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thick leaf ground cherry, yellow nightshade groundcherry

nightshade family

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, annual or perennial, vines, shrubs, or trees, sometimes rhizomatous or tuberous, usually hermaphroditic, sometimes gynodioecious or dioecious in Lycium [andromonoecious or dioecious in some Solanum].
Stems

erect, branching from near base and at most nodes, branches widely spreading, distinctly zigzag, slender, 1–4(–10) dm.

erect or decumbent to prostrate, sympodially branched, glabrous or pubescent, hairs simple, dendritic, or stellate, stalked-vescicular in Quincula, sometimes peltate scales in Solanum, glandular or eglandular, glands sessile or stalked and sometimes multicellular, sometimes prickly or spiny.

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 and spiral or unequally paired, simple or pinnately compound [trifoliolate], estipulate, petiolate or sessile;

blade margins entire, dentate, lobed, or divided.

Inflorescences

terminal, axillary, extra-axillary, in forks of dichotomous branches in some Datura and Solanum, racemose, paniculate, umbellate, or glomerulate cymes, in fascicles of 2–6, or solitary flowers.

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.

bisexual or rarely unisexual, radially symmetric or, less frequently, bilateral;

perianth and androecium hypogynous;

sepals (4 or)5, connate for almost full length to only at base, tubular, campanulate, obconic, or cyathiform [urceolate], with 3–10 equal or unequal lobes, lobules, or teeth, 5–10 veins extending into appendages that protrude below a truncate rim in Lycianthes, bases of lobes sagittate to cordate in Nicandra;

calyx usually persistent in fruit and often accrescent, circumscissile and leaving a basal remnant that is slightly enlarged in some species of Datura, sometimes spreading, reflexed, or expanded to partially or completely enclose fruit, sometimes becoming membranous, sometimes inflated and bladderlike;

petals (4 or)5, connate for almost full length to only at base, stellate, campanulate, campanulate-rotate, rotate, funnelform, salverform, or urceolate [crateriform, cylindric], margins nearly entire or shallowly to deeply 4–10-lobed, lobes sometimes widely flaring or reflexed, corona present between tube and limb in Nectouxia;

stamens (2 or 4 plus staminodes or)5, variously adnate to corolla from near base nearly to rim, with appendage at point of fusion in Cestrum, equal or unequal in length, anthers sometimes connivent, basi-, dorsi-, or ventrifixed, with unequal thecae in Bouchetia, Browallia, and Hunzikeria, connective sometimes thickened, with 2 dorsal wings in Nectouxia, dehiscence longitudinal or poricidal;

pistil 1, 2–5-carpellate;

ovary superior, nectary disc sometimes present at base, [1 or]2–5[or 10]-locular, [1 or]2–many ovules per locule, placentation axile;

style 1, stigma rounded-truncate, capitate, or 2–5-lobed.

Fruiting calyces

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

Fruits

usually berries or 2–4-valved capsules, dehiscence septicidal, circumscissile, or irregular [septifragal, septicidal-loculicidal], sometimes drupaceous in Lycium [schizocarp of nutlets in Nolana].

Seeds

1–many, whitish to straw-colored to dark brown or black, often flattened and discoidal or reniform, sometimes globose, ovoid, prismatic, or angulate, with hyaline margin in Oryctes, with white caruncle in some species of Datura, intermixed with sclerotic granules in Calliphysalis and some Solanum;

embryo straight or curved, endosperm usually abundant.

2n

= 24.

Physalis crassifolia

Solanaceae

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]
nearly worldwide
[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.)

Genera ca. 97, species ca. 2500 (27 genera, 151 species, including 2 hybrids, in the flora).

Solanaceae occur on all continents except Antarctica, but their greatest diversity is in the Neotropics. Their closest relative is the Convolvulaceae, from which they can be distinguished by growth form (when herbaceous vines, not with twining stems), the usually 2-locular ovary with numerous ovules, and the absence of laticifers with milky sap. Members of the Solanaceae exhibit a sympodial growth pattern, resulting in unusual placement of leaves, branches, and flowers (A. Child 1979). The foliage of some Solanaceae is fetid or pungent (reminiscent of rotting potatoes or with a sharp, tomato-like smell). The vegetative portions of most members of the family are poisonous due to the presence of tropane and steroidal alkaloids.

Despite the toxicity of its foliage, many Solanaceae species are of worldwide economic importance, such as tomato (Solanum lycopersicum), eggplant (S. melongena), potato (S. tuberosum), chili and bell peppers (Capsicum spp.), tobacco (Nicotiana tabacum), and Petunia. The family also includes many species of small-scale crop, medicinal, or horticultural importance. The most well known of these in the flora area are tomatillo (Physalis philadelphica), belladonna (Atropa belladonna), goji berry (Lycium barbarum and L. chinense), Chinese lantern plant (Alkekengi officinarum), jimsonweed (Datura spp.), jessamine (Cestrum spp.), and Calibrachoa.

The fruit in the Solanaceae is typically a berry or capsule (with circumscissile dehiscence in Hyoscyamus), but may be hardened and drupaceous in some Lycium. The berry-fruited taxa may have a thick, juicy pericarp, or one that is dry and thin, ultimately shattering into irregular pieces. Many genera have an accrescent calyx. The most enlarged of these may become rigid and spinescent (Hyoscyamus), inflated and almost closed around the berry (Alkekengi, Calliphysalis, Physalis, Quincula), loose and enveloping all (Nicandra) or at least the basal half of the berry (Chamaesaracha), or reflexed or flaring (Jaltomata). The brightly colored, juicy berries of many species are animal dispersed. The inflated, bladderlike calyces in several genera also aid in dispersal (short distance by water or ground level by wind).

The showy flowers of Solanaceae are pollinated by bees, flies, butterflies, moths, or hummingbirds foraging for nectar and/or pollen. Solanum does not produce nectar, and pollen is gathered by vibration (buzz pollination) or manipulation of the anthers. Flowers of Cestrum, Datura, and some Nicotiana emit a strong, sweet fragrance in the evening and are moth-pollinated.

Some Solanaceae grown as ornamentals have escaped and persisted for short periods but have not become established in the flora. Nierembergia hippomanica Miers var. coerulea R. Millan has been collected from disturbed sites in Texas (1958, 1962, 1998), and N. scoparia Sendtner was found along a roadside in Georgia in 1947. Nierembergia Ruiz & Pavon is a South American genus. Brugmansia suaveolens (Humboldt & Bonpland ex Willdenow) Sweet, also native to South America, was found in a ruderal yard (2001) and wooded ravine (1983) in Florida. Brugmansia has been treated as part of Datura, but it can be distinguished from that genus by its woody habit, pendulous flowers, and elongated, spineless fruits.

Since the early 1990s, our understanding of the circumscription of the Solanaceae, as well as phylogenetic relationships within the family, have been clarified using both morphological and molecular characters, particularly chloroplast DNA sequence data (W. G. D’Arcy 1991; A. T. Hunziker 2001; T. R. Martins and T. J. Barkman 2005; R. G. Olmstead and L. Bohs 2007; Olmstead and J. D. Palmer 1991, 1992; Olmstead et al. 1999, 2008; T. Särkinen et al. 2013). Based on molecular data, the family is considered monophyletic with the inclusion of Nolana Linnaeus f., Schizanthus, and Sclerophylax Miers. The subfamilies Solanoideae Burnett and Cestroideae Burnett, traditionally recognized on the basis of morphology alone, have been shown to be nonmonophyletic. Some generic concepts have been revised; changes relevant to the flora area are discussed under the various genera.

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

Key
1. Corollas campanulate, rotate, stellate, or urceolate; fruits berries (hardened or drupaceous in some Lycium).
→ 2
2. Flowers and fruits borne in subumbellate, umbellate, or cymose clusters.
→ 3
3. Flowers and fruits borne in umbellate clusters; anthers dehiscing longitudinally; calyces enlarged, with flaring lobes in fruit.
Jaltomata
3. Flowers and fruits borne in subumbellate or cymose clusters; anthers dehiscing by pores (sometimes expanding to longitudinal slits with age); calyces only slightly enlarged in fruit.
→ 4
4. Shrubs; calyces 10-veined; leaves simple, margins entire.
Lycianthes (in part)
4. Herbs, vines, subshrubs, shrubs, or trees; calyces 5-veined; leaves simple or pinnately compound, margins entire, dentate, or lobed.
Solanum
2. Flowers and fruits solitary or in fascicles of 2–8.
→ 5
5. Stems creeping (rooting at nodes); anthers dehiscing by pores.
Lycianthes (in part)
5. Stems erect, decumbent, prostrate, or scandent; anthers dehiscing by longitudinal slits.
→ 6
6. Shrubs or scandent, perennial herbs; calyces not, or only sometimes, enlarged in fruit.
→ 7
7. Scandent, perennial herbs; flowers and fruits solitary; corollas urceolate.
Salpichroa
7. Shrubs; flowers and fruits solitary or in fascicles of 2–8; corollas rotate, subrotate, or campanulate.
→ 8
8. Plants without spines.
Capsicum
8. Plants spinescent.
Lycium (in part)
6. Herbs erect or decumbent, annual or perennial; calyces enlarged in fruit.
→ 9
9. Corollas campanulate; calyx lobes enlarged and flaring in fruit, not enclosing berries.
Atropa
9. Corollas broadly campanulate, campanulate-rotate, rotate, or urceolate; calyx or calyx lobes enlarged in fruit and completely or partly enclosing berries.
→ 10
10. Calyx lobes with sagittate to cordate bases, lobes expanded and surrounding berries.
Nicandra
10. Calyx lobes without lobed bases, basal portions expanded and com­pletely or partially enclosing berries.
→ 11
11. Plants sparsely to densely covered with stalked, white vesicles.
Quincula
11. Plants glabrous or pubescent with simple or branched hairs.
→ 12
12. Stems erect, usually unbranched; corollas white, lobed; fruiting calyces drying orange-red or bright red, inflated and completely enclosing berries.
Alkekengi
12. Stems erect to ascending or decumbent to prostrate, branched; corollas white, cream, yellow, greenish, or deep purple, unlobed or with obscure or shallow lobes or teeth; fruiting calyces drying brown, enlarged or inflated and completely or partially enclosing berries.
→ 13
13. Flowers and fruits solitary; fruiting calyces inflated and completely enclosing berries.
Physalis
13. Flowers and fruits borne singly or in fascicles of (1–)2–6; fruiting calyces expanded but not inflated, mostly or completely enclosing berries.
→ 14
14. Stems decumbent to ± prostrate; fruiting calyces shorter than berries; berries dry.
Chamaesaracha
14. Stems erect to ascending; fruiting calyces just shorter than or exceeding berries; berries fleshy.
→ 15
15. Corollas broadly campanulate; stamens slightly unequal; fruiting calyces enclosing berries; seeds intermixed with sclerotic granules.
Calliphysalis
15. Corollas rotate; stamens equal; fruiting calyces shorter than or just exceeding berries; seeds not intermixed with sclerotic granules.
Leucophysalis
1. Corollas funnelform, salverform, or tubular; fruits capsules or berries (hardened or drupaceous in some Lycium).
→ 16
16. Plants woody (subshrubs, shrubs, small trees, or lianas).
→ 17
17. Shrubs, spinescent.
Lycium (in part)
17. Subshrubs, shrubs, small trees, or lianas, without spines.
→ 18
18. Inflorescences axillary panicles; fruits berries.
Cestrum
18. Inflorescences axillary or terminal cymes, racemes, glomerules, or solitary flowers; fruits capsules.
→ 19
19. Shrubs or small trees, 0.2–6(–10) m; stamens 5, inserted near base of corolla.
Nicotiana (in part)
19. Shrubs or subshrubs to 1.5 m; stamens 4 (5th much reduced or absent), inserted in distal half of corolla.
→ 20
20. Corollas 5-lobed or sometimes appearing 4-lobed due to fusion of 2 abaxial lobes; capsules 2-valved; seeds prismatic or rounded on back and excavated ventrally.
Browallia (in part)
20. Corollas 5-lobed; capsules 4-valved; seeds reniform and finely wrinkled.
Hunzikeria
16. Plants herbaceous.
→ 21
21. Corollas bilaterally symmetric (or slightly bilateral); fruits capsules.
→ 22
22. Leaf margins usually deeply pinnately dissected or coarsely toothed or lobed.
→ 23
23. Leaf margins coarsely toothed to shallowly lobed; corolla lobes shorter than to as long as tube; capsules circumscissile.
Hyoscyamus
23. Leaf margins pinnate-pinnatifid; corolla lobes longer than tube and often deeply dissected; capsules septicidal.
Schizanthus
22. Leaf margins entire or irregularly crenate or undulate.
→ 24
24. Stems erect; stamens 5.
Nicotiana (in part)
24. Stems erect to ascending, decumbent, or procumbent; stamens 4, sometimes with reduced or sterile 5th.
→ 25
25. Stems sprawling or procumbent; corollas funnelform; calyces not accrescent; capsules hemispheric, 2-valved.
Calibrachoa
25. Stems erect or ascending to decumbent; corollas funnelform or salver­form; calyces accrescent; capsules ellipsoid or ovoid, 2- or 4-valved.
→ 26
26. Corollas funnelform; capsules ellipsoid, 4-valved.
Bouchetia
26. Corollas salverform; capsules ovoid, 2-valved.
Browallia (in part)
21. Corollas radially symmetric; fruits capsules or berries.
→ 27
27. Corollas funnelform with 5 long-acuminate lobes; fruits 4-valved or irregularly dehiscing capsules, sometimes with prickles or tubercles.
Datura
27. Corollas funnelform, salverform, or tubular, lobes not long-acuminate; fruits berries or 2–4-valved capsules, not prickly.
→ 28
28. Plants glabrous.
→ 29
29. Plants prostrate to ascending; inflorescences solitary or fascicled flow­ers; corolla lobes acute to acuminate; fruits berries.
Jaborosa
29. Plants usually erect; inflorescences cymose, forming false racemes or glomerules; corolla lobes rounded to deltate; fruits capsules.
Nicotiana (in part)
28. Plants glandular-pubescent.
→ 30
30. Stamens equal or slightly unequal; fruits berries.
→ 31
31. Corollas salverform, with corona between tube and limb; berries nar­rowly ovoid, juicy.
Nectouxia
31. Corollas tubular; berries globose, dry.
Oryctes
30. Stamens unequal; fruits capsules.
→ 32
32. Flowers and fruits in cymose clusters (appearing as false racemes or glomerules); calyx lobes deltate or triangular.
Nicotiana (in part)
32. Flowers and fruits solitary; calyx lobes linear.
Petunia
Source FNA vol. 14. FNA vol. 14. Author: Janet R. Sullivan.
Parent taxa Solanaceae > Physalis
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
Alkekengi, Atropa, Bouchetia, Browallia, Calibrachoa, Calliphysalis, Capsicum, Cestrum, Chamaesaracha, Datura, Hunzikeria, Hyoscyamus, Jaborosa, Jaltomata, Leucophysalis, Lycianthes, Lycium, Nectouxia, Nicandra, Nicotiana, Oryctes, Petunia, Physalis, Quincula, Salpichroa, Schizanthus, Solanum
Synonyms P. crassifolia var. cardiophylla, P. crassifolia var. versicolor, P. versicolor
Name authority Bentham: Bot. Voy. Sulphur, 40. (1844) Jussieu
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