Herbs or vines (elsewhere also shrubs and trees). Leaves opposite, simple, in ours unlobed; stipules present between the leaves or represented by a line or membrane between the leaf bases. Flowers in ours perfect, regular, ours 5-merous. Sepals free or basally united, valvate or imbricate in bud. Corolla fused, salverform to tubular or campanulate. Stamens epipetalous, usually alternate with the corolla lobes. Ovary superior to partially inferior, in ours free of the calyx, composed of 2 carpels and bilocular. Fruit of our material capsular.
20+ genera and about 600 species from the tropics to the temperate regions; 3 genera and 6 species in TX; 3 genera and 4 species here. This treatment reflects the removal of some genera, including our Polypremum, to the Buddlejaceae
Some, including Gelsemium are ornamental; others are poisonous (Mabberley 1987).
1. Plants twining woody vines ................................................................................1. Gelsemium
1. Plants herbaceous ...................................................................................................................2
2(1) Corolla funnelform, more than 6 mm long; style 1; flowers solitary in the axils and a few terminal ....................................................................................................................2. Spigelia
2. Corolla urceolate, less than 4 mm long; styles 2; flowers in cymes with one-sided
branches ..................................................................................................................3. Mitreola
3 species, 2 of the SE. U.S. and 1 of SE. Asia to Malaysia; we may have the 1 species found in TX.
1. G. sempervirens (L.) Jaume St.-Hil. Carolina Jessamine, Yellow Jessamine, Poor Man's Rope, Evening Trumpetflower. Trailing or high-climbing viney shrub, twining left to right; stems slender, wiry, red-brown, smooth. Leaves semievergreen in our area, ovate to lanceolate or elliptic, to 7.5 cm long and 3 cm broad, acute to acuminate, basally rounded to cuneate, glabrous; petioles 2 to 7 mm long; stipules small and deciduous. Flowers fragrant, 5-merous, solitary or in cymose clusters of up to 6 in the axils; pedicels short and with scaly bracts. Sepals free, lanceolate, 3 to 5 mm long, obtuse to subacute; corolla yellow, funnelform, 2.5 to 3.5 cm long, the lobes ca. 7 to 10 mm long, spreading; stamens attached to the lower portion of the flaring tube, anthers oblong, sagittate; style slender, 2-cleft, each arm divided and thus appearing 4-cleft, flowers heterostylic. Capsule elliptic-oblong, with 2 valves and 2 locules, 1.4 to 2 cm long, 0.8 to 1.2 cm broad, flattened perpendicular to the partition, apically rounded and abruptly beaked; seeds many, brown, apically winged, body papillose. Open sandy woods or wood edges in E. TX; reported from Brazos Co. (TAES 48551; H. B. Parks, s.n., Mar. 9, 1946), but this specimen likely from cultivation as Parks frequently omitted mention of cultivation. Possibly present in the wild in E. portions of Grimes, Leon, and Madison Cos. in pine-sweetgum communities. FL to TX, N. to SE. VA, TN, and AR. Feb.-Apr. [Authority frequently given as (L.) Ait. f.].
Cultivated for its deliciously fragrant flowers, this plant has also been used medicinally to treat migraines and neuralgia (Mabberley 1987), but it is toxic. Most poisonings result from its medicinal use, with severe intoxications involving muscular weakness, spasms, or convulsions, but there are cases of children having been poisoned by sucking the nectar from the blossoms (Lampe 1985). Honey made from the blossoms is reported to be toxic (Tull 1987).
Herbaceous perennials. Leaves simple, opposite, united at the base by stipules or a stipular line. Flowers solitary or in 1-sided cymes (sometimes spike-like), 5-merous. Sepals united at the base, calyx lobes slender. Corolla funnelform or salverform, the lobes relatively short. Anthers linear. Style 1, pubescent on the upper portion, jointed near the middle. Capsule bi-lobed and bilocular, splitting into the 2 component carpels at maturity.
About 50 species of tropical and subtropical Amer., with 1 naturalized in the Old World; 3 species in TX; 1 here.
Several species (e.g. S. anthelmia and S. marilandica) are medicinal, having uses as vermifuges. Some (including the medicinal ones) are poisonous (Lampe 1985; Mabberley 1987).
1. S. loganioides (T. & G. ex Endl. & Fenzl) A. DC. Texas Pink-root. Perennial from a group of slender fibrous roots; stems 1 to several from the base, slender, spreading, to ca. 3 dm tall; herbage more or less glabrous. Leaves ovate to elliptic-lanceolate or sometimes obovate or oblanceolate (especially on the lower portion of the stem), thin-textured, (1.5)2.5 to 5 cm long, to 2 cm broad, apically obtuse to acute, basally cuneate, glabrous or the margins and sometimes the nerves slightly scabrous, short-petiolate, well-spaced; stipules very small. Flowers terminal and in the axils of the upper leaves. Sepals linear-subulate, ca. 5 mm long, 1-nerved, margins scabrous-serrulate, apically acute to obtuse; corolla funnelform to salverform, ca. 12 mm long, white, lobes ca. 3 mm long; anthers and style included; stigma capitate. Pedicels elongating in fruit; capsule bi-globose, slightly compressed perpendicular to the septum, 3 to 4 mm long. Wooded slopes and floodplain woods along river courses in S. TX; endemic; in our area known from along the Old River in Burleson Co. May-Aug, ours primarily May. [S. texana (T. & G.) A. DC.; Coelostylis loganioides T. & G. ex Endl. & Fenzl.; C. texana T. & G.].
Annual or perennial herbs. Stems often slightly quadrangular. Herbage glabrous or essentially so. Leaves opposite, entire; stipules between the leaves, small. Inflorescences pedunculate terminal cymes, branched, the branches helicoid (1-sided) cymes. Flowers small, regular, 5-merous. Sepals united only at the base, ovate to elliptic, several-veined. Corolla longer than the calyx, globose-funnelform or urceolate, white or tinged with pink or blue, in ours 1 to 2 mm long. Stamens included. Gynoecium briefly united to the calyx at the base, the 2 carpels united for 1/2 or more their lengths, divergent to ascending; styles 2. Capsule exserted, strongly 2-horned or miter-shaped, dehiscent along the inner side of each carpel. Seeds many, widely ellipsoid, grooved on one side.
About 6 species of tropical to warm-temperate regions; 2 species in TX, both present here. Texas material was formerly in Cynoctonum; Mitreola is apparently the earliest valid name. This treatment is based, in part, on the work of Nelson (1980).
1. Leaves tapered to a petiole, at least some blades 3 cm long or more; capsules 3 to 4 mm long, exterior mostly smooth ............................................................................1. M. petiolata
1. Leaves sessile or with a petiole less than 1 mm long, blades rarely more than 2 cm long; capsules 2 to 3 mm long, exterior papillose .................................................2. M. sessilifolia
1. M. petiolata (J. F. Gmel.) T. & G. Lax Hornpod. Annual from a cluster of fibrous roots; stems simple or more commonly branched, erect, 1.3 to 7.5 dm tall. Leaves elliptic to elliptic-lanceolate, lance-ovate, or ovate, 2 to 6(8) cm long, apically acuminate to obtuse, basally tapered to a petiole 3 to 15 mm long or the upper leaves sometimes sessile, entire, thin-textured, glabrous; stipules represented by small ovate-triangular lobes. Cymes long-pedunculate, branched, ultimate branches helicoid and secund (1-sided), moderately densely flowered, elongating in fruit so the mature fruit mostly well-spaced and not touching. Sepals ovate to elliptic, 0.8 to 1.2 mm long, basally united; corolla white or the tube slightly bluish; carpels separate about 1/2 their length, divergent. Capsule exserted from the calyx, 3 to 4 mm long, externally smooth or with a few scattered papillae, mitriform (miter-shaped); seeds 0.3 to 0.5 mm long, minutely pebbled, shiny. Moist soil of seeps and ditches and around ponds, streams, and lakes. E. and Cen. TX; SE. VA, TN, and AR, S. to FL and TX. May-Oct.; collected in fruit as late as Nov. [Cynoctonum mitreola (L.) Britt.; M. petiolata (Walt.) T. & G.].
2. M. sessilifolia (J. F. Gmel.) G. Don Wand Hornpod. Annual from a cluster of fibrous roots; stems erect, simple or with a few erect branches above, 1 to 5 dm tall. Leaves often ascending, ovate or oval to broadly elliptic or suborbicular, 0.5 to 2.5 cm long, 0.3 to 2 cm broad, apically acute to obtuse, basally rounded, firm-textured, margin usually papillose as seen with a lens, sessile or with a very short petiole to 0.5 mm long. Cymes compact (moreso than in M. petiolata), ultimate branches secund, dense in flower and fruit. Sepals more or less ovate, ca. 1 mm long, prominently keeled, basally briefly united; corolla white, slightly longer than the calyx; carpels united about 1/2 their length, ascending. Capsule 2 to 3 mm long, externally conspicuously and densely papillose as seen with a lens; seeds smooth, shining, 0.2 to 0.3 mm long. Seepage slopes, bogs, wet savannahs, and along woodland streams. E. TX; FL to TX, N. to SE. VA. June-Oct., our collections primarily Aug.-Oct. [Cynoctonum sessilifolium (Walt.) J. F. Gmel. or Walt. ex J. F. Gmel.].
Ours herbs, elsewhwere mostly trees and shrubs. Leaves usually opposite, entire as in ours or toothed or lobed, stipules usually represented by a line between petiole bases. Flowers variously arranged, perfect or functionally unisexual, usually 4-merous. Sepals united, calyx sometimes parted very deeply. Petals united, corolla usually regular, lobes mostly imbricate. Stamens epipetalous, alternate with the corolla lobes. Gynoecium bilocular, superior to half-inferior, style solitary, terminal, stigma capitate or 2-lobed, ovules many, placentation axile. Fruit usually a septicidal capsule. Seeds sometimes winged.
About 10 genera and 150 species; 3 genera and 7 species in TX; 1 species here.
These plants have often been treated as part of the Loganiaceae.
A monotypic genus.
1. P. procumbens L. Polyprim, Polly-prim, Juniperleaf. Taprooted perennial herb to ca. 3 dm tall; stems several to many from the base, well-branched, radially ascending or repent, ribbed; herbage more or less scabrous. Leaves linear-subulate, 1 to 2.5(3) cm long, 0.5 to 2(2.5) mm broad, acute, ascending to widely spreading, each pair connected at the base by an inconspicuous stipular line. Flowers solitary, terminal and in the axils of the leaves, 4-merous, sessile, or with pedicels less than 0.5 mm long. Calyx with a broad, scarious-margined base, lobes lance-subulate, ca. 2 to 3 mm long, dorsally keeled; corolla white, essentially rotate, about as long as the sepals, bearded in the throat, lobed ca. 1/3 its length, lobes suborbicular, ca. 1 mm long; stamens short, included, anthers globose; style 1, short, stigma entire, ovoid. Capsule ovoid, 1.5 to 2.5 mm long, slightly flattened and notched apically, 2-celled and 2-valvedl; seeds many, yellow, more or less square in outline, with microscopic pits. Usually in sandy soil of fields, roadsides, dunes, pastures, openings in woods, etc. E. 2/3 TX; FL to TX, N. to NY, NJ, E. PA, and SE. MO; S. to Colombia and the W. Indies. Apr.-Nov.
Some studies place Polypremum in or near the Scrophulariaceae (see Zomlefer 1994).
Ours annual or perennial herbs (elsewhere also sometimes woody). Leaves opposite, whorled, or rarely alternate, simple, sessile to petiolate, estipulate, ours commonly glabrous or nearly so. Flowers solitary or in cymose or corymbose inflorescences, terminal and/or axillary, perfect, regular, 4- to 12-merous. Sepals usually united at least briefly, in ours persistent. Petals more or less united, corolla tubular or salverform to campanulate or rotate, often with nectary scales or pits within. Stamens epipetalous, as many as the corolla lobes and alternate with them. Ovary superior, bicarpellate, unilocular (or bilocular through intrusion of the parietal placentae toward the middle), style 1, elongate to obsolete, stigma usually relatively large, entire to bifid. Fruit in ours a 2-valved, septicidal, usually unilocular capsule with many seeds.
74 genera and about 1,200 species worldwide, especially common in temperate and subtropical regions and the montane tropics; 8 genera and 22 species in TX; 3 genera and 3 species here, with 1 more perhaps to be looked for.
The family includes many ornamentals, especially in Gentiana, Exacum, and Sabatia. Some taxa have medicinal uses (Mabberley 1987).
1. Corolla blue, purple, lavender, or white; rarely pink; anthers straight or merely recurved in age ..........................................................................................................................1. Eustoma
1. Corolla pink or rose; anthers twisted or strongly curved in age .............................................2
2(1) Corolla lobes as long as or shorter than the corolla tube; anthers becoming twisted with old age ...............................................................................................................2. Centaurium
2. Corolla lobes longer than the corolla tube; anthers merely curved or rolled in old age .........
..................................................................................................................................3. Sabatia
Annual or short-lived perennial from a taproot, usually with a basal rosette. Stems erect or ascending, leafy, herbage more or less glaucous. Leaves sessile, clasping, entire. Flowers usually 5-merous (occasionally 4-merous), showy, long-pedicelled, solitary or in cymose panicles. Calyx lobes elongate, keeled. Corolla campanulate, the lobes convolute in bud and erect in flower, apically entire to erose or apiculate, nectary glands or appendages none. Stamens inserted on the corolla throat, anthers oblong, versatile, strongly or slightly recurved in age. Style slender, semi-persistent, stigma conspicuously 2-lobed. Capsule ellipsoid, 2-valved and many-seeded.
A genus of 3 species of the S. U.S. to N. S. Amer.; 2 in TX; 1 here. Kartesz (1998) combines the two N. American species.
1. E. grandiflorum (Raf.) Shinners (=Eustoma exaltatum (L.) Salisb. ex G. Don subsp. russellianum (Hook.) Kartesz, comb. nov. ined.) Showy Prairie-gentian, Bluebells, Lira de San Pedro. Annual or short-lived perennial; stems 1 to several from the base, erect, 25 to 70 cm tall; internodes 1.4 to 6 cm long. Leaves elliptic-oblong to elliptic-lanceolate, lanceolate, lance-ovate or sometimes ovate, 3-veined, 1.5 to 8 cm long, 0.3 to 5 cm broad, glaucous. Inflorescence cymose-paniculate, flowers 2 to 6 per cluster; pedicels to 6 cm long. Calyx lobes linear-lanceolate or subulate, 1.2 to 2.3 cm long, 2 to 3 mm broad; corolla very showy, generally blue-purple, occasionally pink or whitish, often with a darker eye, deeply lobed, lobes elliptic-obovate, 3 to 5 cm long, 1.5 to 2.4 cm broad (3 or more times as long as the calyx), apically rounded to truncate, commonly apiculate and very slightly erose; anthers 4 to 5.5 mm long, recurved in age, filaments 10 to 15 mm long; style about as long as the ovary, stigma massive, 2-lobed, the lobes ca. 5 mm long. Capsule to 2 cm long. Moist areas of prairies and fields and around stock tanks and ponds. Throughout much of TX but sadly much less common in our area now than formerly; SW. SD, NE, and E. CO, S. to TX and Mex. June-Sept. [E. russellianum with the authority given variously as (Hook.) Sweet, (L.) Griseb., or (Hook.) G. Don--the latter is used by Kartesz (1998)].
Names have been given to the various color forms: f. grandiflorum (typical blue-purple), f. fisheri (Standl.) Shinners (white), f. bicolor (Standl.) Shinners (white with a purple tinge on the lobes, f. roseum (Standl.) Shinners (pink), and f. flaviflorum (Cockll.) Shinners (yellow).
Often cultivated for the very showy blossoms which last several days as a cut flower. Often sold under the name Lisianthus and offered in various colors. This plant was popularized about the time it became scarce in parts of its range; it is possible that plants and seeds were over-collected from the wild rather than purchased.
Annual herbs, generally low and well-branched but sometimes tall and showy (not ours). Herbage in TX material glabrous. Flowers 4- or 5-merous. Calyx deeply lobed, the lobes narrow and appressed to the corolla tube; corolla salverform or funnelform the tube slender and the limb 4- or 5-lobed, pink (as ours) or white. Stamens inserted on the corolla throat, anthers exserted, spirally twisting after dehiscence. Style 1, slender, stigma capitate or lobed. Capsule fusiform to oblong-ovoid. Seeds small.
30 species of the N. hemisphere; 4 in TX; 1 here.
Some are cultivated for ornament (Mabberley 1987).
1. C. texense (Griseb.) Fern. Lady Bird's Centaury. Stem erect, simple below and often corymbosely branched above, 1 to 3 dm tall. Cauline leaves linear to linear-lanceolate or linear-oblanceolate, 5 to 30 mm long, to 4 mm broad, reduced upwards and the uppermost only subulate bracts. Pedicels 5 to 12 mm long, about as long as the calyx; flowers pink. Calyx 8 to 10 mm long, the lobes 7 to 9 mm long; corolla tube 9 to 11 mm long, about twice as long as the lobes, lobes linear-oblong to oblong-elliptic, 3 to 7 mm long, to 1.5 mm broad. Seeds light brown. Usually in dry calcareous soils of hillsides and barrens, also sometimes in depressions on granite; Ed. Plat. and Blackland Prairies; in our area known from calcareous sandstone in Grimes Co. and (oddly) collected once along the Highway 6 bypass E. of Bryan; MO and TX. Jun.-Aug. [Erythraea texensis Griseb.].
The common name honors former First Lady, Lady Bird Johnson, founder of the National Wildflower Research Center and long-time lover of Texas wildflowers.
Annual or perennial herbs. Stems erect. Herbage glabrous. Leaves opposite and in some species also basal. Inflorescence terminal, cymose. Flowers 4- to 12-merous, ours usually 5-merous, usually pink (rarely white), often with a paler eye and/or star in the center. Sepals basally united, calyx lobes slender, longer than the tube. Corolla rotate. Stamens epipetalous, inserted on the upper rim of the corolla tube, alternate with the lobes, filaments slender, anthers bright yellow, coiled after dehiscence. Style 1, slender, stigma bifid. Capsule ovoid to cylindric. Seeds many.
17 species of N. Amer. and the W. Indies; 7 in TX; 1 here with one more to be looked for.
The flowers are generally showy and some species are cultivated for ornament (Mabberley 1987).
1. S. campestris Nutt. Prairie Rose-gentian. Annual; stems slender, unwinged but somewhat 4-angled, erect, 1 to 3.5(5) dm tall, usually simple in the lower half and if branched above, the branches 1 per node and simple, internodes 1 to 4.4 cm long. Leaves ovate to lance-ovate, ovate-elliptic, or oblong-elliptic, 0.8 to 2.5(4.5) cm long, 0.5 to 1.2(2) cm broad, with 1 or 3 strong nerves rom the base, apically generally acute, base rounded, clasping, and sessile. Inflorescences loose cymose clusters forming a somewhat corymbose arrangement; flowers generally 5-merous (occasional aberrant individuals 4- or 6-merous), subtly fragrant. Calyx tube to 8 mm long, pentagonal, the lateral nerves well-developed and usually with thin, narrow wings that extend to the sinuses between the lobes, lobes linear to lanceolate, 1 to 2.8 cm long, longer than the tube, longer or shorter than the corolla lobes; corolla tube 2/3 or more enclosed by the calyx, lobes broadly ovate to elliptic or spatulate, 1.2 to 2.3 cm long, 7 to 15 mm broad, obtuse to acute, bright rose-pink with a paler spot basally, flowers often with a white eye and yellow star; stigma lobes greenish, turning yellow with age, 5 to 8 mm long. Capsule to 9 mm long. Common in prairies, fields, roadsides, waste places, etc. in both dry and moist soils. E. 1/2 TX and SW. following the coast; IL S. to MS, W. to IA, KS, OK, and TX. Apr.-July. [S. formosa Buckl.].
NOTE: S. angularis (L.) Pursh occurs in E. TX., including on the Blackland Prairies and Post Oak Savannah. It has not been seen by the author from this area, but ought to be looked for. It can be distinguished by its winged stems and unwinged calyx tube which covers 1/3 or less of the corolla tube; branches are usually 2 per node.
Ours annual or perennial herbs, vines, or shrubs (elsewhere also trees), commonly with milky sap. Leaves evergreen or deciduous, opposite, alternate, or sometimes whorled, simple, entire, often revolute; stipules inconspicuous or lacking. Flowers solitary and axillary or in racemes or corymbose or thyrse-like cymes, perfect, regular, ours 5-merous except for the gynoecium. Sepals united, if only briefly, calyx lobes usually imbricate, sometimes with glands or appendages within. Corolla fused, varying in shape from salverform or tubular to urceolate or campanulate, sometimes with appendages in the throat, lobes imbricate or convolute in bud. Stamens epipetalous, alternate with the corolla lobes, anthers introrse, sagittate, sometimes sticky and lightly adhering to the stigma. Nectary glands sometimes present subtending the gynoecium. Gynoecium superior, in ours of 2 unilocular carpels with axile placentation, free below and united only above by the style; stigma usually relatively large, variously shaped. Fruit 2 (or 1 by abortion) cylindrical to fusiform follicles, each few- to many-seeded. Seeds glabrous or with a coma.
A large, diverse family of 215 genera and 2,100 species, primarily tropical but a few temperate; 8 genera and 21 species in TX; 6 genera and 9 species here. Though old, the treatment of Woodson (1938) is useful for descriptions and distribution information.
Sometimes treated to include the Asclepiadaceae, with which it shares many features. Thorne takes this view of the two families--(for a presentation, see Zomlefer (1994).
Many species are poisonous and/or have medicinal properties, e.g. species of Ravolfia (source of reserpine) and Catharanthus (source of several promising anti-leukemia drugs). Many genera include ornamentals, notably Allamanda, Nerium, Plumeria, and Vinca (Mabberley, 1987).
1. Leaves alternate .....................................................................................................1. Amsonia
1. Leaves opposite or whorled .....................................................................................................2
2(1) Plants twining vines; corolla pale yellow ..............................................2. Trachelospermum
2. Plants erect to trailing, not vining, herbs or shrubs; corolla white to blue, red, or pink, usually not yellow .....................................................................................................................3
3(2) Plants cultivated shrubs, occasionally persisting or escaping ................................3. Nerium
3. Plants trailing to erect annual or perennial herbs ....................................................................4
4(3) Stems trailing; plants evergreen herbs; corolla blue-purple to red- purple ..............4. Vinca
4. Stems erect; plants herbs; corolla white to pink or red, not blue ............................................5
5(4) Corolla less than 1 cm broad; native perennial ................................................5. Apocynum
5. Corolla more than 1 cm broad; cultivated annual .......................................6. Catharanthus
Perennial herb from a woody, erect or creeping rootstock. Leaves alternate or crowded enough to appear verticillate, linear to lanceolate or broadly elliptic, petiolate to sessile, margin often revolute. Inflorescences terminal or occasionally lateral thyrsiform cymes with several to many flowers, dense to pen, often not much surpassed by the leaves, flowers held erect to drooping; bracts inconspicuous. Sepals united in the basal 1/5 to 1/3, calyx lobes more or less equal, without appendages. Corolla salverform, blue, orifice of tube constricted in some species but open in ours and the tube enlarged at the point of stamen attachment, orifice densely pilose and the corolla retrorsely pubescent within, glabrous to pubescent externally, lobes spreading to erect, linear to ovate, appendages none. Anthers ovate to triangular, the connective not enlarged, connivent over the stigma, included. Style filiform, stigma depressed-capitate, with a cup-like wing just below the summit; nectary none. Follicles straight to curved, continuous or constricted between the seeds, erect to pendulous. Seeds in 1 row per follicle, ends truncate, coma none.
About 20 species of N. Amer. and Japan; Hatch, et al. (1990) listed 9 species for TX, but Kartesz (1998) merges two of these; 3 species are to be expected in our area. Despite its age, the works of Woodson (1928, 1938) are very useful for characters and species distributions.
Some species have ornamental value (Mabberley 1987). According to Tull (1987), TX plants are poisonous, though they are not listed by the AMA (Lampe 1985).
1. Calyx and corolla both completely glabrous externally (sometimes the calyx with a few weak hairs); leaves usually 6 cm or less long, sessile or subsessile ...................1. A. ciliata
1. Calyx or corolla or both with at least some pubescence externally; leaves to 15 cm long, petiolate (at least the upper and middle ones) ........................................................................2
2(1) Calyx glabrous; corolla villous to glabrate; leaves dull on both sides, thin-textured; follicles erect ...................................................................................................2. A. tabernaemontana
var. salicifolia
2. Calyx at least sparsely pilose or villous; corolla villous; leaves shiny above, subcoriaceous; follicles spreading to pendulous ..........................................................................3. A. illustris
1. A. ciliata Walt. Fringe Slimpod. Stems from woody rootstocks 0.3 to 0.8 cm in diameter, to 3.5 dm tall, pilose to glabrate or glabrous. Leaves alternate but commonly crowded enough to appear nearly verticillate, sessile or subsessile with petioles less than 3 mm long (at least on the upper stem), oblong-lanceolate to linear-lanceolate or filiform, often narrower upwards, to 6(8) cm long and 1.7 cm broad, tapered to the base, apically obtuse to acute, margin often ciliate. Cymes primarily terminal, dense, scarcely exceeding the foliage, with several to many medium-blue flowers; pedicels 3 to 5 mm long, puberulent to glabrate. Calyx lobes ovate to lanceolate, narrowly triangular, or linear-lanceolate, long-acuminate, 0.5 to 2.5 mm long, glabrous or with a few weak, villous hairs only on the narrow, scarious margins; corolla completely glabrous externally, the tube often darker than the lobes or sometimes greenish, 6 to 10 mm long, 1 mm broad at the base, lobes 3.5 to 11 mm long, oblong to broadly ovate, somewhat spreading. Follicles slender, erect, 6 to 11 cm long, not constricted between the seeds, glabrous; seeds 5 to 11 mm long. Calcareous hills and grasslands, along railroads, and in sandy-loam prairie and cedar woods. Ed. Plat., N. Cen., and S. Cen. TX; NC, S. and W. to FL and TX. Mar.-May.
A quite variable species. According to Woodson (1928), present in our area, but no collections seen from our counties by the author. Two varieties are present in TX.:
var. texana (A. Gray) Coult. Texas Slimpod. Leaves well-spaced, commonly strongly dimorphic, the upper oblong-lanceolate to elliptic, much narrower than the lower, which are elliptic-lanceolate to linear-lanceolate, 2 to 6 cm long, 0.3 to 1.7 cm broad, shiny above, dull below. [A. texana (Gray) Heller].
var. filifolia Woods. Leaves crowded, subverticillate, linear-lanceolate to filiform, 2 to 5 cm long, 0.5 to 4 mm broad. [A. ciliata Walt. var. tenuifolia (Raf.) Woods., and listed as such by Kartesz (1998)].
2. A. tabernaemontana Walt. var. salicifolia (Pursh) Woods. Willow Slimpod, Willow Amsonia. Stems 3 to 11 dm tall, few or several from a woody rootstock 0.7 to 1 cm in diameter, old stem bases often present, branches (if any) in the upper 1/5 to 1/2 of the plant, 1 per node, ascending. Leaves alternate and well-spaced, thin-textured, not dimorphic as to size, lanceolate to narrowly oblong-elliptic, 6 to 15 cm long, 1.2 to 2.5 cm broad, acute to acuminate, basally oblong to acute, entirely glabrous or finely pubescent below, dark green and dull on both sides, short-petiolate; petioles 2 to 10 cm long, or sometimes the lowermost leaves sessile. Inflorescences terminal or axillary and subterminal, dense to loose, scarcely exceeding the foliage; flowers several to many; pedicels 3 to 6 mm long. Calyx lobes narrowly to widely triangular, 0.5 to 2 mm long, glabrous, margins narrowly scarious; corolla light blue or the tube darker than the lobes or sometimes greenish, villous on the upper 1/2 of the tube and the middles of the lobes, varying to nearly glabrous and with only a few hairs, tube 6 to 8 mm long, ca. 1 mm broad at the base, lobes spreading, 4 to 9 mm long, spatulate to broadly lanceolate. Follicles erect, 8 to 13 cm long, slender, not constricted between the seeds, glabrous; seeds 6.5 to 9 mm long. Sandy soils around ponds and streams in E. TX; PA to GA, SW. to TX. Mar.-May [Includes var. gattingeri Woods.].
Kartesz (1998) lists also for TX a var. tabernaemontana (A. glaberrima Woods. of Hatch, et al [1990]). This variety (or species) has an entirely glabrous corolla. It is found in extreme SE. TX and is not expected in our area.
NOTE: A. tabernaemontana, A. illustris, and A. repens (the latter apparently not present in our area) are very closely allied. Some TX. material is not readily referable to a particular species. Characters of pubescence, traditionally used to separate the species, are not always definitive and the complex could benefit from intensive study.
3. A. illustris Woods. Blue-star. Perennial from a stout, woody rootstock 1 to 1.5 cm in diameter; stems solitary or few (often bases of old stems present), 6 to 9(12) dm tall, glabrous, branches (if any) in the upper 1/5 to 1/3 of the plant, usually 1 per node and ascending. Leaves not thin-textured, usually sub-coriaceous or firm, lustrous above and dull beneath, well-spaced, not dimorphic, lanceolate to elliptic-lanceolate, (4)5 to 12 cm long, (0.8)1 to 2 cm broad, acute to acuminate, basally acute, glabrous or slightly villous on the midnerve above, glabrous to slightly villous below, short-petiolate; petioles (1)2 to 8 mm long or the lowermost leaves sessile. Cymes terminal or axillary and subterminal, dense, scarcely surpassing the foliage, with many flowers; pedicels 2 to 8 mm long. Calyx lobes narrowly to broadly triangular or lanceolate, acuminate, 0.5 to 3 mm long, sparsely to densely villous or pilosulous, margins narrowly scarious; corolla more or less pilose externally on the upper 1/2 of the tube and the middles of the lobes (usually manifestly so in local individuals), light blue, the tube often darker than the lobes or tinged with green or yellow, tube 6 to 8 mm long, ca. 1 mm in diameter basally, lobes lanceolate, elliptic-lanceolate, or narrowly oblong, 5 to 10 mm long, spreading. Follicles 8 to 14 cm long, widely spreading to pendulous at maturity, slender, constricted between the seeds or not, glabrous; seeds 7 to 10 mm long, dark red-brown, with rows of corky tubercles. Our most common species. Moist or wet soil of meadows, swamps, bottomland woods, and along streams and ditches. E. 1/3 TX; S. MO and E. KS to TX. Mar.-June.
See NOTE at A. tabernaemontana, above.
20 species, most in the E. hemis. from India to Japan; 1 in the SE. U.S. and present here.
Some are cultivated for ornament. In TX, T. asiaticum and T. jasminoides are especially common, both used as ground covers. These two have shiny ovate leaves and white flowers. Neither is known to escape in our area.
1. T. difforme (Walt.) Gray American Star Jasmine, Climbing Dogbane, Climbing Star Jasmine. Deciduous woody twining vine or herbaceous perennial; stems reddish; herbage glabrous to puberulent. Leaves opposite, petiolate or nearly sessile, blades quite variable in shape, elliptic to obovate-elliptic, lanceolate, ovate, or occasionally even suborbicular, sometimes heterophyllous and variable on any one plant, ca. 4 to 12 cm long, 1 to 7.5 cm broad, acuminate (sometimes abruptly so) to apiculate, basally cuneate to rounded; stipules tiny. Inflorescences in alternate axils or sometimes appearing terminal, thyrsiform or corymbose, flowers usually many, relatively small; pedicels 4 to 7 mm long. Calyx lobed nearly to the base, lobes ovate-lanceolate, acuminate, 3 to 4 mm long, more than twice as long as the tube, the tips sparsely barbellate; corolla salverform or slightly funnelform, pale yellow (or greenish), the tube 5.5 to 6.5 mm long, ca. 1 mm broad at the base and slightly expanded above, the 5 lobes oblong-obovate, 3 to 4 mm long, spreading, convolute in bud; stamens and stigma included, stamens inserted halfway down the corolla tube, filaments short, anthers connivent and more or less stuck together around the stigma, connectives enlarged, narrowly 2-lobed; style elongate, stigma fusiform; gynoecium subtended by 5 free or more or less coalescent nectary glands. Follicles paired, slender, terete, sometimes slightly constricted between the many seeds, 10 to 23 cm long, glabrous; seeds truncate, comose. On shrubs and trees along streams and wood edges and in weedy areas. E. TX; in our area usually in dense bottomlands and not often collected; DE to FL and TX, W. to IL, IN, MO, and OK. Apr.-June.
2 species from the Mediterranean to Japan; 1 cultivated and sometimes persisting in parts of TX.
Members of this genus are deadly poisonous--see notes following our species.
1. N. oleander L. Common Oleander, Laurel Rosa, Rose Bay. In our area a shrub to ca. 6 m, commonly dying to the ground in severe freezes; stems usually several to many from the base and the form more or less globose; herbage essentially glabrous; sap not usually milky. Leaves opposite or many of them in whorls of 3 or 4, short-petiolate, coriaceous, oblong-lanceolate, to 30 cm long and 3.5 cm wide, the pinnate venation very strong and even, upper surface very shiny. Inflorescence more or less corymbose, the flowers white rose, red, or yellow, in some cultivars "double" (with 10 petals). Calyx lobes lanceolate to ovate-lanceolate, 4 to 6 mm long, acuminate, somewhat leaf-like; corolla funnelform, glabrous externally, the tube 8 to 12 mm long, ca. 1.5 mm broad at the base, expanded above into a conic-campanulate throat 9 to 10 mm long and ca. 7 mm broad at the orifice, commonly with 5 laciniate-dentate scales; lobes more or less obovate to obovate-oblong, 2 to 3.5 cm long, spreading; stamens borne about the middle of the tube, anthers lobed basally and awned apically, connivent above and loosely cohering to the stigma. Follicles 2 (or 1 by abortion), brown, rather sturdy, 8 to 15 cm long; seeds many, flattened, puberulent, apically comose. Native to the Medit. and E. Asia; cultivated for ornament; sometimes shortly persisting in old landscapes but not (as far as is known) naturalizing in our area.
All parts of this plant are extremely toxic--one leaf can be enough to kill an adult, and even water in which the flowers have been placed is toxic. The toxic principles are cardioactive glycosides similar to digitalis (Lampe 1985). The smoke from burning wood is also potentially toxic, and there are reports (possibly anecdotal) of children being poisoned from using the branches as hotdog roasting sticks.
Perennial herbs or subshrubs. Stems erect to trailing. Leaves opposite. Flowers solitary in the axils of alternate leaves, 5-merous except for the gynoecium. Calyx lobes narrow, without appendages. Corolla funnelform or salverform, the tube cylindrical, throat hairy or thickened, appendages none, lobes twisted to the left, in ours blue-purple or white. Stamens inserted on the corolla, not connivent, anthers with the connective prolonged into an apical appendage. Gynoecium subtended by 2 nectaries nearly equal in size to the carpels at anthesis. Follicles terete, slender, many-seeded. Seeds slightly flattened, without a coma.
7 species of Eur. to N. Afr. and Cen. Asia; 2 species cultivated for ornament and sometimes persistent.
Some are used medicinally in their native regions (Mabberley 1987).
1. Calyx lobes glabrous; leaves narrowed at the base .............................................1. V. minor
1. Calyx lobes ciliate; leaves broadly rounded, truncate, or subcordate at the base ...................
. ................................................................................................................................2. V. major
1. V. minor L. Common Periwinkle. Trailing evergreen herb. Leaves with petioles 1 to 2 mm long; blades firm to subcoriaceous, elliptic, 1.5 to 6 cm long, 8 to 25 mm broad, apically obtuse to broadly acute, base more or less cuneate to broadly acute, shiny above, glabrous or commonly minutely glandular puberulent below. Flowers with pedicels 15 to 33 mm long, glabrous. Calyx lobes ovate-lanceolate, acute, to 3 mm long, glabrous; corolla bright blue (rarely white), the tube 3 to 6 mm long, ca. 1.5 mm broad at the base, throat conic-campanulate, 5 to 7 mm long, 2 to 3.5 mm in diameter. Follicles 2 to 7 cm long, slender, uncommon. Native of Eurasia; widely cultivated; naturalized in the N. temperate region; possible in our area but probably not a permanent member of our flora. Early spring.
2. V. major L. Bigleaf Periwinkle. Trailing evergreen herb; stems to 3 m long. Leaves long-petiolate, blades ovate (or the very lowest suborbicular), basally rounded to truncate or cordate, to 7 cm long and 5 cm broad, ciliate on the margin, otherwise glabrous. Flowers blue to lavender (rarely white). Calyx lobes narrowly linear, ca. 1 cm long, ciliate; corolla tube ca. 15 mm long, narrow below, opening into a conic-campanulate throat, commonly with a white pentagonal ring at the orifice, lobes asymmetrical, to ca. 2 cm long. Follicles ca. 5 cm long. Sandy soil, usually along streams, at edges of woods, or in open woods. E. and Cen. TX; not common; when found in our area usually directly traceable to cultivation. Native of Europe; sometimes escaping cultivation. Feb.-Apr.
Herbaceous perennials from rhizomes. Stems erect to ascending, commonly dichotomously branched. Leaves opposite or occasionally whorled, sessile to petiolate, variously shaped, sometimes mucronate, margin often revolute; stipules small, inconspicuous. Inflorescences terminal and axillary corymbose cymes, the floral bracts small to rather leafy; flowers (in ours) small and pale. Sepals united in the basal 1/3 to 2/3, without appendages. Corolla cylindrical to campanulate or urceolate, the tube relatively short, limb with 5 equal, spreading to reflexed lobes that are twisted to the right, each lobe with a triangular appendage at the base. Anthers narrowly triangular, fertile only in the upper 2/3, connivent above the stigma and lightly adhering to it; connective enlarged, 2-lobed, filaments flattened, apically enlarged, villous. Style short, clavate, stigma conical or ovoid-fusiform; gynoecium subtended by 5 free, ovoid nectaries alternate with the stamens. Follicles separate or stuck together at the tips, spreading to pendulous, terete. Seeds many, truncate, slender-fusiform, overlapping, with a coma.
7 species of the temperate Americas; 4 listed for TX by Hatch, et al. (1990); 3 listed by Kartesz (1998); 1 here.
1. A. cannabinum L. Indian Hemp Dogbane, Prairie Dogbane. Stems from a stout rhizome or rootstock, 2 to 10 dm tall; herbage variously glabrous to pubescent or villous, often glaucous; branches ascending to spreading, mainly in the upper 1/2 of the plant, alternate or opposite. Leaves mostly opposite, short-petiolate to sessile, ascending or slightly spreading, blades ovate to oblong-elliptic or lanceolate, 1.5 to 14 cm long, 0.3 to 4.5(7) cm broad, rounded to acute and usually apiculate apically, acute to cordate basally, generally glabrous above, glabrous to pilosulous or villous below and sometimes glaucous. Cymes usually dense, terminal; floral bracts linear to lanceolate, scarious, from inconspicuous to rather leafy and obvious, often early deciduous; flowers erect to drooping, white to greenish. Calyx lobes lanceolate to ovate-lanceolate or linear, 1.2 to 3(3.5) mm long, about as long as the corolla tube, glabrous; corolla urceolate to narrowly campanulate or short-cylindrical, 2.6 to 4.7 mm long, 1.5 to 3 mm broad at the apex, lobes ca. 1/3 to 1/2 as long as the tube, erect or slightly spreading. Follicles widely spreading to pendulous at maturity, sometimes coherent at the tips, straight to curved, 7 to 19(22) cm long, glabrous; seeds 3 to 6 mm long, coma (1.3)1.6 to 3.7 cm long, white to tawny. Moist or wet sandy or clay soils of bogs, ditches, stream courses, and along rivers, or sometimes open woods or fields. E., Cen., and N. Cen. TX; Can. and ME to WA, S. to FL, TX, and AZ; also N. Mex. Apr.-Aug. [Includes var. glaberrimum A. DC., var. cannabinum, and var. pubescens (R. Br.) A. DC.; A. pubescens R. Br. var. hypericifolium (Ait.) A. Gray; A. sibiricum Jacq. and its var. cordigerum (Greene) Fern. and var. salignum (Greene) Fern.; A. cordigerum Greene; A. suksdorfii Greene var. angustifolium (Woot.) Woods. Sometimes separated from A. sibiricum, but in large parts of its range (e.g. the Great Plains), there is complete overlap in "distinguishing" characters (GPFA 1986).
The root of this plant was used medicinally by plains tribes to treat constipation, dropsy, and ague and as en emetic, antisyphilitic, and general tonic. White settlers in the NW. U.S. learned of it from Native Americans and used it as a diuretic, cathartic, febrifuge, and purgative. It is still used in Appalachia in various remedies. The main chemical constituents are cardiac glycosides which may have anti-tumor properties (Tull 1987; Kindscher 1992). The plant is actually toxic to humans and animals, but because it is unpalatable, cases of severe poisoning are rare (Kindscher 1992). It is not listed by the A.M.A. as a poisonous plant (Lampe 1985). Woody fibers in the outer stem can be used like hemp for cordage. Mesquakie tribes used them for sewing (Kindscher 1992). The sap has a significant hydrocarbon content and might prove useful in fabricating synthetic rubber or for some other use (Tull 1987).
8 species of the tropics, 7 found only in Madagascar; 1 occasionally self-seeding/escaping from cultivation in Texas.
1. C. roseus (L.) G. Don f. Madagascar Periwinkle, Vinca. Perennial, but cultivated in our area as an annual; stems erect, to 7.5 dm, usually shorter. Leaves opposite, oblong-lanceolate, ca. 2.5 to 5 cm long, shiny. Flowers solitary or 2 to 3 in axillary cymes, 5-merous except for the gynoecium, perfect. Corolla salverform, in shades of pink, rose, red, fuchsia, or white, sometimes with a darker or lighter eye or star in the center, to ca. 4 cm broad, tube ca. 2.5 cm long, throat filled with bristly hairs; anthers sessile in the throat, without terminal appendages; style slender. Follicles to ca. 4 cm long; seeds 15 to 30 or more. Native from Madagascar to India; cultivated as a summer annual (plants are quite heat-tolerant); very occasionally self-sowing outside plantings; not long persisting and usually immediately traceable to cultivation; naturalized elsewhere in the tropics nearly worldwide. Summer. [Vinca rosea L.].
Ours herbaceous perennials, subshrubs, or herbaceous vines; sap milky. Leaves usually opposite, sometimes whorled, occasionally alternate, usually entire, estipulate or the stipules minute and/or deciduous. Inflorescences usually axillary and/or terminal umbellate cymes or sometimes the flowers solitary or paired. Flowers perfect, regular. Sepals 5, usually imbricate, more or less connate at the base (often only briefly), often reflexed. Corolla sympetalous, 5-lobed or -cleft, the tube usually short. A 5-lobed corona or crown often present between the corolla and androecium and adnate to either or both, variously shaped and often nectariferous. Stamens 5, inserted on the corolla tube, usually near the base, in our material the filaments united into a tube around the style, anthers united around and coherent to the stigma, the stamens, style, and stigma together forming the gynostegium; each anther sometimes with a terminal scarious or petaloid membrane) which is an outgrowth of the connective) and/or two lateral, wing-like margins; in ours pollen produced in waxy masses called pollinia, the pollinia from adjacent half-anthers of separate stamens connected via translator arms joined to central gland or corpusculum. Gynoecium superior, of two carpels united only by the massive stigma in the gynostegium, stigmatic surface usually reached through the slit formed by the edges of the wing-margins of 2 adjacent anthers. Fruit a pair of follicles, commonly only 1 developing. Seeds many, flattened, usually comose.
347 genera and 2,850 species of the tropics and subtropics (a few are temperate); 5 genera and 59 species in TX; 3 genera and 14 species here.
This is a very large family, closely allied to the Apocynaceae and grouped with it by some taxonomists (e.g. Thorne; see Zomlefer 1994). Many taxa are cultivated for ornament, including species of Asclepias, Hoya, Ceropegia, Stapelia, etc. (Mabberley 1987), with forms ranging from bedding plant to hanging-basket vine to succulent. Most taxa have some sort of very elaborate or complex insect pollination (Mabberley 1987).
1. Corona of 5 erect or spreading fleshy hoods, U-shaped or tubular in cross-section, adnate to the staminal column; plants prostrate to erect perennials, never twining; flowers in umbelliform cymes ...............................................................................................1. Asclepias
1. Corona of 5 to 15 distinct bladelike appendages or a fleshy, irregularly lobed disk or cup; plants twining vines, OR if herbaceous perennials then the flowers in axillary pairs or racemose ..................................................................................................................................2
2(1) Corona 1 row of 5 distinct laminar (blade-like) appendages, each apically bifid into 2 linear lobes which are free or partially fused, more or less as long as the corolla lobes .......
..........................................................................................................................2. Cynanchum
2. Corona 2 rows of laminar appendages shorter than the corolla or a single fleshy cup or disk, sometimes lobed .............................................................................................3. Matelea
Ours perennial herbs, often from an enlarged and more or less woody rootstock, almost always with milky sap. Stems 1 to many, prostrate to erect. Leaves opposite, alternate, whorled, or approximate (nearly opposite), sessile to petiolate, blade variously shaped. Inflorescences in ours terminal or axillary umbellate cymes. Calyx divided almost to the base, of 5 equal lobes, usually reflexed, with few to many minute glandular scales within. Corolla rotate, the lobes reflexed or spreading, sometimes erect, valvate in bud. Corona of 5 fleshy spreading to erect hoods, U-shaped in cross section or tubular, attached to the sessile or stipitate staminal column and subtending the anthers; horns often present, protruding from the hoods, needle-like to falcate or tongue-like, erect or incurved. Anther head more or less pentamerous, short-cylindric to truncate-conic or depressed-spheric; anthers 2-celled, terminal appendage ovate to deltoid, petaloid, lateral margin-wings more or less prominent, corneous (with the texture of thin horn), enclosing the 5 stigmatic chambers; corpusculum narrowly ovate; pollinia pendulous from the translator arms, flattened, asymmetrically spatulate. Follicles 2 or only 1 by abortion, fusiform to ovoid, terete or slightly angled. Seeds many, usually comose, rarely naked.
About 120 species of the W. Hemisph.; a few naturalized in the E. Hemisph.; 36 in TX; 9 in our immediate area. Though rather old, the work of Woodson (1954) is useful for descriptions, diagrams, and distribution information.
The flowers of some are quite showy and several species are commonly grown as ornamentals, e.g. A. curassavica with red and yellow flowers. Some species have medicinal value or produce a usable latex or fibers (Mabberley 1987). Others are the preferred food of butterfly larvae. Some are toxic to livestock, but severe intoxications are rare as they apparently are eaten only if other food is unavailable (GPFA 1986).
1. Hoods widely separated from the anther head at the base and then upright or spreading; horns none ..............................................................................................................1. A. viridis
1. Hoods closely appressed to the anther head at the base; horns present or absent. ............2
2(1) Horns none or vestigial and included well within the hoods (some hoods are pointed--look closely) ......................................................................................................................................3
2. Horns present, exserted from the hoods .................................................................................4
3(1) Leaves primarily alternate or a few subopposite, linear, 1 to 5 mm broad; base of hood with wide lateral lobes subtending the anther wings; anther wings arched, connivent over the anther head ......................................................................................2. A. engelmanniana
3. Leaves opposite (occasionally a few alternate), linear to ovate or suborbicular, usually more than 5 mm broad; base of hood without lobes; anther wings not arched over the anther head ......................................................................................................3. A. viridiflora
4(2) Hoods shorter than to equalling the anther head (or only about 1 mm longer); horns usually conspicuously surpassing the hoods ...........................................................................5
4. Hoods obviously extended above the anther head (at least 1/3 longer), the apex often spreading; horns not much surpassing the hoods ..................................................................6
5(4) Main stem leaves opposite, broadly ovate to elliptic or suborbicular, basally sessile and auriculate ..................................................................................................4. A. amplexicaulis
5. Main stem leaves usually whorled, filiform ...................................................5. A. verticillata
6(5) Hoods about twice as long as the anther head, 7 to 10 mm long, tips expanded above the middle, spreading, lobed ........................................................................6. A. oenotheroides
6. Hoods without the above combination of characters, usually shorter than 8 mm long, and not expanded above the middle ...............................................................................................7
7(6) Stems villous to hirsute with hairs 1 to 2 mm long; sap not milky ...................7. A. tuberosa
7. Stems glabrous to tomentose with hairs less than 1 mm long; sap milky .............................8
8(7) Leaves broadly ovate to narrowly lanceolate, acute to acuminate; peduncle longer than 1 cm; corolla dark red to purple or lavender .............................................................8. A. rubra
8. Leaves varying from broadly oval to oblong or quadrangular, narrower and smaller above; peduncle shorter than 1 cm; corolla pale green to yellow ................................9. A. obovata
NOTE: A. linearis Scheele has been collected from Milam Co., just outside our area. It may eventually be found here. It has opposite, linear-filiform leaves, axillary inflorescences, greenish flowers with hoods about as long as the anther head and horns slightly longer and gently curved, and mature pedicels erect.
1. A. viridis Walt. Antelope-horn, Oblong-leaved Milkweed. Perennial from a stout, woody, elongate or subfusiform rootstock; stems solitary or paired, decumbent to ascending, simple or occasionally branched below, slender to stout, (1.5)2.5 to 6.5 dm tall, glabrous or slightly pubescent above. Leaves subopposite or alternate, petioles 3 to 10 mm long; blades ascending to spreading, lanceolate to oblong or ovate, 4 to 13 cm long, 1 to 6 cm broad, apically acute to obtuse or occasionally emarginate, base tapered to rounded, truncate, or subcordate, glabrous to sparsely puberulent. Inflorescences 1 to 5(7), usually terminal or subterminal and also lateral at the uppermost nodes; peduncles 0.3 to 6 cm long, sparsely and minutely puberulent; pedicels relatively slender, 1 to 3 cm long, pubescent like the peduncles; flowers several to many, 10 to 15 mm tall. Calyx lobes lanceolate to narrowly ovate, 2.8 to 5.5 mm long, essentially glabrous to puberulent; corolla lobes pale green, elliptic-lanceolate to ovate, 13 to 17 mm long, the tips ascending, glabrous; gynostegium sessile, column 2.5 to 3.2 mm tall, 1 to 1.6 mm broad; hoods pale pink-purple or edged with green, abruptly widespread from the base and then ascending, the free portion clavate, 4 to 6 mm long, rather fleshy, closed or essentially so, apex rounded, cucullate (hooded), not extending as long as the anther head, margins puberulent, a small appendage inconspicuous within; horns none; anther head truncate-conic, 2.7 to 3.2 mm tall, 2.7 to 3.4 mm broad; anther appendages ca. 0.6 mm long; anther margin wings obtusely angled in the upper 1/3, rounded in the lower 1/3, ca. 2.4 mm long; corpusculum ca. 0.3 mm long; pollinia ca. 1.3 mm long. Mature pedicels strongly deflexed, follicles erect, broadly fusiform to ovoid, 6 to 13 cm long, 1 to 3 cm wide, minutely puberulent to glabrate; seeds oval to broadly obovate, 7 to 8 mm long, coma white or pale tan, 3 to 4 cm long. Sandy or rocky soil of roadsides, prairies, hillsides, dry woods, etc. E. 1/2 TX; TN and OH to NE, S to FL and TX. Mar.-Sept. [Asclepiodora viridis (Walt.) A. Gray].
Sometimes mistaken for A. asperula (Dcne.) Woods., which is superficially similar but which has narrower, long-acuminate leaves. It is found to the west of our area and apparently does not occur here.
The comas were formerly twisted and used as candle wicks (Ajilvsgi 1984).
2. A. viridiflora Raf. Green Antelope-horn, Green Milkweed. Herbaceous perennial from a slender, woody, vertical rootstock; stems usually solitary or sometimes paired, simple or rarely branched at the base, slender to stoutish, sometimes zig-zagging from node to node above, 1 to 6(10) dm tall, minutely puberulent to tomentose or glabrate. Leaves opposite or nearly so (occasionally some alternate), petioles short, 0.1 to 0.5 cm long or leaves sessile; blades extremely variable in shape, linear to lanceolate, ovate, or suborbicular, ascending to wide-spreading, 4 to 13(14) cm long, (0.2)0.8 to 5(6) cm wide, acute to obtuse or emarginate, often mucronate, basally acute to rounded, inconspicuously puberulent to glabrate or tomentulose. Inflorescences terminal and in the axils of the leaves in the upper 1/2 of the plant, hemispherical; peduncles 0.2 to 2 cm long; pedicels slender, 0.5 to 1.5(2) cm long, peduncles and pedicels minutely puberulent to villous or tomentose; flowers 20 to 80, usually crowded, 9.5 to 12.5 mm tall. Calyx lobes green or tinged with purple, ovate-lanceolate or lanceolate, 2.1 to 4 mm long, minutely puberulent; corolla lobes strongly reflexed, pale green, elliptic-lanceolate, 5.7 yo 6.5(7) mm long, sparsely puberulent dorsally; gynostegium sessile, glabrous, pale green; column 1.2 to 1.5 mm tall, 1.2 to 1.4 mm broad; hoods green, drying darker and purplish or brownish, oblong or oblong-lanceolate, attached to the gynostegium in the lower 1/3, deeply saccate, erect, appressed to the gynostegium, 2.9 to 5 mm long, ca. 1 mm shorter than the anther head, fleshy, opening appressed to the anther head, apex rounded and flat, margins with a small lobe at the base; horns none; anther head truncate-fusiform, 3 to 4 mm tall, 2.7 to 3.1 mm broad; anther apical appendages ca. 1 mm long, white; marginal wings obtusely angled and most obvious above the middle, ca. 3 mm long; corpusculum ca. 0.3 mm long; pollinia ca. 2.1 mm long. Mature pedicels deflexed, follicles erect, narrowly to broadly fusiform, apically attenuate, 7 to 15 cm long, 1.5 to 2 cm broad, puberulent to glabrate; seeds oval to broadly obovate, 6 to 7.5 mm long, coma 3 to 5 cm long, pale tan. Roadsides, prairies, plains, calcareous outcrops, hillsides, etc, in sandy or rocky (often calcareous) soils. Infrequent in scattered populations throughout much of TX; S. Can. to CT and MT, S. to GA, TX, an AZ; also NE Mex. Apr.-Aug.; our collections mostly Jun.-July. [Includes var. lanceolata (Ives) Torr., var. linearis (A. Gray), and var. ivesii Britt; Acerates viridiflora (Raf.) Pursh or (Raf.) Eaton].
The Lakota Sioux made medicines from the pulverized roots and gave it to children for diarrhea. It was also used in a tea believed to stimulate milk production in nursing women (Kindscher 1992).
3. A. engelmanniana Woods. Engelmann's Milkweed. Perennial from a rhizome or a rootstock, this elongate to sub-globose; stems usually solitary and simple, occasionally with a few branches, 3 to 12(14) dm tall, glabrous or else sparsely pubescent in vertical lines. Leaves alternate or only irregularly approximate (subopposite), sessile, loosely spreading, narrowly linear, (5)10 to 20 cm long, 1.5 to 5 mm broad, firm to slightly succulent, base and apex acute, glabrous to sparsely puberulent. Inflorescences scattered, several to many in the axils of the upper leaves, quite crowded, with many flowers, nearly spherical (at least in bud); peduncles 0.1 to 2(4) cm long; pedicels moderately slender, 0.8 to 1.5 cm long, villous or appressed-pubescent; flowers 7 to 10 mm tall. Calyx lobes green or tinged with purple, ovate-lanceolate, 2 to 3.5 mm long, puberulent or essentially glabrous; corolla lobes reflexed, pale green tinged with purple, elliptic-lanceolate, 4.5 to 6 mm long, glabrous; gynostegium yellowish, with a short stipe; column 1.3 to 2 mm long, 1.2 to 1.5 mm broad; hoods yellow-green, oblong, the lower 1/2 attached to the gynostegium, erect, 2.3 to 3.2 mm long, more or less fleshy, the opening appressed to the anther head and essentially hidden, apex truncate to retuse, flat, ca. 1 mm shorter than the anther head, deeply saccate, the base auriculate with lobes that subtend the anther wings; horns absent; anther head depressed-spheric, 1.8 to 2.1 mm tall, 2.7 to 3 mm broad; anther apical appendages ca. 1 mm long, white; anther wings arched, ca. 2 mm long; corpusculum ca. 0.4 mm long; pollinia ca. 1.3 mm long. Mature pedicels deflexed, follicles erect, fusiform, apically attenuate, 6 to 12 cm long, 1.3 to 1.8 cm thick, puberulent to glabrous; seeds oval or broadly obovate, 7 to 9 mm long, coma white to pale tan, 3 to 4 cm long. Prairies, open sandy hills, swales, washes, floodplains, etc. W. 2/3 TX, E. to about Robertson Co. in our area; Great Plains from NE W. to SE. UT, S. to TX, AZ, and N. Mex. May-Sept. [Acerates auriculata Engelm.; Asclepias auriculata (Engelm.) Holz., not H.B.K.].
4. A. amplexicaulis Sm. Blunt-leaved Milkweed. Herbaceous perennial from a deep-seated rhizome and a simple or branched crown; stems usually 1, simple, sturdy, (2)4 to 10 dm gall, glabrous and often glaucous. Leaves primarily opposite, sessile, blades broadly ovate or elliptic to broadly oblong or oblong-lanceolate, spreading, 4 to 12 cm long, 1.8 to 8 cm wide, thick or subcoriaceous, apex obtuse to rounded, often mucronate, base broadly cordate and more or less clasping, surfaces glabrous and more or less glaucous, margins usually puberulent and undulate. Inflorescence usually 1 and terminal, or rarely also some in the axils of the uppermost leaves; peduncle stout, (1)6 to 20(30) cm long (shorter if lateral); pedicels relatively slender, (2)3.5 to 4.5 cm long, puberulent; flowers 18 to 35(60), 15 to 18 mm tall. Calyx lobes lanceolate to ovate-lanceolate, 3 to 5.5 mm long, green or tinged with purple, glabrous; corolla reflexed, lobes green, commonly with some purple, lanceolate to elliptic-lanceolate, 9 to 11 mm long, glabrous; gynostegium stipitate, glabrous; column cylindric, 1.5 to 2.1 mm tall, 1.7 to 2.5(3) mm broad; hoods pale purple or rose, attached at the base, erect or slightly spreading but not widely separated from the anther head, oblong, 5 to 5.5 mm long, slightly fleshy, open and tubular-hooded, apex rounded to truncate or slightly and irregularly toothed, extending beyond the anther head, base slightly saccate; horns adnate to the hood in the lower 1/2 to 2/3, oblong, abruptly narrowed to a subulate tip, arched over the anther head, to about half again as long as the hood; anther head more or less cylindric, 3.5 to 4.5 mm tall, 3 to 3.2 mm broad; anther apical appendages ca. 1.5 mm long; anther wings basally acute, prominently spurred, ca. 5 mm long; corpusculum ca. 0.5 mm long; pollinia ca. 1.4 mm long. Mature pedicels deflexed, follicles erect, fusiform or slightly falcate, 9 to 16 cm long, 0.8 to 2 cm thick, puberulent to glabrous, glaucous; seeds broadly ovate, 6 to 9 mm long, coma pale tan or white, 2.5 to 6 cm long. Mostly on sandy or gravelly soils of woods, prairies, clearings, pastures, roadsides, old dunes, railways, etc. E. 1/3 TX S. to Rio Grande Plains; not especially common here but known at least from Brazos and Robertson Cos.; NH to MN and NE, S. to N. FL. and TX. Apr.-June.
One glycoside from this plant, amplexoside, has been shown to inhibit cell growth in human cancer (Kindscher 1992).
5. A. verticillata L. (Eastern) Whorled Milkweed. Perennial from a shallow cluster of fibrous roots, each nearly as thick as the stem; stems single or few, slender, simple or with a few branches, these often sterile, (2)3.5 to 9 dm tall, usually puberulent in decurrent vertical lines from the leaf bases. Leaves usually in whorls of 3 or 4(6) or occasionally only nearly whorled or opposite, sessile to subsessile, filiform to linear, 1.5 to 8 cm long, 0.5 to 1.5(3) mm broad, firm or somewhat leathery, spreading or erect, acute, basally narrowly acute, margin usually revolute, glabrous to puberulent. Inflorescences single or paired (occasionally 3) in the axils of the upper leavess; peduncles slender, 1 to 4.5 cm long; pedicels filiform, 5 to 11 mm long, puberulent; flowers 6 to 20, 5 to 7.5 mm tall. Calyx green or tinged with purple, lobes linear-lanceolate to ovate or narrowly triangular, 1.2 to 2.5 mm long, sparsely villous to glabrous; corolla rotate, lobes reflexed, white or tinged with purple or green, elliptic, 3.5 to 4.5 mm long, glabrous; gynostegium stipitate, glabrous; column more or less cylindrical, 0.7 to 1.1 mm tall, 0.5 to 0.8 mm wide; hoods greenish-white, broadly oblong, erect, 1.4 to 2 mm long, not fleshy, open in the upper portion, apex rounded, flat or slightly recurved, not surpassing the anther heads, margins entire, base not saccate; horns slender, pointed, attached in the lower 1/3 of the hood, arched over the anther head, 1.5 to 2 times longer than the hoods; anther head cylindrical, 1.5 to 1.8 mm long, 1.3 to 1.5 mm broad; anther apical appendages ca. 0.5 mm long; anther marginal wings abruptly rounded basally, minutely notched, ca. 1.4 mm long; corpusculum ca. 0.2 mm long; pollinia ca. 1 mm long. Mature pedicels straight to curved, not deflexed, follicles erect, narrowly fusiform, 7 to 10.5 cm long, 0.6 to 0.8 cm broad, sparsely puberulent or glabrous; seeds oval or broadly ovate, 5 to 6 mm long, coma white, 2.3 to 3.5 cm long. Usually in dry soils of prairies, open woods, thickets, dunes, roadsides, fencerows, etc. E. 1/2 TX; S. Can., MA to MT, S. to FL, TX, and AZ. Apr.-Aug.(Sept.)
This species is poisonous to livestock (GPFA 1986; Tull 1987). It was used by the Lakota tribes to stimulate milk production in nursing women (Kindscher 1992).
6. A. oenotheroides Cham. & Schlecht. Side-Cluster Milkweed, Hierba de Zizotes. Perennial from a thick rootstock or rhizome; stems few to several from the base, simple or branched from the base, moderately stout, 1 to 4.5 dm long or tall, ascending or decumbent, minutely puberulent. Leaves opposite or nearly so, petioles 0.5 to 2.5 cm long; blades ovate to oblong, deltoid, or occasionally slightly rhombic, 2.5 to 12 cm long, 1 to 6 cm wide, ascending to spreading, apically acute to rounded and often mucronate, base acute to obtuse or truncate, thin but firm, minutely puberulent, especially the undersurface. Inflorescences few to several, in the axils of the upper 1/2 to 2/3 of the stem; peduncles (0)0.1 to 1 cm long, minutely puberulent; pedicels slim, 1 to 2.5 cm long, puberulent; flowers ca. 6 to 18, 14 to 19 mm tall. Calyx lobes green(ish), ovate or ovate-lanceolate, 2 to 4 mm long, puberulent; corolla reflexed, lobes greenish-white or pale yellow, elliptic-lanceolate, 8 to 14 mm long, glabrous; gynostegium very shortly stipitate; column cylindric or broadly obconic, 1.5 to 1.7 mm long, 2.1 to 2.3 mm broad; hoods light greenish-cream, very slenderly oblong in the lower half, strongly flared above, attached in the lower 1/2 and spreading above, 7 to 10 mm long, a little fleshy, the upper portion open, apex flat, erose or repand, margin lobed in the upper 1/5, surpassing the anther head by as much as 3.5 to 4.5 mm, base deeply saccate; horn attached to the lower 3/4 of the hood, curved over the anther head, ca. 1.1 times longer than the hood; anther head truncate-conic, 2.5 to 3.7 mm tall, 3 to 4 mm broad; anther apical appendages ca. 1 mm long; anther marginal wings arched, very minutely notched at the midpoint, ca. 2.7 mm long; corpusculum ca. 0.3 mm long; pollinia ca. 1.6 mm long. Mature pedicels deflexed, follicles erect, broadly fusiform or ovoid, apiculate, 7 to 9 cm long, 1.5 to 2 cm wide, smooth, puberulent to glabrate; seeds obovate or oval, 6 to 8 mm long; coma pale tan, 2 to 3 cm long. Primarily on clay or rocky soils of hills, mesas, salt marshes, fields, roadsides, thickets, dunes, etc. W. 1/2 TX; common in our area; OK to SW. NM, S. to S. TX; also Mex. and Cen. Amer. Throughout the year under favorable conditions; ours mostly spring-fall.
7. A. tuberosa L. Butterfly Milkweed, Butterfly Weed, Pleurisy Root, Orange Milkweed, Chiggerflower. Perennial from a deep, thick rootstock; crown branched; sap not milky; stems 1 to many, simple or with a few branches (usually only in the inflorescence), rather stout, 3 to 9 dm tall, conspicuously villous to hirsute, the hairs usually 1 to 2 mm long. Leaves alternate or those just below the inflorescence subopposite or opposite, crowded, ascending to spreading; petioles 1 to 5 mm long; blades rather variable in shape, linear to ovate-lanceolate or oblanceolate, in our material usually narrow, (2)5 to 11 cm long, (0.4)0.7 to 3 cm broad, apex rounded to acute or acuminate, base cuneate to truncate or broadly cordate, firm, sparsely to densely villous or hirsutulous, especially below, margin often revolute, sometimes obscurely so, occasionally minutely crisped. Inflorescences 1 to several, terminal and subterminal on helicoid branches; cymes umbelliform; peduncles (0)0.1 to 3 cm long; pedicels slender, 12 to 19 mm long, pubescent; flowers 6 to 25, 11 to 15.5 mm tall. Calyx green or tinged with purple, lobes linear to lanceolate, 1.9 to 3.7 mm long, villous; corolla reflexed, orange or occasionally reddish or yellow, drying red, lobes elliptic to lanceolate, 5.5 to 8.5 mm long, glabrous; gynostegium stipitate, glabrous; column obconic, 1 to 2 mm long, 1 to 1.5 mm broad; hoods orange or rarely yellow, lanceolate, hooded, attached near the base, very slightly spreading, 4 to 5.8 mm long, not fleshy, open in the upper portion, apex narrow and rounded, slightly recurved, surpassing the anther head by 1.5 to 2 mm, margins slightly lobed in the lower 1/2, base not saccate; horns acicular (awl-like or subulate), attached to the lower 1/4 of the hood, arched over the anther head, 0.7 to 1.1 times the length of the hoods; anther head cylindric or truncate-conic, 2.2 to 2.6 mm tall, 2.1 to 2.4 mm broad; anther apical appendages ca. 0.5 mm long; marginal wings basally acute, without notches or spurs, ca. 1.6 mm long; corpusculum ca. 0.2 mm long; pollinia ca. 1.2 mm long. Mature pedicels deflexed, follicles erect, fusiform, 8 to 15 cm long, 1 to 1.5 cm broad, smooth, puberulent; seeds broadly oval, 5 to 7 mm long, coma white, 3 to 4 cm long. Open woods, dry fields, thickets, hillsides, canyons, dunes, prairies, etc. Over much of TX, especially the E. 2/3; most of the E. U.S., W. to AZ and NM. Apr.-Sept.
Hatch, et al. (1990) and the GPFA (1986) recognized 2 subspecies: subsp. interior Woods., with leaf bases deeply cordate [A. tuberosa L. var. interior (Woods.) Shinners and forma lutea (Clute) Steyerm.] and subsp. terminalis Woods. with leaf bases obtuse to truncate. Kartesz (1998) lists the two subspecies combined under subsp. interior. Our plants seem to have leaf bases mostly obtuse to truncate or only slightly cordate. I agree with the GPFA that these subspecific designations are of doubtful utility.
This plant is reportedly poisonous to livestock (GPFA 1986). Plains tribes used the raw root for lung disorders and also to treat wounds or sores. There a ritual associated with its gathering and preparation. Dakota tribespeople used the plant as an emetic (Kindscher 1992). It is also reported to have laxative properties and to be useful for heart trouble (Ajilvsgi 1984). The plant is a good addition to a butterfly garden as it is a favorite nectar source for a variety of species.
8. A. rubra L. Red Milkweed. Perennial from a tuberous-fusiform rootstock; stems slender, usually single and simple, 4 to 10 dm tall, glabrous or sparsely and minutely pilose in decurrent lines from the nodes. Leaves opposite, sessile or nearly so; blades narrowly lanceolate to broadly ovate, 5 to 16 cm long, 1.5 to 4(6.5) cm broad, acute to acuminate, base rounded to truncate or subcordate, thin but firm, glabrous, glaucous beneath, margin revolute. Inflorescences terminal and lateral, usually 2 to 4 from the (often) leafless upper nodes, commonly paired when terminal; peduncles ca. 10 to 30 cm long; pedicels filiform, 1 to 2 cm long, glabrous or minutely puberulent; flowers ca. 15 to 18 mm tall. Calyx green, lobes narrowly triangular, 2 to 3 mm long, glabrous; corolla reflexed, dull red, purple, or lavender, lobes oblong, acute, 8 to 9 mm long; gynostegium stipitate, pinkish-cream to purplish; column cylindric, ca. 2 mm tall and broad; hoods orangeish, lanceolate, acute or blunt, 6 to 7 mm long, slightly lobed at the base, not saccate, open in the upper portion; horns subulate, pointed, exserted from about the middle of the hood, not surpassing the hood, arched over the anther head; anther head narrowly conic, ca. 3 mm long and as broad; anther appendages less than 1 mm long; anther marginal wings broadest and slightly lobed below the middle, tapered above. Mature pedicels deflexed, follicles erect, 8 to 12 cm long, ca. 15 mm broad, narrowly fusiform, glabrous; seeds broadly oval, ca. 7 mm long, coma white, ca. 4 cm long. Bogs, wet meadows, marshes, and low pine woods, E. TX; known in our area at least from Robertson Co.; NJ S. to GA and FL, W. to PA and TX. May-Aug.
9. A. obovata Ell. Perennial from a deep, slender rootstock; stems simple or sparingly branched, 1.5 to 5(7) dm tall, tomentulose. Leaves opposite, petioles to 1 cm long; blades variable in size and shape, the lower ones oblong or quadrate to broadly oval, usually smaller and narrower above and sometimes the lowermost also smaller and narrower, ca. 3 to 9 cm long, 1 to 4 cm broad, firm, densely tomentulose, particularly below, apex acute to rounded, sometimes apiculate, base obtuse to truncate or rounded, margin commonly undulate. Inflorescences solitary and terminal or commonly also in the axils of the uppermost leaves, rather dense; peduncles from obsolete to ca. 5 mm long, densely tomentulose; pedicels slender to stout, 8 to 10 mm long, tomentose; flowers to ca. 15 mm long. Calyx green, lobes ovate-lanceolate, 4 to 5 mm long, minutely pilose; corolla yellowish-green (or tinged with purple), reflexed, lobes 8 to 10 mm long, ovate-lanceolate, minutely pilose externally, glabrous within; gynostegium short-stipitate, pale greenish-orange; column obconic, ca. 1.5 mm tall and 2 mm broad; hoods pale greenish orange, broadly oblong-oval, ca. 6 mm long, apex rounded and sometimes notched, the apex at least not fleshy, wide open apically, somewhat gibbous basally, slightly shorter than the anther head; horns projecting from the hood about halfway up, subulate, curved sharply over the anther head and about as long as the hoods; anther head truncate-conic, ca. 3 mm tall and 4 mm broad; apical anther appendages ca. 1 mm long, truncate; anther marginal wings broad and slightly lobed at the base, narrow above. Mature pedicels deflexed, follicles erect, to 12 cm or more long, fusiform, more or less tomentose. Sandy pine and oak woods, savannahs, fields, roadsides, etc. SE TX; SC to FL, W. to TX. May-Sept.
Perennial herbaceous twining vines from thick rootstocks; sap milky. Stems few to several. Leaves opposite, petiolate, blades triangular to ovate, entire, base rounded to cordate, usually with a few subulate glands on the midrib; stipules small, subulate, commonly deciduous. Inflorescences usually corymbose or umbellate axillary cymes, few- to many-flowered. Calyx lobes spreading, with linear to triangular glandular scales within, near or at the sinuses. Corolla rotate to short campanulate or funnelform, slightly to strongly spreading, white to dark purple, yellowish, or yellow-green. Corona a fleshy, shallowly lobed disk or cup or (as in ours) composed of 5 petaloid appendages which are apically bilobed. Gynostegium sessile to stipitate. Anther head depressed-spheric to conic; anther apical appendages ovate to suborbicular, petaloid; wings of anther margins strongly to shallowly angled at the base, conspicuous or inconspicuous, corneous (with the texture of thin horn); corpusculum elliptic to linear, red-brown; pollinia spatulate to oblong, nearly terete. Follicle fusiform, terete, smooth.
About 55 species of the tropics and temperate regions; 5 in TX; 1 here.
1. C. laeve (Michx.) Pers. Blue-vine, Sand-vine, Smooth Swallow-wort, Smooth Anglepod. Twining or trailing vine; stems simple or branched, to 3 m long or more, glabrescent or villous in lines. Petioles 1 to 9 cm long; blade triangular-lanceolate to deltoid or broadly ovate, (2)4 to 11 cm long, (1.5)2 to 10 cm broad, base cordate with a deep, wide sinus, the basal lobes rounded and commonly incurved, apex acuminate or acute, sometimes caudate or apiculate, margin minutely revolute, glabrous to sparsely villous or strigose, especially on the nerves; sometimes small, suborbicular leaves present in the axils. Inflorescences few to many, axillary, umbellate to corymbose (sometimes abbreviated racemes); peduncles 0.3 to 5 cm long, usually shorter than the petiole of the subtending leaf; pedicels slender, 3 to 12 mm long, villous; flowers 5 to 8 mm across. Calyx green or tinged with purple, lobes ovate-elliptic or lanceolate to ovate, 1.5 to 3 mm long, puberulent to sparsely villous, margins scarious; corolla rotate, whitish to cream-colored, the lobes narrowly oblong to oblong-lanceolate, 4 to 7 mm long, glabrous; corona petaloid, of 5 free, erect appendages 5 to 6 mm long, almost as long as the corolla lobes, each broadly ovate below and abruptly narrowed in the upper 1/2 into 2 free or somewhat fused linear lobes 1.5 to 2 times as long as the gynostegium; gynostegium stipitate (often only obscurely so); column ca. 0.5 mm tall, anther head conic; flowers dimorphic with regard to column and anther head, either column obconic and visibly distinct from the anther head which is 1.5 to 2 mm tall, 1.8 to 2 mm broad, with anther wings 0.5 to 0.6(0.7) mm long, or column cylindrical and smooth-fitting into the anther head which is 2.5 to 3 mm tall, 2 to 2.5 mm broad, with anther wings 1.5 to 2 mm long; anther apical appendages ca. 1 mm long; corpusculum 0.2 to 0.3 mm long; pollinia ca. 0.4 mm long. Follicles slender fusiform to lanceolate, slightly angled, 8 to 15 cm long, 1.5 to 2(3) cm thick, sparsely puberulent to glabrous; seeds 7 to 9 mm long, obovate, coma 3 to 4 cm long, white. Silty clay or sandy soils of moist low woods or fields, sometimes climbing on fences or shrubs. N. Cen. TX to S. Cen. coast; present in our area, but judging from collections, not as
common now as formerly; PA to NE, S. to GA and TX. June.-Sept. [Gonolobus laevis Michx.; Ampelamus albidus (Nutt.) Britt.; Enslenia albida Nutt.].
Herbaceous perennials or shrubs. Stems few to many from thick rootstocks, simple or branched, prostrate or twining to suberect. Leaves opposite, petiolate, blade cordate to ovate or suborbicular, basally cordate, entire, often with glands on the midrib; stipules none. Inflorescences axillary corymbose to umbellate cymes, or else flowers paired in the axils. Calyx lobes spreading, commonly with subulate to tubular glands on the inner surface, near the base or in the sinuses. Corolla rotate to campanulate, lobes convoluted in bud, slightly to strongly spreading, white to brown, purple, or greenish, often strongly reticulate-veined. Corona disk- or cup-shaped, thin or fleshy, variously lobed, with or without strap-shaped appendages within. Gynostegium sessile or with a short-stipe; anther head discoid, anthers partially hidden under the flattened stigma. Anther apical appendages suborbicular to ovate, petaloid; lateral wings inconspicuous, straight to curved; corpusculum narrowly elliptic to rhombic, red-brown; pollen in oblong to obovate pollinia. Follicles plump, fusiform, smooth or tuberculate, terete to angled. Seeds comose.
About 130 species, primarily in tropical S. Amer.; 13 TX; 4 here.
1. Stems twining, or at least the tips ...............................................................1. M. gonocarpos
1. Stems prostrate to more or less erect, not twining .................................................................2
2(1) Inflorescences on well-developed peduncles; corona of 2 rows of thin appendages .............
.........................................................................................................................2. M. parviflora
2. Inflorescences without peduncles, the pedicels arising directly from the axils; corona of a single fleshy, lobed disk ...........................................................................................................3
3(2) Pedicels shorter than or equalling the adjacent petiole; corolla usually pubescent within .....
...............................................................................................................................3. M. biflora
3. Pedicels (except perhaps the lowest) longer than the adjacent petiole; corolla glabrous within .........................................................................................................4. M. cynanchoides
1. M. gonocarpos (Walt.) Shinners Anglepod. Perennial, high-climbing twining or trailing vine; stems wiry, simple or branched, clothed with (sometimes sparse) hairs ca. 1 mm long, mixed with minute pale to purplish hairs less than 1 mm long. Leaves opposite, petioles 2 to 12 cm long, pubescent like the stem; blades wide-spreading, (4)6 to 17 cm long, 4 to 11 cm broad, broadly ovate to suborbicular, cordate, ovate-elliptic, or oblong-quadrate, base deeply cordate, the rounded lobes sometimes overlapping, apex abruptly acute or abruptly acuminate, thin-textured, sparsely and minutely hirsute, also with shorter hairs 0.1 to 0.2 mm long beneath. Inflorescences umbelliform, axillary, few to several; peduncles about equalling the pedicels, 1 to 2.5 cm long; pedicels slim, 1 to 3 cm long, usually glabrous; flowers usually many, conical and slightly twisted in bud, 1.7 to 2 cm across when open. Calyx green lobes lanceolate, 2.5 to 5 mm long, glabrous or with the apices ciliate; corolla rotate or very shortly campanulate, divided to near the base, lobes yellow to brownish- or greenish-purple, linear-lanceolate, obtuse to subacute, to 14 mm long, glabrous; corona a fleshy, flat disk, irregularly toothed or lobed, much shorter than the anther head; gynostegium short-stipitate; column obconic, 0.2 to 0.3 mm tall, ca. 1.3 mm broad; anther head discoid, ca. 0.5 mm tall, 0.2 mm broad; anther apical appendages ca. 0.2 mm long; anther marginal wings ca. 0.2 mm long, discoid; corpusculum ca. 0.2 mm long; pollinia ca. 0.5 mm long, widely spatulate. Follicles fusiform, (7)9 to 15 cm long, 2 to 2.8 cm broad, strongly 5-angled, glabrous; seeds ovate or obovate, 8 to 10 mm long, margin thin and erose, coma 3.5 to 4.5 cm long, white. Climbing on trees and shrubs, usually in thickets and along woodland streams. E. 1/2 TX; fairly common here; VA to KS and OK, S. to FL and TX. May-July. [Gonolobus gonocarpos (Walt.) Perry; Vincetoxicum gonocarpos Walt.; Gonolobus suberosus (L.) R. Br.].
2. M. parviflora (Torr.) Woods. Perennial from a thick rootstock; stems 1 to several, simple or sparingly branched, prostrate and spreading; herbage more or less white pilose and said to be malodorous. Leaves opposite, petioles to 15 mm long; blades cordate-ovate to suborbicular or broadly ovate-lanceolate, base truncate to rounded or shallowly cordate, apex broadly obtuse to acute, to 5.5 cm long (usually shorter) and about as wide, major veins often arising from the base. Inflorescences more or less racemose, axillary and/or terminal or sometimes flowers in scattered clusters; peduncles to 7 cm long; pedicels to 5 mm long. Calyx green, lobes linear-oblong to elliptic-lanceolate, obtuse, 2 to 3 mm long; corolla divided to near the base, greenish or brownish, lobes ovate to linear-oblong or elliptic, obtuse, reflexed or strongly spreading, scarious-margined, puberulent only within at the very base, 3 to 5 mm long; corona appendages in 2 rows, the outer row slightly to much longer than the stamen column, longer than wide, the apex of each with 2 prongs or points or occasionally some with only a single point, inner row shorter. Follicles to 9 cm long and 2 cm broad, ellipsoid, puberulent and more or less muriculate. Sandy soils of open woods, prairies, and mesquite plains; known from near Flynn in Leon Co.; Mostly on the Rio Grand Plains; perhaps endemic. Mar.-Aug. [Vincetoxicum parviflorum (Torr.) Heller].
3. M. biflora (Raf.) Woods. Two-flower Milkvine. Perennial from a thick vertical rootstock; stems several to many, prostrate to suberect, 1 to 4 dm long, simple or branched; herbage pilose with hairs to 1 mm long and also with minute, pale glandular hairs ca. 0.1 mm long. Leaves petiolate, 0.3 to 2.5 cm long, blades suborbicular-ovate to triangular ovate, base subtruncate or shallowly to deeply cordate, apex acute to broadly obtuse, 1.5 to 5 cm long, 1 to 3 cm broad. Flowers mostly paired in the axils of the upper leaves; peduncles none or to only 2 mm long; pedicels 0.5 to 1 cm long; flowers 1.3 to 1.7 cm across. Calyx green, lobes ovate to lanceolate, 2 to 3.5 mm long, villous; corolla rotate, red-purple to deep brown internally, sometimes greenish beneath, lobed to near the base, lobes elliptic-oblong or oblong-lanceolate, spreading, 5.5 to 7 mm long, moderately to densely pilose on the inner surface; corona a fleshy disk with 5 broad, inflexed lobes, the lobes triangular below, elongate above into an oblong, truncate apex, much surpassing the anther head; gynostegium sessile, glabrous; column absent; anther head discoid, ca. 0.5 mm tall, ca. 1.8 mm broad; anther apical appendages ca. 0.4 mm long; anther marginal wings ca. 0.3 mm long; corpusculum ca. 0.3 mm long; pollinia ca. 0.3 mm long, obovate. Follicles broadly fusiform to ellipsoid, not strongly angled, 7 to 10 cm long, 1.5 to 2.5 cm broad, moderately to densely pubescent, muricate; seeds orbicular, 9 to 10 mm long, coma 2.5 to 3.5 cm long, white to tan. Sandy, rocky, or clayey soils of open woods and grasslands; in our area often associated with outcrops. N. Cen. TX, S. to the Ed. Plat. and NW. to the S. Plains; also in OK. Mar.-June. [Vincetoxicum biflorum (Raf.) Heller].
4. M. cynanchoides (Engelm.) Woods. Milkvine. Perennial from a thick vertical rootstock; stems few to several, prostrate to suberect, simple or with a few branches below, 1.5 to 5 dm long; herbage pilose with hairs to 1 mm long and also very small pale to purplish glandular hairs ca. 0.1 mm long. Leaves with petioles 0.3 to 2.5 cm long; blades suborbicular-ovate to broadly lance-ovate, 1 to 6.5 cm long, 1 to 4 cm broad, base shallowly to deeply cordate, apex obtuse to acuminate. Flowers mostly paired in the axils of the upper leaves; peduncles usually none or only 0.2 to 1 cm long; pedicels slim, 0.2 to 1 cm long; flowers 0.8 to 1.2 cm across. Calyx green, lobes lanceolate or ovate to elliptic, 2.5 to 3.5 mm long, villous; corolla rotate, brownish or dark maroon above, green to brown below, deeply lobed, lobes lanceolate to ovate or ovate-elliptic, obtuse, with more or less scarious margins, 3.5 to 4 mm long, glabrous within, pilose to glabrescent externally; corona a shallow cup-shaped fleshy disk, broadly and shallowly 5-lobed, lobes little if at all longer than the anther head, each lobe with an ascending, fleshy, tongue-like protuberance within which is shorter than the lobe; gynostegium short stipitate, glabrous; column obconic, 0.2 to 0.4 mm tall, ca. 1 mm broad; ; anther head discoid, ca. 0.7 mm tall, ca. 1.8 mm wide; anther apical appendages ca. 0.3 mm long; anther marginal wings ca. 0.3 mm long; corpusculum ca. 0.3 mm long; pollinia ca. 0.4 mm long, semicircular. Follicles ellipsoid to fusiform, 7 to 8 cm long, 2 to 2.5 cm broad, sparsely to moderately pubescent, more or less muricate, not strongly angled; seeds orbicular, ca. 1 cm long, coma 2 to 2.5 cm long, tan. Sandy soil, usually in open woods and on roadsides. NE TX., S. to the coast, W. to the S. Plains; also OK. Apr.-Aug. [Vincetoxicum cynanchoides (Engelm.) Heller].
Ours annual or perennial herbs or shrubs, sometimes vine-like (elsewhere also trees). Leaves alternate or sometimes in fascicles, occasionally in approximate pairs, simple to odd-pinnately compound; stipules none. Inflorescences terminal, subterminal, axillary, opposite the leaves, or on the internodes, the arrangement cymose, paniculate, umbellate, racemose, or sometimes flowers solitary. Flowers perfect, regular or essentially so, 4- to 6-merous, ours usually 5-merous except for the gynoecium. Calyx synsepalous, usually with 5 teeth or lobes, rotate to campanulate or tubular, usually persistent in fruit, sometimes accrescent or inflated at maturity. Corolla sympetalous, rotate to campanulate, tubular, funnelform, or urceolate, the limb with 5 lobes or teeth or sometimes entire, the lobes valvate or imbricate, usually plicate (pleated) in bud. Stamens usually 5, free, inserted on the corolla and alternate with the lobes, sometimes connivent around the style, anthers opening by longitudinal slits or terminal pores. Gynoecium superior, of 2 united carpels; style 1, terminal, stigma entire or 2-lobed, locules 2 or sometimes 4 because of false septa, sometimes lobed; ovules several to many. Fruit with axile placentation, a berry or capsule. Seeds with the embryo lying near the periphery, often curved, endosperm well-developed.
This is a large family with about 90 genera and 12,600 species nearly worldwide; especially common in S. Amer. 18 genera and 78 species in TX; 8 genera and 24 species here.
The family is important for food crops, including potatoes (Solanum tuberosum), eggplant (S. melongena), tomato (Solanum lycopersicum L. = Lycopersicon esculentum), and peppers (Capsicum spp.). Many species have an alkaloid chemistry, including tobacco (Nicotiana tabacum) and the poisonous members Hyoscyamus (Henbane), Atropa (Belladonna), Datura (Jimsonweed), and Mandragora (Mandrake). These and others also have medicinal uses. Many taxa are grown as ornamentals, including Brugmansia, Cestrum, Nicotiana, Schizanthus, etc. (Mabberley 1987).
NOTE: In addition to the taxa included in the key below, several others deserve mention:
Solanum capsicastrum Link, False Jerusalem Cherry, is S. Amer. native grown for ornament and occasionally encountered as an escape. See NOTE at Solanum.
Solanum lycopersicum L. (=Lycopersicon esculentum Mill.), the cultivated tomato, is sometimes found near old homesites or in dumps, but is not a permanent member of our flora. It has pinnately compound leaves and yellow corollas with recurved lobes.
Bouchetia erecta DC. is found just S. of our area and may someday be found here. It is a low herb with ascending stems, white funnelform flowers ca. 12 to 18 mm long, and capsules. Sheets from our area identified as Bouchetia have proved to be misidentified.
Petunia parviflora Juss. is scattered throughout TX in moist or wet soils of beaches and mudflats, but no specimens from this area have been seen. It is a prostrate herb, rooting at the nodes, with fleshy, spatulate leaves ca. 1 cm long, red-purple funnelform flowers ca. 6 to 7 mm long, and capsules. Cultivated petunias are of hybrid origin and may be found as occasional waifs, but do not persist or escape in our area.
1. Fruit a berry; corolla rotate, campanulate, urceolate, or sometimes funnelform ..................3
2(1) Corolla 6 cm or more long; capsule prickly .............................................................1. Datura
2. Corolla less than 6 cm long; capsule smooth .....................................................2. Nicotiana
3(2) Plants shrubs, usually with thorny branches ...........................................................3. Lycium
3. Plants herbaceous, or if shrubby then without thorns (prickles may be present) ..................4
4(3) Calyx enlarged in fruit to enclose all or nearly all of the berry, not spiny ...............................5
4. Calyx not enlarged in fruit, or if enclosing the berry then spiny ..............................................6
5(4) Calyx inflated and bladdery at maturity, commonly angled; corolla without tomentose pads on the lower part of the lobes ................................................................................4. Physalis
5. Calyx closely fitted around fruit, not inflated or angled; corolla with tomentose pads on the lower part of the lobes, alternate with the filaments ................................5. Chamaesaracha
6(4) Corolla urceolate; plants more or less climbing; anthers dehiscent by longitudinal slits ........
............................................................................................................................6. Salpichroa
6. Corolla rotate; plants not usually climbing, or if so, then anthers dehiscing by terminal pores .........................................................................................................................................7
7(6) Anthers dehiscent by terminal pores .....................................................................7. Solanum
7. Anthers dehiscent by longitudinal slits ................................................................8. Capsicum
Ours annual or perennial herbs with rank-smelling foliage. Stems erect, ascending, or decumbent. Leaves alternate, blades ovate to elliptic, entire to sinuate-pinnatifid, glabrous to variously pubescent. Flowers large, showy, in ours pale, solitary, produced on short pedicels in the forks of the stem, usually erect. Calyx tubular or angled-cylindric, 5-toothed, dehiscent circumscissilely above the base after anthesis, the remaining disk enlarging somewhat and persisting beneath the capsule. Corolla funnelform to tubular, much longer than the calyx, the limb 5- or 10-toothed, convolute-plicate in bud. Stamens 5, equal, anther dehiscence longitudinal. Style about as long as the anthers, stigma capitate, 2-lobed or -lipped; ovary 2-celled or 4-celled because of a false septum. Capsule erect or nodding, dehiscent regularly by 2 or 4 apical valves or else splitting irregularly, the surface usually prickly. Seeds many, flattened.
As treated here without Brugmansia (flowers nodding, some taxa woody), 8 species of S. N. America, but some widely naturalized; 4 listed for TX (Kartesz 1998) with the synonymy somewhat confused; 3 collected locally.
Members of the genus have a powerful alkaloid chemistry. Some were/are used as ritual or sacred hallucinogens by Native Americans (Mabberley 1987). All parts of the plants are potentially poisonous, including the nectar. Most reported poisonings involve the deliberate use of seeds or leaves in an attempt to produce intoxication (Lampe 1985). Some species are cultivated for their showy flowers (Bailey, et al. 1976).
1. Fruiting pedicels erect; capsule regularly dehiscent via 4 valves; corolla 6 to 8 cm long .......
......................................................................................................................1. D. stramonium
1. Fruiting pedicels recurved; capsule irregularly dehiscent; corolla longer than 10 cm ...........2
2(1) Plants cinereous with minute curved and/or appressed hairs, the lower leaf surfaces remaining pubescent and velvety in age .............................................................2. D. wrightii
2. Plants villous or glandular-villous (especially new growth), but the lower leaf surfaces quickly becoming glabrate except for the major veins .........................................3. D. inoxia
1. D. stramonium L. Jimson Weed, Jamestown Weed, Tolache. Taprooted annual; stems simple or with spreading branches mostly in the upper region, to 1.5 m tall; herbage sparsely pubescent to glabrate. Leaves with petioles 2 to 9 cm long, ca. 1/2 the length of the blade; blades ovate to elliptic or lance-ovate, 5 to 25 cm long, 2.5 to 15(20) cm broad, acuminate, base cuneate to subtruncate, margin irregularly sinuate-dentate or sinuate-laciniately lobed. Pedicels 0.5 to 1.5 cm long; calyx tubular, 3.5 to 5 cm long, with 5 unequal teeth 5 to 10 mm long, the persistent calyx disk reflexed; corolla white or tinged with violet, 6 to 8(10) cm long, the limb 3 to 5 cm broad, the 5 lobes ending in subulate teeth 3 to 8 mm long; stamens 5 to 8 cm long, anthers white or violet, 3.5 to 5 mm long, sparsely pubescent. Mature pedicel and capsule erect, capsule ovoid, 3.5 to 5 mm long, finely puberulent to glabrate, the surface densely covered with prickles usually 3 to 5(9) mm long (occasionally nearly smooth), regularly dehiscent via 4 valves; seeds black, subreniform, 3 to 4 mm long, rugulose and finely pitted. Waste places, cultivated areas, bottomlands, etc., the seeds capable of long dormancy, often weedy. Widespread in temperate and tropical regions of the world. Apr.-Nov.
Plants with violet or lavender flowers and violet anthers have been designated var. tatula (L.) Torr. The color difference is the result of a single gene mutation (Mabberley 1987). Some sources feel that distinction at the varietal level is not justified (e.g. GPFA 1986).
The plants contain stramonium, a substance used in the treatment of asthma (Mabberley 1987).
2. D. wrightii Regel. Indian Apple, Sacred Datura. Perennial from a rootstock; stems erect, widely branched, to 1.5 dm tall; herbage cinereous with minute appressed and/or curled hairs, velvety to the touch and the leaf undersurfaces remaining so even in age, sometimes also glaucescent. Leaf blades ovate, to ca. 15(20) cm long, obtuse to acute or short acuminate, base commonly asymmetrical and more or less truncate or rounded, margin irregularly sinuate-repand; petiole shorter than to almost as long as the blade. Flowers erect or slightly nodding, pedicels to ca. 3 cm long. Calyx 7 to 12 cm long, with 5 unequal teeth, densely cinereous, the persistent disk reflexed; corolla white, commonly tinged with lavender or violet, to ca. 15(20) cm long, the limb spreading, to ca. 15 cm broad, the 5 lobes with teeth 1 to 1.5 cm long; stamens to ca. 15 cm long, anthers white, pubescent, ca. 15 mm long. Mature pedicel reflexed, capsule globose, (2.5)3 to 4 cm broad, irregularly dehiscent, the surface puberulent and densely prickly, prickles less than 1 cm long; seeds many, light brown, orbicular-reniform with a cord-like margin, 5 to 6 cm broad, smooth or minutely pitted. Mostly in sandy soils of floodplains, bottomlands, bogs, and so on. E. TX to the T.P.;
TX to CA and N. Mex. May-Nov. [D. meteloides of TX authors, e.g. DC., but not D. meteloides Dunal; D. metel L. var. quinquecuspida Torr.].
NOTE: Some botanists consider D. wrightii and D. inoxia (below) to be conspecific (see, for example, GPFA 1986). If this proves to be the case, the name D. inoxia has priority.
3. D. inoxia P. Mill. Indian Apple. Shoots annual from a perennial rootstock; stems widely branched, to 1 m tall or more; herbage (especially the new growth) densely villous and often also with spreading glandular hairs. Leaf blades ovate, to 25 cm long, acute to acuminate, base asymmetrical, truncate to rounded, margin entire to coarsely sinuate-dentate, lower surfaces soon glabrate except for the major veins; petioles shorter than to about as long as the blades. Flowers usually erect, calyx tubular, 8 to 12 cm long, with 5 unequal teeth, the persistent portion reflexed; corolla white, 12 to 15 cm long, the spreading limb to ca. 12 cm broad, with 5 (rarely 10) subulate teeth ca. 1 cm long. Mature pedicel reflexed, capsule globose, 3 to 4 cm in diameter, splitting irregularly, the surface glandular-puberulent and shortly villous, densely prickly, the prickles mostly less than 1 cm long; seeds light brown, 5 to 6 mm broad, reniform. Stream beds, canyons, bluffs, ledges, etc. Ed. Plat., Rio Grande Plains, and the T.P.; included here on the basis of TAMU 010623, which keys to this species; TX and NM , S. to Cen. Amer. Mar.-Nov. [Specific epithet sometimes given as "innoxia"].
See NOTE at D. wrightii, above.
Annual or perennial herbs, shrubs, or small trees, usually heavily scented and viscid-pubescent. Leaves alternate, entire to repand or pandurate, sessile or petiolate. Flowers few to many in panicles or racemes. Calyx tubular-campanulate, with 5 teeth or lobes. Corolla funnelform or salverform, the tube usually relatively long, the limb 5-lobed, usually spreading, plicate in bud. Stamens 5, inserted on the corolla. Stigma capitate. Capsule ovoid to slenderly ellipsoid, blunt or acute, 2-celled, dehiscent by 2 or 4 apical valves. Seeds many, tiny.
About 67 species of the Americas, S. Pacific, Australia, and SW. Afr.; 6 in TX; 2 confirmed from our area.
The most important species is N. tabacum which provides smoking and chewing tobacco in all its forms. One of its principal chemical components is nicotine, which is also used as an insecticide (Mabberley 1987). This and a number of other species (including our own N. glauca) are poisonous, having various alkaloid chemistries that act on the nervous system (Lampe 1985). Other species, notably N. alata, are grown as ornamentals for their showy and/or fragrant flowers.
1. Plants shrubs or shrublike; leaves glabrous and glaucous; flowers yellow ........1. N. glauca
1. Plants herbs; leaves pubescent; flowers mostly white. ....................................2. N. repanda
1. N. glauca Grah. Tree Tobacco, Mustard Tree, Rapé, Gigante, Buena Moza, Tronadora. Shrub or small tree to 8 dm (as much as 3 m farther south); herbage glabrous. Leaves long-petiolate, blades ovate to oblong-lanceolate, 4 to 18 cm long, obtuse to acute, base cuneate to subcordate, entire or slightly repand, more or less leathery, glaucous. Flowers in loose terminal racemes or panicles. Calyx short-tubular to tubular-campanulate, 8 to 12 mm long, with 5 slightly unequal teeth much shorter than the tube, with tiny white dots as seen with strong magnification; corolla 3.5 to 4.5 cm long, tubular and only slightly lobed, slightly constricted just below the limb, limb spreading little if at all, greenish yellow, minutely villose; filaments attached below the middle of the corolla tube, usually not exserted. Capsule ovoid, partially enclosed by the persistent calyx, 1 to 1.2 cm long, acute, dehiscent via 4 apical valves; seeds oblong-quadrangular, red-brown, faintly lustrous, minutely reticulate or pitted. Sandy or clay soils of stream banks, talus slopes, roadsides, and ledges. Coastal and S. TX, W. to the T.P.; known from our area, possibly originally from cultivation; native to S. Amer., naturalized N. to TX and CA. Flowering throughout the year; our collections from Feb. onwards.
This plant is poisonous, the particular toxin being the alkaloid anabasine. Human fatalities have resulted from ingestion of raw or cooked leaves (Lampe 1985; Tull 1987).
2. N. repanda Willd.ex Lehm. Fiddle-leaf Tobacco, Wild Tobacco, Tabaco Cimarrón. Taprooted annual; stem usually single from the base, to ca. 9(17) tall, the upper portion with uncrowded, slender branches; herbage minutely pubescent or glabrate above. Lower leaves more or less rosulate, contracted into a winged petiole, upper leaves cordate-clasping or auriculate, blades to 20 cm long and 10 cm broad, more or less ovate or the lower ones obovate, commonly repand, often pandurate (obovate and pinched in on each side), apices obtuse to acuminate, reduced up the stem into more or less cordate bracts, these often minute or absent at the top of the plant. Inflorescence a loose panicle or raceme, essentially naked, the flowers opening at dusk. Calyx tube short-campanulate, strongly 10-ribbed, with 5 slender lobes as long as the tube, acute to somewhat blunt; corolla long-funnelform, white or sometimes tinged with rose or the veins brown, tube as much as 5 or 6 cm long, slightly expanded in the throat below the limb that spreads to 4 cm across, lobes short, obtuse to acute; anthers positioned in the expanded part of the throat. Capsule ovoid, ca. 1 cm long, dehiscing by 4 apical valves; seeds brown, shiny or dull, reticulate to pitted. Clay or sandy soils along streams, in depressions and on flats, in wooded ravines, thickets, pastures, roadsides, etc.; sometimes weedy. Ed. Plat. and S. TX; known from our area, the weedy collections from the TAMU campus possibly escapes from a greenhouse; also in N. Mex. Feb.-July. [N. roemeriana Scheele].
Small shrubs. Stems usually thorny and with short spur shoots. Leaves mostly fascicled, in TX material elongate, entire, sometimes somewhat fleshy. Flowers axillary, solitary or in small clusters. Calyx campanulate, with 4 to 6 regular or irregular teeth or lobes, sometimes somewhat bilabiate. Corolla campanulate, tubular-funnelform or salverform, the limb with 4 to 7 lobes. Stamens 4 or 5. Stigma 2-lobed or capitate. Fruit a dry or fleshy berry, often red, globose to ovoid, subtended by the persistent calyx which commonly splits irregularly as the fruit enlarges.
About 85 to 100 species of warm temperate areas, especially the Americas; 7 species in TX; apparently 1 here.
The berries of some are edible, but the leaves may be toxic, especially to livestock (Correll & Johnston 1970; Lampe 1985).
1. L. carolinianum Walt. var. quadrifidum (Dun.) C. L. Hitchc. Carolina Wolf-berry. Small shrub, sparingly short-branched and only sparingly thorny; stems erect to slightly spreading, to 1 m tall; herbage glabrous; young branchlets with short thorns to 1 cm long, older branchlets silvery, with spinose branchlets. Leaves rather succulent, usually in fascicles of 3 to 10, essentially spatulate, 0.7 to 2.5 cm long, 1 to 2(5) mm broad, apex rounded to acute, tapered to a sessile base, midvein scarcely or not visible. Flowers usually solitary, pedicels to 3 cm long (usually much shorter). Calyx tube 2.5 to 3.5 mm long, lobes 4, 1/3 as long as the tube, 1 to 2 mm long, usually more or less equal, triangular, obtuse, margins sometimes sparsely ciliate; corolla lavender to purple or blue-violet, 7 to 10 mm long, rotate-campanulate, tube 1 to 1.5 mm in diameter at the top of the ovary, 3 to 5 mm in diameter at the orifice, lobes 4 (or 5), spreading, equalling or longer than the tube, 4 to 6 mm long, ovate, abruptly narrowed at the base, apically rounded to slightly emarginate, glabrous; stamens exserted because the lobes spread, equalling or slightly shorter than the corolla lobes, pilose in the lower 1/3 to 1/4 and the corolla tube sparsely hairy nearby, anthers 1 to 1.5 mm long; style about as long as the filaments or a little shorter. Berry red, ovoid, fleshy, 0.8 to 1.5 cm long, ca. 1 cm in diameter, dark purple in old or dry material; seeds 50 or more. Near ponds, ditches, or marshes, on wet clay or salt flats, and in sandy-gravelly soil on brushy hills. Coastal and S. TX; MS to TX and NE. Mex. Jan.-Nov.
Uncommon in our area and included on the basis of two specimens, one of which is not determined with complete confidence. In identifying Lycium specimens, the length of the calyx lobes and their symmetry at anthesis is important, as is the length of the corolla lobes. In fruiting specimens, the calyx often splits and appears somewhat bilabiate so that the original length of the calyx teeth is not evident. Since the genus is poorly known from our area, the reader is referred to Correll and Johnston (1970) for any local material not matching the above description exactly (i.e., with an asymmetrical calyx, 5 or more calyx lobes, or calyx teeth less than 1/3 the length of the tube). L. berlandieri Dun. var. berlandieri would be the taxon most likely to occur here other than L. carolinianum.
Annual or perennial herbs, some from rhizomes. Stems erect to decumbent, usually branched. Leaves alternate or occasionally 2 or 3 together where internodes reduced, petiolate; blades generally ovate to lance-linear, entire to toothed or sinuate, often irregularly so, rarely more deeply lobed. Pubescence of stem and foliage various, from glabrous to sparsely to densely pubescent with simple, jointed, branched, stellate, or glandular hairs or a mixture of any of these. Flowers, in ours, solitary in the axils, pedicellate, usually nodding at anthesis. Calyx campanulate, 5-toothed, small at anthesis and enlarging greatly to enclose the fruit. Corolla campanulate to rotate, often reflexed, shallowly 5-lobed or entire, usually some shade of yellow, often with dark brown or black spots within or these faint or absent, also commonly pubescent within. Stamens 5, inserted near the base of the corolla tube, erect, not connivent around the style, filaments from filiform to about as broad as the anthers, in some taxa clavate, anthers yellow to blue or violet or sometimes edged or tinged with blue or violet. Style slender, stigma scarcely broader. Mature calyx (in fruit) strongly 5-angled or else 10-angled or nearly round, sometimes indented basally, glabrous to pubescent, usually strongly inflated and wholly enclosing the berry. Berry sessile or short-stipitate, dryish to juicy, globose, bilocular, many-seeded. Seeds yellow, glabrous, often minutely pitted.
About 80 species worldwide, especially common in Mex. and Cen. Amer.; 16 known from TX; 9 here. This treatment follows those of Waterfall (1958) and Sullivan (1985).
Some have brightly colored calyces and are cultivated for ornament, often with common names referring to Chinese or Japanese lanterns. The berries of some are edible, particularly those of P. philadelphica and P. pruinosa, known as tomatillos or husk tomatoes (Mabberley 1987). Several species were grown by Native Americans for food (Kindscher 1987). The unripe berries of some taxa, however, contain solanine glycoalkaloids and probably all unripe fruit should be considered potentially poisonous (Lampe, 1985).
NOTE: The key below is only for our immediate 7-county area; material from elsewhere should be referred to the Correll and Johnston (1970) and to Sullivan (1985) for identification and to a recent checklist such as that of Hatch, et al. (1990) for current nomenclature. Confident identification requires flowers at anthesis, mature capsules, and the presence of underground structures (or absolute knowledge of their form). Powerful magnification is needed to examine the pubescence.
1. Plants taprooted annuals; anthers usually violet or blue; flowers ca. 4 to 10(12) mm long; anthers ca. 1 to 2.5 mm long ...................................................................................................2
1. Plants perennial (roots often not collected); anthers yellow or sometimes tinged with blue; flowers (10)12 to 20 mm long; anthers 2 to 4 mm long .........................................................3
2(1) Stem glabrous; pedicels glabrous or only slightly pubescent; corolla not or only faintly spotted; fruiting calyx 10-angled .......................................................................1. P. angulata
2. Stem sparsely to densely pubescent; pedicels obviously pubescent; corolla dark-spotted; fruiting calyx 5-angled ...................................................................................2. P. pubescens
3(2) Pubescence all or mostly of stellate or branched hairs (simple or jointed hairs may also be present) .....................................................................................................................................4
3. Pubescence mostly of simple straight, jointed, or glandular hairs, branched hairs (if any) small and much less abundant than unbranched ones ..........................................................6
4(3) Hairs primarily jointed, spreading at right angles, some also 1- to 3-branched ......................
...............................................................................................................................3. P. pumila
4. Hairs all or nearly all stellate ....................................................................................................5
5(4) Flowering calyces and lower leaf surfaces densely pale-tomentose, on young leaves the lower surfaces obscured by hairs ..........................................................................4. P. mollis
5. Flowering calyces and lower leaf surfaces sparsely to densely pubescent, but the hairs not obscuring the leaf surfaces ..................................................................5. P. cinerascens
var. cinerascens
6(4) Stems or leaves usually with some hairs 1- to 3 branched .................................3. P. pumila
6. Stems or leaves without branched hairs (jointed hairs may be present). ..............................7
7(6) Midstem with glandular hairs, pubescence usually moderate to dense ................................8
7. Midstem without glandular hairs, nearly glabrous to moderately pubescent with simple hairs ...........................................................................................................................................9
8(7) Flowering pedicels usually 3 to 8(10) mm long; stem pubescence usually of dense, short hairs and perhaps some longer, spreading hairs ..........................................6. P. hederifolia
var. hederifolia
8. Flowering pedicels usually 10 to 15 mm long; stem pubescence usually of short,
glandular hairs mixed with longer, spreading hairs 0.6 to 2 mm long ......7. P. heterophylla
var. heterophylla
9(7) Hairs of the midstem wholly or partly reflexed or decurved, occasionally villous; corolla (10)14 to 18 mm long .....................................................................................8. P. virginiana
9. Hairs of the midstem and petioles antrorse; corolla 10 to 14(18) mm long ..9. P. longifolia
1. P. angulata L. Cutleaf Ground-cherry, Southwest Ground-cherry, Lanceleaf Ground-cherry, Purple-vein Ground-cherry. Taprooted annual; stems 1 to 5(10) dm tall, erect or sometimes decumbent, branched at the base or above, glabrous or with a few antrorse, appressed hairs, especially on new growth. Leaves with petioles ca. 1 to 4(8) cm long; blades varying from ovate to ovate-lanceolate, linear-lanceolate, oblongish, or elliptic, the larger ones (3)4 to 11 cm long, 3.5 to 8 cm broad, margin usually deeply and irregularly toothed or incised- or undulate-toothed, occasionally entire, glabrous or with sparse appressed hairs. Pedicels at anthesis 5 to 40 mm long, elongating to 20 to 45 mm in fruit, glabrous or only sparsely pubescent; calyx at anthesis 3 to 5 mm long, 2 to 4 mm broad at the base, with lobes 1 to 3 mm long; corolla yellowish, without spots or with only indistinct ones, 4 to 10(12) mm long; anthers usually bluish or violet (rarely yellow, at least in dry material), 1 to 2.5 mm long, filaments slender, 3 to 5 mm long. Fruiting calyx 2 to 3.5 cm long, 10-angled or 10-ribbed, usually nearly round, 1.5 to 2.5 cm broad, glabrous or essentially so; berry 10 to 12 mm in diameter, on a stipe ca. 1 mm long; seeds ovate to widely elliptical, nearly smooth. Open woods, disturbed areas, along creeks or streams, on roadsides, in ditches or swales, etc. E. TX, W. through SW.TX, throughout most of the state except the High Plains; tropical Amer., N. to VA, MO, IL, KS, and TX. Flowering throughout much of the year; our collections mostly May-Oct. [Includes var. angulata, var. lanceifolia (Nees) Waterfall, and var. pendula (Rydb.) Waterfall; P. pendula Rydb; P. lanceifolia Nees; P. ramosissima Mill.; P. capsicifolia Dun., P. linkiana Nees].
2. P. pubescens L. Downy Ground-cherry, Tomate Fresadilla. Taprooted annual; stems 8 to 90 cm tall, erect, widely branched, villous and/or viscid to more or less glabrate. Leaves with petioles 1 to 7(10) cm long; blades ovate (to cordate), the larger 3 to 10 cm long, 2 to 4 cm broad, entire or with few to several irregular teeth or repand-dentate, apex acute to acuminate, base acute to rounded, often oblique, sparsely to densely pubescent, with or without sessile glands, sometimes translucent. Pedicels (3)5 to 7(10) mm long at anthesis, elongating slightly in fruit, decidedly pubescent; calyx at anthesis 4 to 10 mm long, 3 to 12 mm broad at the base of the lobes, lobes 1 to 4 mm long, ovoid-deltoid to lanceolate; corolla yellowish, dark-spotted, (6)8 to 10(12) mm long, 10 to 15 mm broad, more or less densely pubescent below the spots inside; anthers bluish or violet, 1 to 3 mm long, filaments 2 to 3 mm long. Fruiting calyx usually strongly 5-angled, (15)18 to 25(30) mm long, (10)13 to 18(20) mm wide, indented at the base, usually softly pubescent (if only sparsely so), the lobes usually triangular or triangular-ovate, sometimes lance-subulate; berry 10 to 18 mm in diameter, sessile or subsessile; seeds broadly elliptic to ovate, minutely pitted. Woods, thickets, river or stream banks, railways, roadsides, etc. in the E. 1/2 of the state.
Several varieties have been described, though some authorities feel that they intergrade too much to be justifiable (e.g. GPFA 1986). If varieties are recognized, our plants belong to the following two:
var. pubescens Plants more or less villous; leaves usually with 5 to 8 teeth on each margin, blades not translucent. S. U.S.; also Mex. and W. I. Summer, ours mostly May-July. [P. barbadensis Jacq.; P. floridana Rydb.].
var. integrifolia (Dun.) Waterfall Plants not as hairy as in var. pubescens; leaf blades commonly entire or with only 3 or 4 weak teeth on each margin; blades usually translucent. E. 1/2 TX; S. U.S.; also Mex., Cen. Amer., and W.I. Apr.-Nov. [P. pruinosa of authors but not P. pruinosa L.].
3. P. pumila Nutt. Prairie Ground-cherry, Low Ground-cherry. Perennial from a deep rootstock, sometimes colonial from the roots; stems erect, 1.5 to 4.5 dm tall, branched; herbage with jointed hairs 1 to 2 mm long, at least some (commonly those of the leaf blades) 1- to 3-branched, spreading at right angles (or rarely herbage subglabrous and merely appressed-pubescent on buds, calyces, and leaf margins). Leaves with petioles more or less narrowly winged; blades ovate to ovate-lanceolate or lanceolate, sometimes slightly rhombic, entire or sometimes slightly and irregularly sinuate-dentate, the larger ones 4 to 8 cm long. Pedicels at anthesis 15 to 30 mm long, elongating to (15)25 to 40 mm in fruit; calyx at anthesis 10 to 15 mm long, with lobes 3 to 5 mm long; corolla yellowish, only faintly spotted, 12 to 20 mm long, 15 to 25 mm across; anthers yellow, 2.5 to 3 mm long. Fruiting calyx (2) 3 to 4 cm long, 15 to 20 mm broad, usually strongly inflated; berry (10)12 to 15 mm in diameter; seeds broadly elliptic, 2 to 2.5 mm long, minutely reticulate or pitted. Open woods, prairies, and roadsides. NE TX., W. to Brazos Co; E. OK, KS, and MO to AR and IL; not especially common in our area. Apr.-June.
4. P. mollis Nutt. Perennial from a deep rhizome, often with slender, shallower rhizomes; stems erect, 1.5 to 5 dm tall, branched; herbage densely tomentose with branched or stellate hairs 1 mm long or less, occasionally also with multicellular, unbranched or branched hairs 2 to 4 mm long, the hairs especially dense on new growth, on the undersides of young leaves and young calyces obscuring the surface beneath. Leaves with petioles 1/3 to 4/5 as long as the blades; blades broadly ovate to nearly deltoid, 2.5 to 7 cm long, 1.5 to 6(7) cm broad, acute, base truncate to cordate or slightly attenuate, margins coarsely and bluntly dentate or irregularly toothed to nearly entire. Pedicels at anthesis 10 to 25(35) mm long, elongating in fruit to 20 to 40(52) mm long; calyx at anthesis 6 to 10(12) mm long, lobes 2.5 to 5.5 mm long; corolla 9.5 to 15(17) mm long, yellow, with pale to dark brown or purple-brown spots, these may be indistinct or blurry-edged; anthers 3 to 4 mm long, yellow, filaments ca. 1/2 as long as the anthers. Fruiting calyx 2.5 to 4(5) cm long, 1.5 to 3(3.5) cm broad, ovoid, acute to acuminate, about 1/2 filled by the berry, indented at the base.
Most or all of our plants belong to var. mollis--Field Ground-cherry--with hairs nonglandular; corolla spots pale to dark brown; fruiting calyces 1.5 to 3 cm in diameter. Disturbed ground, usually in sandy soil. Common. E. 1/2 TX; SE. OK and SW. AR, W. LA, and TX. Mar.-Oct. [P. viscosa L. subsp. mollis (Nutt.) Waterfall var. mollis].
Along the coast there is a second variety, var. variovestita (Waterfall) Sullivan, with the shorter hairs glandular, corolla spots purple-black, and mature calyx 2.5 to 3.5 cm in diameter. [P. variovestita Waterfall]. It is not expected here but intergrades between the two varieties are possible in the extreme southern portion of our area.
5. P. cinerascens (Dunal) A. S. Hitchc. var. cinerascens Beach Ground-cherry. Perennial from a sturdy deep rhizome; stems 0.5 to 5 dm tall, main stem erect, lowest branches often decumbent; herbage sparsely to densely pubescent with stellate hairs 1 mm long or shorter. Leaves with petioles from 1/5 as long to equalling the blades; blades broadly ovate to orbicular, 1 to 6(8) cm broad, acute or obtuse, base truncate to rounded or slightly attenuate, margin dentate to sinuate or undulate. Pedicels at anthesis 10 to 33 mm long, elongating to 15 to 60 mm in fruit; calyx at anthesis (3.5)5 to 9(11) mm long with lobes 1.5 to 4.5 mm long, sparsely to densely stellate- or branched-pubescent; corolla (7)9 to 16 mm long, yellow, the spots dark purple-black and sharp-edged, sometimes bisected by the yellow corolla veins, limb reflexed at full anthesis; anthers yellow or slightly tinged or edged with blue, 2 to 5 mm long, filaments about half as wide as the anthers. Fruiting calyx 10-angled or -ribbed, 1.5 to 3.5(4.5) cm long, indented at the base, 1 to 3.5 cm in diameter, green, pubescent. Mostly in disturbed areas--roadsides, gardens, gravel pits, cemeteries, etc. Throughout most of the state; common in our area. E. NM and TX to OK, KS, MO, AR, and LA, S. to Yucatán; sporadic in the SE. U.S. Flowering throughout the year until frost; our collections mostly Apr.-Oct. [P. viscosa L. subsp. mollis (Nutt.) Waterfall var. cinerascens (Dunal) Waterfall; P. mollis Nutt. var. parvifolia Rydb.; P. viscosa L. var. yucatanensis Waterfall; P. mollis Nutt. var. cinerascens (Dunal) Gray; P. pensylvanica L. var. cinerascens Dunal; P. viscosa L. var. cinerascens (Dunal) Waterfall; etc.].
A second variety, var. spathulifolia (Torr.) Sullivan, occurs along the coast and is distinguished by entire margins, unreflexed corollas, and leaves typically at least slightly narrower than var. cinerascens [P. lanceolata var. spathulifolia Torr.; P. viscosa L. var. spathulifolia (Torr.) Gray]. Not present in our area, but mentioned here to as an update to the treatment given by Correll and Johnston (1970.)
6. P. hederifolia Gray. var. hederifolia Heartleaf Ground-cherry, Ivy-leaved Ground-cherry. Perennial from a somewhat woody base, sometimes producing shoots from lateral roots; stems erect to ascending or prostrate, simple or usually well-branched, (1)2 to 4(7) dm tall; herbage with a mix of mostly short glandular hairs and some longer jointed hairs 0.6 to 2 mm long, the glandular hairs predominate on the stems and major leaf veins below, the margins with more of the longer hairs. Leaves with petioles 2 to 2.5 mm long; blades ovate to ovate-lanceolate or rotund, (1.5)2 to 4(5) cm long, 2 to 4 cm broad, entire to sinuate-dentate or irregularly toothed, base acute to cordate or sometimes truncate. Pedicels at anthesis 2 to 8(13) mm long, elongating to 15 to 30 mm in fruit; calyx at anthesis 4 to 8 mm broad, the tube 3 to 4 mm long, lobes lanceolate to deltoid, 4 to 6 mm long; corolla yellow or slightly greenish, often spotted (sometimes faintly), 10 to 15 mm long, often reflexed; anthers usually yellow, 3 to 4 mm long, filaments flattened and sometimes clavate. Fruiting calyx 2 to 3 cm long, 1.5 to 2.5 cm broad, with 10 ribs or angles; berry 8 to 10 mm in diameter, red-brown or yellowish; seeds yellow or yellow-brown, elliptic or depressed-ovate, 1.7 to 2.3 mm long, finely reticulate or rarely smooth. Usually on desert plains and in mountains, but also sandy areas and open woods. This variety mostly in SW. TX, sporadically E. to our area; also NM, AZ, UT, and N. Mex.; possibly also in MT and CO. Mar.-July, sporadically later. [P. hederifolia Gray var. puberula Gray; P. palmeri Gray].
7. P. heterophylla Nees. var. heterophylla Clammy Ground-cherry. Perennial from a deep rootstock; stem erect, simple or branched, 15 to 90 cm tall; herbage pubescent with a mixture of short, usually viscid-glandular hairs and long jointed hairs 1 to 2(3) mm long, or sometimes only a few of the longer hairs present. Leaves with petioles 3 to 6 cm long; blades primarily ovate, but varying to slightly rhomboid, (3)5 to 10 cm long, 3.5 to 6 cm broad, irregularly sinuate-dentate to entire, base rounded to subcordate, both surfaces pubescent. Pedicels at anthesis 10(18) mm long, elongating to ca. 3 cm in fruit; calyx at anthesis 7 to 12 mm long, 5 to 12 mm wide at the base of the lobes, lobes deltoid or ovate; corolla yellow, spotted but sometimes faintly so, (10)15 to 18(20) mm long, 12 to 18(22) mm broad; anthers yellow or sometimes tinged with violet or blue, usually 3 to 4.5 mm long, filaments often as broad as the anthers, sometimes clavate. Fruiting calyx ovoid, 2.5 to 4 cm long, 2 to 4 cm broad, strongly indented at the base, much-inflated; berry yellowish, (8)10 to 12 mm in diameter; seeds yellowish, ovate to ellipsoid, 2 to 2.5 mm long, minutely punctate. In various habitats--fields, roadsides, waste places, open woods, etc. E. TX, S. to about Kenedy Co.; Que. and N.S. to ND, CO, and UT, S. to FL and TX. Apr.-Oct. [Includes var. villosa Waterfall, var. ambigua (Gray) Rydb., var. clavipes Fern., and var. nyctaginea (Dunal) Rydb.; P. ambigua (Gray) Britt; P. sinuata Rydb.; P. nyctaginea Dunal].
This is one of the species whose fruit was eaten by Plains Indian tribes (Kindscher 1987).
8. P. virginiana P. Mill. Virginia Ground-cherry. Perennial from a deep rootstock or rhizome; stems erect, (1)3 to 6 dm tall, branches ascending; pubescence varying by variety, usually some hairs of the stem retrorse or reflexed. Petioles ca. 3 to 30 mm long; blades ovate to linear-lanceolate or elliptic, rarely cordate, 2 to 5 cm long, margin entire to sinuate-dentate, base usually narrowed and commonly narrowly decurrent on the petiole. Pedicel at anthesis (5)10 to 20 mm long, from as long as to 1.5 times as long as the flower; calyx at anthesis with a tube 3 to 6 mm long and lobes (2.5)4 to 6 mm long; corolla (10)12 to 18(20) m long, yellow, dark spotted; anthers 2 to 4 mm long, yellow or tinged with blue or violet, filaments from 1/3 as wide to as wide as the anthers, narrowed at the apex. Fruiting calyx (2.5)3 to 4 cm long, longer than wide, indented at the base, 5-angled; berry 10 to 15(18) mm in diameter; seeds ovate to broadly elliptic, yellow, 1.7 to 2.2 mm long, minutely pitted or reticulate.
As treated by Hatch, et al. (1990) and Kartesz (1998), 2 varieties in TX, either of which might be found here, but neither common.
var. virginiana Pubescence from short and retrorse to villous, not glandular; leaf blades ovate to lanceolate (rarely cordate), irregularly toothed to sinuate-dentate; corolla mostly 15 to 20 mm long; anthers yellow, sometimes tinged with blue or violet. Open woods, prairies, and disturbed areas in E. TX; E. U.S. and Can.; sporadic in the Cen. Rocky Mts. Apr.-June. [P. intermedia Rydb.; P. monticola C. Mohr].
var. texana (Rydb.) Waterfall Usually with several branches from near the base; herbage glabrous or nearly so; main leaves ovate, usually entire. Endemic to the Gulf Coast and occasionally slightly inland. Mar.-Jun.; sometimes also winter. No specimens from our area seen by the author, but within the range given by Hatch, et al. (1990). [P. texana Rydb.].
9. P. longifolia Nutt. Longleaf Ground-cherry. Perennial from a rhizome; stems simple below and branched above, usually 3 to 6(9) dm tall; herbage pubescent with opaque, generally antrorsely appressed, only obscurely-jointed or septate hairs on the upper stem, pedicels, lower sides of leaves, and nerves of the calyx, commonly glabrate at maturity. Leaves lanceolate, elliptic-lanceolate, or narrowly rhombic or ovate, 4 to 10 cm long, entire to coarsely, irregularly toothed. Pedicels 1 to 4 cm long; calyx at anthesis campanulate, 7 to 10 mm long, with triangular to ovate lobes 3 to 4 mm long; corolla yellow, spotted with purple or reddish purple, 11 to 20 mm long, 10 to 15 mm broad; anthers 3 to 4 mm long, yellow or tinged with blue. Fruiting calyx 5-angled, usually 2.5 to 3.5(4) cm long.
Two varieties, either of which might be found here according to the literature (Correll and Johnston 1970; Cronquist, et al. 1984; GPFA 1986), but neither seen by the author.
var. longifolia Leaf blades lanceolate, elliptic-lanceolate, or narrowly rhombic, tapered to the petiole, only rarely ovate; anthers yellow. Open woods, hills, plains, prairies, deserts, etc. In most of TX; native to the Great Plains, extending to parts of the E. U.S. and W. to the Rocky Mts. and N. Mex. May-Sept. [P. virginiana P. Mill. var. sonorae (Torr.) Waterfall; P. pumila Nutt. var. sonorae Torr.; P. rigida Pollard and Ball; etc.].
var. subglabrata (Mackenzie and Bush) Cronq. Bladder Ground-cherry. Blades thinner-textured, broader, and more ovate than in var. longifolia, and more abruptly tapered to the petiole. Native to eastern deciduous forests and sporadic westward. [P. virginiana P. Mill. var. subglabrata (Mackenzie and Bush) Waterfall; P. subglabrata Mackenzie and Bush; P. macrophysa Rydb.; etc.].
Perennial herbs from taproots or rhizomes. Stems prostrate to ascending. Leaves alternate, simple, entire to pinnatifid, subsessile to petiolate; blade sometimes decurrent on the petiole, more or less glabrous to pubescent with simple, branched, or stellate, hairs, sometimes glandular. Flowers in groups of 1 to 4 in the axils of the leaves, pedicels elongating and becoming curved in fruit. Calyx campanulate, 5-lobed, accrescent in fruit and at least partly enclosing the berry tightly, not inflated, or angled, little if at all veiny. Corolla white or yellowish, sometimes tinged with purple or green, plicate in bud, subrotate, with white glandular or pubescent appendages in the throat, the 5 lobes shallow. Stamens inserted near the base of the corolla, anthers longitudinally dehiscent, yellow, filaments slender, terete. Stigma slender, little if at all broader than the style; ovary bilocular. Berry tightly enveloped (if not fully enclosed) by the mature calyx, whitish, somewhat dry. Seeds few, borne only on the lower part of the placentae, reniform, flattened, rugose or minutely pitted and/or reticulate; embryo markedly curved.
7 species of the W. U.S. and Mex.; all present in TX; 1 known from our area. This treatment is based, in part, on the work of Averett (1973).
The fruits are inedible.
1. C. edwardsiana Averett Perennial from rather woody roots; stems erect or spreading/reclining in age, 7 to 30 cm long, well-branched from the base and above, terete or faintly striate or angled, commonly cinereous or purplish at the base, glabrous to pubescent with short, stellate or occasionally simple or branched hairs, often with glandular hairs. Leaves linear-lanceolate to very narrowly rhombic, usually undulate, especially above the middle, but sometimes with a few shallow lobes, 2.5 to 7 cm long, 7 to 15 mm broad, basally attenuate, subsessile or short-petiolate, the midrib obvious and the smaller veins not so, surfaces more or less glabrous. Flowers solitary or sometimes paired, on pedicels 2 to 2.5 cm long and elongating in fruit, pubescent and with glandular hairs. Calyx 4 to 5 mm long, lobed about 1/2 the length, the lobes acute, pubescent with stellate, simple, and branched hairs; corolla white, drying yellow, sometimes with a purple tint, 10 to 15 mm broad, the lobes pubescent, especially marginally. Fruit 5 to 8 mm broad, enclosed in the calyx. Mostly on limestone outcrops in the Ed. Plat. and also N. Mex., but collected once or twice in Brazos Co. in 1967; possibly not a permanent member of our flora. Summer, but the Brazos Co. collections from April.
17 species of the SW. U.S. to S. Amer.; 1 species cultivated in Texas and occasionally escaping or persisting.
1. S. origanifolia (Lam.) Baill. Cock's-eggs. Perennial from a fleshy root; stems herbaceous, climbing, sometimes slightly woody at the base, freely branched, branches green; herbage more or less odiferous, from pubescent ot nearly glabrate. Leaves nearly opposite, petioles shorter than the blades; blades ovate to ovate-rhombic or suborbicular, 0.5 to 4 cm long, 0.3 to 3.3 cm broad, entire to undulate-crenate. Flowers solitary in the axils, pedicels filiform, 5 to 10 mm long, little shorter than the petioles, pubescent. Calyx ca. 2.5 to 3.5 mm long, the lobes lanceolate, united only basally; corolla white, urceolate, 7 to 10 mm long, 3 to 5 mm broad, the 5 lobes 1 to 2 mm long, reflexed or spreading, margins pubescent, interior of tube with a hairy ring; stamens attached about midway up the corolla tube or above, anthers exserted; style slender, stigma exserted. Berry ovoid-oblong to ellipsoid, yellow or white, with many seeds. Native to S. Amer., cultivated and sometimes persisting or escaping, not at all common in our area. Mar.-Oct. [Author in some older sources listed as (Lam.) Baill.; S. rhomboidea (Gill. & Hook.) Miers; Perizoma rhomboidea (Hooker) Small].
The fruit is said to be edible but with a poor flavor (Correll & Johnston 1970; Mabberley 1987).
Herbs, shrubs, trees, or vines (ours mostly herbaceous) with various habits, sometimes armed with prickles or spines. Herbage glabrous or variously pubescent, sometimes with stellate hairs. Leaves alternate, petiolate, simple to bipinnatifid, the larger ones sometimes accompanied by smaller lateral ones, estipulate; . Inflorescences racemose, cymose, or appearing paniculate, peduncles axillary or terminal and appearing lateral as an axillary shoot elongates. Flowers perfect, usually 5-merous except for the gynoecium, rarely otherwise. Calyx of united sepals, persistent, sometimes accrescent and tightly enclosing the fruit but not inflated. Corolla usually rotate, white, blue, purple, or yellow, regular or rarely slightly irregular, plicate in bud, the lobes valvate or valvate with the margins inrolled. Stamens connivent around the style, filaments short, anthers apically dehiscent by pores or short introrse slits, sometimes dimorphic. Fruit a juicy or semi-dry berry, usually 2-celled, some species with clumps of stone cells attached to the fruit wall or scattered among the seeds.
About 1,400 species nearly worldwide; 24 listed for TX (Hatch, et. al 1990); 6 here. Reference: Schilling (1981).
The genus is important for the potato, S. tuberosum and the eggplant, S. melongena. Recent studies suggest that the tomato belongs to the genus as S. lycopersicum L. . Many taxa are poisonous to humans and/or livestock, some have edible or ornamental fruit, and some are troublesome weeds (Mabberley 1987).
1. Pubescence of simple hairs or plants glabrous; spines or prickles absent; flowers white (sometimes tinged with purple), to ca. 15 mm in diameter .....................................................2
1. Pubescence of stellate hairs; prickles or spines usually present somewhere on the plant; flowers blue, violet, or yellow (rarely white), more than 15 mm in diameter ..........................3
2(1) Plants perennial, varying from shrubby to scandent; leaves generally cordate-hastate; flowers more than 7 mm broad; fruit red at maturity .....................................1. S. triquetrum
2. Plants annual; leaves generally oval to ovate-lanceolate or elliptic; flowers usually less than 7 mm long; fruit black or purple at maturity ........................................2. S. ptycanthum
3(1) Corolla yellow; leaves lobed more than halfway to the midrib; calyx in fruit densely prickly, wholly enclosing the berry ...............................................................................3. S. rostratum
3. Corolla violet or purple-blue (rarely white); leaves entire or lobed less than halfway to the midrib; calyx in fruit with at most a few prickles, not enclosing the berry ................................4
4(3) Lower leaf surface and stem silver- or pale-canescent with dense stellate hairs ...................
.................................................................................................................4. S. elaeagnifolium
4. Lower leaf surface and stem not silver- or white-canescent, hairs usually tawny and not obscuring the surface beneath ................................................................................................5
5(4) Hairs of lower leaf surface with 4 to 8 rays, sessile; calyx 5 to 7 mm long; corolla 2 to 3 cm broad; fruit 1 to 2 cm in diameter .................................................................5. S. carolinense
5. Hairs of lower leaf surface usually with 8 or more rays, at least some stipitate; calyx 8 to 13 mm long; corolla 3 to 5 cm broad; fruit 2.5 to 3 cm in diameter .............6. S. dimidiatum
NOTES: Solanum lycopersicum L. (=Lycopersicon esculentum Miller.), Tomato, is sometimes found as an escape or a waif in our area, especially where sewage has been spread. Leaves divided, flowers yellow; fruits red, varying in size and shape.
S. capsicastrum Link, FalseJerusalem Cherry, is occasionally encountered here as an escape from cultivation. It has leaves oval to oblong and stellate-pubescent below, corollas white, ca. 11 to 14 mm long, and red to orange fruits ca 13 mm in diameter.
S. citrullifolium A. Br. occurs west of our immediate area and may someday be found in the far W. part of our region. It is a prickly plant with irregularly bipinnatifid leaves, violet corollas, fruit enclosed by the prickly calyx, and 4 yellow anthers accompanied by a fifth, larger anther tinged with violet.
1. S. triquetrum Cav. Texas Nightshade, Hierba Mora. Perennial, varying from a small shrub to a scandent, rather vine-line shrub with elongate branches; stems semi-woody, runner-like, or flexible, angled but not actually triquetrous, to 2 m long; herbage glabrous or nearly so. Leaves (0.5)1 to 6 cm long, deltoid-cordate to triangular-ovate, sometimes with 3 or 5 lobes, the middle lobe the largest, occasionally leaves nearly linear, apex acute to acuminate, margin entire except for any lobes, slightly decurrent on the petiole; petiole about equalling the blade or shorter. Peduncles slim; pedicels knobby-jointed at the base, apices thickening in fruit and becoming clavate. Calyx lobes triangular or ovate; corolla 15 to 20 mm across, rotate and star-shaped, white or tinged with violet; anthers yellow. Berries red, 1 to 1.5 cm in diameter. Thickets, hills, slopes, and (in our area at least) commonly weedy in cultivated or disturbed areas. Cen., S., and W. TX; also N. Mex. Flowering throughout the year.
2. S. ptycanthum Dun. ex DC. Black Nightshade, American Nightshade, Hierba Mora Negra, Blueflower Buffalo-bur. Taprooted annual; stems (1)3 to 6(12) dm tall, erect or with divergent branches from the base; herbage glabrous or sparsely strigose (sometimes densely strigose on the new growth), unarmed. Petioles (0.1)3 to 7(10) cm long, usually at least partly winged with the decurrent blade; blade variable in shape from ovate to ovate-lanceolate, lanceolate, elliptic-lanceolate, or triangular-ovate, (2)5 to 10(17) cm long, more or less translucent, apex acute or short-acuminate to obtuse, base cuneate to rounded, subtruncate, or cordate, margin entire to undulate or sinuate-dentate. Inflorescences axillary or lateral, usually umbellate, rarely corymbose, usually strigose, commonly with 2 to 4 flowers; peduncle filiform to stout, 1 to 3 cm long; pedicels (1)3 to 7(10) mm long, often unequal, reflexed soon after anthesis. Calyx campanulate, lobes lanceolate to oblong, acute to obtuse, 1 to 1.5 mm long, often unequal, partially united, not reflexed in fruit; corolla (3)4.5 to 7(9) mm long, star-shaped, white, sometimes tinged with purple or blue or with a yellow star, lobes 3 to 7 mm long, often reflexed; anthers 1.3 to 2 mm long; style 2.5 to 3.3 mm long. Ripe berries purplish-black, shiny or dull, 5 to 9 mm long, commonly becoming detached at the joint of pedicel and peduncle, but sometimes at the receptacle, clumps of stony cells (3)6 to 20(15) per fruit, varying in size; seeds many, 1.5 to 1.9 mm long, flat, minutely pitted or reticulate. Dry or moist open woods, prairies, shores, roadsides, etc.; commonly weedy in cultivated or disturbed areas. Throughout TX; S. Que. to Man., S. to FL and TX. Flowering throughout the year. [Sometimes seen with specific epithet spelled "ptychanthum." S. americanum--in part. Our plants have long gone by the name S. americanum P. Mill., but unfortunately, the lectotype chosen for S. americana does not belong to this species, so the next available name, S. ptycanthum, must be used (Schilling 1981). This species was also previously referred to S. nigrum, part of a rather confusing complex which includes plants now or previously referred to S. interius Raf., S. douglasii Dun., S. nodiflorum Jacq., S. sarrachoides Sendtner, and about 2 dozen other species, with various works treating the taxa differently. S. nigrum L. as now recognized is not present in TX.]
The foliage and immature fruit are poisonous, with fatalities reported in children (Tull 1987), though the mature berries are reputedly edible (Cronquist, et al. 1984; Lampe 1985; GPFA 1986). S. nigrum (strict sense) has been used medicinally since the time of Dioscorides (Kindscher 1992).
3. S. rostratum Dun. Buffalo-bur, Kansas Thistle, Mala Mujer. Taprooted annual; stems erect, widely branching, (2)3 to 7 dm tall, rather densely beset with straight yellow prickles and densely pubescent with stalked or sessile stellate hairs, those of the leaves with an elongate central ray. Leaves with petioles (1)2 to 6 cm long; blades once- or twice-pinnately lobed, parted, or cleft, the lobes rounded, surfaces stellate pubescent and the veins prickly. Inflorescences racemose, with 5 to 15 flowers, in fruit elongating and the pedicels ascending. Calyx nearly obscured by prickles, at anthesis the lobes 2 to 3 m long, about as long as the tube, calyx in fruit enlarging to wholly enclose the berry; corolla yellow, 1.6 to 2.5(3) cm broad, slightly irregular, lobes triangular or broadly ovate, wide-spreading, about equalling the tube, stellate pubescent; stamens and style declined to the lower side of the flower; filaments short, attached near the throat of the corolla, anthers dimorphic--4 yellow, 7 to 9 mm long, the fifth commonly purplish, longer, broader, and with a curved tip. Berry completely enclosed by the calyx, 7 to 10 mm long; seeds silvery dark brown to black, 2.3 to 3.7 mm long, minutely pitted and often cross-wrinkled on the margins. Overgrazed pastures, waste places, stream beds, etc., often weedy. Throughout TX; NE to TX, adventive eastward and northwestward. Apr.-Oct.(Nov.). [S. cornutum Lam.; Androcera rostrata (Dun.) Rydb.
This species is reported to have caused human and/or livestock poisonings (Tull 1987). The Zunis used a tea made from the powdered root to treat nausea (Kindscher 1992).
4. S. elaeagnifolium Cav. Silver-leaf Nightshade, White Horse Nettle, Trompillo, Bull Nettle. Perennial from a deep, more or less woody running rootstock, 1 to 10 dm tall; stems, lower leaf surfaces, and inflorescences pale- or silvery-canescent with dense stellate hairs that obscure the surface below; prickles short and needle-like, sparse or nearly absent to numerous, only a few on the leaves. Leaves with petioles 1 to 3(5) cm long; blades narrowly lanceolate to lance-oblong or nearly linear, 3 to 10(15) cm long, apex usually obtuse, sometimes acute, tapered to the base, margin entire to sinuate or undulate. Inflorescences cymose-racemose, terminal but surpassed by subtending growth and so appearing lateral at the top of the plant, flowers (1)3 to 7; peduncle short; pedicels rather long, slender, recurved in fruit, sometimes sparsely prickly. Calyx 5-angled, tube 4 to 6 mm long, lobes linear, shorter than to a little longer than the tube, commonly reflexed in fruit; corolla blue-purple (sometimes pale or white), (15)20 to 30(35) mm across, lobes ovate, acute, equalling or longer than the tube, pubescent externally; anthers yellow, 6 to 9 mm long, equal, erect, connivent, the filaments short; style deflexed, ovary white-tomentose. Berry globose, to 15 mm in diameter, yellow or eventually turning black; seeds 3 to 5 mm long, ovoid or oblong, shiny brown, nearly smooth. Abundant in overgrazed pastures. MO and KS, S. to LA and TX, W. to AZ; also N. Mex.; adventive elsewhere. Mar.-Oct.; most noticeable in our area in summer and fall. [Includes forma albiflorum Cockerell, applied to rare white-flowered plants; S. texana Engelm. & Gray; S. roemerianum Scheele].
The Zunis chewed the roots of this plant for toothache, and packed it in cavities (Kindscher 1992).
5. S. carolinense L. Carolina Horse Nettle, Ball-nettle. Perennial from an erect rootstock and usually with creeping rhizomes; stems 0.3 to 1 m tall, erect, sparsely branched; spines yellow, slightly flattened, (1)2 to 4(7) mm long; herbage, inflorescences, and calyx with sessile, 4- to 8-rayed stellate hairs, each with or without an elongate central ray, the hairs not dense enough to entirely obscure the surface beneath. Petioles 1 to 3 cm long, usually spiny; blades elliptic-ovate to ovate, (3)5 to 19(15) cm long, 3 to 8(10) cm broad, apically obtuse to broadly acute, base rounded to cuneate, margin usually with a few teeth, sinuate, or even shallowly lobed, main veins usually with a few prickles. Inflorescences cymose-racemose, with 5 to 20 flowers, elongating to form a simple raceme; pedicels reflexed in fruit. Calyx 5 to 7 mm long, lobed about halfway at anthesis, lobes lance-acuminate, 2.5 to 4 mm long, calyx in fruit becoming 8 to 9 mm long; corolla purple, lavender, or occasionally white, 2 to 3 cm across, the lobes triangular or ovate, 6 to 9 mm long, about as long as the tube, spreading or reflexed, stellate pubescent externally and with a few scattered hairs within; anthers yellow, 6 to 9 mm long, equal, erect and connivent. Berry yellow, globose, 1 to 2 cm in diameter, plump at first but becoming wrinkled; seeds 1.5 to 2(3) mm long, flattened, yellowish, nearly smooth. Usually in sandy or open soils of open woods, fields, and waste places; sometimes weedy. E. TX; S. Ont. and MN, S. to TX and N. Mex., E. to the Atlantic; sporadically introduced westward. Apr.-Oct. [White-flowered forms have been treated as forma albiflorum Benke].
Livestock and deer have been poisoned by this species, and there is at least one report of a child having been fatally poisoned (GPFA 1986). The plant produces a suite of alkaloids, including solanine (Kindscher 1992). It is listed as poisonous by the American Medical Association (Lampe 1985).
6. S. dimidiatum Raf. Western Horse Nettle. Perennial from a deep running rootstock; stem 3 to 10 dm tall, commonly branched above, sparsely grayish-cinereous with (5)9- to 12-rayed stellate hairs without a longer central ray; herbage with similar hairs and also with scattered, flattened, yellow prickles 2 to 7 mm long. Petioles rather short, stout; blades ovate to elliptic-lanceolate, 6 to 15 cm long, 5 to 10 cm broad, irregularly sinuate or shallowly 5- to 7-lobed or parted, apically obtuse, base rounded to truncate or cordate, both surfaces with stellate hairs, the lower surface more densely pubescent and with some of the hairs definitely stipitate (stalked), midrib commonly with prickles. Inflorescences terminal racemes, panicles, or 2- or 3-divided cymes, frequently surpassed by the subtending foliage; pedicels recurved in fruit. Calyx 8 to 13 mm long, lobes frequently 6 rather than 5, abruptly acuminate; corolla blue-purple, violet, or occasionally nearly white, (2)3 to 5 cm across, lobes 5(6), broadly ovate, spreading, pubescent externally; stamens 5(6), lanceolate to linear-lanceolate, erect, connivent, equal, 8 to 10(12) mm long. Berries pale yellow at maturity, globose, (1)2.5 to 3 cm in diameter; seeds oval, yellowish brown, 3.8 to 4.3 mm long, minutely rugose. Sandy soils of prairies, waste places, roadsides, fields, etc. Throughout TX except the extreme W. and E. portions; SC to FL, W. to MO, KS, AR, and TX. May-Oct. [White-flowered plants have been designated forma album (Waterfall) Correll; S. torreyi Rydb.].
Said to be poisonous (GPFA 1986; Tull 1987) but not listed by the American Medical Association (Lampe, 1985). However, it is probably safest to treat all leaves and immature fruit of any Solanum as poisonous and the mature fruit as suspect.
10 species of tropical America; 1 species native to TX.
1. C. annuum L. var. glabriusculum (Dunal) Heiser & Pickersgill Chilli Piquín, Bird Pepper, Bush Redpepper. Technically a shrub, but commonly growing as a perennial or even small enough to resemble a taprooted annual, to 3 m tall or long, usually smaller; stems green, brittle, sometimes zig-zagging from node to node; herbage glabrous or very sparsely puberulent. Petioles shorter than the blades; blades ovate to elliptic-lanceolate or lanceolate, apically acute to acuminate, base rounded to tapered, (1.5)2 to 5(6) cm long, to 3 cm broad. Flowers mostly solitary or sometimes paired, pedicels slender, spreading or reflexed, sometimes becoming stouter in fruit. Calyx small, with 5 shallow lobes in flower, becoming cup-like and truncate in fruit; corolla white, rotate, star-shaped, 7 to 10 mm across; anthers usually bluish. Fruit to ca. 15 mm long, ovoid to more or less globose, red or yellowish at maturity, persistent, very pungent. Along arroyos or rivers, in thickets or groves, or in vacant lots. Ed. Plat. and S. TX., infrequent but present in our area. TX to AZ, also S. FL and widespread in tropical Amer. Flowering throughout the year. [C. annuum L. var. minus (Fing.) Shinners and var. aviculare D'Arcy & Eshbaugh].
The berries are very pungent and useful in cooking, but can cause gastric discomfort if eaten in large quantities (Tull 1987). Handling the fruit or seeds can cause pain and redness in exposed skin (Lampe 1985).
This is the wild-growing form of the species. C. annuum also includes most of the cultivated peppers, these belonging to var. annuum and being divided into groups---cayenne or chile, bell, cherry, etc. Var. frutescens (L.) Kuntze [C. frutescens L.] is the tabasco pepper. Capsaicin from peppers is used medicinally as a local stimulant or counter-irritant, usually in very minute amounts (Mabberley 1987).
Ours annual or perennial herbs (elsewhere also shrubs and trees). Stems twining dextrorsely (to the right). Leaves alternate (absent or scale-like in parasitic Cuscuta), simple to lobed or compound, estipulate. Flowers terminal or axillary, solitary, cymose, racemose, or paniculate, commonly subtended by bracts, regular, hypogynous, usually 5-merous (sometimes 3- or 4-merous in Cuscuta). Sepals equal or unequal, free or basally united, imbricate, persistent, sometimes accrescent. Corolla sympetalous, rotate to salverform or funnelform (rarely curved or somewhat irregular and not so in ours), usually with 5 lobes or angles, induplicate in bud and pleated. Stamens 5, epipetalous, alternate with the corolla lobes, free, anthers usually linear to oblong, extrorse. Nectary disk absent or present as a disk, cup, or ring, sometimes lobed. Gynoecium of 2(3) united carpels, usually with as many locules (sometimes 1, 4, or 5); ovules 2 per locule; styles usually slender, solitary and lobed or bifid or else styles 2 and separate or nearly so; stigmas capitate to 2-lobed, clavate, or linear. Fruits usually capsular, dehiscent by valves, transversely dehiscent, circumscissile, or breaking irregularly, in some species indehiscent. Seeds ca. 1 to 4 per fruit, usually fewer than the ovules, glabrous to pubescent, endosperm absent or meager.
As treated here, inclusive of Cuscuta (which is often separated as a monogeneric family on the basis of characters such as pollen and embryo morphology and the position of corolla lobes in bud), ca. 58 genera and 1,650 species worldwide, especially in the tropic and temperate zones; 12 genera and 79 species listed for Texas (Hatch, et al., 1990); 6 genera and 24 species here.
In some areas, generic boundaries are disputed, unclear, or subject to interpretation. E.g. Stylisma was formerly in Bonamia, some Bonamia were formerly in Breweria, Calystegia is sometimes treated in Convolvulus, Petrogenia is sometimes included in Bonamia, and some taxa in Ipomoea have been or can be placed in Merremia, Operculina, Turbina, Xenostegia, etc.
The family has many ornamentals in Ipomoea, Convolvulus, and so on. I. batatas is the sweet potato commonly grown in the southern U.S. Cuscuta includes leafless parasites that can cause crop losses if not controlled (Mabberley 1987).
1. Plants leafless and parasitic, usually without chlorophyll .....................................1. Cuscuta
1. Plants not parasitic, with green leaves ....................................................................................2
2(1) Stems rooting at the nodes; ovary deeply 2-lobed; corolla ca. 2 to 4 mm broad, greenish ....
.............................................................................................................................2. Dichondra
2. Stems not rooting at the nodes; ovary entire or only shallowly 2-lobed; corolla more than 4 mm broad, white to pink, purple, or blue .................................................................................3
3(2) Styles 2, each 2-branched ...................................................................................3. Evolvulus
3. Style 1, sometimes 2-cleft at the apex .....................................................................................4
4(3) Stigmas elongated or flattened, linear, more than twice as long as broad ..4. Convolvulus
4. Stigmas as broad as long or broader .......................................................................................5
5(4) Leaves linear or linear-lanceolate; style minutely 2-cleft (the divisions less than 2 mm long); sometimes style entire--if so, stigma minute, equal in diameter to the style .................
.................................................................................................................................5. Stylisma
5. Leaves broader; style unbranched and uncleft; stigma lobed or entire .................................6
6(5) Stigma with 2 flattened, elliptical or oblong lobes; flowers in dense, leafy-bracted axillary cymes ..............................................................................................................6. Jaquemontia
6. Stigma entire or with 2 to 3 globose or subglobose lobes; flowers 1 to many in terminal or axillary groups .........................................................................................................7. Ipomoea
Strangle Vine, etc.
Rootless (by flowering time) parasitic herbs, attached to the host plant via haustoria, usually without chlorophyll and yellow or orange, but some with small amounts of chlorophyll in various organs. Stems filiform or slender, twining, fleshy. Leaves reduced to very small scales or absent. Flowers small, ca. 1 to 6 mm long, sessile or short-pedicelled, few to many, usually in cymose clusters, but sometimes through branching the clusters compact and the inflorescence form obscure or else the flowers produced endogenously (directly from the host tissue). Flowers (3- or 4-)5-merous, regular, perfect. Sepals united or nearly completely free. Corolla campanulate to cylindric or somewhat funnelform, shallowly or deeply lobed. Stamens alternate with the corolla lobes, epipetalous, a row of scale-like, fringed or fimbriate appendages bridged (attached to the corolla and/or each other) near the filament bases or below. Ovary 2-celled, each locule with 2 ovules; styles rarely united, usually distinct, stigmas capitate to linear. Fruit a capsule, circumscissile near the base or dehiscing irregularly. Seeds ca. 1 to 3 mm long, brown and glabrous, the embryo without cotyledons, small and filiform or with one end enlarged
About 145 species worldwide; 18 listed for TX with one doubtful (Hatch, et al. 1990); 10 species present or highly likely in our area. See page 578a for addendum.
Useful references are Yuncker (1943) and Gandhi, Thomas, and Hatch (1987). These include some rather helpful illustrations.
Some taxa are or have been serious pests in crops, with harvest and spread of the host spreading the parasite's seeds as well. Some taxa grow on only one or a very few hosts, while others are capable of growing on a wide variety of hosts.
NOTE: Confident identification of Cuscuta requires examining the flowers and fruits under a good dissecting scope. In pressing Cuscuta, it is advisable to mount enough to show the host and part of the parasite. The specimen should be padded with foam or folded newspaper so that the fragile flowers and fruits are not crushed. A generous portion of the material should be dried unpressed and stored in an attached fragment packet (Austin 1979).
The small number of collections and the rather high proportion of collections which are misidentified or damaged beyond identification make a definitive list problematic. Complete distributions by county have not been published.
1. Styles united; capsules ca. 1 cm long, circumscissile .......................................1. C. exaltata
1. Styles wholly separate; capsules much less than 1 cm long, not circumscissile ..................2
2(1) Sepals at anthesis free or nearly so; flowers subtended by bracts resembling the sepals .....
...................................................................................................................................................3
2. Sepals at anthesis obviously united; flowers subtended by scale-like bracts not resembling the sepals, or bracts absent .....................................................................................................5
3(2) Flowers usually pedicellate, in loose panicles; sepals and bracts orbicular-ovate ..................
..........................................................................................................................2. C. cuspidata
3. Flowers sessile, in dense, sometimes rope-like clusters; sepals acute .................................4
4(3) Bracts and sepals loose, lanceolate, acute, the tips recurved .....................3. C. glomerata
4. Bracts and sepals tightly appressed, obtuse, the tips erect ..........................4. C. compacta
5(2) Styles relatively short and stout, separated from one another at the base and the opening between them large; sepals often unequal .............................................................................6
5. Styles slender, more or less filiform, the space between their bases small; sepals usually equal .........................................................................................................................................7
6(5) Flowers generally 5-merous........................................................................... 5. C. obtusiflora
var. glandulosa
6. Flowers generally 3- or 4-merous ................................C. polygonorum (see page 578a)
7(5) Flowers mostly 4-merous .......................................................................................6. C. coryli
7. Flowers mostly 5-merous .........................................................................................................8
8(7) Sepals triangular-ovate or lanceolate, acute ....................................................7. C. indecora
8 Sepals ovate, obtuse ................................................................................................................9
9(8) Capsule globose-ovoid, with a thickened ring at the base of the styles; corolla lobes acute ...................................................................................................................8. C. gronovii
9. Capsule depressed-globose or globose, about as wide as long, not thickened around the style bases; corolla lobes obtuse ..................................................................9. C. pentagona
1. C. exaltata Engelm. Tree Dodder. Stems fleshy and stout, to 3 mm broad. Flower clusters spicate-paniculate; flowers sessile or subsessile, fleshy, ca. 4 mm long from the base to the corolla sinuses, each subtended by one oval to ovate, obtuse bract. Calyx enclosing the corolla except for the lobes, deeply divided, the lobes orbicular-ovate, apically rounded, much overlapping; corolla more or less cylindrical, the lobes much shorter than the tube, oval-ovate to nearly orbicular, erect to spreading; staminal scales 1/2 to 2/3 as long as the corolla tube, represented by 2 wings (1 on each side of the filament), toothed in the upper portion, sometimes free and dentate or emarginate; styles usually equal, united fully or partly and appearing 1, but separable with a dissecting needle, about as long as the ovary, stigma(s) flattened or subglobose. Capsule globose-ovoid to conic, to 10 mm long, usually with the withered corolla around the top, circumscissile near the base; seeds 3 to 4 mm long, nearly ovoid, slightly beaked. Usually on woody hosts, including species of Quercus, Juglans, Ulmus, Smilax, Rhus, Vitis, Diospyros, etc. Gulf Prairies and Marshes, Blackland Prairies, and Edwards Plateau; confirmed from our area; TX and one collection known from FL.
2. C. cuspidata Engelm. Cusp Dodder, Cuspidate Dodder. Stems 0.4 to 0.6 m thick. Flowers membranous, pedicels 0.5 to 3 mm long or flowers subsessile in loose or dense paniculate cymes; bracts present more or less throughout the inflorescence (rarely absent), ovate and obtuse to orbicular and acute, sometimes cuspidate. Flowers ca. 3 mm long from base to corolla sinuses; sepals free or slightly united, shorter than the corolla tube, to 2.2 mm long and 2 mm wide, ovate-orbicular, obtuse to acutish, minutely cuspidate, margins thinner and serrulate, sometimes the median line glandular-thickened or appearing bluntly keeled; corolla funnelform with a bulbous base, 4 mm long, lobes shorter than the tube, to 1.7 mm long and 1.2 mm broad, usually more or less acute and cuspidate, occasionally serrulate apically or obtuse, spreading, commonly with a line of translucent, glandular dots along the midline; staminal scales shorter than the corolla tube, oblong, fringed with medium-long segments, bridged to or below the middle; stamens shorter than the corolla lobes, anthers oval-oblong or ovoid and cordate, to 0.8 mm long, shorter than to longer than the filaments, which are to 0.7 mm long; ovary globose or nearly conic; styles to 2.2 mm long, longer than the ovary, slender, slightly unequal, stigmas capitate. Capsule to 2 mm long and 1.7 mm wide, globose or nearly so, with a thick collar or ridge around the opening between the style bases, often dotted with glandular cells, the corolla persistent around or on the top; seeds nearly obovate, ca. 1.4 mm long. Growing on various herbs, but preferring members of the Asteraceae such as Ambrosia, Baccharis, Iva, Helianthus, Liatris, and so on. Throughout much of TX except the Trans Pecos and Ed. Plat.; confirmed from our area; UT and CO, E. to IN and AR. Aug.-Oct. [Includes var. pratensis Engelm. and var. humida Engelm.].
3. C. glomerata Choisy Cluster Dodder, Glomerate Dodder. Stems 0.6 mm in diameter or broader, but often disappearing from between the dense, rope-like, straw-colored inflorescences that wind round the host plant and are up to 6 cm in diameter; flowers usually produced endogenously in 2 parallel rows on opposite sides of the stem, sessile, subtended and intermixed with 2 to 6(many) scarious, lanceolate or oblong, acute or obtuse, serrate or lacerate, cupped bracts with tips squarrose or recurved. Flowers 4 to 5 mm long from base to corolla sinuses, calyx segments free, oblong-oval, obtuse to acute, similar to the bracts but the tips only spreading and not recurved, to 4.7 mm long and 2.5 mm broad; corolla with a cylindrical tube which is bulbous at the base and shorter than the lobes, lobes oblong to lanceolate, acute to somewhat obtuse, sometimes mucronate, spreading or reflexed, apically sometimes mucronate, often with a row of glandular cells along the midline (though perhaps not as obvious as in C. cuspidata); staminal scales shorter than the corolla tube, to 3.5 mm long, nearly as long as the filaments, broadest at or above the middle, oblong, margins fringed mostly above the middle; stamens shorter than the corolla lobes, anthers elliptic to oblong, equal to or shorter than the filaments; styles slender, slightly unequal, much longer than the ovary, to 3.5 mm long; stigmas capitate. Capsule globose, pointed at the apex or with a prominent collar or thickening around the opening between the styles, withered corolla usually persisting around the top; seeds few, only (0)1 to 2 per fruit, ovoid or globose, ca. 1.7 mm long. Growing primarily on members of the Asteraceae, commonly weedy ones, but able to parasitize a number of herbaceous and woody species. Probably in our area; in the prairie regions from MI to SD, S. to MS and TX. Summer-fall. [C. paradoxa Raf.; C. aphylla Raf.].
4. C. compacta Juss. ex Choisy Compact Dodder. Stems stoutish, to 0.6 mm or more in diameter. Flowers sessile or short-pedicellate, in scattered glomerules or usually the glomerules grouped into dense, rope-like clusters, often the flowers originating endogenously; each flower subtended by 2 to 5(10) tightly appressed, ovate-orbicular, fleshy, overlapping bracts. Sepals free, cupped, shorter than the corolla tube, to 3 mm long and 3.2 mm broad, similar to the bracts except perhaps for a marginal fringe of short, slender filaments, the calyx and bracts together somewhat resembling a tree bud with overlapping scales; corolla 4 to 5 mm long from the base to the corolla sinuses, the tube cylindrical, constricted above the ovary, lobes spreading to reflexed, shorter than the tube, to 1 mm long and 1.2 mm wide, oval-oblong, apically rounded to obtuse, incurved, sometimes with a marginal fringe, becoming more or less urceolate in fruit with the base enlarging around the capsule; staminal scales bridged at or about the middle, about reaching the stamens, margins with long fimbriate processes, occasionally smaller scales present on the bridge between the larger ones; stamens exserted, shorter than the corolla lobes, filaments to 0.5 mm long, rather thick, anthers to 0.6 mm long, oval; styles about equalling or longer than the ovary, to ca. 1 mm long, stigmas capitate. Capsule globose-conical, slightly pointed, to 4 mm long and 4.5 mm wide, somewhat thickened around the opening between the styles but without a marked collar, apex commonly with some glandular dots and enclosed by the withered corolla; seeds 2 to 2.6 mm long, globose-ovoid. On various herbaceous and woody hosts, including Carya, Alnus, Rhus, Rubus, Ilex, Vitis, Vaccinium, etc. NH to NE, S. to FL, TX, and AR; almost certainly present in our area. Summer-fall. [Includes var. typica Yuncker and var. adpressa (Engelm.) Engelm.; C. glomerata Choisy var. adpressa (Engelm.) Choisy; some sources recognize other varieties].
5. C. obtusiflora Kunth in H.B.K. var. glandulosa Engelm. Red Dodder. Stems medium to slender, sometimes very thin in dry material. Flowers subsessile (pedicel to 1.5 mm long) in moderately-open, glomerate-cymose clusters; bracts scale-like, dissimilar to the calyx lobes. Flowers ca. 1.5 to 2 mm long from the base to the corolla sinuses, (4-)5-merous, the parts usually dotted with many translucent gland-like cells. Sepals united below, lobes unequal (one usually smaller than the others), ovate and obtuse, not much overlapping, concealing most of the corolla tube; corolla tube campanulate, enlarging in fruit, lobes a little longer than the tube, to 1.3 mm long and 1.2 mm wide, ovate to ovate-oblong, obtuse to acutish, erect to spreading, occasionally incurved; staminal scales about reaching the filaments, only shortly bridged, oblong, apically fringed with slender, medium-long filaments, sometimes truncate or bifid; filaments rather stout, to 0.6 mm long, usually a little longer than the anthers, anthers oval to subglobose, to 0.5 mm long; styles short and subulate rather than filiform, to 1 mm long, sometimes elongating a little in fruit, widely separated at the base and the interstylar opening usually obvious (at least in fruit), stigmas capitate, ovary globose. Capsule depressed-globose,to 3 mm wide, the withered corolla usually persistent at the base; seeds about 1.5 mm long, ovoid. Occasional throughout the state but most common in the SE; confirmed from our area; Gulf States, also Mex. and W.I. The typical variety is S. American. Parasitizing various plants but commonly on Polygonum and so confused with C. polygonorum (which has 4-merous flowers). [C. glandulosa (Engelm.) Small].
6. C. coryli Engelm. Hazel Dodder. Stems medium to slender, 0.4 to 0.6 mm wide. Flowers in cymose-paniculate clusters, sometimes produced endogenously and appearing in dense, glomerulate clusters directly on the stem of the host; pedicels shorter than to longer than the flowers. Flowers ca. 1.5 to 2 mm long from base to corolla sinuses, fleshy, papillate externally, 4-merous (rarely 5-merous); sepals united basally, the lobes about equalling the corolla tube, triangular-ovate, acute, little if at all overlapping, sinuses between the lobes acute; corolla tube cylindric-campanulate, lobes about as long as the tube, ovate-lanceolate to triangular-ovate, erect, margins minutely crenulate and the tips acute, inflexed; staminal scales bridged below the middle, usually reduced to toothed wings on either side of the filament base, rarely free and bifid or toothed; stamens about as long as the corolla lobes, filaments subulate and rather stout, anthers ovoid to oblong; styles shorter than to as long as the ovary, slenderly subulate, divergent in fruit, sometimes widely so, stigmas capitate. Capsule globose but becoming rather depressed-globose at full maturity, the interstylar opening relatively large and thick-margined, corolla persistent around the base but eventually falling; seeds 1.5 mm long, usually 4, flattened-globose. On a wide variety of woody and herbaceous hosts including Salix, Carya, Rhus, Lechea, Callicarpa, Symphoricarpos, Solidago, Helianthus, Aster, etc. Known in our area at least from Robertson Co.; N. S. to MT, S.to FL and AZ. Summer-fall.
7. C. indecora Choisy Two varieties in our area.
var. indecora Pretty Dodder, Showy Dodder, Largeseed Dodder. Stems 0.4 to 0.6 mm in diameter. Inflorescences loose to dense paniculate-cymose clusters. Flowers 2 to 3 mm long from base to corolla sinuses, variable in size and proportions, whitish, smooth or more usually granulate to papillate-hispid; pedicels equal to or shorter than the flowers or sometimes even longer, 2 to 5 mm long. Calyx usually shorter than the corolla tube, the lobes triangular-ovate, overlapping a little basally, acute or obtuse-ish, to 1.5 mm long and wide; corolla campanulate, the lobes shorter than the tube, erect to spreading, triangular-ovate, the tips acute and inflexed; staminal scales reaching the filaments, bridged at or below the middle, ovate to oblong or spatulate, with abundant medium-length fringe; stamens shorter than the corolla lobes or about as long, filaments stout, to 0.7 mm long, about equalling the oblong anthers which are often purplish; styles slender to subulate, about equalling the ovary, to 1.7 mm long, divergent in fruit; ovary globose-ovoid. Capsule depressed-globose, to 4 mm broad, the rim of the interstylar opening thickened, fruit usually enveloped by the withered corolla which eventually splits and falls; seeds ca. 1.7 mm long, mostly oval, with 1 or 2 flat faces. On a wide range of woody and herbaceous hosts of various families. Definitely present in our area; common in the SE. U.S.; FL to CA, N. to MN, MI, and SD; also Mex., W.I., and S. Amer. Summer-fall. [Includes var. hispidula (Engelm.) Yuncker and var. subnuda (Engelm.) Yuncker; C. decora Engelm. and var. indecora (Choisy) Engelm., var. pulcherrima (Scheele) Engelm., and var. subnuda Engelm.].
var. longisepala Yunck. Longsepal Dodder. As described for var. indecora above, but the calyx lobes lanceolate and generally longer than the corolla tube. More or less throughout the E. 1/2 state.
8. C. gronovii Willd. ex R. & S. Three varieties have been listed for TX; 2 possible here. Kartesz (1998) combines both of the following under var. gronovii.
var. gronovii Gronovius' Dodder, Common Dodder. Stems of medium width or sometimes coarse, 0.4 to 0.6 mm broad. Inflorescences loose or dense paniculate-cymose clusters or occasionally formed endogenously; pedicels 0.5 to 4 mm long, usually shorter than the flowers, rarely longer. Flowers 2 to 2.5 mm long from base to corolla sinuses, often with sparse to dense translucent, gland-like cells. Sepals basally united, calyx ca. 1/2 as long as the corolla tube, lobes oval-ovate to suborbicular, obtuse, bases overlapping, margins usually entire or rarely serrulate, to 1.7 mm long and 1.5 mm broad; corolla tube campanulate to globose, lobes shorter than the tube, ovate, obtuse, spreading or reflexed, to 1.5 mm long and 1.5 mm wide; staminal scales shorter than the corolla tube, bridged below the middle, not quite reaching the stamen attachments, oblong, deeply fringed apically and less so basally and on the bridge; filaments to 1 mm long, stout and somewhat subulate, about equalling or longer than the anthers, anthers oval-oblong, to 0.7 mm long; styles 1.5 to 2 mm long, shorter than to equalling the ovary, thickened at the base, stout to somewhat subulate, stigmas capitate; ovary depressed-globose. Capsule globose-conic to obpyriform, enclosed in the withered corolla, to 4 mm broad; seeds 2 to 4, oblong-ovoid to subglobose, slightly flattened. On a wide variety of herbaceous and woody hosts. Very common in the NE. states and Canada; N. S. to ND and Man., S. to FL, TX, and AZ; also W.I. Summer-fall. [C. vulgivaga Engelm. and varieties thereof].
var. latiflora Engelm. Similar to the typical variety, but the flowers smaller, calyx lobes more oblong-oval, overlapping less at the base, and long enough to reach the corolla sinuses; corolla tube broadly campanulate, the throat wider than the tube and the tube tapered toward the base so the withered corolla usually borne at the base of the capsule and the capsule protruding and essentially naked. In the Blackland Prairie and Post Oak Savannah regions.
9. C. pentagona Engelm. Three varieties in TX.:
1. Calyx lobes overlapping and often forming angles at the base; withered corolla enclosing the smooth
capsule to about the middle; corolla not saccate; interstylar opening inconspicuous ...................9a. var. pentagona
1. Calyx lobes scarcely overlapping; withered corolla enclosing most of the capsule, which is papillate apically; interstylar opening relatively conspicuous ...................................................................................................................2
2(1) Calyx and pedicel not papillate .........................................................................................................9b. var. glabrior
2. Calyx and pedicel papillate ..........................................................................................................9c. var. pubescens
9a. var. pentagona Field Dodder, Five-angled Dodder. Stems slender to medium in width. Inflorescences loose, cymose clusters or glomerulate-cymose clusters; pedicels to 1.7 mm long, about equalling to slightly shorter than the flower. Flowers 1 to 2.5 mm long from base to corolla sinuses, enlarging as the fruit develops, smooth or with scattered, translucent, glandular-like cells. Calyx about as long as the corolla tube, the lobes overlapping at the base, sometimes forming strong angles at the sinuses, ovate to oval-ovate or rhomboid, obtuse, to ca. 1.6 mm long and about as wide, sometimes unequal, minutely serrulate; corolla campanulate, lobes lanceolate or triangular-ovate, about equalling or slightly longer than the tube, acute to acuminate, spreading or reflexed but with tips inflexed, to 1.2 mm long and 1 mm broad, sometimes granulate, but not papillate;
staminal scales bridged below the middle, ovate-oblong, reaching the filaments, conspicuously and abundantly fringed, to ca. 1.5 mm long; stamens shorter than the corolla lobes, filaments to 0.6 mm long, equal to or slightly longer than the oval anthers which may reach 0.4 mm long; styles to 0.7 mm long, slender, equalling or slightly shorter than the ovary, stigmas capitate; ovary globose. Capsule depressed-globose to ovoid, enclosed below the middle by the withered corolla, interstylar opening more or less inconspicuous; seeds basically ovoid, ca. 1 to 1.5 mm long. The host plant varies, but the plant sometimes prefers cultivated legumes such as alfalfa. Throughout much of the U.S., from the E. coast W. to MT and CA; also Mex. and W.I. About May-July(Sept.). [Includes var. microcalyx Engelm., var. typica Yuncker; includes C. campestris Yuncker, formerly separated on the basis of a non-angled calyx. The angularity of the calyx has proven to be variable and unreliable (Gandhi, Thomas, and Hatch 1987); C. arvensis Beyr. and var. pentagona (Engelm.) Engelm].
9b. var. glabrior (Engelm.) Gandhi, Thomas, & Hatch Stems smooth, medium in diameter. Inflorescences loose or dense cymose-globular clusters, subsessile or on short pedicels usually no longer than the flowers. Flowers ca. 2 mm long from base to corolla sinuses, white to reddish, often with many translucent, glandular-like cells. Calyx united to about the middle, about as long as the corolla tube or slightly shorter, relatively loose around the corolla, lobes ovate, oval-ovate, or triangular-ovate, usually not overlapping at the base and the sinuses sometimes slightly obtuse, tips obtuse or sometimes acute or acuminate; corolla campanulate to subglobose, often becoming saccate or urceolate between the stamen attachments, smooth or often papillate to some degree, lobes triangular to nearly lanceolate and about as long as the tube, spreading to reflexed and the tips acute to acuminate, inflexed; staminal scales bridged below the middle, oblong-spatulate, with conspicuous fringe; stamens shorter than the corolla lobes, filaments slender, about as long as to slightly longer than the anthers, anthers oval; styles equalling the ovary or a little longer, stigmas capitate; ovary depressed-globose, more or less scabrous-papillate, especially above the middle. Capsule depressed-globose, the interstylar opening relatively large, surface with many translucent, glandular cells and scabrous at least apically, rarely smooth, surrounded nearly completely by the withered corolla, easily broken from the calyx and possibly mistakable for circumscissile; seeds ca. 1 mm long, more or less oval in outline. Parasitic on a wide variety of plants, primarily herbs. Throughout much of TX; definitely present in our area; NM to TX, OK, and LA; S. into Mex. Summer-fall. [C. glabrior (Engelm.) Yuncker and forma pedicellata Yuncker; C. verrucosa Engelm. and var. glabrior Engelm.; C. pentagona Engelm. var. verrucosa (Engelm.) Yuncker, C. arvensis Beyr. var. verrucosa (Engelm.) Engelm.].
9c. var. pubescens (Engelm.) Yuncker Similar to var. glabrior, except the pedicel and all floral parts densely papillate. Mostly W. of our area but known from calcareous outcrops Grimes Co. [C. glabrior (Engelm.) Yuncker var. pubescens (Engelm.) Yuncker; C. arvensis Beyr. var. pubescens Engelm.].
Perennials, stems trailing or creeping, sometimes rooting at the nodes and/or mat-forming. Leaves relatively long-petiolate, commonly erect, blades orbicular-ovate to orbicular-reniform, entire. Flowers small, single or paired in the axils, long-pedicelled. Sepals 5, basally united. Corolla rotate-campanulate or shallowly funnelform, deeply 5-lobed, pale green to white. Ovary deeply 2-lobed. Fruit dehiscent or indehiscent, with 1 to 4 seeds.
9 species of tropical and subtropical areas; in TX; 1 here with another to be looked for. Sometimes placed in the separate family Dichondraceae.
Some species can be grown as groundcovers or lawn substitutes but will not take as much foot traffic as grass.
1. D. carolinensis Michx. Grass Ponyfoot. Stems prostrate to spreading, rooting at the nodes and patch- or mat-forming, sparsely pubescent, to 12 cm tall, but this often only the height of the leaves. Leaf blades suborbicular to reniform, (0.5)1 to 3 cm broad, sparsely pubescent below, green on both surfaces, generally held upright on petioles 1 to 4 cm long, sometimes shallowly cupped; leaf buds often nodding. Flowers solitary, pedicels 1/3 to 2/3 as long as the petioles, straight and upright, not recurved. Sepals 2 to 3 m long at anthesis, 2 to 3 times longer than wide, oblong or linear to spatulate, obtuse to rounded, pubescent externally; corolla white to light green, rotate-campanulate, slightly shorter than the calyx, the lobes rounded; anthers white; the 2 carpels nearly separate, the 2 globose halves side by side, pubescent; styles separate, stigma capitate. Seeds usually 1 per carpel. Damp ground of lawns, roadsides, open woods, etc. S. TX; TX to VA and FL. Feb.-June. [D. repens Forst. and var. carolinensis (Michx.) Choisy.
Sometimes grown as a ground cover.
NOTE: D. recurvata Tharp. & M. C. Johnst. has been reported for Cen. TX but, so far, has not been collected in our area. It may be present in the far W. portion of our region. It has pedicels recurved abruptly near the summit and calyx 2.5 to 3.2 mm long at anthesis, the lobes 1.5 to 2 times longer than wide. D. micrantha Urban is similar, but has a calyx 1.5 to 2 mm long at anthesis and stems mostly less than 1 mm wide. It is often cultivated as a lawn substitute in the south; we are out of its natural range, but it may be present as an occasional escape.
Perennial herbs. Stems usually several to many from the base, prostrate to erect, not twining. Herbage pubescent. Leaves sessile to subsessile, entire. Flowers axillary, solitary or in few-flowered cymes, sessile or pedunculate. Sepals 5. Corolla rotate to broadly and shallowly funnelform, white to lavender, purple, or blue (ours white to lavender, sometimes drying purple). Ovary with 1 or 2 locules; styles 2, each 2-branched, stigmas filiform. Capsule with 1 to 4 seeds.
98 species of tropical and subtropical America (2 extending to the E. hemisphere); 3 in TX; 2 here.
1. Leaves densely pubescent on both surfaces, the hairs loosely appressed to spreading; sepals linear to narrowly lanceolate; corolla lavender to nearly white, drying purple .............
......................................................................................................................1. E. nuttallianus
1. Leaves densely pubescent below, the hairs mostly closely appressed or a few loose, upper surface glabrous or only sparsely pilose; sepals lanceolate to ovate-lanceolate; corolla white, drying cream ................................................................................2. E. sericeus
1. E. nuttallianus J. A. Schultes Hairy Evolvulus. Plants erect to partly decumbent, sometimes suffrutescent; stems several to many from a branched or creeping rootstock, 5 to 25 cm long or tall; herbage densely pilose, hairs rusty to brown or gray, loosely appressed to spreading. Leaves appressed or closely ascending, lanceolate to linear- or elliptic-oblong, 6 to 20 mm long, 1 to 8 mm broad, longer than the flowers, acute to obtuse, basally attenuate. Flowers usually solitary, peduncle none if flowers more than 1 per cluster; pedicels shorter than the calyces, ca. 3 to 4 mm long; bracteoles 1 to 4 mm long, subulate. Sepals linear to narrowly lanceolate, acuminate, 4 to 5 mm long, densely pilose; corolla rotate-campanulate, lavender to nearly white (occasionally blue), drying purple or violet, 8 to 12 mm across; anthers 1 to 2 mm long, filaments about twice as long. Pedicels often recurved in fruit; capsule globose or ovoid, ca. 4 mm in diameter; seeds (1)2, small and brownish. Sandy or rocky soil. Throughout much of TX except the Piney Woods; supposed to be present in our area. AZ to MT and TN. Apr.-July. [Authority sometimes given as R. & S.; E. pilosus Nutt.; E. argenteus Pursh].
2. E. sericeus Sw. Silky Evolvulus, Dwarf Morning Glory. Perennial from a slender, woody rootstock; stems several to many, erect to prostrate or ascending, 6 to 40 cm long, slender, wiry, densely appressed-pubescent. Leaves generally spreading, sessile or subsessile, lanceolate to linear, elliptic, or even oblanceolate, the upper usually much narrower than the lower, 1 to 3 cm long, 1 to 8 mm broad, densely appressed-sericeus or sometimes with a few loose hairs below, glabrous or sparsely appressed-pubescent or pilose above, apex acute to obtuse, base cuneate to attenuate. Flowers generally solitary in the axils, usually sessile or subsessile with the pedicel shorter than the calyx, rarely pedicels longer; bracteoles subulate or narrowly linear, ca. 1 to 4 mm long. Sepals lanceolate or ovate-lanceolate, short-acuminate, appressed-pubescent, ca. 4 to 5 mm long; corolla rotate to campanulate, usually whit or cream, rarely tinged pink or light blue-violet, usually drying cream, ca. 10 to 18 mm across; anthers 1 to 2 mm long, shorter than the slender filaments. Pedicels commonly recurved in fruit; capsule more or less globose or ovoid, ca. 4 to 5 mm long; seeds sometimes as many as 3, brown or olivaceous. Usually in sandy or silty soil in open areas such as prairies, open woods, roadsides, etc. Cen., W., and S. TX; CA to FL, S. to Arg. Apr.-Oct. [Includes forma pedunculatus v. Ooststr. and var. discolor (Benth.) Gray; E. wilcoxianus House. Kartesz (1998) maintains var. sericues (= var. discolor (Benth) Gray )and var. glaberrimus B. L. Robins (= E. macilentus Small; E. sericeus Sw. var. averyi Wood)].
Ours annual or perennial herbs (elsewhere also shrubs). Stems trailing to twining, decumbent, or in some taxa erect, glabrous to sparsely or densely pubescent. Leaves sessile to petiolate, blades sagittate, hastate, or ovate, less often ovate-lanceolate or linear, entire to crenate, lobed, or laciniate, glabrous or pubescent. Flowers axillary, solitary or cymose; peduncle and pedicels developed. Bracts scale-like to linear, elliptic, ovate, or else absent. Sepals nearly equal or the outer shorter than the inner, in ours suborbicular to ovate or elliptic, glabrous or pubescent. Corolla funnelform or funnelform-campanulate, 5-angled or with 5 shallow lobes, white to pink or rose, glabrous or margins puberulent. Stamens included, unequal, 2 shorter than the other 3, base of filaments glandular pubescent; anthers basally auriculate. Ovary ovoid to subglobose, 2-celled, glabrous to pubescent; style 1, stigmas 2, filiform to cylindrical or subulate, more or less flattened, slightly acute. Capsule 2-locular, with 4 valves. Seeds 1 to 4, glabrous.
About 250 species worldwide, especially in temperate regions; 2 species in TX, both present in our area. Plants of the genus Calystegia (ca. 25 species worldwide; 2 in TX, neither apparently here) are sometimes included in Convolvulus but differ in having, among other features, usually conspicuous floral bracts and unilocular fruits.
Several species are terrible weeds, especially C. arvensis, a native of Eurasia that is widely
naturalized. Other species are cultivated for their flowers, notably C. tricolor, whose blossoms are white or purple with a white throat and yellow eye (Mabberley 1987).
1. Calyx 6 to 12 mm long, densely pubescent; plants from a taproot, sometimes dividing, but not forming creeping patches; leaves densely pubescent on both surfaces ..1. C. equitans
1. Calyx 3 to 5 mm long, inconspicuously pubescent or glabrate; plants from a deep, creeping root, often forming large patches; leaves glabrous to inconspicuously puberulent
............................................................................................................................2. C. arvensis
1. C. equitans Benth. Gray Bindweed, Texas Bindweed. Plants from a taproot; stems prostrate, decumbent, or sometimes twining, to 2 m long, densely pubescent. Petioles 0.25 to 5 cm long, generally 1/4 to 3/4 the length of the blade; blades 1 to 7 cm long, 0.2 to 4 cm broad, quite variable in shape, ovate-elliptic to triangular-lanceolate or narrowly oblong, commonly with spreading basal lobes and thus more or less hastate or sagittate, apex rounded to acute, base commonly deeply indented or sometimes truncate, margin rarely entire, usually toothed, lobed, or both, the extreme form being more or less palmatifid with the central lobe the longest, both surfaces densely covered with loosely appressed hairs. Flowers usually solitary in the axils (occasionally paired, rarely in 3's), peduncles 0.5 to 10.5 cm long; pedicels shorter, 5 to 24 mm long, to about twice as long as the calyx; bracts and bracteoles scale-like or subulate, pubescent like the leaves. Sepals oblong to ovate, (5)6 to 12 mm long, 3 to 6 m broad, apically obtuse to somewhat retuse, margins membranous and lighter than the body, surface rather densely appressed-sericeous, in age the bases often strongly shouldered, auriculate, or cordate; corolla funnelform or campanulate, base somewhat tubular, 1.5 to 3 cm long, 5-angled, the angles sometimes extended into slender points, white to pink, commonly with a red(dish) center, pubescent externally between the pleats. Capsule globose to subglobose, 7 to 8 mm broad, glabrous; seeds 1 to 4 per fruit, 4 to 4.5 mm long, black, glabrous and with a granular surface. Most often in sandy or rocky soil of prairies, disturbed areas, outcrops, hills, etc. Throughout most of TX except the Piney Woods; KS and CO to TX and AZ, S. to Mex. and S. Amer. In our area about Apr.-Oct. [C. hermannioides A. Gray; C. incanus of various authors but not C. incanus Vahl].
This plant is sometimes a troublesome weed in crops such as cotton and grains (Kirkpatrick, 1992).
2. C. arvensis L. Field Bindweed, Common Bindweed, Possession Vine. Perennial from a rhizomatous, spreading root; stems decumbent or trailing to twining, to 1 m or more long, glabrous to densely and loosely pubescent. Petioles 3 to 40 mm long, shorter than to equalling the blades; blades 1 to 10 cm long, 3 to 40(60) mm broad, quite variable in shape, oblong-elliptic to deltoid-ovate, ovate, ovate-lanceolate, or narrowly oblong, apex obtuse to acute, base truncate or usually strongly indented, commonly with 2 basal acute to obtuse lobes and so sagittate or hastate, lobes entire or with 2 or 3 teeth, glabrous to more or less (but usually inconspicuously) puberulent, especially below. Flowers in cymes of 2 or 3 or often solitary; peduncles 1 to 9 cm long; pedicels 5 to 18(35) mm long, 1 to 3 times longer than the blades, glabrous to minutely pubescent; bracts elliptic, linear, or obovate, 2 to 3(9) mm long; bracteoles 2 to 4 mm long, linear. Outer sepals elliptic, 3 to 4.5 mm long, 2 to 3 mm broad, glabrous or loosely pubescent; inner sepals suborbicular to obovate, 3.5 to 5 mm long, 3 to 5 mm broad, all sepals with scarious margins, mucronate, ciliate, obtuse or sometimes truncate or emarginate; corolla open-funnelform or campanulate, opening in the morning, 1.2 to 2.5 cm long, white or occasionally pink or lavender-pink, the outside sometimes with vertical bands of pink or brownish-lavender along the main ribs; stamens 8 to 13 mm long, anthers 2 to 3.5 mm long; ovary ovoid, glabrous; style 7 to 10 mm long. Mature pedicels usually recurved in fruit; fruits subglobose to ovoid, 5 to 7 mm broad, glabrous; seeds 1 to 4 per fruit, 3 to 4 mm long, black to dark brown, tuberculate and glabrous. Roadsides, waste places, cultivated areas, railways, etc. Cen. and W. TX to the Panhandle, possibly spreading in the state; native of Eurasia and now a pan-temperate weed. Late Apr.-July. [Strophocaulos arvensis (L.) Small; C. ambigens House].
This plant can be an aggressive weed in field crops (Mabberley 1987). It is listed as a noxious weed in CA. (Hickman 1993). Tull (1987) mentions a yellow dye made from the whole plant.
Herbaceous perennial from single or fascicled roots. Stems several to many, prostrate to ascending, straight or twining, sometimes forming clumps or mats, or sometimes the lower portions appearing rhizomatous, overall to 2 m long or more. Leaves sessile or short-petiolate, the blades entire, variously shaped. Flowers solitary or few in pedunculate axillary clusters. Sepals 5. Corolla funnelform to campanulate, with 5 shallow lobes or 5-angled, white to lavender. Stamens 5, exserted or included. Styles 2, united at the base or nearly to the summit, free branches 2, sometimes short or unequal (rarely 1 obsolete); stigmas small, capitate. Capsule chartaceous, 2-locular, 1- to 4-seeded, longer than wide.
6 species of the S. and E. U.S.; 4 in TX; 1 here. Formerly included by some authors in Bonamia and some taxa with synonyms in Breweria.
1. S. pickeringii (Torr. ex M. A. Curtis) Gray var. pattersonii (Fern. & Schub.) Myint. Bigpod Bonamia. Plants usually from a single woody root, the woody bases of previous seasons' stems often remaining; stems 1 to 2 m long or more, prostrate, trailing or reclining on surrounding plants, often pale purple-brown, sparsely to densely and minutely pubescent and with a few longer hairs. Leaves sessile or essentially so, blades linear, acute to obtuse, entire, 2 to 7 cm long, 1 to 3 m broad, minutely pubescent (at least beneath) to glabrate. Flowers 1 to 5 in bracteate axillary cymes; peduncles 3 to 7 cm long, equalling or exceeding the calyces; pedicels of lateral flowers 4 to 20 mm long, central flowers sometimes sessile; bracts similar to leaves (0.5)1.5 to 2.5 cm long. Sepals ovate to ovate-lanceolate or ovate-orbicular, apically acute to obtuse, 3.5 to 6 mm long, 3 to 5 mm broad, with pale appressed hairs dorsally; corolla campanulate to campanulate-funnelform, 10 to 18 mm long, limb entire or shallowly 5-lobed, pubescent between the pleats; stamens slightly exserted, filaments glabrous or rarely with scattered hairs near the base, anthers basally sagittate, oblong; ovary ovoid, bilocular, villous; styles fused almost to the base of the stigmas, style branches unequal, 1 to 1.5 mm long, stigma capitate. Capsule ovoid, pubescent; seeds 1 or 2, smooth and brown. Sandy open areas such as prairies, open woods, and bog margins. This variety in E. and Cen. TX; E. IA, IL, and KS, S. to TX and LA. May-Sept. [Breweria pickeringii (Torr.) A. Gray var. pattersonii Fern. & Schub.; S. pattersonii (Fern. & Schub.) G. N. Jones].
About 120 species of tropical and subtropical areas; 1 species in TX.
1. J. tamnifolia (L.) Griseb. Hairy Clustervine. Annual; stems twining, climbing, or creeping, sparsely to densely pilose, to 2 m long, but often beginning to flower while still young and erect. Leaves petiolate; blades cordate or ovate to elliptic-ovate, 3 to 12 cm long, 2 to 9 cm broad, acuminate, the larger ones basally cordate, some with bases rounded, margin pilose, surfaces sometimes sparsely pilose. Peduncles axillary, often shorter than the leaves at anthesis, elongating with age and often ending longer than the leaves; inflorescence densely cymose, more or less capitate, 2 to 3 cm broad, subtended by leafy, lanceolate to elliptic bracts. Sepals 5, free, lanceolate to subulate, densely hispid-pilose, ca. 2/3 as long as the corolla; corolla 12 to 16 mm long, 10 to 20 across, funnelform-campanulate, 5-angled, blue; stamens 5; style 1, stigmas 2, ovoid, oblong, or elliptical, more or less flattened; ovary bilocular. Capsule subglobose, 4 to 6 mm long, 4-seeded; seeds ca. 2 mm long, brown to black, glabrous. Roadsides, disturbed or cultivated areas, and streambeds. E. TX; VA to FL, W. to TX, S. to Brazil. July-Oct. [Thyella tamnifolia (L.) Raf.].
Ours annual or perennial herbs (elsewhere also woody vines, shrubs, and some trees). Stems prostrate to erect or trailing, creeping, or twining to the right, glabrous to variously pubescent, some (none of ours) rooting at the nodes. Leaves sessile to petiolate, simple and entire to lobed or else palmately compound or pinnately dissected, base of simple or lobed leaves commonly cordate and apices acute to acuminate, surfaces glabrous or variously pubescent. Flowers axillary or terminal, in cymose, thyrsoid, or rarely racemose inflorescences, sometimes flowers solitary; peduncles and pedicels glabrous to pubescent. Bracts present, varying from scale-like to leafy; bracteoles scale-like. Sepals 5, overlapping, often unequal, from nearly linear to nearly orbicular, variously pubescent. Corolla typically funnelform, sometimes salverform, very rarely campanulate or urceolate, usually glabrous, variously colored, in our species shades of white, pink, purple, blue, or red, usually open for less than a day. Stamens included or exserted, unequal, filaments glandular-pubescent at the base; anthers oblong. Ovary with 2 to 4 locules; style 1, stigma globose or with 2 or 3 globose lobes. Capsule variously dehiscent or indehiscent, with 1 to 4(6) seeds. Seeds glabrous to pubescent.
About 500 species from the tropics to the warm temperate regions; 39 listed for TX (Hatch, et al. 1990); 7 here
The genus includes many ornamentals, among them I. purpurea, I. tricolor, I. violacea, etc. Sweet potatoes, commonly called yams in the southern U.S. (but not to be confused with the true yam, Dioscorea), are I. batatas. some species have medicinal value, mostly as purgatives, and the seeds of some contain hallucinogenic alkaloids (Mabberley 1987).
NOTE: Because they are not persistent members of our flora, the following key does not include species likely to be found only in cultivation or around homesites.
1. Leaves pinnately dissected nearly to the midrib, the segments linear-filiform; corolla salverform, deep red .........................................................................................1. I. quamoclit
1. Leaves entire to palmately lobed or compound; corolla funnelform, white to pink, purple, or blue .......................................................................................................................................2
2(1) Leaves palmately compound ................................................................................2. I. wrightii
2. Leaves simple and entire to lobed, but not divided to the base ..............................................3
3(2) Sepals linear-lanceolate, with elongate, slender, erect to recurved tips much longer than the body; corolla bright blue ............................................................................3. I. hederacea
3. Sepals oblong to ovate, tips straight and shorter than to only slightly longer than the body; corolla white to purple, pink, or reddish, rarely blue ...............................................................4
4(3) Peduncles, pedicels, and usually petioles with reflexed hairs (some spreading hairs may be present as well); sepals lance-oblong ..........................................................4. I. purpurea
4. Peduncles, pedicels, and petioles glabrous or with spreading or ascending hairs; sepals ovate to lanceolate ....................................................................................................................5
5(4) Corolla 5 to 8 cm long and about as wide, white with a lavender to red-purple tube; sepals subequal ...........................................................................................................5. I. pandurata
5. Corolla 1.8 to 5.5 cm long, white or pale rose to purple-rose; sepals often markedly unequal .....................................................................................................................................6
6(5) Corolla white (rarely pinkish), 1.8 to 2.3 cm long ..............................................6. I. lacunosa
6. Corolla rose-lavender to purple-rose, usually with a darker throat (rarely white), 2.8 to 5.5 cm long ......................................................................................................7. I. cordatotriloba
1. I. quamoclit L. Cypress Vine. Annual herb with viny twining stems to ca. 3 m long; herbage glabrous or essentially so. Petioles 0.2 to 4.5 cm long; blades oblong or ovate to elliptic in outline, 1 to 9 cm long, 0.8 to 7 cm broad, pinnately divided nearly to the midrib, with 9 to 19 pairs of segments 0.2 to 1.5 mm broad, about as wide as the midrib, acute and minutely mucronate. Peduncles 1.5 to 14 cm long, bearing (1)2 to 5 flowers in each cyme; bracts narrowly elliptic to ovate, mucronate, 0.6 to 1.5 mm long; bracteoles similar, 0.6 to 1 mm long; pedicels 8 to 25 mm long, thickened below the flower; flowers open throughout the day. Sepals about equal, elliptic-oblong, obtuse and mucronate, the outer ones ca. 4 to 6 mm long, 2 to 3 mm broad and with a subterminal bristle 0.25 to 0.3 mm long, inner sepals 5 to 7 mm long, 3 to 3.8 mm wide, subterminal bristle 0.3 to 0.6 mm long; corolla deep red, essentially salverform, 2 to 3 cm long, the limb 1.8 to 2 cm broad; stamens exserted, 2.5 to 3 cm long, anthers 1.5 to 1.8 mm long; ovary glabrous, 4-locular, style exserted, 2.3 to 3 cm long, stigma lobes 2. Capsule ovoid, glabrous, 7 to 9 mm broad, with a terminal point 4.5 to 6 mm long; seeds black, 4.5 to 5.5 mm long, sparsely tomentose. In our area usually cultivated; known from a few Brazos Co. collections, some of which were probably from cultivation even if not designated as such; probably not a permanent member of our flora; native to S. Amer.; naturalized in the SE. U.S.--VA to KS, S. to FL and TX; also Mex. July-Nov. [Quamoclit pennata Bojer; Q. quamoclit (L.) Britt.; Q. vulgaris Choisy].
This plant is a beautiful addition to the home landscape, especially on arbors and fences. It is attractive to butterflies and hummingbirds.
2. I. wrightii Gray Wright Morning-glory. Annual trailing or climbing vine; stems to 2 m or more, glabrous. Petioles 1.5 to 5 cm long; blades palmately compound, leaflets 3 to 7, rhombic to lanceolate (or linear-lanceolate), acute or else obtuse and mucronate, (1.5)2 to 6 cm long, subequal or unequal, usually entire or the lowermost sometimes 2-lobed. Peduncles slender, variable in length (the lowermost shortest and uppermost longest), commonly curled or coiled and aiding in climbing; flowers usually solitary; bracts paired, minute; pedicels thicker than peduncles. Sepals subequal, 4 to 6 mm long, ovate or ovate-oblong, rounded to obtuse and mucronate; corolla funnelform, 1 to 2 cm long, pinkish lavender or rose, throat darker, drying purplish, 1 to 2 cm long; stamens and style included. Capsule subglobose, ca. 1 cm broad. Silty or alluvial damp soil of creeksides, fencerows, etc. Common in S. and Cen. TX; probably native to India and widely naturalized in warm regions worldwide. June-Oct. [I. heterophylla (Rottb. & Willd.) Voigt; I. pulchella of authors but not I. pulchella Roth. (though many old specimens are labeled as such); I. spiralis House].
3. I. hederacea Jacq. Ivy-leaf Morning-glory. Annual herbaceous twining vine; stem with spreading or subappressed short hairs and spreading longer hairs. Petioles shorter than to equalling the blades, 5 to 12 cm long; blades 5 to 12 cm long and about as wide, cordate-ovate to suborbicular, entire or 3-lobed, apex and lobe apices acute to acuminate, sparsely pubescent above, paler and slightly more densely and more softly pubescent beneath. Peduncles shorter than the subtending leaves, 5 to 10 cm long; flowers 1 to 3(6) per cyme, open in the mornings; bracteoles scale-like; peduncles and pedicels with at least some hairs retrorse. Sepals subequal, base ovate, apex elongate, linear-lanceolate, spreading or recurved, 1.2 to 2.4 cm long, densely hirsute, at least on the basal portion, sometimes enlarging in fruit; corolla funnelform, 1.5 to 5 cm long, 3 to 5 cm across the limb (slightly larger in cultivated material), tube white or pale yellow, limb deep, clear, sky blue, fading red-purple and red-purple in dried material; stamens ca. 2/3 as long as the corolla, white, not exserted; style included, ovary pubescent; nectary white. Capsule subglobose, slightly flattened, 8 to 12 mm broad, shorter than the enclosing persistent sepals; seeds commonly 4, 4 to 5 mm long, pear-shaped and with 2 flat faces and 1 rounded, black or dark brown, densely short pubescent. Fencerows, fields, streambanks, disturbed ground, etc. E. TX, W. to W. Cross Timbers, S. to the Rio Grande, scattered upriver to Big Bend; probably native to the E. U.S., often cultivated and escaping; seeds also spread as a contaminant in crop seeds; ME to SD, FL to Mex.; also W.I. July-Nov. [Includes var. integriuscula Gray; I. desertorum House; I. barbigera Sims or I. barbigera Sweet; Pharbitis barbigera (Sweet) G. Don; Pharbitis hederacea (Jacq.) Choisy].
4. I. purpurea (L.) Roth Common Morning-glory, Mexican Morning-glory, Woolly Morning-glory. Annual herbaceous vine; stems twining; herbage and inflorescence with loose or appressed, usually retrorse trichomes and also with antrorse to oblique or erect hairs to 4 mm long. Petioles 1 to 14 cm long, pubescent like the stem; blades cordate-ovate or shallowly to deeply 3-(5-) lobed, 1 to 11 cm long, 1 to 12 cm broad, base cordate, apex acute to acuminate (rarely obtuse), mucronate, upper surface sparsely pubescent, lower surface paler and more densely pubescent. Peduncles 0.2 to 15 cm long, pubescent like the stems; flowers 2 to 5 per cyme, rarely solitary; bracts linear to lanceolate, 1.3 to 9 mm long; bracteoles similar, to 4.5 mm long; pedicels 5 to 16 mm long, in fruit reflexed and enlarged, to 2.5 cm long. Sepals subequal, the outer narrowly ovate-lanceolate to elliptic, 8 to 17 mm long, (1.5)2.5 to 4.5 mm broad, acute to acuminate, densely pubescent, at least at the base, the hairs with enlarged bases, inner sepals 8 to 15 mm long, 2.5 to 3 mm broad, ovate-lanceolate, acute to abruptly acuminate; corolla funnelform, 2.5 to 5 cm long, ca. 4 to 5 cm broad (larger in cultivated forms), purple, bluish, or white, or sometimes the tube white and the limb colored, glabrous; stamens included, unequal, the longer 18 to 25 mm long and the shorter 13 to 22 mm long, anthers 1.5 to 2 mm long; style included, 14 to 22 mm long, stigma globes 3, ovary glabrous. Capsule globose, ca. 1 cm broad, shorter than the sepals, 6-valved; seeds often 6, black to dark brown, ca. 5 mm long, irregularly angled, tomentulose. Waste places and disturbed areas; probably native to Mex., cultivated and escaping and widespread in the E. U.S. and pantropical. Summer-fall. [Includes var. diversifolia (Lindl.) O'Donell; Convolvulus purpureus L.; I. hirsutula Jacq. f.; Pharbitis purpurea (L.) Voigt].
5. I. pandurata (L.) G. F. W. Mey. Bigroot Morning-glory, Wild Potatovine, Man-of-the-earth. Perennial herb from a much-enlarged root; stems trailing or twining, glabrous to sparingly pubescent. Petioles 1 to 8 cm long, shorter than to about equalling the blades; blades 2 to 10 cm long, 2 to 9 cm broad, cordate-ovate and unlobed to shallowly 3-lobed or pandurate, glabrous or pubescent beneath. Peduncles usually equalling or longer than the leaves, 10 to 20 cm long; flowers 1 to several; bracts small, paired; pedicels ca. 1 cm long, slightly thickened. Sepals somewhat leathery, oblong-elliptic, 12 to 15 mm long, often unequal, the outer shorter, mostly rounded, glabrous or sparsely short pubescent; corolla funnelform, 5 to 8 cm long and about as broad, white, almost always with a purplish-red (or lavender) tube interior; stamens included, unequal, 2 to 3 cm long, anthers 5 to 7 mm long; style included, ovary glabrous. Capsule ovoid, 1 to 1.5 cm long, glabrous; seeds 1 to 4, ca. 4 mm long, angled,with 2 faces flat and 1 rounded, dark brown, the angles with long tawny hairs. Roadsides, fencerows, woods, clearings, riverbanks, fields, etc. E. TX, W. to about Bexar, Blanco, and Johnson Cos.; CT to Ont., S. to FL and TX. June-Sept.
Plains tribes used the root as a cathartic and diuretic, as did doctors among early European settlers (Kindscher 1992).
6. I. lacunosa L. Pitted Morning-glory, (Small) White Morning-glory. Annual herb; stems twining to suberect, 1 to 2(3) m long, glabrous or sparsely pubescent with spreading hairs that have enlarged bases. Petioles equalling the blades or slightly longer, 3 to 8 cm long, sparsely pubescent; blades variously shaped, from cordate-ovate and entire to somewhat angled or toothed to deeply 3- to 5-lobed, 2 to 10 cm long, 1.5 to 10 cm broad, base generally cordate, apex acute to acuminate or obtuse, glabrous to sparsely pubescent above and below. Peduncles mostly shorter than or equalling the petioles, 1 to several per axil, each usually with 1 (to 3) flowers subtended by 2 minute bracts; bracteoles scale-like; pedicels minutely warty. Sepals nearly equal, lanceolate or lance-ovate to oblong, (8)10 to 15 mm long, acute to acuminate, mucronate, sometimes falcate, margins ciliate but otherwise glabrous except sometimes roughened in the upper portion; corolla funnelform, 1.5 to 2(2.5) cm long, white or with a trace of pink, glabrous, lobes of the limb short, obtuse, and mucronulate; stamens included, filaments white, anthers purple; ovary pubescent; nectary white. Capsule subglobose to hemispheric, apically bristly-pubescent, (6)10 to 15 mm broad, with 4 valves; seeds 4, rarely fewer, 5 to 6 mm long, ellipsoidal, angled, with 2 flat faces and 2 rounded and strongly humped, dark brown, glabrous. Ditches, stream banks, damp thickets, roadsides, etc. E. TX, W. and S. to Tarrant, Travis, and Matagorda Cos.; PA and KS, S. to SC and TX. Sept.-Oct.
Capable of hybridizing with I. cordatotriloba, below. It is possible that there may be individuals which are not strictly referable to either species.
7. I. cordatotriloba Dennst. Herbaceous perennial, flowering the first year from seed; stems twining, to several m long, sparsely pubescent with spreading hairs with enlarged bases, sometimes nearly glabrous. Petioles 2 to 12 cm long, about equal to slightly longer than the blades; blades 3 to 10 cm long (often all less than about 6 cm long), and about as wide at the base, variable in shape from cordate-ovate and entire to strongly angled to deeply 3- to 5-lobed, apex and lobes acuminate to cuspidate, upper surface usually sparsely pubescent. Peduncles 1 or 2 per axil, from 5 to 20 cm long, with 1 to 5(8) flowers; bracts minute; pedicels 5 to 20 mm long, weakly winged or muriculate. Sepals more or less coriaceous, ovate-lanceolate or linear-oblong, acuminate or cuspidate, (7)10 to 15 mm long, slightly unequal, variously pubescent to glabrous, sometimes with marginal cilia; corolla funnelform, 2.8 to 5.5 cm long, pink-purple (rarely white), usually with a darker eye; stamens and style included, anthers 1.5 to 3.2 mm long. Capsule subglobose or slightly flattened, 6 to 9 mm broad, apically pubescent; seeds angled, with 2 faces flat and 1 rounded but not humped, glabrous, purple-black, 4 mm long. Roadsides, fields, disturbed areas, thickets, etc. Very common in our area. E. 1/2 to 1/3 TX; NC to FL, W. to AR. Jun.-Oct. [I. trichocarpa Ell., a long-used but unfortunately illegitimate name (Austin, 1988).]
Three varieties; two in TX. Capable of hybridizing with I. lacunosa, above.
var. cordatotriloba Sharppod Morning-glory. Stems and leaves glabrous to moderately hispid-pilose; sepals hispid-pilose, at least marginally. E. 1/3 TX (but absent from near the Red River), W. to Medina and Menard Cos.; SE. U.S. Very common here. [I. trichocarpa Ell. var. trichocarpa; I. carolina (L.) Pursh, but not I. carolina L.; I. carolina sensu Small, not sensu Poir.]
var. torreyana (Gray) D. Austin Stems and leaves glabrous; sepals glabrous. Cen. TX: Dallas, Throckmorton, Sutton, and Val Verde Cos. S. to the lower Rio Grande Valley, rare in E. TX; also Mex. [I. trichocarpa Ell. var. torreyana (Gray) Shinners; I. trifida of some authors, in part, but not I. trifida G. Don; I. trifida var. torreyana Gray; I. trifida var. berlandieri Gray].
One of our most common and showiest late summer and fall wildflowers. Tull (1987) reports a pale yellow dye from the flowers.
Ours annual or perennial herbs (elsewhere also subshrubs). Leaves all opposite, all alternate, or opposite below and passing to alternate above, simple and entire to pinnately divided or compound (some other taxa palmatifid), margins entire to toothed, glabrous to pubescent and/or glandular. Flowers terminal or axillary, pedicellate or sessile, solitary, paired, or in cymes, the cymes often clustered into corymb-like or paniculate inflorescences. Flowers perfect, hypogynous, regular or slightly irregular. Sepals (4)5, united basally, the calyx wholly herbaceous and accrescent in fruit or with herbaceous ribs connected by scarious membranes, these usually ruptured by the developing fruit, calyx lobes linear to deltoid, entire to trifid, sometimes spinose. Corolla (4)5-merous, rotate to salverform, funnelform, or campanulate, usually regular, sometimes slightly zygomorphic. Stamens (4)5, included or exserted, epipetalous within the corolla tube, often at different levels, equal or unequal. Carpels 3, united; style 1, with (2)3 linear stigma lobes, included or exserted. Fruit usually a 3-celled capsule, loculicidally dehiscent (or rarely indehiscent or circumscissile, but not in ours). Seeds 1 to many per cell, variously rounded, angular, or winged, in some taxa viscid or mucilaginous when moistened.
About 20 genera and 275 species of the Americas and Eurasia; 7 genera and 33 species in TX; 3 genera and 5 species here, usually on sandy soils.
Many taxa are cultivated for ornament, including species of Phlox, Ipomopsis, and Cobaea (Mabberley 1987).
1. Leaves essentially entire, all opposite or opposite below and passing to alternate above .....
......................................................................................................................................1. Phlox
1. Leaves pinnately toothed, lobed, or dissected, all alternate ...................................................2
2(1) Corolla red with yellow spots, salverform, the limb flaring abruptly from the tube ..................
.............................................................................................................................2. Ipomopsis
2. Corolla blue-violet, rotate, the short tube widening gradually into the limb .................3. Gilia
Ours annual or perennial herbs (elsewhere sometimes subshrubs). Stems usually erect. Herbage pubescent with glandless or glandular hairs or sometimes glabrous. Leaves opposite or opposite below and alternate above, simple, elliptic to linear, subulate, or ovate, entire or essentially so, sessile or subsessile. Flowers terminal, in bracted corymb-like or paniculate clusters or occasionally solitary, 5-merous, regular. Sepals united for ca. 3/8 to 3/4 their length, connected by a scarious/hyaline membranes, calyx teeth acute to acuminate, cuspidate, or aristate, often pubescent within. Corolla salverform, pink, red, lavender, purple, bluish, or white (rarely yellow), the lobes obtuse to cuspidate/apiculate, truncate, or notched-obcordate. Nectary commonly present. Stamens distinctly uneven, the filaments relatively short and inserted unevenly in the corolla tube, anthers usually included or in some taxa exserted. Ovary with either a short or very long style, stigma lobes 3. Capsule ovoid to oblong or ellipsoid. Seeds 1 to few (or several) per locule (typically seeds fewer than ovules), not mucilaginous or viscid when moistened.
About 67 species in N. Amer. (and 1 in NE. Asia); 12 in TX; 3 here. The genus is very diverse in TX and many species have infraspecific taxa.
Phlox provides some important garden plants. P. stolonifera is a creeping rock-garden plant or groundcover, P. paniculata is the parent of many perennial border plants, and our own P. drummondii boasts many cultivars of showy annuals (Mabberley 1987).
1. Leaves nearly all opposite, those of the midstem very narrow, linear and acuminate, generally less than 5 mm broad; plants perennial; sepals awned or aristate with a bristle to 3 mm long ...............................................................................................................1. P. pilosa
1. Leaves generally opposite below and alternate above, those of the midstem usually proportionately broader and obtuse to acute; plants annual; sepals acute to acuminate, awns (if any) short, to 1.5 mm ..................................................................................................2
2(1) Corolla lobes generally less than 7 mm broad, apically pointed, at anthesis usually not overlapping; upper leaves linear, narrowed to the base; plants delicate and with fine pubescence ......................................................................................................2. P. cuspidata
2. Corolla lobes generally more than 7 mm broad, apically truncate to rounded or minutely apiculate, at anthesis usually overlapping or contiguous; upper leaves sessile or clasping, generally rounded at the base; plants commonly robust and coarsely pubescent .................
......................................................................................................................3. P. drummondii
1. P. pilosa L. Prairie Phlox. Perennial from a slender to stout rootstock; stems 1 to several, branched near the base or above, in TX to ca. 60 cm tall and usually with 7 to 15 nodes; herbage pubescent or glandular-pubescent. Most lower and midstem leaf blades narrowly linear, to ca. 12.5 cm long and less than 5 mm broad, acuminate, upper leaves slightly broader and narrowly lanceolate, up to 1 cm broad, all leaves pilose, especially marginally and along the midrib. Flowers in cymes aggregated into open or compact panicles; pedicels to ca. 2 cm long or sometimes flowers subsessile; inflorescence usually strongly glandular. Calyx 8 to 15 mm long, glandular-pubescent, the sepals united to about the middle or below, the lobes subulate to linear-attenuate, apically aristate with an awn to about 3 mm long; corolla glabrous to glandular- or eglandular-pubescent, pink, purple, or white, sometimes with a darker eye, the tube 8 to 16 mm long, the lobes 8 to 16 mm long, oblanceolate to obovate, obtuse to apiculate, ca. 6 to 8 mm broad, at anthesis often not overlapping; styles (1)1.5 to 4 mm long, united about halfway, stigmas positioned below the anthers. Capsule generally with one seed per locule.
Divided into 9 subspecies; 5 in TX; apparently only 1 here.
subsp. pilosa Downy Phlox. Plants with abundant glandular pubescence in the upper parts; larger leaves 4 to 8 cm long, 3 to 9 mm wide (usually in the narrow end of this range); sepals subulate, 8 to 12 mm long, with awns 1.5 to 3 mm long; corolla tube 1 to 1.6 cm long, pubescent to glabrous. Open woods, moist meadows, hillsides, roadsides, creek bottoms, etc. E. 1/3 TX; WI to NY and CT, S. to FL and TX, W. to KS and OK. Spring. [P. pilosa L. var. virens (Michx.) Wherry; P. vilosissima (Gray) Small (in part); P. agrillacea Clute & Ferris.
2. P. cuspidata Scheele Pointed Phlox. Taprooted annual, generally rather delicate and with fine, fairly sparse pubescence; stem simple or branched, 5 to 55 cm tall. Leaves of the lower 4 to 6 nodes opposite, more or less oblanceolate, becoming alternate upwards and linear, to 35 mm long and 6 mm broad. Flowers in rather loose, asymmetric clusters in a vaguely spiral cyme; pedicels to 6 mm long. Sepals 7 to 10 mm long, united 1/3 to 1/2 their length by pale membranes, apices tipped with awns 1 to 1.5 mm long, with fine, spreading glandular pubescence; corolla purple to pinkish (occasionally white) with a paler, dark-striate eye, drying blue, tube 8 to 15 mm long, pilose with some fine glandular hairs, the lobes ca. 3 to 8 mm wide, spatulate-oblanceolate, in flower generally not overlapping, apically abruptly constricted to a definite point; styles 1.5 to 2.5 mm long, fused to about or below the middle. In moderately acidic sandy and sandy loam soils of roadsides, grasslands, pastures, open oak woods, etc.; in our area, at least, commonly associated with Arenaria and Krigia. Throughout the E. part of TX except the Pineywoods; also adjacent OK. Spring, ca. Feb.-May, the later flowers typically smaller; much less common in wet years.
Three varieties have been described, all of which are possible here (but some plants not assignable with confidence). Kartesz (1998) no longer recognizes varietal distinctions.
var. humilis Whiteh. Plants diminutive, leaves mostly less than 3 mm broad, corolla tube ca. 9 mm long, the lobes only ca. 3 mm broad.
var. cuspidata Some leaves more than 3 mm broad, corolla tube 9 to 12 mm long, the lobes ca. 7 mm long and 5 mm broad, usually sharply pointed. Common in our area.
var. grandiflora Whiteh. Some leaves more than 3 mm broad, corolla tube 11 to 15 mm long, lobes ca. 11 mm long and 8 mm broad.
3. P. drummondii Hook. Drummond Phlox. Taprooted annual, varying in stature from 1 to 5 dm tall; stem simple or branched. Leaves of the lowermost 3 to 5 nodes opposite, with subpetiolar bases, relatively narrow, passing upwards into alternate leaves which are broader and sessile or clasping and rounded at the base, pubescence usually relatively coarse. Flowers in asymmetrical clusters of 2 to 6, these grouped into vaguely spiral inflorescences. Sepals ca. 7 to 12 mm long, united ca. 1/3 their length by pale membranes, usually glandular-pilose; corolla quite variable in color, the lobes at anthesis usually overlapping or at least touching, apically truncate to obtuse; styles 2 to 3 mm long, united to the middle or below, stigma usually positioned below the anthers. Seeds usually 1 per cell. Mostly in neutral to moderately acidic sandy soils of open woods, grasslands, roadsides, and so on. E. TX, rarer W. and NW. Spring.
This taxon has been divided into 6 subspecies or varieties based largely on corolla color; 4 variants are possible here.
1. Flower color in a given population highly variable, with many different colors present (red, pink, white, purple, etc.); stems well-branched, usually tall and stout; leaves relatively short and broad ...............................3a. var. peregrina
1. Flower color in a given population generally varying little; stems not manifestly tall, stout or highly branched .............2
2(1) Corolla blood-red, with a darker eye ring or star, the color persisting as maroon in dried specimen .............................
................................................................................................................................................3b. subsp. wilcoxiana
2. Corolla purple or purple-pink, fading (usually to bluish) on drying ...............................................................................3
3(2) Eye marked with a dark star or ring ......................................................................................3c. subsp. drummondii
3. Eye pale and marked with a slender-armed purple star ...........................................................3d. subsp. mcallisteri
3a. var. peregrina Shinners. Plants tall, stout-stemmed, and well-branched; larger leaves ca. 5 times longer than wide. Corolla tube ca. 15 mm long; corolla color quite variable in any population--red white, pink, purple, pale yellow, etc., often marked with a variously colored eye ring or star--and sometimes monstrosities (e.g., flowers with extra petals) present; quite striking in large groups. A cultivated strain which is supposed to be a hybrid between subsp. drummondii and subsp. wilcoxiana. Planted in gardens and along roadsides and now escaped and naturalized in many places in TX and in warm temperate parts of the world. Often growing in the same area and under the same conditions as subsp. drummondii. [Included by Kartesz (1998) under subsp. drummondii.]
3b. subsp. wilcoxiana (Bogusch) Wherry Goldsmith Phlox. Plants not markedly tall, stout, or well-branched; major leaves 5 to 10 (ave.= 7) times longer than wide. Sepals 8 to 11 mm long; corolla tube 15 to 17 mm long, pilose, limb intense blood red with a darker eye ring or star, the color preserved in dried specimens as maroon. Native to an area bounded by Milam, Jackson, Goliad, and Comal Cos. [P. drummondii Hook. var. wilcoxiana (Bogusch) Whiteh. Some sources (e.g. Correll & Johnston 1970) list P. goldsmithii Whiteh. as a synonym.]
3c. subsp. drummondii Drummond Phlox Plants not obviously stout, tall, or well-branched; major leaves 5 to 10 (ave. = 7) times longer than wide. Sepals 7 to 10 (ave. = 8.5) mm long; corolla tube 12 to 16 mm long, pilose, limb purple-pink, with a dark eye ring or star, drying bluish. S. Cen. TX, S. of a Brazos-Guadalupe Co. line, S. to San Patricio, Jackson, Austin, and Harris Cos. Common in our area. [Some sources (e.g. Kartesz 1998), cite P. goldsmithii Whiteh. as a synonym.]
3d. subsp. mcallisteri (Whiteh.) Wherry Plants not markedly tall, stout, or bushy; herbage not or only sparsely glandular; major leaves 4 to 5 times longer than wide. Pedicels to 15 mm long. Sepals 8 to 11 mm long; corolla tube 13 to 15 mm long, limb light purple to light purple-pink, with a pale eye bearing a narrow-rayed purple star, fading on drying. Primarily in NE. TX, W. to Wichita Co. and S. to Leon Co. [P. drummondii Hook. var. mcallisteri (Whiteh.) Shinners].
Annuals, biennials, perennials (some monocarpic), or subshrubs. Basal leaves usually in a rosette, stem leaves alternate, reduced upwards, entire or pinnately (as in ours) or palmately lobed or dissected, the ultimate segments linear to slenderly oblong, cuspidate to bristle-like, often firm and sharp. Flowers terminal or axillary, solitary or in cymose groups arranged in a dense to loose, corymbose, capitate, thyrsoid, or paniculate inflorescence, each flower usually subtended by a bract, bracts commonly pinnatifid. Sepals equal to subequal, partially united by scarious membranes, acute to acuminate, often sharp-pointed. Corolla regular or slightly irregular, usually essentially salverform, the 5 lobes shorter than the tube, red (as in ours) to pink, lavender, or white, sometimes spotted. Stamens included or exserted, attached unevenly in the corolla tube or in the sinuses between the lobes. Style exserted or included, stigma with 3 lobes; ovules 2 to many per locule. Capsule ovoid to oblong. Seeds usually elongate and angled, rarely ovoid, waxy or weakly viscid or mucilaginous when wet.
24 species of W. N. Amer. and FL; 1 in S. Amer.; 9 in TX; 1 here.
Some are cultivated for ornament.
1. I. rubra (L.) Wherry Standing Cypress, Texas Plume. Biennial from a branched root or taproot; stem single, erect, 1 to 2 m tall, sparsely pubescent, eglandular. Basal leaves in a definite rosette (but often withered by flowering time), cauline leaves finely pinnately divided, 4 to 8 cm long, with 10 to 15 linear-filiform segments 5 to 20 mm long and cuspidate, midrib sparsely pubescent. Flowers held horizontally, in small clusters in an elongate thyrse. Sepals 8 to 9 mm long, united for ca. 3 to 4 mm, the scarious membranes glabrous, the lobes broadly subulate, attenuate, with an awn-like tip, sparsely pubescent; corolla bright scarlet red, the tube stout, only slightly flared, 2 to 2. 5 cm long, lobes ca. 9 to 11 mm long, 4 to 6 mm broad, ovate to elliptic, obtuse to nearly acute, often with white or pale pink spots; stamens exserted, unequal, the lower 3 about 1 to 3 mm longer than the upper 2; style exserted, ovary ovoid. Capsule oblong, 8 to 10 mm long; seeds 10 to 12 per locule, the seed coat swelling when wet but little if at all viscid. Sun or partial shade in dry rocky or sandy soils. Cen. and E. TX; TX to FL, N. to NC and OK. About Apr.-July. [Gilia rubra (L.) Heller; G. coronopifolia Pers.].
This is an extremely striking plant in flower. It is quite worthy of cultivation and is especially suited for borders.
Annual, biennial, or perennial herbs, sometimes suffrutescent. Leaves basal and/or alternate, cauline leaves well-developed or greatly reduced, blades pinnately toothed or incised, pinnatifid, or dissected, the segments thin and herbaceous to acerose, apically cuspidate to rather spiny; petioles either long and slender, short and winged, or absent. Flowers solitary, in uneven pairs, or in small, bracted clusters at the ends of the branches and forming a paniculate inflorescence. Calyx with 5 equal or subequal lobes, these acute to cuspidate, herbaceous, connected by scarious membranes. Corolla rotate to nearly salverform, blue-violet (as in ours) to white or lavender, the throat often yellow or paler. Stamens equal to rather unequal, exserted (as in ours) or included. Style exserted (at least in ours); stigma lobes 3. Capsule oblong to ovoid. Seeds few to several per locule, mucilaginous when wet.
About 25 species of the W. hemisphere, most common in W. N. Amer.; 7 in TX; 1 here. Many species formerly in Gilia have been removed to other genera, including Ipomopsis.
The flowers of some, especially some of the annual species, are showy enough that the plants are cultivated for ornament (Mabberley 1987).
1. G. rigidula Benth. Perennial, somewhat suffrutescent at the base, though often not large and sometimes flowering the first year, 8 to 25(40) cm tall; herbage glandular-puberulent. Leaves deeply pinnatifid to pinnately dissected, 6 to 20(25) mm long, with 2 to 7 segments, the segments 2 to 12 mm long, very narrow and acerose (spine-like) to flat and linear to oblong and cuspidate. Flowers solitary or in loose clusters; peduncles to ca. 25 mm long and the bracts linear to trifid. Calyx ca. 5 to 9 mm long, cylindric to ovoid, the tube 2 to 4(5) mm long at anthesis, sepals united to about or below the middle, the uniting scarious membranes present also as thin margins on the lobes, lobes obscurely awned; corolla rotate, intense blue-violet to purple, throat often yellow, ca. 8 to 25 mm across, open in the sunlight; stamens ca. 5 to 7 mm long, exserted, anthers commonly sagittate; stigmas usually exceeding the anthers. Capsule 3 to 5 mm long; seeds many, small. Dry rocky and sandy soils. Primarily in the W. 1/2 of TX; CO, NE, and KS, S. to OK, TX, NM, AZ and Mex. Spring, sometimes again in the fall. Included here on the basis of one collection from Brazos Co. in 1981 (TAMU 11745). (Over)
There is rather a lot of variation in stature, pubescence, flower size and width of leaf segments. Some sources recognize subspecies (e.g. Correll & Johnston 1970; Kartesz 1998), while others do not (e.g. GPFA 1986). If subspecific taxa are recognized, our plant is probably subsp. rigidula--Plants to 25 cm tall; lower leaves to 2.5 cm long, unevenly dissected into a few, flat, oblong, often dentate divisions; upper leaves with linear or filiform divisions, only slightly acerose; sepals 5 to 7 mm long, united about halfway; corolla 8 to 15 mm across; stamens 5 to 7 mm long. W. 1/2 TX; TX to CO an AZ, S. to N. Mex. Spring, sometimes again in the fall. [If subspecies are not recognized, the species includes subsp. rigidula, subsp. acerosa (Gray) Wherry, and subsp. insignis Brand and synonyms include G. acerosa A. Gray, G. insignis (Brand) Cory & Parks, and Giliastrum acerosum (A. Gray) Rydb.].
Ours annual, biennial, or perennial herbs. Herbage glabrous to pubescent and/or glandular. Leaves alternate or opposite, simple and entire to pinnately lobed or compound, margins often toothed; stipules none. Flowers perfect, regular, in ours 5-merous, sessile or pedicellate, rarely solitary, usually in simple or compound, often scorpioid cymes. Calyx lobes united only at the very base, herbaceous, sometimes accrescent, sometimes appendages present between the lobes. Corolla sympetalous, usually campanulate, sometimes rotate to funnelform (or tubular), in ours usually white or some shade of blue or purple. Stamens 5, equal or unequal, attached at the base of the short corolla tube, alternate with the corolla lobes, included or exserted, commonly with a small scale or gland on either side of the base of each filament. Ovary superior (rarely inferior, and not in ours), unilocular or sometimes appearing partially or wholly bilocular because of intrusion of the parietal placentae; style usually 1, included or exserted, with 2 stigma lobes or cleft nearly to the base, in some taxa styles 2 and separate. Capsule loculicidally to septicidally dehiscent by 2 or 4 valves. Seeds 1 to 20 per cell, smooth to corrugated, pitted, or alveolate.
20 genera and 275 species nearly worldwide (except Australia), especially common in W. N. Amer.; 6 genera and 33 species in TX; 4 genera and 10 species here.
Some taxa are cultivated ornamentals (Mabberley 1987).
1. Plants of streams, ditches, and other wet places; axillary spines present; styles fully distinct; capsule fully bilocular ...............................................................................1. Hydrolea
1. Plants of dry to moist soils; spines none; style 1, lobed to deeply cleft (rarely styles 2); capsule 1-celled or incompletely 2-celled ...............................................................................2
2(1) Leaves entire; placentae intruding so placentation appears axile .............................2. Nama
2. Leaves toothed, lobed, or divided; placentae parietal ............................................................3
3(2) Flowers solitary or 2 to 3 in small, axillary or terminal cymose clusters; sepals alternate with smaller appendages ...................................................................................3. Nemophila
3. Flowers in dense or open, terminal scorpioid cymes; appendages none ...........4. Phacelia
Annual or perennial herbs, usually of marshes, ditches, streambeds, or other wet areas, usually with axillary spines. Leaves entire, alternate or sometimes opposite below and alternate above. Flowers in axillary or terminal cymes or corymbs. Calyx unappendaged, persistent but not conspicuously accrescent. Corolla rotate-campanulate, as long as or longer than the calyx, usually blue, rarely white. Stamens included or exserted, without scales at the base, but the bases dilated. Styles 2(3), completely separate. Capsule globose or ovoid, commonly irregularly dehiscent, with many very small striate or rugose seeds.
20 species, primarily of the tropics; 3 in TX; 1 here.
1. H. ovata Nutt. ex Choisy Hairy Hydrolea. Perennial from a rhizome; stem usually simple below, branched above, pubescent or hirtellous, or sometimes the lower portion becoming glabrate with age. Leaves alternate, very short-petiolate, the blades ovate, (1.5)3 to 6 cm long, (1)1.5 to 2.5 cm broad, acute, entire, minutely pubescent; axillary spines slender, straight, to 1.5 cm long. Flowers showy, ca. 1.5 to 2.5 cm across, in terminal cymes with leafy bracts, aggregated into a somewhat paniculate inflorescence; pedicels slender, pubescent, about equalling the calyx or slightly longer. Calyx shorter than the corolla, hirsute and with longer, slender glandular hairs, the lobes lanceolate, about equal, acuminate; corolla intense blue, deeply divided, the lobes 10 to 14 mm long; stamens exserted, commonly tinged with blue, pollen white; ovary and lower portion of styles glandular-pubescent, upper part of styles glabrous, styles exserted. Capsule globose-ovoid, glandular, short-beaked, enfolded by the calyx. Margins of ponds, streams, stock tanks, ditches, etc. E. and SE. TX; GA to LA and TX, N. to MO and AR. June-Oct. [Nama ovatum (Nutt. ex Choisy) Britt.].
TX material annual or perennial herbs, commonly low and well-branched, pubescent. Leaves alternate, simple, entire to sinuate, not much reduced above. Flowers subsessile, solitary in the axils or in small, terminal, non-scorpioid cymes. Calyx divided to near the base, without appendages, not much accrescent in age. Corolla purple or violet to white, funnelform to campanulate or rarely tubular, usually longer than the calyx. Stamens included, unequal, unequally inserted in the corolla tube, base of each filament dilated or with minute appendages. Ovary ovoid to globose, unilocular, but the placentae commonly intruded and placentation appearing axile; style 1, shallowly to deeply bifid. Capsule with many minute, usually reticulate or sometimes shallowly pitted seeds.
45 species of the SW U.S. and tropical America, with 1 in Hawaii; 12 in TX; 2 here.
1. Leaf blades spatulate to obovate, flat, decurrent on the stem and forming wings ..................
.......................................................................................................................1. N. jamaicense
1. Leaf blades linear-oblong to obovate, flat to strongly revolute, not decurrent on the stem ....
...........................................................................................................................2. N. hispidum
1. N. jamaicense L. Fiddleleaf Nama. Annual herb; stems prostrate to ascending, to 25 cm tall, branched from the base, branches to ca. 50 cm long; herbage strigose-hirsute or hispidulous. Leaves alternate, spatulate to obovate, 8 to 80 mm long, 3 to 8(20) mm broad, flat, decurrent on the stem forming conspicuous wings. Flowers solitary or paired in the upper leaf axils. Calyx deeply divided, lobes linear or linear-spatulate, accrescent and adherent to the ovary in fruit, in age 4 to 8 mm long; corolla white to lavender, more or less tubular, 6 to 7 mm long, equalling or longer than the corolla; stamens included, bases dilated into small scales; styles 2, distinct, slender. Capsule cylindric, 6 to 8 mm long, loculicidally dehiscent, with many tan, alveolate or reticulate, subglobose seeds ca. 0.2 mm in diameter. Moist lawns and under shrubs. E. and SE. TX; E. to FL; S. to Cen. Amer. and W.I. Known in our area from Brazos and Washington Co. collections from the 1940's; no recent collections seen by the author; perhaps overlooked, very rare, or no longer present in our area. Apr.-July. [Marilaunidium jamaicense (L.) Kuntze].
2. N. hispidum Gray Rough Nama, Sand Bells. Annual; stem erect to ascending, (7)10 to 50 cm tall, branched from the base or above; herbage strigose-hispid or hirsute. Leaves mostly alternate, flat to strongly revolute, linear-oblong to obovate, 1 to 7 cm long, 1 to 8 m broad, tapered to the base, not decurrent on the stem. Flowers solitary or in small clusters. Calyx lobes linear to linear-lanceolate, 3 to 7 mm long, subequal, hirsute or hispid; corolla funnelform to campanulate, pink to purple or violet, 8 to 15 mm long, the lobes rounded to obtuse; filament bases only slightly dilated, the adnate portion shorter than the free portion, anthers yellow. Seeds yellow, alveolate. Gravelly sand and sandy soils of various habitats--outcrops, roadsides, vacant lots, etc. Nearly throughout TX, W. to CA, N. to OK, and S. to Mex. Mar.-July. [Marilaunidium angustifolium (A. Gray) O. Ktze.].
This is a highly variable species as to habit, leaf shape, and pubescence. Varieties which were formerly recognized are now known to intergrade; nor can they be separated geographically (GPFA 1986).
Annual herbs. Leaves alternate, oblong to orbicular, pinnately lobed or divided. Flowers solitary or several in loose, non-scorpioid cymes. Calyx deeply divided, the 5 lobes alternating with 5 smaller auriculate appendages, often accrescent in fruit. Corolla either large and blue to purplish or small and white. Stamens included, the base of each filament flanked by a pair of scales. Style divided to about the middle. Capsule globose, with 1 to 4 seeds. Seeds subglobose-ovoid, pitted, with a small elaiosome.
11 species, 9 of W. and 2 of SE. N. Amer.; 2 in TX, both here.
Several species are cultivated for ornament, including N. maculata, Five-spot, which has a white corolla tipped with blue.
1. Corolla blue or purplish, 1 to 3 cm broad; calyx in fruit equaling or longer than the sepals;
terminal flowers often in cymes................................................................. 1. N. phacelioides
1. Corolla white, to 5 mm broad; calyx in fruit shorter than the sepals; all flowers solitary.........
............ ..................................................................................................................2. N. aphylla
1. N. phacelioides Nutt. Large-flower Nemophila. Stems simple to branched, erect or ascending, to 40 cm tall or long, hirsute to glabrescent. Lower leaves oblong in outline, 6 to 8 cm long, 2 to 5 cm broad, pinnatifid to pinnately divided (sometimes almost compound), the divisions 9 to 11, entire to toothed, with petioles shorter than or equalling the blades, upper leaves ovate to orbicular in outline, all leaves hirsute to hispid. Flowers solitary in the axils or in small terminal cymes, pedicellate. Calyx lobes oblong to ovate-lanceolate, 6 to 9 mm long, 2 to 3 mm broad, each sinus with a smaller, spreading-erect auriculate appendage, all hispid to hirsute, calyx accrescent in fruit; corolla rotate, 10 to 30 mm broad, lobes about twice as long as the tube, rounded to emarginate, lobes and throat blue to purple, the tube forming a paler eye; stamens included, about equalling the tube, equally inserted near the base of the tube, base of each filament with a pair of partly free, fimbriate appendages; ovary ovoid to globose, unilocular, style divided to about the middle. Capsule globose, ca. 5 to 9 mm broad, hispid; seeds usually 4, globose, reddish-brown, pitted. Sandy soils of open woods, hillsides, roadsides, etc., sometimes where somewhat shady or slightly moist. E. and SE. TX; E. OK and W. AR to SE. TX. Mar.-May.
2. N. aphylla (L.) Brumm. Smallflower Nemophila. Stems weak, sparingly hispid. Leaves long-petioled, ovate to orbicular in outline, 1 to 3.5 cm long and 1 to 2.5 cm broad, pinnately lobed, the 3 to 5 segments ovate to obovate and entire or themselves ternately lobed, mucronate, pubescent. Flowers all solitary, opposite or beside the leaves on slender pedicels. Calyx lobes lanceolate, 1 to 3 mm long, pilose to hispid; corolla white, to ca. 3(5) mm long, campanulate; stamens included. Capsule globose, 3 to 5 mm broad, at maturity much longer than the weakly accrescent calyx, pubescent; seeds 1 or 2, light brown, ellipsoid or with one flat surface. Uncommon and easily overlooked in wet woods. E. TX; DE to FL, W. to KY, AR, and TX; possibly MO and OK as well. Mar.-May. [N. microcalyx (Nutt.) Fisch. & Mey.].
Taprooted annual, biennial, or perennial herbs. Stems erect to decumbent, simple or well-branched. Herbage glabrous to pubescent or glandular-pubescent. Leaves primarily alternate, sometimes also basal, petiolate to sessile, entire to pinnately lobed, pinnatifid, or pinnately dissected. Inflorescences terminal or subterminal single to compound scorpioid cymes, the flowers pedicellate to nearly sessile. Calyx lobed to near the base, appendages none, lobes linear to lanceolate, accrescent in fruit or not. Corolla blue to purple, lavender, or white, rotate to campanulate (as in ours) or tubular, usually longer than the calyx. Stamens equal or nearly so, included or exserted, attached near the base of the corolla tube, bases pubescent and flanked by either a pair or scales or a gland bordered by parallel flaps or filament bases merely dilated, anthers turning inside out at maturity. Ovary globose-ovoid, unilocular, style 1, included or exserted, shallowly bifid to divided for 1/3 to 3/4 its length. Capsule globose-ovoid. Seeds 2 to many, oblong to globose, rounded or flattened, pitted or cross-wrinkled, sometimes ridged.
150 species, mostly of W. N. Amer., some in the E. U.S. and S. Amer.; 15 in TX; 5 here.
The genus includes many taxa cultivated for their flowers (Mabberley 1987).
1. Plants robust, viscid-pubescent; inflorescence a dense, compound scorpioid cyme; filaments with a pair of scales at the base but no glands; seeds 1 to 4 per capsule, each with a ridge separating 2 hollows .....................................................................1. P. congesta
1. Plants rather delicate, not viscid pubescent; inflorescence a dense to lax simple scorpioid cyme; filament bases flanked by a gland bordered by parallel flaps but no scales; seeds usually 6 or more per capsule, without a pronounced ridge ...................................................2
2(1) Branches, pedicels, and apex of ovary glabrous or with a very few short, stiff, spreading hairs ........................................................................................................................2. P. glabra
2. Branches, pedicels, and apex of ovary definitely pubescent ..................................................3
3(2) Basal leaves in a rosette, only shallowly toothed or lobed; pedicels in age shorter than the calyces, usually stiffly erect ............................................................................3. P. strictiflora
3. Basal laves not or only weakly rosulate, pinnatifid to pinnately divided; pedicels equalling or longer than the calyces, spreading to ascending or reflexed .............................................4
4(3) Stem leaves dentate or shallowly lobed; seeds usually 10 to 15 per fruit ...4. P. patuliflora
4. Stem leaves deeply lobed to pinnatifid; seeds usually 6 to 8 per fruit ...............5. P. hirsuta
1. P. congesta Hook. Spike Phacelia, Blue Curls. Annual (or biennial); stem erect, simple or well-branched, 1 to 10 dm tall; herbage glandular and variously pubescent to hispid. Leaves oblong to ovate in outline (1)3 to 9(12) long, 1 to 4 cm broad, once or twice pinnatifid or pinnately compound, the terminal 3 lobes often not completely separated and longer than the lower segments, margins rather bluntly toothed, surfaces glandular and/or strigose. Flowers in dense, terminal compound scorpioid cymes, densely pubescent; pedicels short, 3 mm long or less. Calyx lobes more or less linear to narrowly oblanceolate, 3 to 5 mm long, ca. 1 mm broad, glabrous to hispid or setose; corolla campanulate, blue to purple, 4 to 7 mm long, 3 to 6 mm across, lobes rounded, commonly pubescent; stamens exserted beyond the corolla by 2 to 4 mm, base of filaments with paired scales but no glands; ovary subglobose to ovoid, puberulent to glandular, style 7 to 8 mm long, exserted, parted about 3/4 its length. Capsule ovoid, ca. 2 to 3 mm long, with 2 or 4 brown seeds, each with a ventral ridge with a furrow on either side. Sand, gravel, and rocky soils throughout much of the state except the Pineywoods and Post Oak Savannah; not too common in our area--more common to the W.; AZ and NM to OK and TX; also N. Mex. Mar.-June. [Includes var. dissecta Gray; the former var. rupestris Macbr., with white flowers, is now treated as P. rupestris Greene (Hatch, et al. 1990)].
2. P. glabra Nutt. Smooth Phacelia. Rather delicate annual, slightly succulent; stems 5 to 40 cm tall, simple and erect in small individuals or commonly branched near the base and the branches erect or ascending; stems and inflorescence glabrous or with only a very few stiff, spreading hairs. Basal leaves petiolate, oblong to oblong-oval, 1.5 to 4 cm long, 0.5 to 1.5 cm broad, pinnatifid to pinnate, with 2 to 4 pairs of lobes or leaflets and a larger terminal 3- or 5-lobed lobe or leaflet, cauline leaves mostly sessile and clasping, oval to broadly ovate in outline, deeply lobed with 1 to 5 pairs of acute to obtuse lobes, margins usually hirsute-ciliate and sometimes sparsely strigose on either or both surfaces. Inflorescences simple axillary or terminal scorpioid cymes with 5 to 15 flowers; pedicels glabrous (or with a few spreading hairs), at anthesis spreading or spreading ascending, 6 to 12 mm long. Calyx lobes usually slightly unequal, narrowly oblong to oval, 2 to 4 mm long, 0.5 to 2 mm broad, margins hirsute-ciliate, surfaces usually glabrous, in fruit erect or ascending; corolla blue-lavender with a white eye and 2 small purplish dots at the base of each of the rounded lobes, 5 to 12 mm across, the lobes sparsely pilose externally; stamens about equalling the corolla, 3 to 5 mm long, filaments densely pubescent in the lower 1/2, basal gland with flaps wholly adnate to the corolla; style included, 3 to 5 mm long at maturity, divided about halfway, apex of ovary glabrous or with a very few hairs; ovules usually 4 per placenta. Capsule globose, 3 to 4 mm in diameter; seeds 4 to 8 per fruit, angled-ovoid, ca. 2 mm long, brown, finely alveolate and areolate. Wood edges, vacant lots, prairies, roadsides, etc. NE part of TX; to AR, OK, and LA; common in our area. Spring. [Cosmanthus nemophiloides Kunth].
The small flowers are not individually showy, but a solid patch of this plant is quite lovely.
3. P. strictiflora (Engelm. & Gray) Gray Annual; stem single and erect or branched and the branches erect to decumbent, 5 to 30 cm tall; herbage hirsute, sometimes finely so, and the inflorescence region canescent to hirsute. Basal leaves in a definite rosette, petiolate, usually basally cuneate, narrowly oblong to oblanceolate or oval in outline, 1 to 6 cm long, 0.5 to 3 cm broad, toothed to shallowly or deeply pinnately lobed, with 1 to 6 pairs of lobes, margins and upper surface sparsely hirsutulous, lower surface glabrate, or else both surfaces densely strigose, stem leaves smaller, sessile, orbicular to linear-oblong in outline, varying from merely dentate to deeply pinnately lobed with 2 to 6 pairs of teeth or lobes. Inflorescences simple terminal cymes with 3 to 20 flowers; pedicels at and after anthesis strongly ascending (sometimes slightly spreading), 2 to 10 mm long, usually equalling or shorter than the calyx. Calyx lobes linear to oblanceolate, 5 to 15 mm long, 1 to 4 mm broad, pubescent, calyces of lower flowers of the cyme sometimes strongly accrescent; corolla rotate-campanulate, purple to lavender, 8 to 20 m across, 7 to 10 mm long, the lobes obovate and minutely crenulate, externally pilose; stamens included, 5 to 7 mm long, filaments pubescent in the lower 2/3, flaps of basal glands wholly adnate to the corolla; style included, ultimately 5 to 12 mm long, ovary globose, apically densely hirsute. Capsule globose-ovoid, 3 to 6 mm broad, with 8 to 14 ovules per placenta; seeds 10 to 20, black, ca. 2 mm long, finely alveolate and areolate. Sandy soils of woods, wood edges, and fields. E. TX; OK to TX, MS, and AL. Mar.-May.
This is a highly variable species. Constance (1949) described 4 varieties, which are still recognized by Hatch, et al. (1990) and Kartesz (1998). Some sources, however, (e.g., GPFA 1986) contend that the 4 varieties are sympatric and intergrade too much to be recognized. If varieties are recognized, our plants are probably assignable to the following three:
var. strictiflora Prairie Phacelia. Stem spreading-hirsute, inflorescence loosely hirsute; foliage dull green, not succulent; basal rosette usually withering early, the leaves pubescent beneath, lobed or divided; cauline leaves lobed to almost pinnatifid; calyces all alike in fruit. Cen.-E. TX.
var. connexa Const. Similar to var. strictiflora, but the stems finely strigose, inflorescence canescent; calyx lobes linear to lance-linear. NE. TX and adjacent OK.; especially common in fallow fields.
var. lundelliana Const. Foliage bright green, slightly succulent; basal rosette persistent, the leaves glabrous beneath and shallowly toothed; cauline leaves broad and shallowly toothed; calyces of lower flowers of cymes markedly accrescent. W. OK and N. Cen. TX, S. to about Waco.
4. P. patuliflora (Engelm. & Gray) Gray Annual; stem usually branched from the base, branches spreading to stiffly ascending, 8 to 60 cm tall; herbage densely spreading-hirsute to finely strigulose, the inflorescence region loosely hirsute to strigulose and canescent, rarely somewhat glandular. Basal leaves usually not rosulate, not persistent, blades basally truncate to cuneate, oblong to oval (occasionally orbicular) in outline, 2 to 10 cm long, 1 to 4 cm wide, pinnately lobed to pinnatifid or compound, with 1 to 7 pairs of lobes or leaflets and a larger terminal leaflet, all divisions dentate or lobed, the terminal leaflet often trilobed, both surfaces strigulose to hirsute, cauline leaves primarily sessile, oblong to orbicular, dentate or shallowly lobed with 3 to 6 pairs of teeth or lobes. Inflorescences simple scorpioid cymes with 5 to 30 flowers; pedicels at and after anthesis wide spreading to weakly ascending or reflexed, 3 to 20 mm long, in fruit longer than the calyx and spreading to ascending. Calyx lobes narrowly oblong to lanceolate or obovate, 5 to 12 mm long, 1 to 4 mm wide, equal or nearly so, surfaces strigulose to glabrate and margins hirsute; corolla lavender to purple-violet, usually with a white eye, rotate to campanulate or broadly campanulate, 8 to 20 mm across, lobes obovoid and minutely crenate, sparsely pilose externally; stamens included, 5 to 8 mm long, filaments pubescent in the lower 2/3, flaps of basal glands wholly adnate to the corolla; style included, ultimately 5 to 8 mm long, divided 2/3 its length, basally hirsute, ovary apically hirsute. Capsule 4 to 6 mm broad, ovules 6 to 12 per placenta; seeds 10 to 15 per capsule, angled-ovoid, ca. 2 mm long, finely alveolate and areolate. Mostly in sandy alluvial soils of woods, river terraces, and plains. Cen. and SE. TX; also N. Mex. Mar.-May.
Three varieties are recognized by Moyer and Turner (1994). Our plants are probably all var. patuliflora, Sand Phacelia. Branches mostly decumbent rather than ascending; calyx lobes obtuse; corolla pale lavender to violet; fruiting pedicels spreading to reflexed. Cen. and SE. TX and Rio Grande Valley and Plains and adj. Mex.
5. P. hirsuta Nutt. Hairy Phacelia. Annual; stem simple or more commonly branched at the base or above, 10 to 50 cm tall, branches erect to ascending, stems densely hirsute, the hairs stiff and spreading, inflorescence region hirsutulous and spreading-hirsute. Basal leaves scarcely rosulate, petiolate, oblong in outline, 2 to 4.5 cm long, 0.8 to 2.5 cm broad, pinnatifid to pinnate, with 2 to 4 pairs of lobes or leaflets and a larger trilobed terminal leaflet, cauline leaves usually short-petiolate, sometimes sessile and somewhat clasping, oblong to orbicular, pinnately lobed to pinnatifid (rarely only toothed), with 2 to 4 pairs of lobes, both surfaces strigose. Inflorescences terminal, simple scorpioid cymes with 10 to 25 flowers; pedicels at and after anthesis spreading-ascending to spreading, 3 to 15 mm long. Calyx lobes linear to oblanceolate, 5 to 10 mm long, 1 to 3 mm broad, unequal, strigose an strigulose inside and out, spreading to reflexed at anthesis, in fruit erect to ascending; corolla rotate to campanulate, 8 to 13 mm across, light blue to lavender, with a white eye and 2 purple dots at the base of each orbicular, entire lobe, externally pilose; stamens usually included, 4 to 6 mm long, filaments pubescent in the lower 2/3, flaps of basal glands wholly adnate to the corolla; style included, ultimately 5 to 6 mm long, divided 1/3 to 1/2 its length, pubescent in the lower portion, apex of ovary pubescent. Capsule subglobose, 3.5 to 4 mm broad, ovules usually 4 per placenta; seeds 6 to 8 per fruit, angled-ovoid, ca. 2 mm long, brown, finely alveolate and areolate. Sandy soils of openings in deciduous woods and along wood edges. NE. TX; S. MO and SE. KS through AR and OK, S. to LA and TX. Mar.-May. [P. parviflora var. hirsuta A. Gray; P. dubia (L.) Trel. var. hirsuta Trel.].
Ours annual, biennial, or perennial herbs (elsewhere also shrubs and trees). Stems and leaves often bristly-pubescent or hairy, rarely glabrous. Leaves alternate or basal and alternate, simple, usually petiolate, estipulate. Flowers perfect, usually regular, in ours 5-merous, solitary or more usually in solitary or grouped cymose inflorescences, these commonly one-sided and coiled (scorpioid), with or without bracts between, beside, or opposite the flowers, the inflorescence uncoiling and elongating with age and sometimes appearing racemose in fruit. Sepals 5, free or fused, equal or unequal, usually pubescent, occasionally glabrous, persistent. Corolla sympetalous, primarily salverform or funnelform, often with folds or appendages (fornices) in the upper throat opposite the lobes, lobes rounded to pointed, small appendages present inside the base of the tube. Stamens 5, inserted on the corolla tube alternate the lobes, commonly included. Nectary disk sometimes present subtending the ovary. Gynoecium superior, in ours with 2 united carpels, usually 4-ovulate and developing into 4 single-seeded lobes or 1 carpel suppressed and the remaining carpel developing into 2 single-seeded lobes or sometimes (as in Heliotropium) the gynoecium essentially unlobed; style 1, arising from between the lobes (gynobasic) or sometimes terminal (as in Heliotropium); in taxa outside our area, other gynoecium variations occur; a low to pyramidal or columnar receptacle (gynobase) often prominent, particularly in fruit. Fruits in our species mericarp-like nutlets, usually separating from each other and falling individually, their surfaces varying in pubescence and ornament. Seeds with scanty or no endosperm.
About 154 genera and 2,500 species worldwide; 18 genera and 73 species in TX; 8 genera and 16 species here. A useful reference for descriptions and family distributions is the work of I. M. Johnston (1964).
In the Boraginaceae, subfamily and generic classifications are based on features of the mature gynoecium. Confident identification to species is also greatly facilitated by the presence of mature fruit.
The family has a number of important ornamental genera, including Myosotis (Forget-me-not), Heliotropium (Heliotrope), Echium, Anchusa, and so on. Some of the woody taxa (e.g. Cordia) provide timber. Some taxa are dye plants (e.g. Alkanna) and some are potherbs or have medicinal value (e.g. Symphytum--Comfrey and Borago--Borage) (Mabberley 1987).
1. Ovary deeply 2- or 4-lobed; style gynobasic (arising from between the lobes) .....................2
2(1) Calyx with at least a few uncinate (hooked) hairs, strongly irregular; nutlets keeled all the way around, otherwise smooth .............................................................................2. Myosotis
2. Calyx without uncinate hairs, regular or irregular; nutlets keeled or grooved on the ventral surface and/or rough, pitted or hairy ........................................................................................3
3(2) Nutlets pubescent, attached apically-laterally, spreading at maturity .........3. Cynoglossum
3. Nutlets glabrous, attached basally or laterally, remaining erect and parallel at maturity ......4
4(3) Nutlets attached to the gynobase basally and bearing a small basal scar or attached laterally and with a longitudinal groove or triangular open area; gynobase raised, more or less pyramidal ...........................................................................................................................5
4. Nutlets attached basally, bearing a large basal scar; gynobase low ......................................6
5(4) Nutlets with a ventral groove or triangular open area ......................................4. Cryptantha
5. Nutlets with a small, raised basal attachment scar ............................................5. Amsinckia
6(4) Undersurface of leaves with midvein and lateral veins conspicuous ..........6. Onosmodium
6. Undersurface of leaves with no veins or only midvein conspicuous .......................................7
7(6) Nutlets white, shiny, pitted ...........................................................................7. Lithospermum
7. Nutlets brownish, roughened ........................................................................8. Buglossoides
1. Corolla bright yellow or orange ................................................................................................2
1. Corolla white, blue, purple, or pale yellow ..............................................................................3
2(1) Plants perennial from stout, dark roots; corolla more than 1 cm long ......7. Lithospermum
2. Plants annual from slender taproots; corolla less than 1 cm long ....................5. Amsinckia
3(1) Corolla blue or purple; fruits glabrous ...........................................................1. Heliotropium
3. Corolla white (occasionally tinged with blue or yellow); fruits glabrous to pubescent; if flowers blue then fruits pubescent and stem leaves clasping ................................................4
4(3) Cymes without bracts ................................................................................................................5
4. Cymes with leafy bracts or flowers apparently solitary to few in the axils of the upper leaves ........................................................................................................................................7
5(4) Corolla throat without appendages; style terminal ........................................1. Heliotropium
5. Corolla throat with appendages; style gynobasic ....................................................................6
6(5) Corolla more than 3 mm long; plants more than 25 cm tall .......................3. Cynoglossum
6. Corolla less than 3 mm long; plants less than 25 cm tall ................................4. Cryptantha
7(4) Calyx with at least a few uncinate (hooked) hairs, strongly irregular ..................2. Myosotis
7. Calyx without uncinate hairs, regular or irregular .....................................................................8
8(7) Style terminal; flowers apparently solitary to clustered in the axils of the upper leaves ..........
.........................................................................................................................1. Heliotropium
8. Style gynobasic; flowers in cymes or solitary to few in the axils .............................................9
9(8) Style long-exserted; leaves with midvein and lateral veins obvious below ..6. Onosmodium
9. Style included; leaves with no veins or only midvein obvious below ...........8. Buglossoides
Ours annual or perennial herbs, sometimes somewhat suffrutescent, usually branched, erect to prostrate. Leaves all cauline, sessile to petiolate, variously shaped, linear to deltoid or ovate, glabrous or more commonly pubescent. Flowers usually in definite scorpioid cymes (except H. tenellum with flowers apparently solitary or few in the upper axils), with or without bracts, cymes becoming elongate and spike-like with age. Calyx deeply lobed or the sepals essentially free, equal or unequal, glabrous to variously pubescent. Corolla funnelform to salverform, blue to white, yellow, or purple, without appendages in the throat, but commonly pubescent within. Filaments very short and stamens included. Style 1, terminal on the ovary but sometimes so short that the stigma appears sessile; stigma conic, flattened, bifid, or capitate, mostly sterile and receptive only in a ring around the base. Fruit splitting into 4, 1-seeded nutlets or 2, 2-seeded nutlets.
About 250 species of temperate to tropical regions; 15 species in TX; 7 to be expected here.
The genus includes several popular ornamentals, including H. amplexicaule and H. arborescens, the latter cherry-scented and also used in perfumes. Other species are used in herbal remedies in their native regions (Mabberley 1987).
1. Plants glabrous, rather succulent, usually glaucous ..............................1. H. curassavicum
var. curassavicum
1. Plants hispid to strigose or villous ............................................................................................2
2(1) Mature fruit breaking into 2 2-seeded nutlets; stems and younger parts of the plant with glandular hairs ...........................................................................................2. H. amplexicaule
2. Mature fruit breaking into 4 1-seeded nutlets; plants without glandular hairs .......................3
3(2) Cymes without leafy bracts; leaves usually with veins other than the midvein evident .........4
3. Cymes with leafy bracts or the flowers apparently in the leaf axils ........................................5
4(3) Corolla blue or purple; leaves to 15 cm long, ovate to elliptic; petioles 4 to 10 cm long; plants erect .........................................................................................................3. H. indicum
4. Corolla white or white with a yellow throat; leaves narrowly oblanceolate to elliptic, to 4 cm long; petioles to 1 cm long; plants decumbent to erect ............................4. H. procumbens
5(3) Corolla lobes longer than wide, narrowed at the base; leaves linear; style ca. 1 mm long ....
............................................................................................................................5. H. tenellum
5. Corolla lobes wider than long or scarcely developed; leaves lanceolate to ovate, oblong, or elliptic ....................................................................................................................................6
6(5) Corolla 15 to 20 mm across, 5-angled, scarcely lobed; flowers appearing scattered along the stem ...............................................................................................6. H. convolvulaceum
6. Corolla 6 to 17 mm across, star-shaped, with 5 triangular, acute lobes; flowers and fruit definitely racemose ......................................................................................7. H. racemosum
1. H. curassavicum L. var. curassavicum Seaside Heliotrope, Chinese Pulsey, Quailplant, Cola de Mico. In our area usually perennial from a rhizome-like taproot (in colder areas annual); stem prostrate to decumbent, succulent and more or less rubbery, to 4 dm long or tall, ca. 5 mm thick, well-branched, branches ascending; herbage glabrous, often glaucous. Lower leaves opposite to subopposite or alternate, upper leaves alternate, all thickish and succulent, oblanceolate or narrowly oblanceolate, 2 to 4(6) cm long, 4 to 8(20) mm broad, obtuse to rounded or sometimes acute, basally tapered. Cymes solitary or usually clustered at the ends of the branches and on axillary peduncles, bractless, densely-flowered, tightly coiled when young, looser and raceme-like with age, to 10 cm long, the axis usually rather broad and flattened; flowers pedicellate to subsessile. Calyx lobed nearly to the base, lobes (1)2 to 3 mm long at anthesis, shorter than the nutlets when mature, subequal or equal, lanceolate to oblong; corolla usually white (sometimes bluish or with a yellow eye), 1.2 to 3.5 mm long, 4 to 7 mm broad, funnelform to nearly salverform, the tube about equalling the limb, the lobes rounded; style very short to absent, stigma flat and wide, about as wide as the ovary, persistent. Fruit shallowly 4-lobed, 2 to 2.5 mm tall, slightly flattened laterally, held within the accrescent calyx, glabrous and smooth or faintly ribbed, outer layer of fruit wall thickened, buoyant and helping seed dispersal by water. In damp, often saline soils, commonly sands, and around ponds, beaches, and salt flats. Throughout the state; DE to FL, W. to OK, TX, and NM; also W.I., Mex, Cen. Amer., and N. S. Amer.; also along the W. coast of S. Amer; introduced in Europe; another variety found in much of the W. U.S. Flowering throughout the year.
2. H. amplexicaule Vahl Violet Heliotrope. Perennial from a deep, woody rootstock; stems several from the base, 2 to 5 dm tall, ascending or decumbent; herbage more or less densely minutely pubescent and usually also villous or hispid, the younger portions and the inflorescence glandular-pubescent. Leaves oblong to lanceolate, 2 to 9 cm long, to 18 mm broad, tapered to a short petiole or sessile, apically acute to rounded, margins entire or with a few weak, blunt teeth, upper leaves more sessile and more densely tawny-pubescent. Cymes in clusters of 2 to 5 at the tips of the branches, on naked peduncles 1 to 10 cm long, without bracts, dense at first, uncoiling and becoming elongate. Calyx sessile, lobes linear-lanceolate, nearly equal, 2.5 to 5 mm long, shorter than the corolla, usually glandular pubescent, persistent, but spreading after the nutlets fall; corolla blue to purple or rarely white, ca. 4 to 8 mm across, with short, broad lobes, tube ca. 5 mm long. Fruit separating into 2, 2-seeded nutlets or mericarps, these 1.5 to 2.5 mm tall, 2 to 3 mm long, slightly laterally compressed, glabrous, the surface rough and/or rugose-tuberculate. Frequently cultivated and sporadically escaping or adventive in waste places, fields, etc. Native to Urug. and Argen. Apr.-Aug. [Cochranea anchusaefolia (Poiret) Guerke].
This plant is grown for its ornamental flowers. It also has some use in fertility-regulating medicines (Mabberley 1987).
3. H. indicum L. Indian Heliotrope, Turnsole, Alacrancillo. Taprooted annual herb; stems erect, usually simple or branched above, to 1 m tall but often flowering when much smaller, hirsute to hispid-bristly, often hollow. Lower leaves opposite, upper alternate, all with petioles 4 to 10(12) cm long, blades ovate to nearly deltoid, 2 to 6 cm long, acute to obtuse, basally acute or obtuse to cordate, sometimes slightly decurrent on the petiole, thin-textured, rugose, glabrate to sparsely hispid-scabrous, margins undulate to repand. Cymes usually solitary at the stem tips, rarely paired, bractless, at anthesis to 15 cm long and the flowers crowded in 2 rows, elongating with age to ca. 30 cm long, the axis hirsute. Calyx sessile, sepals 1.5 to 2 mm long, shorter than the mature fruit, linear to linear-lanceolate, unequal, hirsute; corolla blue to violet (rarely white), salverform, the tube longer than the calyx, ca. 3 mm long, constricted at the throat, limb 2 to 4.5 mm across, puberulent or strigose externally; style ca. 1 mm long, stigma capitate. Fruit miter-shaped (squarish at the bottom, with rounded angles meeting at a point), separating at first into 2 nutlets but later into 4, each with 1 or 2 ribs dorsally, faintly reticulate, glabrous, ca. 2 mm long. Moist or seasonally moist areas--ditches, river bottoms and banks, ponds, lakes, creeks, etc. E. 1/2 TX, W. to Dallas, Bexar, and Hidalgo Cos.; VA to W. MO and SE. KS, S. to FL and TX; in warmer parts of the Americas, to N. Argen.; also the Old World Tropics, but probably native to this hemisphere. June-Nov. [Tiaridium indicum (L.) Lehm.].
4. H. procumbens Mill. Fourspike Heliotrope. Taprooted annual; stems 1 to several, erect to decumbent or sprawling, with ascending branches, 1 to 5 dm tall or long; herbage moderately to densely strigose or appressed villous-hispid, foliage pale- or gray-green canescent. Leaves many, with petioles to 1.5 cm long; blades varying from oblanceolate to elliptic, to ca. 4 cm long and 2 cm broad, flat or revolute, basally cuneate, apex rounded to obtuse, often only the midvein obvious, sometimes the laterals visible, upper and lower surfaces similar in appearance. Inflorescences usually many, terminal and axillary, solitary or paired, on peduncles 1 to 3 cm long, definitely coiled, bractless, dense, 3 to 10 cm long at anthesis, elongating to as much as 15 cm in fruit. Calyx in flower 1 to 1.2 mm long, divided nearly to the base, the lobes unequal, some subulate and some lance-ovate, all doubling in size in fruit, becoming 2 to 2.5 mm long and more unequal; corolla white, sometimes with a yellow throat, drying yellowish, the tube ca. 1.5 mm long, slightly shorter than the calyx at anthesis, the limb rotate, 2 to 3 mm broad, the lobes triangular and acute. Fruit strigose, smooth, slightly broader than long, 1 to 2 mm broad, splitting into 4, 1-seeded nutlets. Usually in damp areas or places subject to flooding, weedy on shores and the margins of ponds and ditches. Primarily in the Coastal Plain, N. to Jefferson, Hays, Brazos, Bexar, and Brewster Cos.; FL to TX, SE. OK, and AR; also W.I., Cen. Amer., and tropical S. Amer. Apr.-Nov. [Many older specimens bear the name H. inundatum Sw., which is listed as a synonym of H. procumbens Mill. var. procumbens. No varieties were described by Correll and Johnston (1970); it is presumed that what we have is the typical variety].
5. H. tenellum (Nutt.) Torr. Pasture Heliotrope. Taprooted annual; stem erect, 2 to 5 dm tall, usually simple at the base and sparingly branched in the upper 1/2, stem and branches slender, strigose. Leaves sessile, linear, 1 to 3(5) cm long, 1 to 3(5) mm wide, tapered to both ends, whitish-strigose, usually densely so, the margins revolute. Flowers in very loosely-spaced racemose cymes, the bracts so similar to the leaves that the flowers appear solitary or grouped in the axils of the upper leaves and at the branch tips. Calyx lobes linear with revolute margins, very unequal, at least 1 of them ca. 4 mm long, equalling or exceeding the corolla and foliaceous, becoming at least 5 mm long in fruit (sometimes as much as 1 cm long), twice as long as the shortest; corolla funnelform, white with a yellow eye, ca. 6 mm long, the tube shorter than all of the sepals, the limb 3 to 6 mm broad, pleated, with deeply-cut obovate to elliptic lobes, tube glabrous within and strigose externally, the limb strigose externally; style ca. 1 mm long, stigma bifid. Fruit ca. 1.5 mm tall, 3 mm broad, shorter than the calyx, splitting into 4, 1-seeded nutlets, the nutlets rather blunt, sparsely to moderately appressed-pubescent or strigose. On gravelly or rocky soils of outcrops, open areas, woods, or brushlands; in our immediate area commonest in native prairie and on limestone and calcareous sandstone outcrops; E. 2/3 TX; KY to SE. KS, S. to TX, AL, and GA; also N. Mex. May-Oct. [H. nuttallii House; Lithospermum tenellum Nutt.; Lithococca tenella (Nutt.) Small].
6. H. convolvulaceum (Nutt.) Gray Bindweed Heliotrope. Annual from a slender to stout taproot; stems 1 to 4 dm tall, branched at the base and above, branches prostrate to ascending or stems simple below and with elongate, ascending branches above; herbage strigose, younger parts especially densely so and canescent. Leaves with petiole shorter than the blade; blades to 4 cm long and 1.5 cm broad, ovate to lanceolate, apically acute to obtuse, base cuneate to rounded, both sides strigose, the hairs sometimes from mineralized discoid or pustule-like bases, margins entire, flat or revolute. Cymes at the branch tips, very short, not elongating, the flowers appearing to be scattered among the leaves, true bracts few or none. Calyx at anthesis 4 to 6 mm long, enlarging to 6 to 8 mm long at maturity, sepals narrowly lanceolate to subulate-linear, slightly unequal with the 2 largest broader and 0.5 to 1 mm longer than the shortest; corolla open mornings and evenings, salverform to broadly funnelform, white with a yellow throat, (8)15 to 20 mm across, the tube much longer than the calyx, constricted below, 8 to 11 mm long, strigose externally on the ribs; style elongate and slender, glabrous, stigma included, capitate, often with a terminal conical appendage, strigose. Fruit 3 to 4 mm tall, 2 to 2.5 mm broad, separating into 4 nutlets (sometimes apparently 2), united in pairs, the shared sides flat and reniform, the exterior sides rounded and strigose, attachment scar with a keel on either side. Sandy soils and dunes. W. TX, E. along river valleys to Grayson, McLennan, and Brazos Cos.; specimen seen by author is a very old one from along the Brazos River (H.B. Parks, 1948, TAMU 7974); this plant may be a sporadic or impermanent member of our flora; UT, WY, NE, CO, and KS, S.to AZ, TX, and OK; also N. Mex. June-Oct. [Euploca convolvulacea Nutt.]
7. H. racemosum Rose & Standl. Taprooted annual to 40 cm tall; stems simple at first but later with upright to decumbent or sprawling branches, usually ending spreading or decumbent; herbage strigose, the young parts densely so. Leaves with petioles shorter than the blades; blades lanceolate or sometimes oblong to elliptic, 1.5 to 4 cm long, to 11 mm broad acute, basally cuneate, strigose on both surfaces, the hairs with bulbous or discoid bases, margins entire, flat to slightly revolute. Flowers in fairly well-defined racemose, bracted cymes 5 to 10 cm long, usually forming the distal 1/2 of the branches. Calyx 4 to 5 mm long at anthesis, enlarging to 5 to 6 mm long in fruit, strigose, the lobes slightly unequal, linear to linear-lanceolate, acuminate; corolla open at night or on cloudy days, fragrant, white, funnelform, 6 to 17 mm across, the tube 4 to 8 mm long, strigose, limb star-shaped, with 5 triangular, acute lobes, inflexed folds from the sinuses extending down weakly into the throat, ribs pubescent externally; style slender, stigma with a small disk, a short, columnar central portion, and a terminal tuft of hairs. Fruit ca. 2 mm tall, 3 to 3.5 mm long, 2 mm thick; nutlets 4, sometimes coherent and appearing 2, grouped in pairs, the shared faces reniform and the outer surfaces rounded, strigose. Deep sands of the TX Coastal Plain, N. to Travis Co.; endemic. (Apr.)May-Oct.(Dec.) [H. convolvulaceum (Nutt.) Gray var. racemosum (Rose & Standl.) I. M. Johnst.].
Included here on the basis of several very old collections from Brazos Co. (H. B. Parks, s.n., Jan, 1949, TAES 151680-151682). Probably either not a permanent member of our flora or only sporadic.
Annual or perennial herbs. Herbage glabrous to strigose. Leaves alternate, basal leaves (if any) petiolate, stem leaves sessile. Cymes produced at the branch tips and solitary in the upper axils, slender, uncoiling and elongating to appear raceme-like, with or without bracts; mature pedicels spreading or ascending. Calyx divided equally or unequally to below the middle, in ours the lobes lanceolate or triangular, strigose to hirsute, in our species with few or many uncinate (hooked) hairs. Corolla white or slightly bluish (in other species also blue or rose), small, less than 4 mm long, salverform to nearly funnelform, with a short tube, the lobes convolute or contorted in bud, rounded, throat appendages (fornices) prominent. Stamens included or exserted. Style included, stigma disk-shaped. Ovary 4-ovulate. Nutlets 4, ovoid-elliptic, attached to the flat or high gynobase by a basal-lateral scar, smooth and shiny, sharply keeled all the way around.
50 species of temperate regions; 2 in TX; both here.
Several of the blue-flowered species, especially M. sylvatica and M. scorpioides, are cultivated for their small but pretty flowers (Mabberley 1987).
NOTE: The following two species are recognized by Hatch, et al. (1990) and by Kartesz (1998). However, as pointed out by Steyermark (1963), the characters traditionally used to distinguish these taxa overlap in every particular. The extremes in variation are easily recognizable, but many specimens fall somewhere in between. These two may someday be proven to belong to one highly variable species.
1. Fruiting calyces 4 to 7 mm long, closely spaced, ca. 1 cm or less apart,
with only a few uncinate hairs, mostly at the base ................................................1. M. verna
1. Fruiting calyces 5 to 9 mm long, usually 1 to 5 cm apart, uncinate hairs rather abundant all over the calyx ............................................................................................2. M. macrosperma
var. macrosperma
1. M. verna Nutt. Early Scorpion-grass, Southern Forget-me-not. Annual or winter annual; stem 5 to 40 cm tall, simple or often branched, usually in the inflorescence but sometimes with branches from many of the axils; herbage hirsute-puberulent to hirsute-strigose or -villous. Leaves mostly 1 to 5 cm long and 2 to 10 mm broad, the lowermost more or less rosulate, short-lived, oblanceolate and petiolate, becoming more elliptic and sessile upwards, but only moderately reduced in size, pubescent with bulbous- or disk-based hairs. Inflorescence rarely more than half the height of the plant, cymes often leafy-bracted below or up to the middle, to ca. 15 cm long, at anthesis densely flowered. Pedicels in fruit erect to suberect, up to half as long as the calyx, spaced about 1 cm or less apart; calyx irregular, the upper 3 lobes shorter than the lower 2, 2.5 mm long at anthesis, enlarging to 4 to 7 mm long in fruit, densely hispidulous, with only a few hairs (usually at the base) uncinate; corolla white, 2.5 to 3 mm long, 1 to 2 mm broad, the lobes exceeding the calyx; style about reaching the top of the nutlets. Nutlets 1.2 to 1.5 mm long. Usually in upland sites--rocky or open woods, banks, slopes, and prairies. NE. TX, S. to about Travis and Harris Cos.; N. Eng. to FL, W. to B.C., CA, ID, and TX. Mar.- May. [Formerly often listed as M. virginica (L.) B.S.P., but this misapplied.].
2. M. macrosperma Engelm. var. macrosperma Spring Forget-me-not, Big-seed Scorpion-grass. Taprooted annual or biennial; stem 2 to 8 dm tall, usually simple only when young and thereafter branched at least above the middle, sometimes also below or basally, spreading-hispidulous and often strigose above. Basal leaves forming a short-lived rosette, stem leaves plenty, little reduced upwards, 2 to 8 cm long, 0.5 to 1.7 cm broad, hispidulous, the hairs with disk-like or bulbous bases. Cymes initially coiled, becoming elongate and raceme-like and erect, to 40 cm long, usually with a few bracts at the base, at anthesis rather loosely flowered. Pedicels in fruit ascending or outcurved, 2 to 10 mm long, the lowermost 1 to 5 cm apart in fruit; calyx at anthesis 2.5 to 3 mm long, irregular, the 3 upper lobes shorter than the lower 2, at maturity 5 to 9 mm long, slightly accrescent, falling with the nutlets from the pedicels, rather densely covered throughout with uncinate hairs; corolla white or sometimes bluish, 2.5 to 3 mm long, 2 to 2.5 mm broad, exserted from the calyx (at least the lobes). Nutlets 1.4 to 2.2 mm long. Usually in moist or rich sites, especially bottomlands. E. and NE. TX, SE. to the coast; MD to FL, W. to IL, MO, and TX. Mar.-May. [M. verna Nutt. var. macrosperma (Engelm.) Chapm.; sometimes formerly listed as M. virginica (L.) B.S.P. var. macrosperma (Engelm.) Fern, but apparently the M. virginica misapplied.].
Biennial or perennial (rarely annual) herb. Leaves alternate, basal and/or cauline, basal and lower ones petiolate, the upper sessile. Cymes solitary or grouped at the stem tips, bractless or bracted only at the base, scorpioid and coiling when young, elongating and rather raceme-like in fruit, flowers short-pediceled. Calyx lobed to below the middle, sepals equal, somewhat enlarged in fruit, spreading or reflexed. Corolla cylindrical or funnelform, white, blue, purple-red, or lilac, the lobes broadly rounded, imbricate in bud, commonly overlapping at the edges, throat with prominent, trapezoidal, oblong, or subulate appendages. Anthers included or slightly exserted. Style gynobasic (not always obviously so in developing ovaries), ovules 4. Nutlets (1 to) 4, rather divergent from the convex to pyramidal gynobase, attached with a small or large medial to apical scar (but sometimes appearing attached basally), the surface usually densely covered with glochidiate hairs, sometimes
with an elevated margin and often with a free, subulate portion appressed to the style and obscuring it.
About 55 species of temperate and warm regions; 2 in TX; 1 uncommon here.
A few species have ornamental or medicinal uses (Mabberley 1987).
1. C. virginianum L. Blue Hound's Tongue, Wild Comfrey. Perennial from a large root; stem erect, unbranched, 3 to 8 dm tall, stem and herbage conspicuously spreading-hirsute. Basal leaves with blades elliptic-oblong, 10 to 20 cm long, tapered at the base and decurrent on long petioles, stem leaves sessile, some definitely clasping, some narrowed below and abruptly expanded at the base, leaves progressively smaller upwards. Cymes (1)3(4) at the tip of the stem, 10 to 20 cm long at maturity. Calyx at anthesis 3 to 4 mm long, with lobes expanding to 2.5 mm long in fruit; corolla white to light blue, 8 to 12(16) mm broad, the tube 1.5 to 3 mm long, lobes to ca. 4 mm long, overlapping and the sinuses closed. Fruiting pedicels 5 to 15 mm long, recurved. Nutlets 4, ca. 5 to 7 mm long, uniformly bristly except for the attachment scar, without a raised margin; style 1 to 2 mm long, obscured by the nutlets. Usually in upland deciduous woods, but known from near a bog in Leon Co. Que. and N. B. to CT and NY, S. to FL, W. to B.C., MN, and TX. Mar.-Apr.
Kartesz (1998) lists 2 varieties. If varieties are recognized, our plants are probably var. virginianum.
NOTE: The Asian species C. zeylanicum (Vahl) Thunb. ex Lehm., Ceylon Hound's Tongue, is sporadically introduced in E. TX. It may someday be found here. It is a perennial with appressed, silky hairs, tawny-haired cymes, corolla only 4 to 5 mm long, and nutlets 2.5 to 4 mm long.
Annual, biennial, or perennial herbs, sometimes suffruticose. Stems 1 to several. Herbage usually with dense, conspicuous, stiff pubescence, sometimes the hairs with pustule-like, silicified bases. Leaves basal and/or cauline, sometimes opposite below, alternate above, sessile or the lower petiolate, usually without apparent lateral veins, reduced upwards and grading into the bracts of the inflorescence. Cymes single or paired at each branch tip, dense and definitely coiled when young, elongating and becoming spike- or raceme-like, occasionally grouped to form a thyrse- or panicle-like inflorescence; bracts present or absent, sometimes present only below, easiest to see in the elongated cymes; flowers sessile or shortly pediceled. Calyx divided almost completely, the lobes equal, erect or connivent, linear to lanceolate or oblong, with hairs inside and out, persistent and closely investing the nutlets at maturity. Corolla tiny or conspicuous, but seldom truly showy, salverform to funnelform, usually with a shortly cylindrical tube, with or without scales inside at the base, limb commonly white, with appendages (fornices) in the sometimes-yellow throat, lobes imbricate in bud, rounded. Stamens attached in the corolla tube below the middle, filaments very short. Ovary deeply 4-lobed, style gynobasic, stigma capitate, ovules 2 to 4. Nutlets 4 or else 1 to 3 (by abortion), dissimilar or all alike, erect, ovate to triangular, often angled, attached for most of the ventral length to the usually columnar, pyramidal, or subulate gynobase, the ventral attachment scar usually a groove forked at the bottom and open or closed at the base or sometimes a triangular opening, outer surface smooth to roughened, with or without a margin.
About 100 species of W. N. Amer.; 15 in TX; apparently only 1 here.
Some species provide forage for sheep (Correll & Johnston 1970).
Positive identification requires mature nutlets.
1. C. mexicana (Brandeg.) I. M. Johnst. Mexican Cryptantha. Taprooted annual; stems several to many from the base, repeatedly dichotomous, branches low, spreading or sometimes erect, 5 to 20 cm tall; herbage hispid and sometimes also sparsely strigose. Leaves oblong-lanceolate to slenderly oblong, to 4.5 cm long and 6 m broad, gradually reduced upwards, obtuse to acute, hispid, the spreading hairs from disk-like mineralized bases. Stems floriferous to below the middle, often even to the base, cymes many, definitely coiled, to 15 cm long, densely or loosely flowered, with many obvious bracts, pubescence of unexpanded tips often pale yellowish, at least when dry. Calyx at anthesis to 2.5 mm long, lobes lanceolate, enlarging to 4 mm long and 2.5 mm broad at maturity and becoming broadly ovoid, eventually deciduous; corolla small, 2 to 2.5 mm long, white, the lobes ascending, apically rounded, appendages in throat puberulent, ca. 2 mm tall, trapezoid-shaped. Nutlets 4, triangular-ovate, to 1.3 mm long and 0.9 mm wide, all alike but sometimes the abaxial one more firmly attached, ventral surface with a large triangular open area representing the attachment scar, surface covered with coarse, low, rounded warts. Usually in caliche or limestone-based soils of the Trans Pecos, E. to Howard, Tom Green, Edwards, McMullen, and Cameron Cos.; TX, S. into N. Mex.; common along highways and possibly expanding its range. Apr.-June. [In the past, confused with C. albida (H.B.K.) I. M. Johnst. and not distinguished from it. Old, mislabeled specimens may exist.].
Included here on the bases of one specimen (TAES 46829) collected in Brazos Co. by H. B. Parks. It was identified as C. texana (A. DC.) Greene but has the nutlets characteristic of C. mexicana. C. texana is found in S. TX N. to Bastrop Co. and with an outlying population in Somervell Co. It is possible this species too may be found here. It can be distinguished from C. mexicana by inflorescence without bracts; nutlet surface with many minute warts, ventral surface with a vertical groove divided at the bottom.
Taprooted annuals. Stem simple or branched. Herbage usually bristly-hispid. Leaves basal and cauline, the rosette leaves sometimes short-lived, stem leaves alternate, linear to ovate, smaller than the basal leaves, usually without obvious veins, hairs often with pustule-like bases. Cymes terminal on the branches and in the axils of the upper leaves, strongly coiled when young, elongating with age, without bracts or with a few bracts in the lower portion; pedicels very short. Calyx lobed nearly to the base, the lobes in ours erect, lanceolate to oblong. Corolla tubular to salverform, yellow to orange and the throat sometimes marked in red, tube cylindrical, glabrous, without appendages inside at the base, lobes imbricate in bud, spreading, throat appendages absent (as in ours) to well-developed, if present then closing the throat. Stamens included, attached near the mouth of the tube or farther down, filaments short. Style filiform, included, stigma capitate, emarginate, included; ovules 4. Nutlets 4, ovoid-trigonous, strongly keeled on the ventral surface, the small attachment scar near the base of the ventral side raised and somewhat caruncle-like; gynobase pyramidal, about half as tall as the nutlets; cotyledons each 2-parted.
About 50 species of W. N. Amer.; 3 in TX (Hatch, et al. 1990); 1 known from our area.
1. A. menziesii (Lehm.) A. Nels. & J. Macbr. Smallflower Fiddleneck. Stem simple or branched form the base, branches usually decumbent, 1.5 to 7 dm tall; herbage with spreading bristly-hispid hairs and little or no finer appressed pubescence except sometimes in the inflorescence. Leaves usually not providing a basal rosette, oblong or broadly linear or the upper ones sometimes elliptic or even lanceolate (rarely ovate), 4 to 10(12) cm long, to 2 cm broad, bristly pubescent with mostly pustule-based hairs. Cymes with no bracts or with only a few at the base, elongating greatly in fruit. Sepals nearly entirely free, 5 to 10 mm long in fruit, the lobes slenderly to widely lanceolate, obtuse bristly pubescence sometimes reflexed in the lower portion; corolla light yellow with no red markings, 4 to 7 mm long, the tube slightly or not exserted from the calyx, limb 1 to 3 mm broad, lobes tiny; stamens inserted just at the apex of the tube. Nutlets triangular-ovoid, green to gray, brown, or black, 2 to 3.5(5) mm long, the dorsal and lateral ridges strongly toothed, ventral attachment scar slightly elevated, ovate to rounded, ca. 1 mm long, surface tuberculate to muricate, often rugose. Grasslands and dry areas of Cen. TX; AK to CA and NE, E. to ID, UT, and as far E. as TX; introduced eastward outside the home range. Mar.-May. [A. micrantha Suksd.; A. idahoensis M. E. Jones; Echium menziesii Lehm. Kartesz (1998) lists this species and A. intermedia Fisch. & Mey. combined under A. menziesii.].
Included here on the basis of a specimen found as a weed in a clover field on the A&M campus in 1946. (TAES 59450); quite possibly not a persistent member of our flora.
Taprooted perennial herbs. Stems branched. Herbage hispid-hirsute. Leaves primarily cauline, alternate, entire, with the midvein and laterals quite obvious beneath. Cymes terminal, with leafy bracts, coiled when young, elongating and becoming racemose with age, the flowers sessile or with short pedicels. Calyx deeply 5-lobed, shorter than the corolla, the lobes sometimes unequal. Corolla white, greenish white, or pale yellow, tubular to funnelform, externally pubescent, glabrous within, lobes erect, acute to acuminate, the sinuses thickened and inflexed. Stamens included, filaments very short, anthers sagittate, included but barely so--about reaching the corolla sinuses, dehiscing before the corolla is mature. Ovary 4-lobed, style filiform, conspicuously exserted, persistent. Nutlets usually only 1 or 2 at maturity, ovoid, smooth to pitted, white (as in ours) to brownish-white, basally attached to an almost flat gynobase, the attachment scar relatively small.
About 15 species of N. Amer; 4 species listed for TX (Hatch, et al. 1990), but three of these treated as subspecies of a single species by Kartesz (1998) and other sources (e.g., Gleason and Cronquist 1963; GPFA 1986). It does seem easiest to "lump" our plants together, otherwise specific determination is difficult and dependent upon having an absolutely perfect and complete specimen at the right stage of development.
NOTE: The other species found in TX, O. helleri Small, is reported from the Post Oak Savannah and Blackland Prairies by Hatch, et al. (1990). However, it seems to be confined to the Edwards Plateau and extreme central portion of the state.
1. O. molle Michx. Stems solitary or several, 3 to 12 dm tall, branched above or below. Hairs of 2 sorts--erect or spreading bristles and softer, appressed pubescence. Basal rosette leaves persistent or not, cauline laves sessile, lanceolate to elliptic or ovate, 5 to 13 cm long, 1 to 3 cm broad, with 5 to 7 nerves prominent below; . Cymes simple or forked. Sepals linear-lanceolate, linear-oblong, or lanceolate, 3.5 to 12 mm long, hirsute externally, strigose within; corolla nearly tubular, 7 to 18(20) mm long, with erect, pointed, greenish lobes. Nutlets 1 to 4, sometimes constricted near the base, smooth or pitted, white or brownish. Throughout much of TX except the Pineywoods, Trans Pecos, and High Plains; S. Ont. and NY to MN, S. to NC, LA, and TX, S. to NE, NM, and UT Apr.-June.
5 subspecies; 3 reported from TX; 2 here.
subsp. bejariense (DC. ex A. DC.) Cochrane Bejar Marbleseed. Stem usually solitary, 6 to 9 dm tall, branched above, the branches ascending, whitish hispid-pubescent and the branches also softly appressed- pubescent. Basal leaves persistent (usually withered) until anthesis, leaves lanceolate or the lower ones oblanceolate-spatulate, the largest leaves to 13 cm long and 1.4 cm wide. Calyx lobes linear-lanceolate, 8 to 10 mm long, acute, appressed pubescent and the base long-ciliate; larger corollas 14 mm long, very finely pubescent externally or appearing glabrous, lobes 3 to 4 mm long, triangular, long-acuminate, sometimes bent, with a few spreading hairs apically. Nutlets ca. 3 mm long, obtuse to acute, slightly to strongly constricted at the base. Dry open woods and hillsides. E. TX, W. to Tom Green and Uvalde Cos., E. to Gonzales and Cherokee Cos.; endemic. Mar.-June; collected in fruit in the fall. [O. bejariense DC. ex A. DC.; O. molle Michx. var. bejariense (DC. ex A. DC.) Cronq.].
subsp. occidentale (Mack.) Cochrane Western Marbleseed. Stems usually several in a tuft from the base, 3 to 6 dm tall (rarely taller), branched above or often from near the base, branches spreading to suberect; herbage with silvery-white or sometimes yellowish appressed to spreading pubescence usually less than 2 mm long, pubescence of branches usually appressed. Basal leaves usually not persisting, cauline leaves usually ca. 5 cm long, 1.5 cm broad, acutish, pubescence usually soft, appressed, and little if at all papillose, but sometimes (usually N. of TX) the hairs spreading and with papillose bases. Bracts 6 to 24 mm long, similar to the leaves, oaten 2-ranked; pedicels in fruit to ca. 6 mm long. Calyx lobes 6 to 12 mm long, lanceolate and acute to more or less obtuse, rarely linear-oblong, canescent, hairs somewhat spreading; corolla 11 to 20 mm long, canescent externally, lobes 3 to 4 mm long, broadly triangular, a tuft of hairs at the apex of each lobe inconspicuous. Nutlets 3.5 to 4 mm long, more or less acute, not constricted at the base, scarcely if at all pitted, dull. Prairies, rock woods, bottomlands, and hillsides. NE. and N. Cen. TX, S. to Bexar, Victoria, and Jackson Cos.; IL to Man. and Sask., S. to NM, UT, and TX. Apr.-June. [O. occidentale Mack.; O. molle Michx. var. occidentale (Mack.) I. M. Johnst.].
Annual or perennial herbs (as ours), some (not ours) somewhat shrubby. Herbage variously pubescent. Leaves basal and cauline, sessile, variously linear to ovate. Flowers in leafy-bracted scorpioid cymes or solitary in the axils of the upper leaves, in ours yellow, in other species also white or violet, sometimes heterostylous or plants producing cleistogamous flowers. Calyx divided nearly all the way, lobes narrow. Corolla longer than the calyx, salverform to tubular or funnelform, the lobes imbricate in bud, spreading at anthesis, throat with appendages, pubescent patches, or glandular areas. Stamens inserted on the corolla tube, filaments short, anthers included or slightly exserted, usually with apiculate connectives. Style filiform, stigmas paired, ovules 4. Nutlets 4 or rarely fewer, in some species the showy chasmogamous blossoms essentially sterile and nutlets produced almost entirely by late-season cleistogamous flowers, each nutlet erect, ovoid or angled, with a broad basal attachment scar, the surface usually white or whitish, porcelain-like and very hard, smooth, roughened, or warty; gynobase flat to very broadly pyramidal.
About 40 species of N. Amer. and another 20 in the E. hemisphere; 10 in TX; 2 here.
Some taxa are dye plants or ornamentals. Others have medicinal uses (Mabberley 1987). Native Americans used the roots of Texas species as a source of purple dye (Correll & Johnston 1970). The outer layer of the root is so full of dye that specimens often stain the herbarium paper on which they are mounted.
1. Largest leaves at midstem and the lowermost very reduced; corolla yellow-orange .............
......................................................................................................................1. L. caroliniense
1. Largest leaves at the base of the plant, basal cluster present; corolla lemon yellow .............
..............................................................................................................................2. L. incisum
1. L. caroliniense (Walt. ex J. F. Gmel.) MacM. Carolina Gromwell, Puccoon. Perennial from a stout, vertical, purple-dye producing taproot; stems few to many, 2 to 4.5 dm tall, simple or branched above, minutely appressed pubescent and also with spreading hairs 1 to 2 mm long. Basal rosette none, lowermost leaves quite reduced, cauline leaves numerous and crowded, linear to lance-linear, 2 to 6 cm long, 3 to 12 mm broad, the largest borne about midstem, strigose with papilla-based hairs, herbage commonly darkening on drying. Cymes terminal, solitary or clustered, few-flowered, coiled at first but straightening and elongating to 1 to 2(3) dm long; bracts ovate to elliptic, acute, much longer than the calyces so that the flowers appear solitary or clustered in the axils of the upper leaves; pedicels 3 to 5 mm long, erect. Calyx strigose, at anthesis 6 to 8 mm long, the lanceolate, herbaceous lobes very unequal, enlarging to 8 to 10(17) mm long at maturity; corolla salverform, bright orange-yellow, 11 to 22 mm long, lobes entire, sparsely hispidulous-villous externally, throat appendages weakly indented; flowers heterostylous: corolla tube of short-styled flowers longer than the calyx, in long-styled flowers about as long as the calyx; stamens attached about the middle of the corolla tube in long-styled flowers, near the top of the tube in short-styled flowers; style exserted to the top of the corolla tube in long-styled flowers, only to the middle of the tube in short-styled flowers. Nutlets white, shiny, ovoid, ventrally keeled, 3 to 3.5 mm long, smooth except for a few pits and slots around the basal keel. Sandy soils of roadsides, prairies, and open woods. Pineywoods, W. to the W. Cross Timbers and S.to the Coastal Plain, also localized populations in the Panhandle; NY to SD and NV, S. to FL and TX; also Mex. Mar.-May. [L. carolinianum Lam.; Batschia caroliniensis (Walt. ex J. F. Gmel.].
Hatch, et al. (1990) did not recognize varieties, but Kartesz (1998) does. If varieties are recognized, it is assumed that ours are var. caroliniense.
2. L. incisum Lehm. Narrowleaf Gromwell, Puccoon. Perennial from a stout, purple-dye producing taproot; stems 1 to many, erect, to 4 dm tall, simple below and usually branched above later in the season; herbage strigose, the hairs usually tightly appressed, but sometimes loosely so. Lowest leaves in a basal cluster, these commonly dried but persistent at flowering time, but sometimes absent by then, to 12 cm long and 1 cm broad, usually (but not always) wider than the lower stem leaves, acute, stem leaves many, linear to lance-linear, 1.3 to 6 cm long, 3 to 10 mm broad, gradually reduced upwards, strigose, the hairs sometimes papilla-based, leaf bases sometimes hispid-ciliate. Spring flowers: in leafy-bracted terminal clusters, with pedicels 1 to 4 mm long, erect, chasmogamous, usually functionally sterile, calyx 3 to 15 mm long, the slender lobes unequal; corolla salverform, usually lemon-yellow (occasionally tinged with orange or green), strigose externally, the tube 15 to 35(48) mm long, limb 9 to 20 mm across, the lobes usually erose or fimbriate, occasionally entire, throat with weakly to strongly indented appendages; stamens positioned at the top of the tube; style 1 to 3 cm long, stigma slightly to strongly exserted. Late spring and summer flowers: produced when the plants are well-branched and very leafy, the basal rosette leaves usually gone, and the stem leaves present are usually only 2 to 3 cm long and 1.5 to 3 mm broad; cleistogamous and very fertile, small and inconspicuous among the bracts, terminal; pedicel ultimately 2 to 4 mm long, typically recurved; calyx much longer than the corolla, 2 to 6 mm long at anthesis and subsessile, enlarging at maturity to 5 to 7(8) mm long; corolla closed, 1 to 6 mm long or absent, tubular, glabrous except for the hairy summit; styles all short. Nutlets white, lustrous, constricted near the base, sometimes pitted. Plants vary in height, bushiness, and corolla size across the range, being shorter, bushier, and more often with basal leaves westward and northward in TX, the nutlets smaller and more pitted here than to the E., and chasmogamous flowers smaller and less fertile westward. Usually in sandy soils, but in varying habitats--prairies, fields, open woods, and disturbed areas. Throughout most of the state; Ont. to B.C., S. to UT, TX, MO, and IN; also N. Mex. Collections with chasmogamous flowers primarily Feb.-early April. [L. linearifolium Goldie; L. brevifolium Engelm. & Gray; L. mandanense Spreng; L. angustifolium Michx.; Batschia linearifolia (Goldie) Small].
About 15 species native to Eurasia; 1 species widely introduced/naturalized in the U.S.; present in our area.
Sometimes included in Lithospermum.
1. B. arvensis (L.) I. M. Johnst. Taprooted annual; stems 2 to 7 dm tall, usually shorter, simple or with a few branches above; herbage pale-strigose. Leaves 2 to 5 cm long, 2 to 10 mm wide, the basal leaves in a rosette, commonly absent or withered by flowering time, oblanceolate to spatulate, cauline leaves lanceolate or linear and acute to obtuse, only the midrib obvious below. Cymes racemose, with leafy bracts so that the flowers appear solitary in the axils of the upper leaves; pedicels ca. 1 mm long. Calyx lobes subulate to linear, hispid, in fruit the lobes erect, ascending, 8 to 13 mm long and the short tube becoming pale and somewhat papery; corolla white to blue, purple, or yellow, 5 to 8 mm long, scarcely exserted from the calyx, limb to 4 mm across, gradually expanded, puberulent, without appendages, lobes ascending, ovate and obtuse. Nutlets light brownish-gray ca. 3 mm long, trigonous-ovoid, with a prominent ventral keel, rugose, tuberculate, rough, or pitted, basal attachment scar broad. Open
woods, fields, waste places, etc., commonly in sandy soil. E. 1/2 TX; native of Eurasia, introduced in the U.S., Afr., Asia, and S. Amer. Feb.-June. [Lithospermum arvense L.].
Ours herbs, shrubs, and small trees (elsewhere also larger trees and woody vines). Stems and branches often quadrangular. Herbage glabrous to variously pubescent. Leaves usually opposite, simple or palmately compound, deciduous, estipulate. Inflorescences terminal or axillary spikes, heads, cymes, panicles, etc., often subtended by involucral bracts, individual flowers often subtended by bractlets. Flowers pedicellate or sessile, usually perfect (sometimes imperfect or plants polygamous), hypogynous, usually slightly to strongly irregular. Calyx of (4)5 sepals united at least basally, sometimes zygomorphic, tubular to campanulate or salverform, commonly persistent and accrescent. Corolla of (4)5 lobes, more or less irregular (sometimes nearly regular or bilabiate), salverform with a well-developed tube and limb or else funnelform and the tube widening gradually into the limb. Stamens (2)4 or 5, inserted on the corolla tube, alternate with the lobes, if 4 often didynamous; staminodes present in some taxa. Nectary disk sometimes present or nectary present among corolla lobes. Carpels most often 2, united, gynoecium initially 2-locular but usually a false septum present, producing 4 locules, each uniovulate; some taxa with more locules or else, as in Lantana, 1 carpel aborting and the fruit 2-celled, sometimes the false septum not developing and ovules 2 per locule; placentation axile; style 1, terminal. Fruit usually dry, schizocarpic, separating into 4, 1-seeded nutlets OR fruit more or less fleshy and drupe-like, 2- or 4-celled, with (1)2 or 4 stones, sometimes dehiscent at maturity.
This is a large family whose limits are not well defined. Several small families can be split off from the core of the Verbenaceae on the basis of gynoecium and fruit anatomy (e.g. Avicenniaceae, Phrymaceae, etc.). However, at what taxonomic level these differ from Verbenaceae (s.s.) seems to be a matter of taste.
As treated here (without the Avicenniaceae, etc.) about 71 genera and 1,780 species, mostly tropical and a few subtropical and temperate; 14 genera and 71 species in TX; 6 genera and 21 species here.
The family includes a number of important ornamentals in Verbena, Clerodendron, Caryopteris, etc. Some of the tropical genera are important for timber (e.g. Tectona--teak). Some taxa are used in regional herbal medicines. Others may be weedy (Mabberley 1987).
1. Leaves palmately compound; plants trees or large shrubs ........................................1. Vitex
1. Leaves simple; plants shrubs, subshrubs, or herbs ................................................................2
2(1) Flowers in dichotomously branched cymes; fruit fleshy, bright pink-purple ......2. Callicarpa
2. Flowers in spikes or racemes that are variously arranged; fruit dry or else dark blue or black if fleshy .............................................................................................................................3
3(2) Fruit fleshy, drupe-like; flowers yellow to orange, red, or pink--only lavender or white in cultivated varieties ...................................................................................................3. Lantana
3. Fruit dry and nutlet-like; flowers purple, lavender, pink, blue, or white--only bright red, pink, or yellow in cultivated varieties .................................................................................................4
4(3) Plants upright shrubs; flowers white ........................................................................4. Aloysia
4. Plants herbvs (sometimes woody at the base); flowers blue, purple, pink, or white .............5
5(4) Inflorescences terminal, variously arranged; fruit a cluster of 4 nutlets; plants erect to prostrate ..................................................................................................................5. Verbena
5. Inflorescences axillary; fruit a cluster of 2 nutlets; plants prostrate or creeping .......6. Phyla
Ours trees or shrubs (elsewhere also scandent plants). Leaves opposite, palmately (1-)3- to 7-foliolate, leaflets petiolulate or sessile, entire to incised or lobed. Inflorescences basically cymose, variously arranged, axillary or terminal, rarely (and not in ours) in heads; bractlets usually small and linear, but sometimes longer than the calyx. Calyx campanulate to tubular-funnelform, with (3)5 lobes or teeth, usually slightly irregular. Corolla blue, violet, white or yellowish, funnelform or salverform, tube cylindric, slightly to strongly curved, limb oblique, nearly 2-lipped, the upper part bifid, the lower part 3-lobed, the posterior 2 lobes exterior and usually smaller. Stamens 4, didynamous, often exserted. Ovary bicarpellate, eventually 4-locular and 4-ovulate; style briefly bifid. Mature calyx commonly accrescent, flat to cup-like, scarcely enclosing the fruit. Fruit drupe-like and more or less fleshy, with a hard endocarp, the 4 seeds without endosperm.
About 250 species of tropical to temperate regions; 2 cultivated in TX and escaping; both recorded from our area.
The trees that Americans are familiar with are small and grown for their ornamental flowers. Some tropical species, though, are important timber plants (Mabberley 1987).
1. Leaflets mostly 5 to 9, entire to merely undulate; cymules essentially sessile, usually unbranched ...............................................................................................1. V. agnus-castus
1. Leaflets mostly 3 or 5, usually dentate, serrate, incised, or pinnatisect; cymules usually with a definite stalk and obviously branched ....................................................2. V. negundo
1. V. agnus-castus L. Common or Lilac Chaste-tree, Indian Spice, Hemp-tree, Monk's Pepper-tree, Wild-pepper, Abraham's Balm, Chaste Lamb-tree, Sage-tree, Wild Lavender, Chastity Tree. Shrub or small tree to 5 m; branches densely short-pubescent and resinous-punctate, more or less aromatic. Petioles to 7.5 cm long; blades (3-)5- to 7-(9-) foliolate, the leaflets elliptic-lanceolate to oblanceolate, to 15 cm long and 2(4) cm broad, the 3 middle leaflets largest and usually petiolulate, all tapered to both ends, entire to undulate-repand, green above, white or gray tomentose and glandular below. Flowers in small cymules grouped into terminal or axillary (or supra-axillary) pyramidal-paniculate clusters, the cymules usually sessile and unbranched; peduncle 1.5 t 8 cm long, it and the axis white-puberulent and resinous; bractlets 1 to 4 mm long, linear-setaceous. Calyx widely campanulate, 2 to 2.5 mm long, densely pale-puberulent, the rim with 5 small, obtuse teeth; corolla slightly zygomorphic, the tube 6 to 7 mm long, puberulent externally above the calyx, limb to 13 mm wide, the lobes 2 to 3 mm long. Creek and river bottoms, fencerows, old homesites, etc. Native to S. Eur. and Asia; widely cultivated and escaping; found escaped or naturalized nearly throughout TX except the High Plains and Trans-Pecos; S. U.S., N. to MD. Apr.-Oct., sometimes in several flushes or as moisture permits.
Several forms exist in cultivation.
var. agnus-castus is the typical variety with lavender or lilac flowers and is the most common here.
var. caerulea Rehd. has blue flowers.
forma alba (West.) Rehd. has white flowers.
The twigs are used in basketwork in some regions. The fruit can be used as a pepper substitute, which accounts for many of the common names. White flowered plants are a traditional symbol of chastity (Mabberley 1987).
2. V. negundo L. Shrub or small tree to 4 m; branchlets short-pubescent. Petioles to 6 cm long, puberulent; leaflets 3 or 5, subequal or the middle 3 somewhat larger, all petiolulate or only the 3 larger with stalks, oblong or elliptic to lanceolate, to 11 cm long and 4 cm broad, long-tapered to the apex, tapered to the base, margin irregularly dentate to serrate with a few teeth, varying to deeply and irregularly incised or pinnatisect and divided about halfway to the midrib, upper surface glabrate to minutely puberulent, lower surface appressed-puberulent (sometimes hard to see on older leaves). Inflorescences small, stalked, branched cymules paired and grouped into large paniculate, simple or branched inflorescences to 42 cm long and 15 cm broad, canescent throughout; peduncle 4.5 to 7 cm long, it and the axis densely white puberulent; bractlets many, linear, 2 to 8 mm long. Calyx widely campanulate, 2 to 3 mm long, densely pubescent, the 5 teeth (long-) apiculate; corolla violet to purple or bluish, tube ca. 3 mm long, limb to 7 mm broad. Occasionally escaped from cultivation; native to China. Summer. (Over)
var. heterophylla (Franch.) Rehd. has leaflets mostly only 2 to 7 cm long, deeply and irregularly pinnatisect or incised, often to halfway to the midrib. Known from a Brazos Co. specimen (TAES 16266, from 1919), but possibly only from cultivation--the collector, H. B. Parks, frequently failed to include such designations on his labels. [Includes var. incisa (Lam.) Clarke].
var. intermedia (P'ei) Moldenke has leaves dentate to serrate. Escaped in S. and SE TX; known from Grimes Co.
About 150 species of tropical and subtropical regions; 1 in TX.
Some, including ours, are cultivated for their ornamental fruit (Mabberley 1987).
1. C. americana L. American Beautyberry, French Mulberry, etc. Shrub to 3 m tall, well-branched but not dense; twigs densely stellate-scurfy or tomentose, light brown. Leaves opposite, thin, ovate to elliptic, 8 to 23 cm long, 3.5 to 15 cm broad, acute to acuminate, tapered to the petiole, serrate or crenate-dentate, at least along the middle of the sides, pale stellate-scurfy, especially below and when young, becoming glabrate above, light green. Cymes axillary, 1 to 3.5 cm long, usually shorter than the subtending petiole, dense, and with many small flowers, several to many times dichotomous; peduncle 3 to 10 mm long, pubescent like the twigs or glabrate; pedicels 0.4 to 1.2 mm long, scurfy or glabrous; bractlets small, subulate or setaceous. Calyx campanulate to obconic, (1)1.6 to 1.8 mm long, 1 to 1.5 mm in diameter, nearly truncate, with 4 very small teeth, slightly puberulent-granulose; corolla white to pale pink or bluish, funnelform, small, the tube 2.6 to 2.9 mm long, the 4 lobes 1.3 to 1.5 mm long, blunt. Fruit in dense clusters, very showy, bright pink-purple to lilac, violet, or rose, 3 to 6 mm in diameter, globose, fleshy, each 4-seeded, the seeds ca. 2.3 mm long. Dry or moist woods, thickets, wet slopes, bottomlands, fencerows, swamp margins, etc., common as an understory plant in local woods. E. TX, W. to Tarrant, Kendall, and Bexar Cos.; MD to MO, S. to NC, TN, AR, OK, and TX; also N. Mex. Flowers Jun.-fall; fruit conspicuous in autumn.
Sometimes cultivated--and deservedly so--for the showy fruit. A white fruited form exists but is not found in our area outside of cultivation. Some sources say the fruit is safe to eat, while some say it is not. It is not listed by the AMA as poisonous (Lampe 1985), but it is unpalatable and probably should not be considered edible (Tull 1987).
Herbs or shrubs, erect to scandent or decumbent; branches in ours angled. Herbage scabrous and hirsute or tomentose, often aromatic or unpleasantly scented. Leaves opposite, ovate to ovate-lanceolate, dentate to crenate. Inflorescences spikes, cylindrical or commonly contracted and head-like, usually pedunculate and axillary. Blossoms yellow, orange, red, purple, white, etc, each subtended by an ovate to lanceolate bract. Calyx small, truncate and entire or with small teeth. Corolla salverform, regular to zygomorphic, 4-lobed or sometimes somewhat 5-lobed or obscurely bilabiate, the lobes obtuse to retuse. Stamens 4, included, didynamous, inserted about the middle of the corolla tube. Ovary unicarpellate, bilocular, each cell uniovulate; style included, stigma oblique or sublateral, bilobed. Fruit usually fleshy and drupe-like, in ours blue-black, the endocarp stony, 2-celled or splitting into 1-seeded pyrenes; seeds without endosperm.
About 150 species of tropical America and Africa; 6 in TX; 2 here.
Many species are cultivated for their showy flowers. Some, e.g. L. camara, are weedy when introduced and compete aggressively with native plants. The fruit of some species is edible (Mabberley 1987).
1. Leaves (or at least the basal ones) coarsely serrate-dentate with outward-pointing teeth, all relatively small, mostly to 5 cm long and 2 to 4 cm wide; lower inflorescence bracts often larger than the upper; native species ....................................................1. L. urticoides
1. Leaves with regular, abundant, small, forward-pointing teeth, the larger blades mostly 5 to 11 cm long; inflorescence bracts all the same size; introduced species ..........2. L. camara
1. L. urticoides Hayek Common Lantana, Texas Lantana, Hierba de Cristo, Bunchberry, Calico Bush. Shrub (or perennial herb where colder) 0.5 to 2 m tall; bark gray or brown, unarmed to densely armed with recurved prickles; herbage glabrous to densely hirsute, the younger parts hispidulous to hirtellous. Petioles 2 to 12 mm long, sparsely to densely pilose-hirsute; blades ovate to subrotund-ovate, acute to obtuse, base truncate to subtruncate or cuneate-attenuate to the petiole, (2.5)3 to 5(9) cm long, (1.5)2 to 4(6.5) cm broad, margin coarsely serrate from tip to near the base, the teeth relatively large, irregular, wide-spreading, acute to obtuse, the upper surface short-strigose and very scabrous, often somewhat bullate (blistered) on drying, lower surface varying from densely hirtellous to hispidulous, puberulent, or subglabrate. Peduncles 3 to 9 cm long, usually slightly longer than the subtending leaves, subglabrous to more usually hispidulous-hirsute; floral bractlets narrowly lanceolate to oblong, 4 to 9 mm long, 1 to 2 mm broad, acute, sparsely to densely strigillose, often unequal, the innermost about as long as the corolla tubes and the outer longer, and sometimes broader, forming an involucre-like structure about as long as the corolla tube; heads hemispherical, 2 to 3 cm broad, not elongating. Corollas yellow, dark yellow, orange, red, or some combination of these, the tube 7 to 10 mm long, densely pubescent externally, limb 5 to 9 mm broad. Fruit to ca. 6 mm in diameter, black or deep blue. Fields, woods, swamps, roadsides, chaparral, etc. Throughout much of TX except the NW.; also NM, AZ, and CA; N. Mex.; introduced in NC and MS. Often cultivated. Spring-fall. [Includes var. hispidula Moldenke; L. horrida Kunth in H.B.K. (sensu Moldenke. not Kunth) and var. latibracteata Moldenke; L. scorta Moldenke; L. notha Moldenke].
The fruit is reportedly edible (Moldenke 1942; Correll and Johnston 1970) but the unripe fruit can be dangerous to children, while the fruit and foliage are also said to be toxic to livestock (Tull 1987).
2. L. camara L. West Indian Lantana, Alfombrilla-hediona. Shrub to ca. 2 m tall; branches unarmed or with a few prickles, commonly hispid, young branchlets usually densely spreading-hirsute. Petioles 5 to 20 mm long, glandular-puberulent and spreading-hispidulous; blades ovate to oblong-ovate, (2)5 to 11(12) cm long, mostly 2.5 to 7 cm broad, acute to short-acuminate (occasionally obtuse), base acutely narrowed or rounded to a cuneate-acuminate portion, margin crenate to serrate with many regular, appressed or forward-pointing teeth, more or less rugose-reticulate and scabrous above, more or less densely short pubescent blow, mostly on the veins. Peduncles 2 to 8 cm long, shorter than to longer than the subtending leaves, more or less appressed pilose-puberulent or the hairs sometimes spreading; floral bractlets oblong to narrowly lanceolate, 4 to 7 mm long, 1 to 1.5 mm broad, all equal, about half as long as the corolla tubes, subulate to acute (rarely a few larger ones present), strigose; heads hemispheric, ca. 2 to 3 cm broad, not elongating. Calyx ca. 3 mm long; corollas showy, the outer ones yellow and becoming orange or dark red, the inner ones yellow and changing to orange, tube ca. 10 mm long, puberulent externally, the limb 6 to 9 mm wide. Fruits ca. 3 mm broad, black. Native from FL and Berm., through the W.I.; less common in Cen. and S. Amer.; widely cultivated (sometimes weedy) and escaping in S. and Cen. TX; introduced in tropical Asia and Afr. Blooming nearly throughout the year. [Includes var. mista (L.) Bailey, var. mutabilis (Hook.) Bailey, var. aculeata (L.) Moldenke, var. flava (Medik.) Moldenke, var. hybrida (Neubert) Moldenke, var. nivea (Vent.) Bailey, and var. sanguinea (Medik.) Bailey; L. aculeata L.; L. tiliifolia of authors, not of Cham.].
The fruit is poisonous (Lampe 1985). Cultivated forms have many colors, the heads solid or mixed yellow, white, pink, lavender, red, etc. Butterflies are attracted to the blossoms.
Shrubs, usually erect and the foliage sweetly scented. Leaves opposite, deciduous, entire to toothed. Inflorescences slender racemes or spikes, each flower subtended by an inconspicuous or conspicuous bractlet. Calyx with 4 slender, subequal teeth, tube angled, tubular-campanulate and usually pubescent. Corolla salverform, zygomorphic and bilabiate, lobes of the lower lip about equalling the upper lip. Stamens 4, didynamous, inserted about the middle of the corolla tube, included. Ovary unicarpellate, bilocular, each cell uniovulate. Fruit dry and separating at maturity into 2 thin-walled cocci.
37 species of SW. N. Amer., Mex., and S. S. Amer.; 3 in TX; 1 here.
Some species are cultivated for ornament or used as an aromatic or flavoring (Bailey, et al. 1976; Mabberley 1987).
1. A. gratissima (Gill. & Hook.) Troncoso Whitebrush, Common Bee-brush, Palo Amarillo, etc. Plants slender, to 3 m tall, well-branched; twigs stiff, gray, grayish-puberulent, sometimes spiny at the tips, wood yellow. Leaves opposite-decussate, often with axillary fascicles of smaller leaves; petiole 1 to 3 mm long or wanting; blades narrowly oblong-elliptic to lance-oblong, 3 to 27 mm long, 2 to 8 mm broad, apically obtuse to acute or sometimes minutely mucronate or emarginate, tapered to the base, entire, upper surface minutely strigillose and scabrous-pustulate, lower surface densely grayish-puberulent and glandular-punctate. Inflorescences usually 1 or 2 per node, longer than the leaves, 2 to 7 cm long, many-flowered; flowers white or tinged with violet, vanilla-scented, subtended by lanceolate, acuminate, densely puberulent, deciduous bractlets 1 to 1.5 mm long. Calyx tubular-campanulate, 2.5 to 3 mm long, 4-ribbed, densely white-hirsute and with glands beneath the hairs, the 4 lobes linear-subulate and unequal; corolla tube longer than the calyx, externally glabrous, limb ca. 3 mm broad, throat loosely villous internally. Dry areas: grasslands, bluff, washes, woods, outcrops, etc. Throughout much of TX except Plains Country; very uncommon in our area, but 1 specimen known from Brazos Co. (TAMU 020773)--possibly cultivated; also NM, Mex., and S. Amer. Mar.-Nov. A. ligustrina (Lag.) Small; A. lycioides Cham.; Lippia lycioides (Cham.) Steud., L. ligustrina (Lag.) Britt.].
The Brazos Co. specimen is var. gratissima, with narrow, entire leaves. Var. schulzae (Standl.) L. Benson has larger, broader leaves, usually with 2 to 8 teeth.
The foliage is reportedly a good browse for sheep, though the flowers are said to be toxic to horses. The flowers are a source of nectar that makes good honey (Powell 1988). The leaves and flowers can be made into a tea (Tull 1987).
Ours annual or more usually perennial herbs. Stems and branches procumbent to ascending or erect, often quadrangular, glabrous to variously pubescent. Leaves opposite, simple, very rarely entire, usually dentate or serrate to pinnately incised, cleft, lobed, or pinnatifid, glabrous to variously pubescent, sessile or petiolate. Inflorescence basically a spike, dense to loose, sometimes flat-topped and appearing somewhat umbellate, sometimes elongating greatly in age and the fruits remote; spikes solitary at the branch tips, cymosely in 3's, or clustered and more or less paniculate, each flower subtended by a bractlet, these usually rather inconspicuous but sometimes foliose and longer than the calyx. Calyx cylindrical, with 5 angles, 5 ribs, and 5 unequal teeth, little changed in fruit. Corolla salverform or funnelform, the tube straight or curved, the limb flat and spreading, slightly 2-lipped or zygomorphic, lobes 5, usually unequal. Stamens 4, didynamous, attached in the upper 1/2 of the tube, usually included, anther connective with or without a glandular appendage. Ovary bicarpellate, with 4 uniovulate cells; style 1, terminal, the apex 2-lobed, one lobe sterile and smooth, the other papillose and stigmatic. Fruit separating at maturity into 4 nutlets, usually enclosed by the calyx, both the exterior and the commissural (shared, interior) faces variously and often diagnostically sculptured.
About 250 species of temperate and tropical America, only 2 or 3 in the Old World; 34 species in TX; 13 known from our area.
Some species have medicinal value, but more familiar are the species with showy flowers and cultivated as garden ornamentals. V. x hybrida and related cultivars are the common garden verbenas, available in a wide variety of colors.
NOTE: Our plants can be divided into two sections, Verbenaca and Glandularia. The two groups are nearly universally recognized, the only disagreement being at what level they should be split. Umber (1979) argued for maintaining Glandularia as a separate genus, an approach adopted by Kartesz (1998). However, many floras (e.g., GPFA 1986) retain all the taxa in Verbena. The two genera (or sections) may be separated by the characters noted in the following key. These characters are small but consistent. However, since these features are not easily discernable on many specimens under normal field and classroom conditions, all our plants are presented as Verbena and synonyms are provided in the species entries so that the user may apply names in Glandularia if desired.
a. Style several times longer than ovary; nutlets with a basal cavity; sterile style lobe
extending well beyond the stigmatic surface; inflorescence generally of 1 to 3 spikes; calyx usually twice as long as the nutlets and contorted beyond them; anther connective usually with a glandular appendage; corolla usually relatively large and showy ..................................
....................................................................................................genus or section Glandularia
a. Style equalling to only 3 times longer than the ovary; nutlets without a basal cavity; sterile style lobe usually not extending beyond the stigmatic surface; spikes generally aggregated into cymes or panicles; calyx seldom more than twice as long as the nutlets and not contorted beyond them; anther connective without a glandular appendage; corolla usually small and not singly showy ..............................section Verbenaca or genus Verbena
1. Spikes more than 6 mm broad (excluding the corollas), dense in both flower and fruit; flowers usually (but not always) singly showy ..........................................................................2
1. Spikes generally less than 6 mm broad (excluding the corollas), dense or loose; flowers usually (but not always) small and not singly showy ...............................................................7
2(1) Leaves merely serrate ............................................................................................1. V. rigida
2. Leaves pinnately incise, cleft, or pinnatifid ...............................................................................3
3(2) Leaves two or more times pinnately divided, the central portion about as narrow as the ultimate divisions .......................................................................................................................4
3. Leaves pinnately incised, cleft, lobed, or once divided, the central portion wider than the ultimate divisions .......................................................................................................................5
4(3) Bractlets 1/4 to 1/2 the length of the calyx; central portion of leaf ca. 1 mm broad ................
...........................................................................................................................2. V. pulchella
4. Bractlets slightly shorter than to longer than the calyx; central portion of leaf more than 1 mm broad ....................................................................................................3. V. bipinnatifida
var. bipinnatifida
5(3) Floral bractlets much exceeding the calyces ..................................................4. V. bracteata
5. Floral bractlets shorter than to only slightly longer than the calyces .....................................6
6(5) Corollas 2 to 7 mm broad; calyx ca. 6 mm long ..................................................5. V. pumila
6. Corolla more 7 or more mm broad; calyx ca. 10 to 13 mm long ...............6. V. canadensis
7(1) Leaves (at least some) trifid, cleft, divided, or strongly incised ...............................................8
7. Leaves merely serrate ............................................................................................................10
8(7) Stems (and leaves) glabrous or minutely scabrous or pubescent ................7. V. officinalis
subsp. halei
8. Stems (and leaves) densely pubescent, canescent, or hirsute ..............................................9
9(8) Leaf blades (at least some) 3-cleft or 3-lobed; plants generally 5 to 10 dm tall ..8. V. xutha
9. Leaf blades narrowly elongate, not 3-cleft, merely incised to subpinnatifid; plants usually less than 5 dm tall ..........................................................................................9. V. canescens
10(7) Spikes compact in flower and fruit, the calyces overlapping; leaves toothed mostly above the middle ...............................................................................................................................11
10. Spikes becoming loose and open in fruit, the calyces scarcely overlapping; leaves toothed to near the base ......................................................................................................................12
11(10) Leaves all tapered to the base ...................................................................10. V. brasiliensis
11. Leaves (at least some) cordate or subcordate and clasping ...................11. V. bonariensis
12(11) Mature calyx prolonged beyond the nutlets, forming a point ............................12. V. scabra
12. Mature calyx lobes merely slightly arched above the nutlets, the effect essentially blunt ......
.........................................................................................................................13. V. urticifolia
var. urticifolia
NOTES: The common garden verbena, V. x hybrida Groenl. & Ruempler, occasionally escapes or persists in TX. It has shallowly pinnately lobed, rather densely pubescent leaves, and variously colored corollas to 2.4 cm across.
The species listed in above key are those known from our area. Hybrids between closely related species may be common (as noted at species entries) and are often intermediate between parents in appearance. It is also not impossible that one or another of the species found mostly to our south and west may be found here someday. Material absolutely not referable to the species outlined above should be compared with the key presented by Correll & Johnston (1970) and checked against the nomenclatural changes presented by Kartesz (1998).
1. V. rigida Spreng. Tuber Vervain. Rhizomatous perennial; stems 2 to 6(7) dm tall, simple below and branched above, strongly 4-angled, scabrous. Leaves sessile, basally subcordate, somewhat clasping, blades lanceolate to oblanceolate or oblong, ca. 4 to 11 cm long, 0.7 to 3 cm broad, acute to acuminate, sharply serrate with coarse, sometimes spreading teeth, margins of older leaves frequently revolute, scabrous and slightly hispid on both surfaces, veins impressed on the upper surface and raised below, uppermost pair(s) of leaves commonly bractlike and much narrower and more nearly entire. Spikes usually short, 1 to 6 cm long, dense in flower and fruit, more than 1 cm broad, excluding corollas, usually cymosely arranged in 3's or in ternately-branched cymes and ultimately in 3's, the lateral spikes pedunculate, the central spikes sessile or pedunculate; floral bractlets lanceolate, acuminate-subulate, longer than the calyx, glandular-pubescent and ciliate. Calyx 3 to 5 mm long, sometimes tinged with purple, lobes acute, tips mucronulate-subulate, in fruit incurved and connivent over the nutlets, glandular; corolla purple to magenta, drying blue-violet, tube 2 to 3 times longer than the calyx, pubescent, limb (4)5 to 7 mm wide. Nutlets 3-angled, ca. 2 mm long, the upper 1/2 raised-reticulate and the lower portion striate, the commissural faces muricate-scabrous. Mostly on roadsides and in pastures and prairies. E. TX, W. to about Fayette and Travis Cos.; native to Parag. and Brazil; introduced in the W.I. and S. U.S. from GA to FL and W. to TX. Apr.-Oct. [V. venosa Gill. & Hook.].