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FNA | Family List | FNA Vol. 14 | Solanaceae

23. Physalis Linnaeus, Sp. Pl. 1: 182. 1753; Gen. Pl. ed. 5, 85. 1754.

Ground-cherry, husk-tomato, tomaté [Greek physa, bladder or bellows, and alis, belonging to, alluding to inflated fruiting calyx]

Janet R. Sullivan

Margaranthus Schlechtendal

Herbs [shrubs], annual or perennial, taprooted or rhizomatous, glabrous or pubescent, hairs simple, forked, or dendroid-stelliform, sometimes multicellular and appearing jointed, glandular or eglandular. Stems erect to weakly decumbent, branching. Leaves alternate, sometimes geminate, petiolate or sessile; blade simple, margins entire, dentate, or sinuate. Inflorescences axillary, solitary flowers [fascicles of 2–5]. Flowers 5-merous, (pendent or nodding); calyx campanulate with 5 broadly to narrowly triangular lobes, accrescent and inflated in fruit, becoming reticulate-membranous and bladderlike with narrow orifice and completely enclosing berry; corolla yellow or pale cream-yellow to nearly white, often with 5 large spots or smudges or star-shaped tinge of color in throat, and mat of white hairs at base of throat, radial, rotate or campanulate-rotate, limb sometimes widely flaring or reflexed at maturity, 5-angulate or obscurely 5-lobed, (P. solanacea dark purple or rarely yellowish or greenish with large purple spots, urceolate, with 5 shallow teeth) [campanulate with open limb]; stamens inserted at base of corolla tube, equal; anthers basifixed, oblong to narrowly elliptic, dehiscing by longitudinal slits; ovary 2-carpellate; style straight, slender, sometimes expanding distally; stigma minutely capitate or truncate. Fruits berries, globose, juicy [somewhat dry], completely enclosed by inflated fruiting calyx. Seeds reniform, flattened [oblique-triangular]. x = 12, 24.

Species ca. 90 (24, including 1 hybrid, in the flora): North America, Mexico, West Indies, Bermuda, Central America, South America; introduced in Asia, Africa, Pacific Islands, Australia.

Physalis is recognizable by the fruiting calyx that enlarges and inflates to completely enclose the berry, and pendent or nodding flowers borne singly at each node; most members also have an unlobed, yellow or cream-yellow, campanulate-rotate corolla. Physalis solanacea is atypical in having an urceolate corolla that is mostly or completely lurid purple; the molecular phylogenetic analysis by M. Whitson and P. S. Manos (2005) showed that it is part of a clade of more typical species. Their study also supported treatment of Calliphysalis and Alkekengi as genera distinct from Physalis.

The variable morphology of Physalis species has resulted in many misidentified herbarium specimens. Hair morphology is important in identifying most taxa.

Two species occur as waifs in the flora area: Physalis minima Linnaeus, native to the New World tropics, has been collected as a weed in sugarcane fields and sandy open pastures in Louisiana; and P. ixocarpa Brotero ex Hornemann [including var. parviflora (Waterfall) Kartesz & Gandhi and var. immaculata (Waterfall) Kartesz & Gandhi], native to Mexico, has been collected sporadically from disturbed sites in the United States and Canada.

Several species of Physalis are commonly cultivated for their edible berries: P. philadelphica (tomatillo) has fruits with a flavor reminiscent of tomato and is used in Mexican-style salsa; P. minima (pygmy groundcherry) has fruits similar to a cherry tomato; and P. peruviana Linnaeus (cape gooseberry) and P. grisea (strawberry tomato) have sweet fruits that are used in pies, jams, sauces, and as a garnish. The berries of many wild-growing species are eaten as well. Fruits of P. minima and P. peruviana are also used worldwide as an antispasmodic, diuretic, antiseptic, sedative, and analgesic. Unripe fruits and foliage of Physalis species contain solanine and other solanidine alkaloids and are toxic if ingested in large quantities.

After this manuscript was completed, Physalis macrosperma Pyne, E. L. Bridges & Orzell was published describing plants similar to P. heterophylla but with larger fruiting calyces and seeds. Further study is needed to clarify the disposition of this taxon.

SELECTED REFERENCES Martínez, M. 1998. Revision of Physalis section Epeteiorhiza (Solanaceae). Anales Inst. Biol. Univ. Nac. Autón. Méx., Bot. 69: 71–117. Martínez, M. 1999. Infrageneric taxonomy of Physalis. In: M. Nee et al., eds. 1999. Solanaceae IV: Advances in Biology and Utilization. Kew. Pp. 275–283. Menzel, M. Y. 1951. The cytotaxonomy and genetics of Physalis. Proc. Amer. Philos. Soc. 95: 132–183. Rydberg, P. A. 1896b. The North American species of Physalis and related genera. Mem. Torrey Bot. Club 4: 297–374. Seithe, A. and J. R. Sullivan. 1990. Hair morphology and systematics of Physalis (Solanaceae). Pl. Syst. Evol. 170: 193–204. Sullivan, J. R. 1985. Systematics of the Physalis viscosa complex (Solanaceae). Syst. Bot. 10: 426–444. Sullivan, J. R. 2004. The genus Physalis (Solanaceae) in the southeastern United States. Rhodora 106: 305–326. Sullivan, J. R., V. P. Patel, and W. Chissoe. 2005. Palynology and systematics of Physalis (Solanaceae). In: R. C. Keating et al., eds. 2005. A Festschrift for William G. D’Arcy: The Legacy of a Taxonomist. St. Louis. Pp. 287–300. Waterfall, U. T. 1958. A taxonomic study of the genus Physalis in North America north of Mexico. Rhodora 60: 107–114, 128–142, 152–173. Whitson, M. and P. S. Manos. 2005. Untangling Physalis (Solanaceae) from the physaloids: A two-gene phylogeny of the Physalinae. Syst. Bot. 30: 216–230.

1 Corollas urceolate, 2.5–4.5 mm; pedicels 1.5–3 mm in flower, 3–5 mm in fruit; fruiting calyx 5-ribbed   22 Physalis solanacea
+ Corollas rotate to campanulate-rotate, 5–20 mm; pedicels 2–46 mm in flower, 3–60 mm in fruit; fruiting calyx 10-ribbed or sharply 5-angled.   (2)
2 (1) Anthers strongly twisted after dehiscence; fruiting calyx filled, and often burst, by berry   19 Physalis philadelphica
+ Anthers not twisted after dehiscence; fruiting calyx loosely enclosing, or nearly filled, by berry.   (3)
3 (2) Plants pubescent, hairs 2- or 3-branched or dendroid-stelliform, or glabrous except for dendroid-stelliform hairs on leaf margins and calyx.   (4)
+ Plants pubescent, hairs simple, or glabrous except for simple hairs on pedicels and calyx, or glabrous.   (12)
4 (3) Hairs predominantly 2- or 3-branched.   (5)
+ Hairs predominantly dendroid-stelliform.   (6)
5 (4) Hairs to 0.5 mm, appressed, giving plants a grayish appearance; pedicels 3–11(–17) mm in flower, 10–15(–20) mm in fruit   10 Physalis fendleri (in part)
+ Hairs 0.5–2 mm, divergent; pedicels 8–46 mm in flower, 15–55 mm in fruit   21 Physalis pumila (in part)
6 (4) Plants glabrous except for leaf margins and calyx.   (7)
+ Plants pubescent.   (8)
7 (6) Leaves sessile; blade linear-lanceolate (sometimes folded along midrib), 0.2–0.8(–1) cm wide   3 Physalis angustifolia
+ Leaves sessile or petiole to 1/10 blade; blade narrowly spatulate to linear-lanceolate, 0.2–1.5(–2) cm wide   9 Physalis × elliottii (in part)
8 (6) Leaves sessile or petiole to 1/10 blade; blade elliptic or spatulate to linear-lanceolate.   (9)
+ Leaves petiolate, petioles 1/5 to as long as blade; blades orbiculate to broadly ovate or elliptic.   (10)
9 (8) Leaf blades 1–4 cm wide; corollas with dark purple-black spots; coastal Louisiana, Texas   6 Physalis cinerascens (in part)
+ Leaf blades 0.2–1.5(–2) cm wide; corollas with pale brown, ochre, or green spots or smudges; Florida   9 Physalis × elliottii (in part)
10 (8) Leaf blades broadly elliptic to ovate, base rounded, margins usu­ally entire or rarely shallowly sinuate   24 Physalis walteri
+ Leaf blades orbiculate to ovate, base truncate to slightly attenuate, margins usually dentate or sinuate, sometimes entire.   (11)
11 (10) Plants sparsely to somewhat densely pubescent, hairs to 1 mm   6 Physalis cinerascens (in part)
+ Plants densely pubescent, hairs to 1 mm (obscuring plant surface on younger growth), sometimes also with 2–4 mm branched or simple hairs   17 Physalis mollis
12 (3) Annuals, taprooted; fruiting calyces sharply 5-angled or 10-ribbed.   (13)
+ Perennials, rhizomatous; fruiting calyces 10-ribbed.   (19)
13 (12) Corollas rotate; flowering pedicels (13–)20–34(–40) mm   1 Physalis acutifolia
+ Corollas campanulate-rotate; flowering pedicels 2–17(–22) mm.   (14)
14 (13) Corollas without spots or smudges, or only tinged purple; fruiting calyces 10-ribbed.   (15)
+ Corollas with 5 large, dark purple-black spots or smudges; fruiting calyces sharply 5-angled.   (16)
15 (14) Plants glabrous or sparsely pubescent, hairs eglandular; leaf blades narrowly elliptic-ovate to linear-lanceolate; pedicels 7–17(–22) mm in flower, 15–30 mm in fruit   2 Physalis angulata
+ Plants sparsely to densely pubescent, hairs intermixed glandular and eglandular; leaf blades broadly ovate to orbiculate; pedicels 4–7 mm in flower, 5–10 mm in fruit   16 Physalis missouriensis
16 (14) Leaf margins coarsely dentate, teeth 10+ per side; pedicels (10–)15–35 mm in fruit   7 Physalis cordata
+ Leaf margins entire, irregularly crenate-dentate, or coarsely den­tate, teeth fewer than 10 per side; pedicels 5–15 mm in fruit.   (17)
17 (16) Plants villous, hairs intermixed with stalked and sessile glands; leaf blades gray-green, usually drying orange or with orange patches   11 Physalis grisea
+ Plants ± glabrous to villous, hairs glandular and/or eglandular; leaf blades green, drying green or grayish-brownish.   (18)
18 (17) Pedicels noticeably stout, especially in fruit; fruiting calyces nearly spheric   18 Physalis neomexicana
+ Pedicels slender; fruiting calyces always noticeably longer than wide   20 Physalis pubescens
19 (12) Plants becoming suffrutescent, pubescent, hairs divergent, to 0.5 mm; flowering calyces (3–)4–7(–8) mm   8 Physalis crassifolia
+ Plants remaining herbaceous, glabrous to densely pubescent, hairs appressed or not, 0.5+ mm; flowering calyces 5–14 mm.   (20)
20 (19) Leaf blades broadly ovate to orbiculate; plants often glandular.   (21)
+ Leaf blades ovate to elliptic, broadly lanceolate, or oblanceolate; plants not glandular.   (23)
21 (20) Pedicels 4–8(–13) mm in flower, 5–15 mm in fruit   12 Physalis hederifolia
+ Pedicels (8–)9–17(–25) mm in flower, 15–30(–35) mm in fruit.   (22)
22 (21) Plants glabrous to villous, hairs antrorse, to 1 mm, sometimes also simple, jointed, divergent, 1–2 mm; often with slender, shallowly buried rhizomes   4 Physalis arenicola
+ Plants villous, hairs divergent, 1–2 mm, sometimes also with shorter glan­dular hairs; all rhizomes stout and deeply buried   13 Physalis heterophylla
23 (20) Plants pubescent, hairs 1–3 mm; anthers usually dark purple to blue, rarely yellow   5 Physalis caudella
+ Plants glabrous or pubescent, hairs to 0.5 mm; anthers yellow or with blue or purple tinge.   (24)
24 (23) Plants pubescent, hairs mostly simple with some 2- or 3-branched intermixed   10 Physalis fendleri (in part)
+ Plants glabrous or pubescent, hairs simple only.   (25)
25 (24) Plants glabrous or sparsely strigose, hairs antrorse, to 0.5 mm   15 Physalis longifolia
+ Plants sparsely to densely pubescent, hairs divergent, 1–1.5 mm, and antrorse or retrorse, to 0.5 mm.   (26)
26 (25) Stems erect; leaf blade margins coarsely to shallowly dentate or entire; hairs divergent and retrorse   23 Physalis virginiana
+ Stems erect to decumbent; leaf blade margins entire or sinuate; hairs divergent and antrorse.   (27)
27 (26) Leaf blades oblanceolate; eastern coastal plain   14 Physalis lanceolata
+ Leaf blades elliptic-ovate to ovate-lanceolate; Great Plains   21 Physalis pumila (in part)

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