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.

Distribution

North America, Mexico, West Indies, Bermuda, Central America, South America, introduced in Asia, Africa, Pacific Islands, Australia.

Discussion

Species ca. 90 (24, including 1 hybrid, in the flora).

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

None.

Lower Taxa

Physalis acutifolia, Physalis angulata, Physalis angustifolia, Physalis arenicola, Physalis caudella, Physalis cinerascens, Physalis cordata, Physalis crassifolia, Physalis fendleri, Physalis grisea, Physalis hederifolia, Physalis heterophylla, Physalis lanceolata, Physalis longifolia, Physalis missouriensis, Physalis mollis, Physalis neomexicana, Physalis philadelphica, Physalis pubescens, Physalis pumila, Physalis solanacea, Physalis virginiana, Physalis walteri, Physalis ×elliottii

Key

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