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thick leaf ground cherry, yellow nightshade groundcherry

ground-cherry

Habit Herbs perennial, becoming suffrutescent, rhizomatous, rhizomes often just below soil surface, vertical, stout, puberulent, hairs divergent, to 0.5 mm, some glandular, appearing ± glabrous without magnification. 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, branching from near base and at most nodes, branches widely spreading, distinctly zigzag, slender, 1–4(–10) dm.

erect to weakly decumbent, branching.

Leaves

petiolate;

petiole mostly as long as blade;

blade broadly ovate to deltate, 0.8–3.3(–4.5) × 0.8–3.3(–4.5) cm, base cordate, sometimes slightly unequal, margins entire to unevenly coarsely dentate, sometimes thick and slightly succulent.

alternate, sometimes geminate, petiolate or sessile;

blade simple, margins entire, dentate, or sinuate.

Inflorescences

axillary, solitary flowers [fascicles of 2–5].

Pedicels

8–24(–33) mm, (11–)14–30(–35) mm in fruit.

Flowers

calyx (3–)4–7(–8) mm, lobes 1–3 mm;

corolla pale yellow with yellow or greenish-brown smudges or tinge, campanulate-rotate, 8–14 mm;

anthers yellow, not twisted after dehiscence, 1.5–3 mm.

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.

Fruiting calyces

loosely enclosing berry, 10-ribbed, 14–30(–40) × (10–)15–20(–25) mm.

Fruits

berries, globose, juicy [somewhat dry], completely enclosed by inflated fruiting calyx.

Seeds

reniform, flattened [oblique-triangular].

x

= 12, 24.

2n

= 24.

Physalis crassifolia

Physalis

Phenology Flowering year-round in areas without frost, mostly Mar–Apr.
Habitat Gravelly or sandy slopes, washes, roadsides, mesas, canyons.
Elevation 100–1700 m. (300–5600 ft.)
Distribution
from FNA
AZ; CA; NV; UT; Mexico (Baja California, Baja California Sur, Chihuahua, Sinaloa, Sonora) [Introduced in Australia]
[WildflowerSearch map]
[BONAP county map]
from USDA
North America; Mexico; Central America; South America; West Indies; Bermuda [Introduced in Asia, Africa, Pacific Islands, Australia]
[BONAP county map]
Discussion

In Physalis crassifolia, the corolla limb is widely flaring and reflexed when the flower is fully open, and the flowers are more nodding than fully pendent. Some herbarium specimen labels indicate that plants flower the first year. Physalis greenei Vasey & Rose, not validly published, has been misapplied to some representatives of P. crassifolia. Physalis crassifolia is widespread in Arizona, but it is restricted in California to south-southeastern counties as far north as Inyo, in Nevada to Clark and Lincoln counties, and in Utah to Washington County.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

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.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

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.
P. 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.
P. 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.
P. fendleri
5. Hairs 0.5–2 mm, divergent; pedicels 8–46 mm in flower, 15–55 mm in fruit.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. 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.
P. neomexicana
18. Pedicels slender; fruiting calyces always noticeably longer than wide.
P. 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.
P. 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.
P. 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.
P. arenicola
22. Plants villous, hairs divergent, 1–2 mm, sometimes also with shorter glan­dular hairs; all rhizomes stout and deeply buried.
P. 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.
P. 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.
P. fendleri
24. Plants glabrous or pubescent, hairs simple only.
→ 25
25. Plants glabrous or sparsely strigose, hairs antrorse, to 0.5 mm.
P. 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.
P. virginiana
26. Stems erect to decumbent; leaf blade margins entire or sinuate; hairs divergent and antrorse.
→ 27
27. Leaf blades oblanceolate; eastern coastal plain.
P. lanceolata
27. Leaf blades elliptic-ovate to ovate-lanceolate; Great Plains.
P. pumila
Source FNA vol. 14. FNA vol. 14. Author: Janet R. Sullivan.
Parent taxa Solanaceae > Physalis Solanaceae
Sibling taxa
P. acutifolia, P. angulata, P. angustifolia, P. arenicola, P. caudella, P. cinerascens, P. cordata, P. fendleri, P. grisea, P. hederifolia, P. heterophylla, P. lanceolata, P. longifolia, P. missouriensis, P. mollis, P. neomexicana, P. philadelphica, P. pubescens, P. pumila, P. solanacea, P. virginiana, P. walteri, P. ×elliottii
Subordinate taxa
P. acutifolia, P. angulata, P. angustifolia, P. arenicola, P. caudella, P. cinerascens, P. cordata, P. crassifolia, P. fendleri, P. grisea, P. hederifolia, P. heterophylla, P. lanceolata, P. longifolia, P. missouriensis, P. mollis, P. neomexicana, P. philadelphica, P. pubescens, P. pumila, P. solanacea, P. virginiana, P. walteri, Physalis × elliottii
Synonyms P. crassifolia var. cardiophylla, P. crassifolia var. versicolor, P. versicolor Margaranthus
Name authority Bentham: Bot. Voy. Sulphur, 40. (1844) Linnaeus: Sp. Pl. 1: 182. (1753): Gen. Pl. ed. 5, 85. (1754)
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