature and the vernacular generic name coincides with the generic name given by academic science. In fact, Physalis comes from the Greek physa which means bladder ( BAILEY, 1938: 657; SOUKUP, 1970: 261; PARODI, 1980: 945). The name “kamambu’i” (the particle “i” is a diminutive that means “small”) is usually given to P. viscosa , because of its smaller size ( HASSLER, 1909: 145; BODENBENDER, 1941: 18; SCHULZ, 1976: 42; PIN & al., 2009: 152).

The vernacular nomenclature and the classification systems of the hunter-gatherer peoples are of great interest because they contain an invaluable body of observations on nature and culture. Very little has been investigated on this topic among the indigenous peoples of the Gran Chaco. Regarding Physalis it can be seen that there are primary names and also compound names. The primary names probably suggest certain meanings, although we have little information in that respect. Such is the case of the “qotoñí” (Toba-Pilagá) or “makani” (Maká) names given to Physalis . As for the compound and descriptive names we resort to the phytonymy of the Wichí people, who give Physalis the following names: “wahat te’lhui” (= sábalo eyes; sábalo, a fish), “wuq’ute lhui” (= owl eyes), “wi’yes te’lhui” (= cavy eyes; cavy, a small rodent). With such descriptive names they compare the fruits surrounded by the calyx with the morphology of the eyes of the animals mentioned. The name “p’oh p’oh” is an onomatopoeic, primary name, and alludes to the explosion of the closed accrescent calyx produced when it is crushed or struck ( MARANTA, 1987: 186; ARENAS, 2003: 286).