LOGANIACEAELOGANIACEAE

Logania Family


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



LOGANIACEAE GELSEMIUM1. GELSEMIUM Juss. Yellow Jessamine


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).



LOGANIACEAE SPIGELIA2. SPIGELIA L. Pink-root, Worm-grass


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.].



LOGANIACEAE MITREOLA3. MITREOLA L. Miterwort, Hornpod


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. petiolataLOGANIACEAE MITREOLA 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. sessilifoliaLOGANIACEAE MITREOLA 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.].






BUDDLEJACEAEBUDDLEJACEAE

Buddleja Family


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.



BUDDLEJACEAE POLYPREMUM1. POLYPREMUM L.


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).






GENTIANACEAEGENTIANACEAE

Gentian Family


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



GENTIANACEAE EUSTOMA1. EUSTOMA Salisb. Prairie-gentian, Catchfly-gentian


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.



GENTIANACEAE CENTAURIUM2. CENTAURIUM Hill Centaury


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.



GENTIANACEAE SABATIA3. SABATIA Adans. Rose-gentian


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.






APOCYNACEAEAPOCYNACEAE

Dogbane Family


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



APOCYNACEAE AMSONIA1. AMSONIA Walt. Blue-star, Slimpod, Amsonia


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. ciliataAPOCYNACEAE AMSONIA 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. tabernaemontanaAPOCYNACEAE AMSONIA 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. illustrisAPOCYNACEAE AMSONIA 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.




APOCYNACEAE TRACHELOSPERMUM3. TRACHELOSPERMUM Lem. Climbing Dogbane


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.




APOCYNACEAE NERIUM3. NERIUM L.

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.



APOCYNACEAE VINCA4. VINCA L. Periwinkle


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. minorAPOCYNACEAE VINCA 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. majorAPOCYNACEAE VINCA 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.



APOCYNACEAE APOCYNUM5. APOCYNUM L. Dogbane, Indian Hemp


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).



APOCYNACEAE CATHARANTHUS6. CATHARANTHUS G. Don f.


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.].






ASCLEPIADACEAEASCLEPIADACEAE

Milkweed Family


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



ASCLEPIADACEAE ASCLEPIAS1. ASCLEPIAS L. Milkweed


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. viridisASCLEPIADACEAE ASCLEPIAS 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. viridifloraASCLEPIADACEAE ASCLEPIAS 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. engelmannianaASCLEPIADACEAE ASCLEPIAS 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. amplexicaulisASCLEPIADACEAE ASCLEPIAS 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. verticillataASCLEPIADACEAE ASCLEPIAS 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. oenotheroidesASCLEPIADACEAE ASCLEPIAS 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. tuberosaASCLEPIADACEAE ASCLEPIAS 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. rubraASCLEPIADACEAE ASCLEPIAS 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. obovataASCLEPIADACEAE ASCLEPIAS 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.



ASCLEPIADACEAE CYNANCHUM2. CYNANCHUM L. Swallow-wort, Sand-vine


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.].



ASCLEPIADACEAE MATELEA3. MATELEA Aubl.


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. gonocarposASCLEPIADACEAE MATELEA 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. parvifloraASCLEPIADACEAE MATELEA 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. bifloraASCLEPIADACEAE MATELEA 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. cynanchoidesASCLEPIADACEAE MATELEA 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].






SOLANACEAESOLANACEAE

Nightshade or Potato Family


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 capsule; corolla funnelform to salverform or tubular ..................................................2

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




SOLANACEAE DATURA1. DATURA L. Jimson Weed, Thorn-apple, Stramonium


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. stramoniumSOLANACEAE DATURA 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. wrightiiSOLANACEAE DATURA 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. inoxiaSOLANACEAE DATURA 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.



SOLANACEAE NICOTIANAL2. NICOTIANAL. Tobacco


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. glaucaSOLANACEAE NICOTIANAL 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. repandaSOLANACEAE NICOTIANAL 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].



SOLANACEAE LYCIUM3. LYCIUM L. Wolf-berry, Desert-thorn, Squaw-berry


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.



SOLANACEAE PHYSALIS4. PHYSALIS L. Ground-cherry


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. angulataSOLANACEAE PHYSALIS 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. pubescensSOLANACEAE PHYSALIS pubescensL. 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. pumilaSOLANACEAE PHYSALIS 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. mollisSOLANACEAE PHYSALIS 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. cinerascensSOLANACEAE PHYSALIS 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. hederifoliaSOLANACEAE PHYSALIS 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. heterophyllaSOLANACEAE PHYSALIS 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. virginianaSOLANACEAE PHYSALIS 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. longifoliaSOLANACEAE PHYSALIS 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.].



SOLANACEAE CHAMAESARACHA5. CHAMAESARACHA A. Gray False Nightshade


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.




SOLANACEAE SALPICHROA6. SALPICHROA Miers


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).




SOLANACEAE SOLANUM7. SOLANUM L. Nightshade


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. triquetrumSOLANACEAE SOLANUM 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. ptycanthumSOLANACEAE SOLANUM 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. rostratumSOLANACEAE SOLANUM 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. elaeagnifoliumSOLANACEAE SOLANUM 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. carolinenseSOLANACEAE SOLANUM 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. dimidiatumSOLANACEAE SOLANUM 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.



SOLANACEAE CAPSICUM8. CAPSICUM L. Pepper


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).






CONVOLVULACEAE</STRONG>CONVOLVULACEAE

Morning Glory Family

(Including Cuscutaceae--Dodder Family)


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



CONVOLVULACEAE CUSCUTA1. CUSCUTA L. Dodder, Love Vine, Devil's Gut, Witches' Shoelaces,

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

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. exaltataCONVOLVULACEAE CUSCUTA 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. cuspidataCONVOLVULACEAE CUSCUTA 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. glomerataCONVOLVULACEAE CUSCUTA 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. compactaCONVOLVULACEAE CUSCUTA 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. obtusifloraCONVOLVULACEAE CUSCUTA 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. coryliCONVOLVULACEAE CUSCUTA 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. indecoraCONVOLVULACEAE CUSCUTA 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. gronoviiCONVOLVULACEAE CUSCUTA 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. pentagonaCONVOLVULACEAE CUSCUTA 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.].



CONVOLVULACEAE DICHONDRA2. DICHONDRA J. & G. Forst. Ponyfoot


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.





CONVOLVULACEAE EVOLVULUS3. EVOLVULUS L. Evolvulus


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. nuttallianusCONVOLVULACEAE EVOLVULUS 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. sericeusCONVOLVULACEAE EVOLVULUS 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)].



CONVOLVULACEAE CONVOLVULUS4. CONVOLVULUS L. Bindweed


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. equitansCONVOLVULACEAE CONVOLVULUS 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. arvensisCONVOLVULACEAE CONVOLVULUS 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.



CONVOLVULACEAE STYLISMA5. STYLISMA Raf. Bonamia


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].



CONVOLVULACEAE JAQUEMONTIA6. JAQUEMONTIA Choisy


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.].



CONVOLVULACEAE IPOMOEA7. IPOMOEA L. Morning-glory


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. quamoclitCONVOLVULACEAE IPOMOEA 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. wrightiiCONVOLVULACEAE IPOMOEA 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. hederaceaCONVOLVULACEAE IPOMOEA 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. purpureaCONVOLVULACEAE IPOMOEA 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. pandurataCONVOLVULACEAE IPOMOEA 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. lacunosaCONVOLVULACEAE IPOMOEA 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. cordatotrilobaCONVOLVULACEAE IPOMOEA 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.






POLEMONIACEAEPOLEMONIACEAE

Phlox Family


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




POLEMONIACEAE PHLOX1. PHLOX L. Phlox


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. pilosaPOLEMONIACEAE PHLOX 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. cuspidataPOLEMONIACEAE PHLOX 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. drummondiiPOLEMONIACEAE PHLOX 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].




POLEMONIACEAE IPOMOPSIS2. IPOMOPSIS Michx. Ipomopsis


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.



POLEMONIACEAE GILIA3. GILIA R. & P. Gilia


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.].






HYDROPHYLLACEAEHYDROPHYLLACEAE

Waterleaf Family


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



HYDROPHYLLACEAE HYDROLEA1. HYDROLEA L. Hydrolea


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.].



HYDROPHYLLACEAE NAMA2. NAMA L. Nama


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. jamaicenseHYDROPHYLLACEAE NAMA 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. hispidumHYDROPHYLLACEAE NAMA 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).



HYDROPHYLLACEAE NEMOPHILA3. NEMOPHILA Nutt. Baby Blue-eyes


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. phacelioidesHYDROPHYLLACEAE NEMOPHILA 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. aphyllaHYDROPHYLLACEAE NEMOPHILA 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.].



HYDROPHYLLACEAE PHACELIA4. PHACELIA Juss. Phacelia


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. congestaHYDROPHYLLACEAE PHACELIA 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. glabraHYDROPHYLLACEAE PHACELIA 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. strictifloraHYDROPHYLLACEAE PHACELIA 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. patulifloraHYDROPHYLLACEAE PHACELIA 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. hirsutaHYDROPHYLLACEAE PHACELIA 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.].






BORAGINACEAEBORAGINACEAE

Borage Family


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).



Plants with Mature Fruit (more accurate)

1. Ovary entire or only shallowly lobed; style terminal ......................................1.
Heliotropium

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



Plants with Flowers at Anthesis (use if mature fruits not present)


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



BORAGINACEAE HELIOTROPIUM1. HELIOTROPIUM L. Heliotrope, Turnsole


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. curassavicumBORAGINACEAE HELIOTROPIUM 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. amplexicauleBORAGINACEAE HELIOTROPIUM 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. indicumBORAGINACEAE HELIOTROPIUM 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. procumbensBORAGINACEAE HELIOTROPIUM 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. tenellumBORAGINACEAE HELIOTROPIUM 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. convolvulaceumBORAGINACEAE HELIOTROPIUM 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. racemosumBORAGINACEAE HELIOTROPIUM 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.





BORAGINACEAE MYOSOTIS2. MYOSOTIS L. Scorpion-grass, Forget-me-not


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. vernaBORAGINACEAE MYOSOTIS 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. macrospermaBORAGINACEAE MYOSOTIS 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.].



BORAGINACEAE CYNOGLOSSUML3. CYNOGLOSSUML. Hound's Tongue, Beggar's-lice


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.



BORAGINACEAE CRYPTANTHA4. CRYPTANTHA G. Don Cryptantha


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.



BORAGINACEAE AMSINCKIA5. AMSINCKIA Lehm. Fiddleneck


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.



BORAGINACEAE ONOSMODIUM6. ONOSMODIUM Michx. Marbleseed, False Gromwell


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.].



BORAGINACEAE LITHOSPERMUM7. LITHOSPERMUM L. Gromwell, Puccoon


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. carolinienseBORAGINACEAE LITHOSPERMUM 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. incisumBORAGINACEAE LITHOSPERMUM 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].



BORAGINACEAE BUGLOSSOIDES8. BUGLOSSOIDES Moench


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.].






VERBENACEAEVERBENACEAE

Verbena Family


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



VERBENACEAE VITEX1. VITEX L. Chaste-tree


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-castusVERBENACEAE VITEX 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. negundoVERBENACEAE VITEX 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.


VERBENACEAE CALLICARPA2. CALLICARPA L. Beautyberry


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).



VERBENACEAE LANTANA3. LANTANA L. Lantana


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. urticoidesVERBENACEAE LANTANA 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. camaraVERBENACEAE LANTANA 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.



VERBENACEAE ALOYSIA4. ALOYSIA Ort. Bee-brush


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).



VERBENACEAE VERBENA5. VERBENA L. Vervain, Verbena


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



Key to species of Verbena (sensu lato)


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. rigidaVERBENACEAE VERBENA 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.].


2.V. pulchellaVERBENACEAE VERBENA pulchella Sweet (=Glandularia pulchella (Sweet) Troncoso ) Moss Verbena. Perennial; stems 10 to 30 cm tall, decumbent and rooting at the nodes, branches divergent, ascending, to 60 cm long, sparsely antrorse-pilose, becoming glabrate with age, the main internodes 1 to 3 cm long. Petioles 0.5 to 1.5 cm long; blades triangular in overall outline, 2 to 3.5 cm long, 2 to 3 cm broad, 3-parted and the parts pinnatifid, the central portion and the ultimate divisions linear, ca. 1 mm broad, divisions 1 to 3 mm long, entire or dentate, acute to obtuse, margins revolute, upper surface sparsely antrorsely appressed-strigose, undersurface more densely pubescent and with the veins raised, glabrescent with age. Spikes terminal on the branches, solitary, erect to ascending, dense in flower and fruit, elongating to 3 to 4 cm with age, at least 1 cm broad excluding the corollas; flowers sessile, ascending at anthesis, spreading in age; bractlets ca. 1/4 to 1/2 as long as the calyx, 2 to 3 mm long, subovate-lanceolate, dorsally canescent. Calyx (6)8 to 9 mm long, densely antrorsely white-strigose, and with a few scattered dark glands, tube ca. 7 mm long, teeth ca. 1 to 2 mm long, filiform; corolla showy, blue to violet, lilac, or rose-purple, tube ca. 3 to 5 mm longer than the calyx, glabrous, limb ca. 1 cm broad, the lobes about 3 to 4 mm long and wide, obcordate and emarginate, basally ciliate-pilose; anthers with a glandular appendage; style 1 to 1.3 cm long, longer than the calyx, lasting only 1 day. Fruit linear-oblong, 3 to 3.2 mm long, yellowish, separating into 4 nutlets; nutlets with a basal cavity, externally reticulate, muriculate or warty on the commissural faces. Roadsides, waste places, fields, etc. SE. TX, S. and W. to Zapata, Webb, and Hays Cos.; NC to FL and the Gulf Coast; also IL, KY, MO, AZ, and CA; introduced from S. Amer. Mar.-July. [Includes var. gracilior (Troncoso) Shinners; V. tenera Spreng.; V. erinoides Lam. This plant has been presented in many manuals (e.g., by Correll & Johnston 1970 and Radford, et al. 1968) as V. tenuisecta Briq. or G. tenuisecta (Briq.) Small. Apparently, that name was used before it was firmly established that this plant is the same as a S. American species with an earlier valid name.].


3.V. bipinnatifidaVERBENACEAE VERBENA bipinnatifida Nutt. var. bipinnatifida (=Glandularia bipinnatifida (Nutt.) Nutt. var. bipinnatifida) Dakota Vervain, Small-flowered Vervain, Mexican Vervain, Common Vervain, Wild Vervain, Pink Vervain, Moradilla, Ragweed Vervain, Fringed Vervain, etc. Perennial, sometimes flowering the first year or annual; stems prostrate to ascending (rarely erect), 5 to 60 cm long or tall, often rooting at the lower nodes, sparingly or densely branched from the base, moderately to densely hispid-hirsute. Leaves petiolate; blades 1 to 6 cm long and wide, usually strongly bipinnatifid (sometimes merely pinnatifid, 3-parted, or deeply incised), the segments (3)4 to 10(15) mm broad, linear-oblong, appressed-pubescent on both surfaces, margins usually revolute. Spikes solitary at the ends of the branches, pedunculate, 6 to 20 cm long, dense at anthesis, elongating in fruit but usually remaining fairly crowded, more than 1 cm wide excluding the corollas; floral bractlets 0.6 to 1.2 times as long as the calyx, linear-subulate or lanceolate, glandular or eglandular and with long, stiff hairs; flowers with a sweet fragrance or unscented. Calyx 7 to 10 mm long, eglandular to densely glandular, usually hispid or hirsute, the lobes unequal, 1 to 4 mm long; corolla some shade of purple (drying purple or blue-violet), tube ca. 1.5 times as long as the calyx, limb 6 to 15(20) mm broad; anther connective with a glandular appendage, this minute or about equalling the anther cells; style 10 to 20 x longer than the ovary. Nutlets cylindrical, 2 to 3 mm long, black, with a basal cavity, the outer surface raised-reticulate apically and raised-striate basally, commissural face rounded at the top and not reaching the tip of the nutlet, bare or with smooth or papillose ridges. Dry prairies, roadsides, meadows, streamsides, and so on, commonly on calcareous soils. Nearly throughout TX but rare in the NE; variable and widespread in plains, ND and CO, S. to AZ and TX, rare in MO, AR, and LA; also much of Mex. Flowering nearly throughout the year, our collections mostly Mar.-Oct. [Includes var. latilobata Perry; synonyms for the species as a whole include V. ambrosifolia Rydb. ex Small and forma eglandulosa Perry, V. ciliata Benth. and var. longidentata Perry, V. pubera Greene, V. wrightii A. Gray and forma albiflora, V. andrieuxii Schauer, and V. demareei Moldenke; not Glandularia wrightii (Gray) Umber as described by Umber 1979.].

Tull (1987) includes a recipe for a bright yellow dye using the whole plant.

Known to hybridize with V. canadensis; see NOTE at that species below.


4.V. bracteataVERBENACEAE VERBENA bracteata Lag. & Rodr. Prostrate Vervain. Annual or short-lived perennial; stems usually several from the base, well-branched, prostrate to decumbent or ascending, rarely erect, 1 to 5(7) dm long, coarsely hirsute. Leaves 1 to 7 cm long, 0.6 to 3 cm broad, essentially lanceolate to ovate-lanceolate, pinnately incised or 3-lobed, the lateral lobes small, narrow, and spreading and the central lobe larger, cuneate-obovate, usually incised-dentate or cleft, both surfaces hirsute or appressed-hirsute, basally narrowed into a short margined or winged petiole, major veins sometimes rather prominent below. Spikes terminal on the branches, usually solitary, without an apparent peduncle, commonly ascending, 2 to 20 cm long, 1 to 2 cm wide, dense at anthesis, elongating in fruit, conspicuously bracted; floral bractlets much exceeding the calyx (usually 2 to 3 times longer than the calyx), 8 to 15 mm long, spreading-ascending in flower, recurved in age, acute to acuminate, hirsute, the lower lanceolate, leaf-like, and often incised, the upper linear-lanceolate. Calyx 3 to 4 mm long, the lobes short and connivent over the mature nutlets, hirsute, especially on the veins; corolla blue to violet, lavender, lilac, or purple, tube slightly exserted from the calyx, finely pubescent externally, limb 2.5 to 3 mm across. Nutlets 2 to 2.5 mm long, yellow to yellow-brown, trigonous, outer faces raised-reticulate apically and striate below, commissural face reaching all the way to the top of the nutlet, scabrous and muricate or short-papillose. Low waste ground, disturbed or cleared areas, roadsides, and grassy places. Trans-Pecos and Plains Country, E. through the Ed. Plat. to NE. TX; nearly throughout the U.S. and S. Can. but some eastern locations probably introductions. Included here on the basis of old collections from Brazos Co. from the 1940's (e.g. TAES 54623 and 29471); not seen in the field in recent years. Apr.-Oct. in TX. [V. bracteosa Michx.; V. imbricata Woot. & Standl.].

5.V. pumilaVERBENACEAE VERBENA pumila Rydb. (=Glandularia pumila (Rydb.) Umber) Pink Vervain, Hairy Vervain, Wild Vervain. Annual or short-lived perennial; stems usually several from the base, decumbent to ascending (rarely erect), 0.5 to 2(3) dm long, hirsute and commonly finely glandular. Leaves petiolate, blades 1.5 to 3 cm long, more or less triangular-ovate in outline, trifid or pinnatifid, the divisions often lobed or incised, but the central undivided portion much broader than the ultimate divisions, apex obtuse, base cuneate or truncate, often a portion tapered onto the petiole forming a margin, both surfaces appressed-hirsute, the veins impressed above and raised below, margins revolute. Spikes usually solitary at the ends of the branches, sessile or short-pedunculate, dense in flower and usually not elongating much in age, ca. 1 cm broad; floral bractlets about as long as the calyx, linear-lanceolate, hispid-hirsute. Calyx ca. 6 mm long in fruit, pubescent on the veins, often finely glandular, lobes short, subulate; corolla usually pink, sometimes lavender to blue, rarely white, tube 8 to 10 mm long, slightly exceeding the calyx, little if at all pubescent externally, limb only 3 to 5 mm wide; anther connective gland absent or very small. Nutlets grayish-black, 2 to 3 mm long, reticulate in the upper 3/4, the lower 1/4 vaguely striate, commissural face with smooth or papillose ridges. Roadsides, open woods, salt flats, prairies, pastures, river terraces, etc. Throughout most of TX, not as common in the upper panhandle and E. 1/3 state; NM, OK, TX, AR and perhaps LA; also N. Mex. Jan.-Aug. [V. inconspicua Greene].


6.V. canadensisVERBENACEAE VERBENA canadensis (L.) Britt. (=Glandularia canadensis (L.) Nutt.) Rose Vervain. Perennial, sometimes flowering the first year; stems usually several, decumbent or ascending, sometimes rooting at the lower nodes, usually well-branched, glabrate to spreading-hirsute (sometimes irregularly so), 3 to 6 dm long. Leaves quite variable, petiolate; blades ovate to ovate-oblong, triangular-ovate, or lanceolate in overall outline, 2.5 to 9 cm long, 1.5 to 4 cm broad, incised, incised-pinnatifid, or 3-cleft and incised or pinnatifid, in any case the central undivided portion broader than the ultimate lateral divisions, apex acute to acuminate, base truncate or cuneate, usually tapered onto the winged petiole, both surfaces glabrate, appressed-hirsute, or strigose, slightly scabrous. Spikes pedunculate, usually solitary at the ends of the branches, dense in flower, elongating in fruit, more than 10 mm broad excluding the corollas; floral bractlets shorter than or equalling the calyx, linear-attenuate, ciliate and irregularly hirsute. Calyx 10 to 13 mm long, glandular hirsute, the lobes very narrow, subulate-setaceous, unequal, the posterior (uppermost) the shortest; corolla white, blue, rose, lilac, magenta, lavender, purple, etc. or occasionally variegated, tube about twice as long as the calyx, glabrous to finely pubescent or glandular externally, limb 11 to 15 mm broad, lobes emarginate; anther connective usually with a prominent glandular appendage (or this sometimes absent). Mature fruit constricted along the lines of cleavage, separating into 4 nutlets, each 3 to 3.5 mm long, nearly cylindrical, the base slightly broadened and with a cavity, gray-black to black, reticulate in the upper 1/2 to 2/3, striate below, commissural face muriculate-scabrous or papillose. Open roadsides, woods, hills, railway embankments, etc., frequently on sandy or gravelly soil. NE., E., and S. TX, W. to N. and Cen. TX, perhaps introduced in the S. and W. portions of this range; NC, KY, IA, SE. NE, KS, and CO, S. to FL and E. TX; introduced in CT, NY, MI, and MN. Feb.-Oct. [Includes var. atroviolacea Dermen, var. compacta Dermen, var. grandiflora (Haage & Schmidt) Moldenke, var. drummondii (Lindl.) Baxt., and var. lambertii (Sims) Thell.; V. lambertii Sims; G. lambertii (Sims) Small].

NOTE: Said to hybridize with V. bipinnatifida. V. x oklahomensis Moldenke is the name usually applied to the hybrids, but Kartesz (1998) lists this as a synonym of V. canadensis.


7.V. officinalisVERBENACEAE VERBENA officinalis L. subsp. halei (Small) Barber Slender Vervain, Texas Vervain, Blue Vervain, Candelabra Vervain, Standing Vervain. Annual or short-lived perennial; stems 1 to several from a (semi-)woody base, erect, sparingly branched or branched only in the inflorescence, branches ascending, stem and branches glabrous or sparsely strigillose above. Leaves opposite but the larger ones sometimes crowded toward the bottom of the plant, 1.5 to 8(10) cm long, 1 to 4 cm broad, sparsely strigillose on both surfaces, quite variable in shape, the basal and lower leaves oblong to ovate, irregularly incised or dentate, tapered into a petiole about as long as the blade, passing into midstem leaves which have shorter petioles and are once or twice pinnatifid, uppermost leaves merely incised, dentate, or entire, central portion leaf usually becoming narrower upwards, often some or nearly all the leaves absent by flowering time. Spikes slender, less than 6 mm broad excluding the corollas, elongate, arranged in a loose terminal panicle or sometimes only in 3's, flowers usually imbricate at anthesis, becoming more remote with age; floral bractlets about 1/2 the length of the calyces and tightly appressed to them, ovate-lanceolate, ciliate, sparsely and minutely strigillose. Calyx commonly with some purple or bluish coloration, (1)2 to 3.5 mm long, to 4 mm long in fruit but scarcely accrescent, strigillose or hirsute, especially on the nerves, teeth short, unequal, subulate, the orifice more or less truncate. Corolla pale lavender to blue, blue-lavender, purple, or occasionally white or pinkish, blue 2 to 7 mm long, only slightly longer than the calyx, glabrous to sparsely pubescent, limb 4 to 7 mm broad, lobes retuse. Nutlets yellowish to reddish-brown, 1.5 to 2.2 mm long, without a basal cavity, externally reticulate in the upper 1/4 to 1/2, striate basally, commissural face faintly to densely saginoid (with tiny, peg-like projections). Exceedingly common plant of roadsides, fields, open woods, hillsides, meadows, vacant lots, etc., usually in sandy and/or calcareous soil. Throughout much of TX, especially the E., S., and Cen. regions; NC to OK, S. to FL, AL, and TX; also N. Mex. Feb.-Nov., usually with a flush in spring and a second in fall. [V. halei Small and forma albiflora L. I. Davis and forma roseiflora (Benke) Moldenke].

These plants were formerly treated as V. halei and have long been known to be close to V. officinalis, an Old World species. For example, North American plants were listed under V. officinalis by Britton and Brown in the first edition of their Illustrated Flora (1898). Barber (1982) examined the differences and similarities between Old and New World plants and concluded that the two are conspecific. Kartesz (1998) maintains V. halei. (Over.)

The Old World form was once used medicinally to treat eye diseases, supposedly because the wide-eyed flowers suggested such a use (Mabberley 1987). North American folk and pharmacological traditions employed V. officinalis as an emetic and expectorant (Kindscher 1992).


8.V. xuthaVERBENACEAE VERBENA xutha Lehm. Coarse Vervain, Gulf Vervain. Perennial; stems 1 to several from the base, often clumped, erect or rarely procumbent, 6 to 10(20) dm tall, 4-angled, simple or with ascending branches, spreading hirsute-hispid. Leaves essentially sessile, the larger 5 to 12 cm long, more or less ovate in overall outline, incised-pinnatifid or usually at least some trifid (often deeply so), the 3 segments irregularly serrate to incised the 2 laterals smaller than the central and positioned very near the leaf base, sparsely strigose-hispid above, canescent and spreading-hirsute below, especially on the veins, which are rather pale, upper leaves less divided, more or less lanceolate. Spikes elongate, slender, less than 6 mm broad excluding the corollas, dense only at anthesis, rather stiff, loosely panicled at the ends of the branches, short-pedunculate; floral bractlets lanceolate, subulate or acuminate, about as long as the calyx or a little shorter, strigillose and ciliate. Calyx 3 to 4 mm long, strigose-hirsute, the lobes unequal, acuminate-subulate; corolla dark blue to blue-purple, lilac, or violet, tube about equalling or slightly longer than the corolla, rather densely pubescent externally, limb 5 to 9 mm across, pubescent externally on the lower portion, lobes obtuse to retuse. Nutlets reddish-brown, ca. 2 mm long, without a basal cavity, the outer faces reticulate above, striate below, commissural face reaching the top of the nutlet, muricate or muricate-scabrous. Roadsides, railroad embankments, fields, prairies, beaches, etc., usually in dry sandy soil. Throughout much of TX except the Cross Timbers and Prairies and Plains Country; AL, W. to AZ. Mar.-Oct. [V. matthesii Turcz.; V. strigosa Hook.; V. lucaeana Walp.].


9.V. canescensVERBENACEAE VERBENA canescens Kunth in H.B.K. Gray Vervain. Perennial from a woody root; stems usually several from the base, ascending or erect, 4-angled, branched, (10)15 to 46 cm tall, hirsute or canescent. Leaves commonly crowded, mostly sessile and the base semi-amplexicaule or with a subpetiolar base, 1 to 5 cm long, oblong-lanceolate, incised-dentate or subpinnatifid but not trifid, apically acute, margins revolute, upper surface rugose and hirtellous, the hairs commonly with tiny bulbous bases, lower surface canescent-hirsute, with conspicuous veins. Spikes sessile, solitary (or loosely paniculate) at the ends of the branches, in short individuals making up 1/3 to 1/2 the height of the plant, loose or dense, less than 6 mm broad, hirsute or glandular-hirsute; floral bracts broadly ovate to lance-acuminate, variable in length, but often longer than the calyx, ciliate. Calyx 2 to 3 mm long, cylindrical-truncate, the teeth inconspicuous, canescent with spreading hairs, often somewhat glandular; corolla purple to blue or lavender or sometimes pinkish-purple, the tube slightly longer than the calyx, limb 4 to 6 mm across, lobes retuse. Nutlets nearly cylindrical, ca. 2 mm long, the exterior raised-reticulate above and striate at least in the lower 1/2, the commissural face scabrous-muricate, not extending to the top of the nutlet. Sandy or rocky soils, usually where calcareous, in dry hills, ravines, open woods, valleys, and blackland prairies. Trans-Pecos, S. to Cameron Co., E. to Wharton and Waller Cos.; known from Brazos Co., at least formerly; also S. CA and N. Mex. Mar.-Oct. [Includes var. roemeriana (Scheele) Perry].


10.V. brasiliensisVERBENACEAE VERBENA brasiliensis Vell. Brazilian Vervain. Perennial; stems 1 to several from a woody root or crown, 1 to 2.5 m tall, sharply 4 angled, slightly constricted at the nodes, essentially glabrous below, slightly scabrous-pubescent in the upper parts, antrorsely so on the angles. Leaves elliptic or lanceolate (2)4 to 10 cm long, (0.5)0.8 to 2.5 cm broad, acute to acuminate, tapered into a cuneate, subsessile (but not clasping) or petiolar base, margins mostly sharply serrate with spreading teeth, at least above the middle, upper surface strigillose-scabrous with short, often pustule-based hairs, veins impressed, lower surface sparsely pubescent, especially on the raised veins. Spikes less than 6 mm broad, densely flowered, compact at anthesis, but elongating somewhat to ca. 6 cm in fruit, sessile or short-pedunculate, arranged in open cymes, these frequently clustered in loose panicles; floral bractlets lance-subulate, ciliate on the nerves and margins, about as long as the calyces. Calyx 2 to 3.5 mm long or more, the lobes acute and with short subulate tips, all short-pubescent; corollas purple to lilac, not individually showy, tube slightly longer than the calyx, pubescent, limb 2.5(3) mm broad. Sometimes only 2 nutlets developing, brown, 1.5 to 1.9 mm long, externally somewhat rugose, commissural face gray, retrorsely muricate. Roadsides, waste places, coastal prairies, etc., usually in sandy soils and in our area frequently in large stands. Primarily in E. and SE. TX; native to a large portion of S. Amer.; naturalized in the U.S. from VA to FL and the Gulf Coast, W. to MO, OK, and TX; also OR and CA, Jamaica, S. Afr., and elsewhere. May-Oct.


11.V. bonariensisVERBENACEAE VERBENA bonariensis L. South American Vervain, Pretty Vervain. Very similar to V. brasiliensis, above, except the leaves lanceolate, the bases more or less auriculate and amplexicaule (most noticeable on larger leaves); spikes commonly sessile, usually short and little if at all elongating; calyx 3 mm long; corolla tube barely twice as long as the calyx; nutlets trigonous, ca. 2 mm long, predominately striate externally but reticulate toward the tip. Usually in sandy loam soil of flatlands, stream courses, and along rice field fences. E. TX from the Red River to Jefferson Co., W. at least to Robertson Co.; native to Brazil, Parag., Urug., and Argen.; introduced in the U.S. from DA and NJ to VA and FL, W. to OK and TX; also CA, HI, and Puerto Rico. One specimen (TAES 56843) from Newton Co. bears the notation "newly introduced to the Gulf Coast" and is dated 1947. Thus the species has been in TX only about 50 years and may still be increasing its range.


12.V. scabraVERBENACEAE VERBENA scabra Vahl Sandpaper Vervain, Harsh Vervain, White Vervain. Perennial from stolons; stem 1 from the base, 0.3 to 1.5 m tall, hispidulous, simple or usually branched, branches spreading to ascending. Petioles 0.5 to 2 cm long; blades ovate to elongate-ovate, 3 to 13 cm long, to 5 cm broad, acute to somewhat obtuse, basally rounded to nearly truncate or broadly cuneate, margin serrate-dentate for nearly the entire length, upper surface very scabrous and also strigillose, undersurface slightly paler and less scabrous, also with the veins hispidulous. Spikes slender, less than 6 m broad, pedunculate, grouped in rather open panicles, the flowers usually dense, sometimes more remote, spikes elongating and becoming somewhat less dense with age; floral bractlets ovate-acuminate, about half as long as the calyx or less, hispidulous. Calyx hispidulous, in fruit spreading from the rachis at an angle of 45 degrees or more, (1.5)2.5 to 3 mm long, more or less ovoid, the apex acute as the slightly unequal lobes are connivent over the nutlets; corolla blue (to rarely pink or lavender), the tube little if at all exceeding the calyx, limb 2 to 3 mm broad, lobes 1 mm long or less, obtuse; stigmatic surface halfway between 2 nearly equal, obtusish sterile lobes. Nutlets 3-angled, 1 to 1.3 mm long, vaguely striate and reticulate apically, commissural faces reaching the tip of the nutlet, meeting at right angles, muriculate, Usually in rich soil of marshes, swamps, low ground, lakeshores, and stream courses. Throughout much of the S. 1/2 to 2/3 of TX, so probably present in our area; no collections from our area held by TAMU or TAES. NC to FL, W. to AZ and CA; also N. Mex. and W.I. Mar.-Dec.


13.V. urticifoliaVERBENACEAE VERBENA urticifolia L. var. urticifolia White Vervain, Nettle-leaved Vervain. Perennial, usually not stoloniferous; stems 0.5 to 2.5 m tall, simple or more commonly branched at the base, hispid or hirtellous to glabrous. Leaves petiolate; blades broadly lanceolate to oblong-ovate or ovate, 8 to 20 cm long, short-acuminate to acute, basally rounded to broadly cuneate and decurrent on the petiole, coarsely serrate-crenate to doubly so, said to be hirtellous to glabrous above or with scattered pilose hairs below, but our plants definitely scabrous above, the hairs from minute pustules. Spikes slender, less than 6 mm broad, pedunculate, usually rather loosely flowered, 1 to 3 dm long, arranged in loose panicles; floral bractlets ovate-acuminate, 1/2 to 3/4 as long as the calyx, ciliate. Calyx 1.5 to 2 mm long, usually ascending at maturity, pubescent, especially on the veins, lobes subequal, obtuse to subulate, slightly incurved over the nutlets but not prolonged into a point, the overall effect one of bluntness and the nutlets exposed at the tip; corolla white, the tube scarcely longer than the calyx, glabrous or sparsely pubescent below the sinuses, limb 1.5 to 2(4) mm across, the lobes obtuse; stigmatic surface subtended by 1 sterile style lobe. Nutlets ca. 2 mm long, vaguely striate or rugose, commissural faces meeting at right angles, nearly smooth. Low rich or open woods, waste places, bottomlands, streambanks, pastures, and fencerows. E. part of TX, from Bowie and Willbarger Cos. to Newton, Brazoria, and Gonzales Cos; also Wheeler Co. in the Panhandle. Que. and Ont. to NE, S. to FL, W. to TX and OK. Jun.-Oct.



VERBENACEAE PHYLA6. PHYLA Lour. Frog-fruit


Perennial herbs. Main stems creeping or procumbent, branches trailing to ascending, sometimes slightly woody at the base. Herbage nearly glabrate or strigose with more or less cinereous hairs. Leaves opposite, variously shaped, dentate except at the base, flat or pinnately pleated above. Inflorescences 1 to 3 in the axils, usually spike-like, hemispheric or cylindric, elongating in fruit. Flowers small, sessile, each subtended by a small cuneate-obovate or fan-shaped bractlet. Calyx small, membranous, the rim bifid or 4-fid or with 4 teeth. Corolla salverform, tube cylindrical, slightly shorter than the calyx or sometimes longer, limb oblique, slightly 2-lipped, 4-lobed, the lobes often retuse. Stamens 4, didynamous, inserted about the middle of the corolla tube, included or slightly exserted. Ovary 2-locular, each locule 1-ovulate. Fruit small, dry, included within the calyx and sometimes adherent to it, exocarp dry and membranous, inclosing but scarcely distinct from the 2 pyrenes within.

15 species of tropical and warmer regions; 4 in TX; 2 here. Retained by some in Lippia, which differs in having the flowers 4-ranked and the spikes sometimes aggregated into corymbs or panicles.

Though the flowers are small, the clusters can be attractive. Some species, especially P. nodiflora, are grown as ornamental ground covers (Mabberley 1987).


1. Leaves mostly widest above the middle and toothed only near the apex ...1.P.nodiflora

1. Leaves mostly widest at or below the middle and toothed from below the middle to near the apex ...2.P.lanceolata


1.P. nodifloraVERBENACEAE PHYLA nodiflora (L.) Greene Common Frog-fruit, Sawtooth Frog-fruit, Spatulate-leaved Frog-fruit, Turkey-tangle, Weighty Frog-fruit, Capeweed, Matgrass, Hierba de la Virgen Maria, Wedge-leaf Frog-fruit, Texas Frog-fruit, Hoary Frog-fruit. Stems mostly prostrate and rooting at the nodes, to 9 dm long, simple or branched, sometimes tinged with purple, strigillose; branches, if any, decumbent to erect, glabrate to puberulent or strigillose. Petioles obsolete or 1 to 8 mm long and winged; blades often rather thick-textured, quite variable in shape, spatulate to oblanceolate, obovate, elliptic, or cuneiform, 1 to 7.2 cm long, 2 to 25 mm wide, apically rounded to obtuse or subacute, tapered from the middle or above down to the base, margin serrate above the middle or only near the apex, teeth 1 to several per side, spreading or appressed, upper and lower surfaces glabrous to strigillose-puberulent, secondary veins obscure or essentially invisible on both surfaces. Peduncles usually much longer than the subtending leaves, 2 to 11.5 cm long, glabrous to antrorsely appressed-pubescent or strigose; heads initially globose, becoming elongate and cylindric, to 3 cm long and 5 to 10 mm wide; bractlets obovate or slightly rhomboid-cuneate, closely imbricate, ca. 2 to 3 mm long, acute to acuminate or mucronate, sometimes with membranous margins apically, glabrous to finely ciliate or densely strigillose. Corollas white to rosy, sometimes with a yellow eye, tube 2 to 2.5 mm long, limb ca. 2 mm broad, sometimes strigillose externally. Fruit ca. 1 mm long. Lawns, roadsides, pastures, ditches, thickets, riverbanks and bottoms, seashores, etc; often where moist. Throughout TX; PA to FL and KY, W. to OK, CO, NM, AZ, and CA; N. and coastal Mex.; Cen. and S. Amer.; nearly worldwide in tropical and subtropical regions. Mar.-Nov. [Includes var. antillana Moldenke, var. canescens (Kunth) Moldenke, var. longifolia Moldenke, var. repens (Spreng.) Moldenke, var. reptans (Kunth) Moldenke, var. rosea (D. Don) Moldenke, and var. texensis Moldenke; P. incisa Small; Lippia nodiflora (L.) Michx. and vars.; L. canescens Kunth].

This is a highly variable species. Many varieties have been named, but as they intergrade, the current trend is toward combining them all. This is the approach presented by GPFA (1986) and Kartesz (1998) and favored here. Other sources (e.g., Hickman 1993 and Smith 1994) consider this reduction too extreme and recognize a few combined varieties. If such varieties are recognized our plants can usually be assigned to one of the following:


var. nodiflora Leaf teeth usually small, appressed, forward-pointing; leaves generally 2 to 4 times longer than wide; floral bractlets glabrous to ciliate; corolla white or rose. [All of the above synonyms except P. incisa Small].


var. incisa (Small) Moldenke Leaf teeth usually relatively large and spreading; leaves generally 4 to 5 times longer than wide; floral bractlets densely white-strigillose; corolla white with a yellow eye, sometimes tinged purplish in old age. [P. incisa Small].


2.P. lanceolataVERBENACEAE PHYLA lanceolata (Michx.) Greene Northern Frog-fruit, Lance-leaf Frog-fruit. Stems prostrate or procumbent to ascending, (1)2 to 6(10) dm long, simple or sparingly branched, commonly rooting at the nodes, glabrous to white-strigillose, branches sometimes erect. Petioles obsolete or 5 to 10 mm long, strigillose; blades lanceolate to lance-elliptic, oblong, oblong-lanceolate, or ovate, 15 to 75 mm long, 5 to 30 mm broad, widest at or below the middle, acute to subacute, tapered to a cuneate base, serrate from the apex to below the middle, the teeth sharp, both surfaces bright green and somewhat obscurely strigillose, veins conspicuous but flat above, conspicuous and somewhat raised below. Peduncles equalling or usually longer than the subtending leaves, 4 to 9 cm long, sparsely and inconspicuously white-strigillose; heads initially globose, becoming elongate, to 3.5 cm long and 5 to 7 mm across; bractlets imbricate, obovate, 2.7 to 3 mm long, acute, strigillose. Calyx about equalling the corolla tube; corolla pale blue, purplish, or white, 2 to 2.5 mm long. Fruits 0.9 to 1.2 mm long, obovate. Moist soil, marshes, riverbottoms, shores, swales, ditches, low woods, etc. Throughout much of TX, except the Trans-Pecos; Ont. to MN and SD, S. to FL, TX, NM, and CA; also N. Mex. May-Oct. [Lippia lanceolata Michx. and var. recognita Fern. & Grisc.].








LAMIACEAE (LABIATAE)LAMIACEAE

Mint Family


Annual, biennial, or perennial herbs or sometimes small shrubs. Stems typically square in cross-section. Herbage often aromatic. Leaves opposite (rarely basal or whorled), estipulate, simple, often crenate or serrate or occasionally pinnatifid or lobed. Inflorescence of cymules borne in the axils of leaves or bracts, the opposite pairs of cymes forming verticils, verticils one to many, OR inflorescence essentially racemose when the cymules reduced to single flowers. Flowers perfect (rarely unisexual), sometimes with bracteoles subtending the individual flowers. Calyx synsepalous, often at least somewhat 2-lipped, teeth usually 5 (sometimes 10), the 3 upper teeth fused and the lower 2 free, occasionally all the teeth nearly equal or in Scutellaria both lips entire, calyx tube usually enlarged in fruit. Corolla gamopetallous, slightly to strongly 2-lipped, the upper lip of 2 fused petals, erect, usually galeate (hood-like), sometimes short or notched, lower lip of 3 fused petals, usually spreading, the middle lobe sometimes cuplike, occasionally all the petals nearly equal and the corolla almost regular (in Mentha often 4-lobed) or, as in Teucrium, all 5 petals pulled to the bottom of the corolla. Stamens included within the corolla or exserted, usually 4 and didynamous, sometimes only 2, connective between anther cells often enlarged; anther sacs parallel or divergent, in Salvia one cell of each pair rudimentary. Style bifid, in most species gynobasic and arising from below and between the 4 distinct lobes of the ovary, or in some species, notably Trichostema, the lobes of the ovary at least partially united below and the style more nearly terminal. Ovules solitary in each of the 4 cells of the ovary--ovary actually bicarpellate but becoming 4-locular during development. Fruit a cluster of 4 hard-coated cocci or nutlets, sometimes fewer by abortion, usually enclosed by the persistent calyx.

About 3,500 species worldwide in 180 genera. There are 32 genera and about 115 species in TX; 19 genera and 40 species in our area.

The mint family is of great economic importance for many herbs and flavorings--mint, rosemary, thyme, marjoram, oregano, basil, sage, etc.--as well as aromatic and ornamental plants--lavender, bee balm, Salvia, lemon balm, and so on. These plants occasionally escape or remain around old homesites but are not generally persistent members of our flora.

1. Calyx bilabiate, with two entire lips; the upper lip often having a ridge or crest ........................

. ............................................................................................................................1. Scutellaria

1. Calyx regular, OR calyx bilabiate but one or both of the lips not entire, having instead lobes or teeth ............................................................................................................................2

2(1) Stamens with anthers 4 ...........................................................................................................3

2. Stamens with anthers only 2 .................................................................................................16



3(2) Calyx distinctly irregular, deeply divided cleft into two lips, or else appearing one-sided or with very unequal lobes ............................................................................................................4

3. Calyx regular or only slightly irregular, not deeply divided or one-sided (upper and lower teeth sometimes unequal) .......................................................................................................9


4(3) Flowers in racemes or spikes ..................................................................................................5

4. Flowers in whorls, clusters, panicles, or cymes ......................................................................7


5(4) Corolla less than 4 mm long; anther sacs divergent; herbage often purplish .........2. Perilla

5. Corolla longer than 4 mm; anther sacs parallel; herbage green. ..........................................6


6(5) Stem pubescent below the inflorescence; upper corolla lobe bifid and erect; nutlets pubescent ...............................................................................................................3. Brazoria

6. Stem glabrous below the inflorescence; upper corolla lobe entire and hood-like; nutlets glabrous ...............................................................................................................4. Warnockia


7(4) Flowers in panicles or axillary cymes; stamens manifestly arched or coiled; corolla blue, pink, purple, or white .......................................................................................5. Trichostema

7. Flowers in dense, sessile or nearly sessile clusters in the axils of the upper leaves or bracts; stamens not greatly arched or coiled; corolla color various .......................................8


8(7) Corolla violet or white; flowers in 3's, each group subtended by an orbicular bract; flower cluster elongated and spike-like in appearance ....................................................6. Prunella

8. Corolla orange-yellow or red; flower clusters globular .........................................7. Leonotis


9(3) Style not strongly gynobasic, cocci united for about 1/3 their length .............5. Trichostema

9. Style gynobasic, cocci largely separate at maturity ..............................................................10


10(9) Calyx tube obscurely or not at all nerved; calyx slightly or strongly inflated at maturity; anther sacs parallel; inflorescence a spike ....................................................8. Physostegia

10. Calyx tube usually with 5-13 distinct nerves; calyx not inflated at maturity; anther sacs parallel or divergent, if parallel, the inflorescence not a spike .............................................11


11(10) Uppermost leaves or bracts clasping or connate-perfoliate; nutlets trigonous in cross- section ......................................................................................................................9. Lamium

11. Upper leaves petiolate or sessile, but not clasping; nutlets generally terete in cross-section but sometimes trigonous ........................................................................................................12


12(11) Nutlets wrinkled or roughened; flowers in a terminal bracteate spike; flowers apparently with 1 lip (the lower) ............................................................................................10. Teucrium

12. Nutlets smooth; flowers in verticils, heads, axillary cymes or clusters, or non-bracteate spikes; flowers 2-lipped or nearly regular ..............................................................................13


13(12) Plants white woolly-pubescent; calyx teeth hooked apically ..........................11. Marrubium

13. Plants not white-woolly, pubescence various (sometimes whitish-canescent but the hairs very short); calyx teeth not hooked ........................................................................................14


14(13) Flowers axillary or in corymbs, cymes, or heads ....................................12. Pycnanthemum

14. Flowers in a series of verticils, forming a spike-like inflorescence .......................................15


15(14) Stamens of equal length; corolla slightly bilabiate to nearly regular; anther sacs parallel

.................................................................................................................................13. Mentha

15. Stamens didynamous; corolla strongly bilabiate; anther sacs divergent ............14. Stachys


16(2) Each anther with one normal and one rudimentary or reduced sac; flowers in loose, spike- like inflorescences, the flower clusters subtended by minute bracts .....................15. Salvia

16. Each anther each with two normal anther sacs; flowers verticillate or in axillary cymes or clusters; bracts or leaves of the inflorescence usually larger and more obvious ................17


17(16) Plants with a basal rosette of lyrate-pinnatifid leaves ..............................................15. Salvia

17. Plants without the above habit ...............................................................................................18


18(17) Stamens included completely within the corolla tube .....................................16. Rhododon

18. Stamens exserted beyond the corolla tube ...........................................................................19

19(18) Flowers in axillary cymes, though these may be few-flowered and paired at the nodes to esemble verticils; lower calyx teeth longer than the upper ...............................17. Hedeoma

19. Flowers in true verticils; calyx teeth equal .............................................................................20


20(19) Corolla more than 6 mm long .............................................................................18. Monarda

20. Corolla less than 6 mm long ...............................................................................19. Lycopus



LAMIACEAE SCUTELLARIA1. SCUTELLARIA L. Skullcap


Annual or perennial herbs or (some, but not ours, subshrubs), from taproots and/or rhizomes or underground stolons. Stems square, procumbent to erect, simple to branched. Herbage not aromatic. Leaves sessile to petiolate, entire to crenate or serrate. Flowers 1 to 3 in the axils of the foliage leaves or in bracted axillary or terminal racemes; pedicels ca. 2 to 4 mm long. Calyx roughly campanulate, zygomorphic, bilabiate, both lips entire, upper lip slightly galeate and with a concave crest or appendage often longer than the lip itself; calyx accrescent and closing in fruit or sometimes splitting at maturity, often with some purple pigment. Corolla zygomorphic, bilabiate, the tube ascending and curved, the upper lip hooded, entire or slightly notched, lower lip spreading or convex, notched or entire, all blue to purple, rarely white or pink, lower lip often with a whitish patch bearing darker spots. Stamens 4, didynamous, exserted from the throat and ascending under the upper lip; anthers ciliate, those of the upper stamens 2-celled and cordate, those of the lower stamens 1-celled. Style gynobasic, ovary 4-lobed. Cocci yellow to brown or black, spherical to ovoid, ca. 1 to 2 mm in diameter, papillate or tuberculate.

About 300 species worldwide; 15 in TX; 5 here.

The calyx crest (described by some as "tractor-seat shaped") and the entire calyx lips distinguish this genus from all our other native mints. Many species have lovely flowers and would make good additions to the home garden.


1. Flowers in axillary or terminal racemes; flowers subtended by bracts dissimilar to the leaves ........................................................................................................................................2

1. Flowers in the axils of the foliage leaves .................................................................................3


2(1) Midstem leaves oblong to linear-lanceolate, entire, tapered to the base ...1.S.integrifolia

2. Midstem leaves deltoid-ovate, more or less dentate-serrate, basally essentially truncate .....

. ...2.S.ovata


3(1) Plants from rhizomes with bead-like thickenings; cocci with a small wing, skirt, or raised band plus peg-like papillae ...3.S.parvula

3. Plants from taproots; cocci without a wing, skirt, or band; papillae, if any, not peglike .........4


4(3) Major stem leaves deltoid-ovate or -cordate, strongly crenate-dentate; calyx with stiff cilia on the margin and nerves; cocci papillate ...4.S.cardiophylla

4. Major stem leaves ovate, usually only the lowermost crenate, if any; calyx pilose more or less overall; cocci with overlapping platelets ...5.S.drummondii



1.S. integrifoliaLAMIACEAE SCUTELLARIA integrifolia L. Rough Skullcap. Herbaceous perennial, somewhat woody near the base, not stoloniferous; stems 1 to several from the base, often forming clumps, simple to branched, (1.5)3 to 8 dm tall, spreading-pilose below and with incurved hairs above, sometimes also glandular. Leaves 3 to 8 pairs below the inflorescence, lowermost leaves deltoid-ovate to ovate, basally truncate to broadly cuneate, apically acute to obtuse, crenate, 0.7 to 3.5 cm long, 0.2 to 2 cm wide, petioles 0.8 to 2.5 cm long, leaves becoming narrower up the stem, the uppermost leaves lanceolate to narrowly elliptic or oblong, entire to crenate, punctate, 2 to 6 cm long and up to 1.6 cm broad, narrowed to petioles 0 to 10 mm long, all leaves pubescent. Inflorescence of 1 to 5 terminal racemes 3 to 20 cm long, each with 6 to 40 flowers; flowers subtended by linear-lanceolate, leafy bracts, bracts reduced up the axis; pedicels short. Calyx 2.5 to 3.5 mm long in flower, 5 to 8 mm long in fruit, puberulent; corolla blue to violet (rarely white to rose), often lighter below, 1.3 to 2.5 cm long, minutely pubescent, the lips nearly equal. Cocci dark brown to black, subglobose, with flattish papillae that somewhat overlap one another. Moist places along wood edges and clearings. E. TX; N. Eng., OH, KY, and MO, S. to FL and TX. Apr.-June. [Includes var. hisida Benth. and forma rhodantha Fern.].


2.S. ovataLAMIACEAE SCUTELLARIA ovata Hill Eggleaf Skullcap. Perennial with slender rhizomes; stem simple or branched, erect to ascending, 1.8 to 6(8) dm tall, middle and upper stem villous to pilose with capitate glandular or eglandular hairs. Petioles 0.8 to 4(6) cm long, 3/10 to 8/10 the length of the blades; blades thin or slightly thicker, flat to slightly puckered, cordate to ovate, those of midstem leaves (3)5 to 8(12) cm long, (2.5)3.5 to 6(7) cm broad, basally cordate with a deep sinus, margin crenate-serrate, pubescence variously glandular or eglandular, sometimes glabrescent. Flowers in terminal racemes, in the axils of ovate bracts 4 to 9 mm long and 3 to 6 mm wide--much shorter than the leaves, sometimes racemes produced at the uppermost 2 or 3 nodes. Calyx glandular, in flower 3 to 4 mm long, elongating to 6 to 8 mm in fruit, in fruit the crest ca. 4 mm tall; corolla 10 to 23 mm long, blue with a white lower lip, the tube ca. 2 mm broad at the base, flaring to 5 mm broad at the throat, lateral lobes about as long as the upper lip, lower lip ca. 1 cm wide. Nutlets buff to brown, orange, or black, ovoid, the surface covered with conical papillae. Open woods and brushlands. E. and SE TX; MD to TN, W. to KS and TX; also E. Mex. Apr.-Jun.

Nine subspecies, distinguished primarily by pubescence characters, are listed for N. America by Kartesz (1998). Three of these, subsp. ovata, subsp. bracteata (Benth.) Epl., and subsp. mexicana Epl., are listed for TX. However, the species is highly variable and the subspecies intergrade, leaving quite a bit of doubt as to their distinctiveness. In fact, the subspecies are not recognized in some major floras (e.g., Steyermark 1963; Radford, et al. 1968; GPFA 1986; Smith 1994). An older treatment of the variation in this species, involving 3 varieties distinguished by bract and calyx proportions and leaf textures, is used in some manuals (e.g., Steyermark 1963), but is similarly difficult to apply.


3.S. parvulaLAMIACEAE SCUTELLARIA parvula Michx. Small Skullcap. Perennial herb from fibrous roots and spreading moniliform rhizomes, the small, bead-like tubers elliptic to fusiform, whitish, to ca. 1.5 cm in diameter; stems (0.8)1 to 3 dm tall, usually erect, branched at the base or simple, glabrescent to appressed-scabrescent or capitate-pilose; foliage and calyx often with some purple color, especially after drying. Leaves slightly thickened, generally ovate, basally rounded, truncate or cordate, pilose to glabrous, the lateral veins anastomosing before reaching the margins in pilose plants, lowermost leaves ovate to rotund, 7 to 12 mm wide and with petioles 2 to 15 mm long, midstem leaves sessile to short-petiolate, ovate to deltoid-ovate, 8 to 20 mm long and 4 to 15 mm broad, irregularly crenate-subdentate, sometimes glandular. Flowers 2 per node in the upper 1/2 of the plant, in the axils of the foliage leaves; pedicels 3 to 4 mm long, retrorsely hirtellous. Calyx 2 to 3 mm long in flower, (3)4 to 7 mm long in fruit, sometimes with sessile glands, scutellum arising at a 45 degree angle and wider than the calyx; corolla blue to violet, 6 to 11 mm long, the upper lip galeate, lower lip 4 to 7 mm broad, hirtellous in 2 lines. Cocci ovoid, ca. 1 mm in diameter, dull dark brown, covered with conical, rounded tubercles or papillae and with a dentate crest or band on one margin. Sand or clay soils, wood margins, open woods and roadsides, etc. E. and N. Cen. TX; E. 1/2 of the U.S. and Can.--Que. S. to FL and W. to the Great Plains. Mar.-June.

Three varieties occur in TX, all of which seem to be present here.


var. australis Fassett Southern Small Skullcap Stem pubescence of long, glandular-capitate hairs ca. 1/2 as long as stem is wide, plus some short, curved to ascending eglandular hairs; leaves broadly ovate to ovate, those of the midstem 1 1/4 times longer than wide, usually with longish, spreading capitate hairs on both surfaces, especially along veins, lateral veins anastomosing to form 1 submarginal vein, margins little if at all revolute. [S. australis (Fassett) Epl.].


var. leonardii (Epl.) Fern. Leonard's Small Skullcap Stems glabrate to antrorsely appressed-scabrous (sometimes also with glandular-capitate hairs near the base); leaves ovate to lanceolate, those of the mid-stem about 2.2 times longer than wide, lateral veins rarely branching or anastomosed, margins manifestly revolute. [S. leonardii Epl.; S. ambigua Nutt.].


var. parvula Stems densely glandular-pubescent, the hairs about 1/3 as long as the stem is wide, usually also with short, retrorse, eglandular hairs; leaves ovate to narrowly ovate, those of the midstem ca. 1.7 times longer than wide, veins on the lower surface densely eglandular-pubescent, veins sub-anastomosing near the margins, margins revolute; calyx and lower leaf surface with sessile glands.


4.S. cardiophyllaLAMIACEAE SCUTELLARIA cardiophylla Engelm. & Gray Annual from a weakly-defined taproot and many fibrous roots; Stems erect, stiff, 1 to several from the base, 4 to 6 dm tall, branched above the middle, branches strict and ascending, lower internodes often quite elongated, stems and branches with recurved and appressed hairs, especially along the angles, sometimes also with some spreading glandular hairs on the lower portions. . Lowermost leaves early deciduous, midstem leaves with petioles about equalling the blades, blades deltoid, slightly cordate to truncate basally, obtuse to acute apically, blades of larger leaves crenate-dentate, 2 to 5 cm long, leaves becoming smaller and more sessile and entire up the stem, uppermost leaves subsessile and nearly entire, lower leaf surface hispidulous with spreading hairs, upper surface glabrous, margins usually ciliate. Flowers in the axils of upper, reduced leaves; pedicels with pubescence like that of the stem. Calyx ca. 4 to 4.5 mm long at anthesis, enlarging to 5 to 6 mm at maturity, retrorsely pubescent and ciliate along the margins, scutellum concave, ca. 3.5 mm tall; corolla blue-purple, the lower lip often whitish with purple spots, ca. 13 to 22 mm long, pubescent externally, the tube pilose inside, lips glabrous, upper lip galeate. Cocci shallowly papillate, black. Generally in sandy or rocky soils along wood edges and in clearings, damp areas, oil fields, etc. E. 1/4 TX W. to Brazos, Robertson, Freestone, and Kaufman Cos.; also AR, LA(?). Apr.-Oct.


5.S. drummondiiLAMIACEAE SCUTELLARIA drummondii Benth Drummond's Skullcap. Taprooted annual herb 5 to 20(30) cm tall, larger plants branched at the base with branches procumbent at their bases; herbage glandular-pilose, sometimes densely so, especially the younger parts or sometimes (especially the lower portions) also with finer retrorse or spreading eglandular hairs. Lowermost leaves with petioles about equalling the blades, blades ovate and the margins subcrenate, midstem leaves with shorter petioles, blades ovate or less commonly elliptic, the larger ones 1.1 to 1.5(2.4) cm long and 0.6 to 1(1.6) cm wide, entire, rounded-acute apically, uppermost leaves usually entire and subsessile (petioles only 1 to 2 mm long). Flowers 2 per node in the axils of the leaves in the upper 1/2 to 2/3 of the plant; pedicels 2 to 3 mm long. Calyx pilose and more or less glandular, 2 to 4 mm long at flowering, enlarging to ca. 5 mm long in fruit, scutellum rising at a 45 degree angle, ca. 3 mm tall; corolla pubescent externally, violet to blue-violet, 5 to 15 mm long, tube ca. 1 mm in diameter basally, throat pilose, tube more or less glabrous within, upper lip galeate, lower lip, 3 to 5 mm broad, glabrous or with 2 patches of hairs, usually also with a white patch which is purple-spotted. Cocci dark brown to black, ca. 1.3 mm in diameter, ovoid, covered with flattened, overlapping conical papillae and lamellae (especially the lower surface.) Sandy, gravelly, rocky, or clayey soils or on rock ledges, also thickets, brush, etc. Most of TX; also OK, NM, Mex. Feb.-July, occasionally caught flowering in Oct. in our area.

Our plants are probably all var. drummondii. [S. helleri Small].

This charming little plant would probably be wonderful in a rock garden.






LAMIACEAE BRAZORIAE2. BRAZORIAEngelm & Gray Rattlesnake Flowers


Taprooted annual herbs. Stems 1 to several from the base, simple or branched below, pubescent below the inflorescence. Leaves sessile or petiolate, the lower often with winged petioles, typically oblong to oblanceolate, denticulate to nearly undulate or nearly entire, upper leaves oblong to elliptic, usually sessile and slightly clasping. Flowers in spikes or racemes, these upright, terminal or both terminal and axillary. Calyx sparsely to densely pubescent, campanulate and bilabiate, more or less inflated and closed at maturity, two lower lobes with 3 to 5 teeth. Corolla rose, pale pink, purple, or lavender, bilabiate, upper lip bifid, lower lip 3-lobed, the lobes usually notched, throat with an annulus of hairs. Stamens 4, the upper pair longer, each anther with 2 parallel sacs. Cocci 3-angled, with curved hispidulous hairs.

This genus is restricted to 2 species in TX; 1 here. This treatment follows M. W. Turner (1966).


1. B. truncata (Benth.) Engelm. & Gray Rattlesnake Weed. Plants simple or branched below, any branches immediately ascending. Floral bracts mostly 4 to 8 mm long, acuminate to attenuate, ciliate. Corolla 15 to 22(25) mm long, the two upper lobes lavender and the three lower paler lavender, annulus in throat usually perpendicular to axis of corolla. Lower lip of calyx inflexed in fruit. Usually on sandy soils, mostly in localized populations under xeric conditions.

There are two intergrading varieties, both present in our area (Turner 1996). Plants intermediate between the two are known from Robertson Co.


1. Mature inflorescences densely flowered, with lower internodes mostly 5 to 6 mm long; lower lobes of calyx usually woolly-tomentose basally, with hairs to 2 mm long ...........................................................................1a. var. truncata

1. Mature inflorescences loose and interrupted,with lower internodes mostly 8 to 13 mm long; lower lobes of calyx lightly pubescent to canescent, the hairs to only 0.2(1) mm long ........................................................1b. var. pulcherrima


1a. var. truncata Rattlesnake Flowers. Plants usually 20 to 35 cm tall, Lower leaves oblanceolate or spatulate, to 5 to 9(10) cm long, 1 to 2 cm broad, narrowed into winged petioles. Spikes densely flowered, the internodes mostly 5 to 6 mm long; bracts ovate-orbicular, apiculate and with long marginal hairs. Calyx ca. 5 mm long at anthesis, enlarging to 8 to 9 mm at maturity when it becomes flattened on the lower side and gibbous at the base on the upper side, with sparse, scattered glandular hairs, upper lobes mucronate to apiculate, lower lobes usually basally woolly-tomentose or at least densely pubescent, many of the hairs to 2 mm long; upper corolla lobes ca. 9 mm long, the 3 lower ca. 6 mm long; annulus usually 2 to 3 mm above ovary; style 1 to 1.5 mm long. Sandy soils from S. Cen. to E. Cen. TX, N. to Burleson Co., where uncommon. Apr.-May.


1b. var. pulcherrima (Lundell) M. W. Turner Plants mostly 3 to 6 dm tall. Basal leaves withering early, mostly 5 to 9(11) cm long, 1 to 2(3) cm broad, stem leaves usually 5 to 8 pairs below the inflorescence, long-tapering spatulate, to about 13 cm long and 2.5 cm wide (smaller on smaller plants), remotely crenate-denticulate along the distal margin, acute to obtuse, lower leaves petiolate and upper leaves clasping or with winged petioles, all leaves glabrous except for short pubescence at the base on the upper surface. Racemes loose and interrupted, the lower internodes chiefly 8 to 13 mm long, axis pubescent with short glandular and eglandular hairs; bracts to 6 mm long, ovate-lanceolate to lanceolate, acute to acuminate-subulate, conspicuously ciliate along the margins. Upper calyx lobes rounded to apiculate, both upper and lower lobes with scattered eglandular and capitate-glandular hairs, most only ca. 0.2 mm long, occasional hairs to 1 mm long; upper corolla lobes ca. 10 mm long, the lower 3 lobes 7 to 8 mm long; annulus usually 3 to 4(5) mm above the ovary; style 1.5 to 2 mm long. Sandy soils of roadsides and open fields in NE. Cen. TX--Leon, Anderson, Henderson, Freestone, and Houston Cos. Apr.-June. [B. pulcherrima Lundell].



LAMIACEAE WARNOCKIA3. WARNOCKIA M. W. Turner


A monotypic genus once placed as a part of Brazoria (Turner, 1996).


1. W. scutellarioides (Engelm. & Gray) M. W. Turner Brazos Mint, Prairie Brazoria. Annual, stems erect, 15 to 30(75) cm tall, branched, stems and branches below the inflorescence mostly glabrous. Leaves mostly 3 to 5 cm long, 5 to 10 mm broad, sessile, somewhat clasping, oblong to elliptic, rounded to acute, serrate above the middle, teeth often curved, lower leaves sometimes oblanceolate and tapered to a winged petiole. Spikes or racemes to as much as 20 cm long, terminal or axillary and terminal, sometimes arranged in a panicle, somewhat loose, lower internodes 5 to 10 mm long; bracts ovate to lanceolate, acute to acuminate, 3 to 6 mm long, ciliate-margined. Calyx 3 to 6 mm long, with a few capitate-glandular hairs, some eglandular hairs mostly to 0.1 mm long also present, markedly unequally bilabiate, the three upper lobes ovate and apiculate, broader than the two lower, lanceolate to apiculate ones, calyx enlarging to 5 mm in fruit when the upper lip arches backwards and the lower lip arches upwards, nearly closing the throat; corolla red purple to purple, pink, or flesh-color, ca. 0.7 to 1.5 cm long from the base of the tube to the tip of the lower lip, minutely pubescent externally, upper lip entire and nearly galeate, lower lip trifid, the lobes emarginate, throat without an annulus of hairs; stamens 4, didynamous, anthers purplish, margins with small laciniations or teeth; style 1 to 1.5 mm long, bilobed, glabrous, branches linear-lanceolate. Nutlets ca. trigonous, 1.2 to 1.5 mm long, glabrous and somewhat granular. Calcareous or clay soils of W. Cen. TX (one population known from Tyler Co.); in our area from at least Brazos, Burleson, Grimes, and Washington Cos.; also OK, Mex. Mar.-May. [Brazoria scutellarioides Engelm. & Gray].



LAMIACEAE PERILLA4. PERILLA L.


Six Asian species with one naturalized in TX.


1. P. frutescens (L.) Britt. Common Perilla, Beefsteak Plant. Annual; stems 2 to 8 dm tall, erect, branching, pilose or retrorsely pubescent to glabrous; herbage aromatic, often purple or purple-tinged. Leaves ovate or sometimes to suborbicular (rarely elliptic), 5 to 13(15) cm long, ca. 2 to 8 cm wide, glabrous or sparsely pubescent, primarily on the veins, margins serrate or crenate or incised, apices acute or abruptly short-acuminate, basally cuneate or attenuate to the petiole (less commonly rounded or nearly truncate); petioles 1 to 7(8) cm long, usually sparsely pubescent. Flowers solitary in the axils of small bracts, forming 1-sided, spikelike racemes to ca. 15 cm long; peduncles 1 to 5 cm long; bracts 3 to 6 mm long, ovate to elliptic, usually folded, generally entire, acute to acuminate apically and rounded at the base, sessile or nearly so; pedicels 1 to 3 mm long. Calyx campanulate, 10-nerved, bilabiate, 2 to 3 mm long at anthesis, 7 to 12 mm long and with the tube slightly inflated on the lower side at maturity, villous externally, sparsely villous within, upper lip erect, with 3 triangular to elliptic teeth united for about 1/3 their length, the 2 teeth of the lower lip narrowly triangular, about equalling the upper lip; corolla white to lavender, tubular, 2.5 to 3.5(6) mm long, slightly longer than the calyx, slightly zygomorphic, the two lips about equal, the 5 lobes broadly rounded, sometimes pubescent externally and the lower lobes bearded within; stamens 4, about equal, about as long as the corolla, straight, exserted from the corolla, not connivent. Cocci globose, 1.2 to 2 mm in diameter, reddish brown or sometimes darker, reticulate. Damp, wooded areas, along woodland streams, and in seepage areas. E. TX; native to India; naturalized from FL to LA and TX, and from N. Eng. and NY to OH, IN, MO, and KS. Some sources recognize varieties.

The seeds yield an edible oil that is also used in paints and inks; also used for waterproofing paper. Cultivated in India and Asia; sometimes weedy where naturalized (Mabberley 1987).



LAMIACEAE TRICHOSTEMA5. TRICHOSTEMA L. Blue Curls


Taprooted annual (ours) or perennial herbs. Stems well-branched, branches opposite, obscurely angled, variously pubescent (often stipitate-glandular). Leaves petiolate to subsessile, linear to elliptic-ovate, entire or toothed. Inflorescence a panicle of paired, bracteate racemes or helicoid cymes, with 1 flower in the axis between each pair, occasionally the entire inflorescence racemose (usually not ours); floral bracts similar to the leaves though smaller. Calyx regular or irregular, if zygomorphic then the upper "lip" with 3 more or less equal teeth joined for about 2/3 of their length, the lower lip generally shorter than the upper and 2-toothed; at maturity (after flowering) the flowers often resupinating and the lower lip appearing as the upper. Corolla blue-violet, in some species rose-pink or white, 5-lobed, with the lowermost lobe lip-like and up to 1.5 times the length of the others 4 which are subequal, united, and somewhat galeate or at least arched, corolla pubescent externally. Stamens 4, inserted near the throat, ascending under the upper lip, straight to curved or coiled, anther sacs divergent at maturity. Style not strongly gynobasic, generally curved in the same way as the stamens, upper stigma lobe often somewhat shorter than the lower. Cocci obovoid, united for ca. 1/3 their length, rugose to reticulate or alveolate, dull, olive or brown to blackish.

About 15 species in N. America; 4 in TX; 2 here (but one sometimes given its own genus.)


1. Calyx definitely irregular, the lower teeth shorter and the flower turned upside down (resupinate) ...1.T.dichotomum

1. Calyx nearly regular, the teeth about equal and the flowers not resupinate ............................

...2.T.brachiatum


1.T. dichotomumLAMIACEAE TRICHOSTEMA dichotomum L. Forked Blue Curls, Bastard Pennyroyal. Annual; stems usually branched, to 1 m tall but often less, at least the main stems pubescent below the inflorescence, the hairs short and spreading or downward-curled, some longer, pilose hairs often present, especially at the nodes, occasionally also with capitate glands, rarely only pilose. Petioles 0 to 15 mm long; blades oblong to ovate or elliptic or elliptic-lanceolate, 2 to 7 cm long and 0.4 to 2.5 cm wide, entire or crenate to rarely lobed, acute to obtuse, basally cuneate, leaf surface with short hairs curled toward the apex. Peduncles to 5 cm long, pedicels to ca. 5 mm long, both with stalked, capitate glands, usually also with short, spreading hairs, also with or without longer hairs 2 to 3 times the length of the glandular hairs; bracts of the inflorescence often leaf-like. Calyx strongly zygomorphic, 2.7 to 6 mm long, enlarging to 4.6 to 8.9 mm in fruit and becoming inverted so that the 2 small teeth are uppermost, the upper 3 teeth fused for 1/2 to 2/3 their length, about 3 times as long as the lower 2 teeth; corolla falling easily and often absent in herbarium specimens, bluish, the tube not prolonged beyond the lower calyx teeth, lower lip to 7 mm long, commonly spotted; stamens exserted, 6 to 16 mm long, arched or more commonly coiled with the style through about 1 full turn. Cocci 1.5 to 2.9 mm long, glabrous, distinctly alveolate or reticulate. Sandy soils, open woods, stream banks, etc. E. TX; N. Eng. S. to NC, SC, and FL, W. to MN, KS, OK, and TX. July-Oct.

Noteworthy both for its fall-flowering and its charming flowers with watch-spring stamens, this plant would be a good choice for the shade garden.


2.T. brachiatumLAMIACEAE TRICHOSTEMA brachiatum L. Flux-weed, False Pennyroyal. Annual; stems 1.5 to 5 dm tall, simple below, branched in the upper 1/2 to 2/3 and the lower stem commonly leafless, pubescence below the inflorescence of spreading or recurved hairs, usually also pilose and occasionally also stipitate-glandular. Leaves sessile or with poorly defined petioles to 1 cm long; blades elliptic to lanceolate or linear-elliptic, 1 to 4(5) cm long, 3 to 10(12) mm broad, acute, entire or irregularly serrate or weakly lobed, strongly 3-nerved, with stipitate or sessile glands, pubescence of short hairs curled toward the apex. Flowers usually solitary in reduced axillary cymes, sometimes 2 or 3 per cyme; peduncle usually less than 2 cm long; pedicels 1 to 15 mm long; floral bractlets linear-elliptic, 1 to 4 mm long. Calyx campanulate, to as much as 8 mm long over all, tube shorter than the teeth, variously pubescent, regular or nearly so, obscurely 10-nerved, the teeth triangular-lanceolate, 1.5 to 3.7 mm long in flower, 2.5 to 4.9 mm long in fruit; corolla bluish or rosy, bilabiate, the tube straight, scarcely exserted from the calyx, 1.6 to 3.5 mm long, slightly expanded in the throat, lower lip ligulate and deflexed, 1.3 to 2.1 mm long, slightly longer than the upper 4 teeth which are spreading or ascending; stamens straight to slightly curved, only slightly exserted, 2.3 to 4.7 mm long. Cocci ovoid, 2 to 3 mm long, sparsely pubescent at least apically with short curled hairs, reticulate with prominent ridges, and commonly with a few stipitate glands. Sandy or limey soils. Primarily on the Ed. Plat.; known from calcareous outcrops in Grimes Co.; NY to MN and SE. SD, S. to FL and TX, also AZ. July-Oct. Treated by Kartesz (1998) as Isanthus brachiatus (L.) B.S.P.



LAMIACEAE PRUNELLA6. PRUNELLA L. Self-heal, Heal-all


About 7 species found worldwide. We have the one found in TX.


1. P. vulgaris L. Common Self-heal, Heal-all, Carpenter Weed. Perennial herb from a short caudex or slender rhizome; stems 1 to several from the base, erect, ascending, or decumbent, often clustered, glabrous to pilose or villous, 1 to ca. 8 dm tall. A tuft of basal leaves commonly present, leaves ovate to ovate-lanceolate or elliptic, or the middle and lower leaves elliptic-oblong, 3 to 7(10) cm long and (0.5)1 to 3(4) cm wide, glabrous to lightly villous, acute to obtuse or sometimes rounded apically, basally cuneate to rounded, crenate to serrate (sometimes remotely or irregularly so), leaves just below the inflorescence often very narrow and nearly entire; petioles 0.5 to 4 cm long, those of the lower leaves the longest, reduced upwards and the upper leaves often sessile. Inflorescence spikelike in appearance, flowers in small cymules of 3, each cluster subtended by a bract; bracts orbicular to reniform, abruptly cuspidate, glabrous or villous-hirsute on the veins, markedly ciliate on the margins, 5 to 15 mm long and 7 to 14 mm wide, often edged or tinged with purple; bracteoles beneath each flower none or, if present, minute and subulate. Calyx sometimes purple, bilabiate, 10-nerved, 6 to 15 mm long, glabrous or villous externally, the throat naked and closed in fruit, tube 2.5 to 3.5 mm long, upper lip truncate or with 3 short teeth or mucronate lobes united for 3/4 or more of their length, the middle lobe longer than the laterals, lower lip 2-lobed, the lobes united for about 1/3 their length, lanceolate to narrowly deltoid, ciliate, about equalling the upper lip; corolla 1 to 2.3 cm long, violet (rarely white), glabrous to sparsely villous externally, glabrous or with a ring of hairs within, bilabiate, tube 7 to 12 mm long, upper lip arched, galeate, usually unlobed, lower lip spreading or deflexed, shorter than the upper, 3-lobed, the 2 lateral lobes entire and the median one erose, fringed, or ciliate, often paler than the rest of the corolla; stamens 4, didynamous, the lower pair the longer, ascending under the upper lip, filaments 2-pronged at apex, one prong bearing the anther and the other sterile, anther cells divergent; style with 2 equal lobes. Cocci ovoid, smooth or ribbed or carunculate, lustrous, yellow-brown to dark brown, ca. 1.7 to 2.2 mm long. Open woods, low meadows, roadsides, pastures, etc. E. 1/4 TX; native to Eurasia and widespread in N. America, occurring in all regions of the U.S. and Canada where there is sufficient moisture. Apr.-Jun. in TX.

Those who recognize subspecific taxa (some recent authors do not) list 2 for TX: subsp. vulgaris (includes var. hispida Benth.), with stems and lower leaf surfaces usually densely pubescent, and subsp. lanceolata (= var. lanceolata (Benth.) Fern.) with narrower leaves than the typical form. The vast majority of our local material seems to belong to the latter at whatever level it is recognized.

This plant has been used in teas for bathing cuts and bruises and for treating sore throats (Mabberley 1987).



LAMIACEAE LEONOTIS7. LEONOTIS L.


About 40 species native to Africa. At least 2 species have become naturalized in N. America. We have the 1 found in TX.


1. L. nepetifolia (L.) Ait. f. Lion's Ears, (Poodle Mint). Annual herb 0.5 to 2 m tall (sometimes more); stems simple or freely branched, soft-pubescent to canescent. Leaves ovate to ovate-deltoid, 5 to 12 cm long and 4 to 10 cm wide, obtuse-apiculate, basally truncate to cordate, sometimes oblique, coarsely crenate (occasionally crenate-dentate), minutely pubescent on both surfaces; petioles 3 to 7 cm long. Inflorescence of 1 to 3, globose, headlike clusters completely encircling the stem, each cluster 4 to 6 cm in diameter, many-flowered, and subtended by two narrow, reflexed leaves, 2 small leaves usually at the top of the stem; flowers opening from the top of each cluster down, so that the upper flowers of the smaller, upper clusters may be open before the lower flowers of the bottom, oldest cluster; pedicels 1 to 2 mm long. Calyx with curled pubescence, 1.5 to 2 cm long, bilabiate, the tube 10-nerved, the lobes 7 to 10, all of different lengths and each aristate or spine-tipped, the upper most lobe much the largest and somewhat hooded; corolla orange-yellow to scarlet (due mostly to the colored, dense pubescence), 2 to 2.5 cm long overall, bilabiate, the tube dilated above and curved, the upper lip erect to galeate, 0.8 to 1.2 cm long, lower lip much shorter than the upper, 3 to 4 mm long, 3-lobed; stamens 4, didynamous, exserted beyond the margin of the upper lip, filaments minutely pubescent; stigma 2-lobed with 1 lobe much longer than the other. Cocci 3-angled, oblanceolate, 3 to 3.3 mm long, depressed-truncate and glandular at the apex, brown. Waste places, roadsides, and cultivated ground. E. and Cen. TX; TN to NC, GA, FL, and TX, W. to MS and AL; native to S. Africa. Jun.-Sept. [Sometimes spelled nepetaefolia].

Not very common in our area, possibly only occasionally escaping cultivation, but quite unmistakable when found. The dry stems with the round balls of empty calyces are interesting in arrangements.



LAMIACEAE PHYSOSTEGIA8. PHYSOSTEGIA Benth. False Dragonhead, Obedient Plant


Perennial herbs from rhizomes, these simple or branched, vertical or horizontal; stems to ca. 2 m tall, erect, generally unbranched below the inflorescence; pubescence, if present, generally only in the inflorescence. Leaves of overwintering rosette often deciduous prior to anthesis, cauline leaves sessile to petiolate (petioles to 6.5 cm), often the upper leaves clasping, blades glabrous, linear to oblanceolate, oblong, or ovate, margins entire to sharply serrate. Inflorescence of 1 to 20 racemes, the axes puberulent to tomentose or glabrous near the base, sometimes also with stalked glands. Flowers sessile or the lowermost short-pediceled, solitary in the axils of small bracts. Calyx regular (or very nearly so), campanulate to campanulate-tubular, with 10 very obscure nerves, externally puberulent to pubescent (rarely sub-glabrous), often also glandular-punctate or stipitate-glandular, interior surface glabrous or with stalked glands, teeth or lobes 5, erect, shorter than the tube, deltoid to lanceolate, acuminate to cuspidate. Corolla much longer than the calyx, bilabiate, tubular below and widening upward, exterior pubescent or tomentose to glabrous, interior primarily glabrous to subglabrous, except near the attachment of the stamens, upper lip erect to slightly galeate or horizontal to spreading, entire to slightly emarginate, lower lip about equalling the upper, 3-lobed, the lobes divergent to reflexed, the median lobe larger than the laterals, color ranging from pure white to deep lavender, usually with darker markings within. Stamens 4, didynamous, the outer pair the longer, ascending under the upper lip parallel to each other or sometimes divergent, exserted from the corolla tube but equalling the upper lip, filaments somewhat coherent by long, tangled hairs; anthers purple to white, glabrous to pubescent and with a few glands on the abaxial surface, sacs parallel, dorsifixed. Ovary deeply 4-lobed, with a single nectary adjacent to two of the lobes and larger than all the lobes; style equalling stamens, stigma lobes equal or subequal. Cocci 1.7 to 4.2 mm long, trigonous, usually smooth.

Species of Physostegia are difficult to circumscribe and cannot be identified by single characters, rather, each has a suite of characters which separates it from other species. Important characters include corolla color, rhizome type (vertical vs. horizontal), leaf margin, leaf base, and the presence or absence of glandular hairs. These characters are much more obvious in fresh material and each seems to have arisen and disappeared, evolving and reverting many times in the history of the genus--hence shared characters do not necessarily reflect interspecific relationships. All of this is complicated by the fact that not all individuals possess all the characters "typical" of the species. Fortunately, Cantino published an excellent monograph (1982) with detailed keys, descriptions, and maps. The following key and descriptions are based on this monograph and on species annotated by Dr. Cantino. NOTE: that corolla measurements are based on herbarium sheets; flowers in fresh material will be a little longer.

Most species have, at one time or another, been placed in Dracocephalum.

According to Cantino, there are 12 species in N. America from Can. to Mex.; 7 in TX; 5 quite likely here.

The common name Obedient Plant stems from the fact that the flowers, if moved laterally in the inflorescence, will "obediently" stay where placed. All have ornamental value and some are grown in gardens--escapes make identifications even more difficult.


1. Calyx and rachis of inflorescence with stalked glands as well as non- glandular

puberulence ...1.P.digitalis

1. Calyx and rachis puberulent but without glands OR if with glands then up to 40 pairs of sterile bractlets present at the base of the raceme .................................................................2


2(1) Leaves sessile or petiolate, but not clasping ...........................................................................3

2. Leaves, or at least one pair, clasping the stem .......................................................................4


3(2) Plants flowering April to June; hairs of raceme axis 0.13 to 0.2 mm long (use 60x

magnification) ...2.P.angustifolia

3. Plants flowering June to October; hairs of raceme axis rarely even 0.1 mm long (use 60x magnification) ...3.P.virginiana

ssp. praemorsa


4(2) Rootstock with one to many elongate, horizontal secondary rhizomes, each terminated by a perennating bud ...4.P.intermedia

4. Rootstock bearing perennating buds or buds on the ends of short vertical rhizomes; horizontal rhizomes absent ......................................................................................................5


5(4) Most or all of the larger leaves sharply serrate; largest leaves on dry specimens not wider than 2.5 cm (slightly wider in fresh material) and rarely less than 5 times longer than wide ...........................................................................................................................................6

5. Most or all of the larger leaves bluntly toothed to entire or largest leaves more than 3 cm wide or less than 3 times longer than wide .............................................................................7


6(5) Corolla usually very pale lavender to white; stem usually with 11 to 20 nodes below the inflorescence; petiolate lower stem leaves usually deciduous before anthesis ......................

...2.P.angustifolia

6. Corolla deep lavender to red-violet; stems usually with 7 to 10 nodes below the

inflorescence; petiolate lower stem leaves frequently (but not always) present at and after anthesis ...5.P.pulchella


7(5) Largest leaves primarily on the lower 1/3 of the stem, those of the upper 2/3 sharply serrate ...5.P.pulchella

7. Largest leaves spread more evenly over the stem, or if mainly on the lower 1/3 of the stem, then leaves of the upper 2/3 bluntly toothed to entire ...1.P.digitalis


NOTE: P. correllii (Lundell) Shinners Occurs to our east and west, but no specimens seen from our area. Widespread but rare from S. LA to NE. Mex. Charcters include extensive horizontal rhizomes, midstem leaves at least partially clasping, usually sharply serrate, upper stem leaves not much reduced, inflorescence and calyces usually with a few stalked glands, and calyx conspicuously glandular-punctate. River bottoms and ditches. Flowering from the third week of June to the end of September. [D. corellii Lundell].


1.P. digitalisLAMIACEAE PHYSOSTEGIA digitalis Small Rhizomes strictly vertical, usually unbranched, to 10 cm long; stem to 2 m, with 9 to 13(16) nodes below the inflorescence. Lower and midstem leaves sessile or the lowermost 1 to 4 pairs petiolate, these petiolate leaves early deciduous, sessile leaves commonly with 1 to 3 pairs of weak veins (in addition to the midvein) arising from the base of the blade, 5 to 17 cm long and 1.5 to 7 cm broad, obovate to elliptic or broadly oblanceolate, less commonly ovate (these broad shapes unique among the species in our area), base rounded to cuneate, commonly somewhat auriculate and clasping, apex obtuse (to acute), becoming acute or less commonly acuminate on the upper leaves, margins subentire to repand or bluntly toothed (rarely some leaves sharply serrate), uppermost leaves much smaller than those of the midstem--leaves of the second pair below the terminal raceme 1 to 3(6) cm long and less than 1/4 as long as to equal to or longer than the internode immediately above, oblanceolate to lanceolate, sessile, basally rounded or sometimes clasping slightly, attenuate, sharply serrate (sometimes bluntly toothed or entire). Racemes 1 to 11, the axes pubescent to tomentose, the hairs mostly 0.2 to 0.3 mm long, and with a few stalked glands sometimes present; floral bracts lanceolate to ovate, attenuate, 3 to 9 mm long, 1.5 to 4 mm wide; flowers 25 to 41 mm long, closely spaced, calyces of adjacent flowers overlapping by half or more their length at anthesis. Calyx occasionally with stalked glands but never conspicuously glandular-punctate, tube at anthesis 4 to 8.5 mm long, lobes primarily attenuate to cuspidate, 1.5 to 4 mm long, calyx in fruit 7 to 13 mm long; corolla pale lavender to whitish, the inside usually spotted with purple, glabrous or nearly so, sometimes sparsely puberulent. Cocci 2 to 3(3.3) mm long, trigonous, the sides concave and smooth. Moist open pinewoods, roadsides. W. LA and E. TX. Not usually seen from our area but probably present where the Pineywoods extend into Grimes Co. Mid. Jun.-late July. [P. obovata of some authors.]


2.P. angustifoliaLAMIACEAE PHYSOSTEGIA angustifolia Fern. Stems 3 to 15 dm tall; rhizomes vertical and unbranched, 2 to 4(10) cm long (rarely branching to produce elongate horizontal rhizomes). Leaves ascending, nearly appressed, lower and midstem leaves sessile or lowermost 1 to 4(7) pairs petiolate, petiolate leaves usually early deciduous; sessile leaves 3 to 21 cm long, 0.3 to 2(2.7) cm wide, lanceolate to oblanceolate, occasionally elliptic to linear, basally cuneate to narrowly truncate, often somewhat auriculate and usually at least slightly clasping, apically acute to attenuate (the lowermost leaves occasionally obtuse), margins sharply serrate, rarely bluntly toothed to entire, leaves of uppermost stem usually much smaller than the midstem leaves, those of the second pair below the terminal raceme (0.7)1 to 5(6) cm long and 1/8 to 2/3 as long as the internode immediately above (rarely longer than the internode), similar to the midstem leaves but generally entire or only remotely serrate. Racemes 1 to 5(8), axes densely pubescent, trichomes 0.13 to 0.2 mm long, some trichomes often to 0.25 mm long; floral bracts lanceolate, attenuate, generally 3 to 6 mm long and 1 to 2.5 mm wide, the lowermost pair sometimes to 12 mm long; flowers (18)22 to 33(36) mm long, closely spaced, at anthesis adjacent calyces overlapping 1/4 to 1/2 or more their length. Calyx glandular-punctate or not, stalked glands absent, tube at anthesis (3.5) 4 to 7 mm long, lobes acute to cuspidate, (1)1.5 to 2.5(3) mm long, calyx in fruit (6.5)7 to 10(10.5) mm long; corolla pale lavender to white (rarely brighter), spotted and sometimes also streaked inside with purple, puberulent to tomentulose. Cocci 2 to 3(3.5) mm long, trigonous, sides concave to slightly convex, smooth. Moist prairies, meadows, ditches, marshes, and river bottoms. SW. GA and AL to Cen. TX, N. to TN, MO, and KS. Early Apr.-3rd week of July (rarely late July.) [P. edwardsiana Shinners, D. virginianum (L.) Benth v. album Nutt.].

Most easily confused with P. pulchella and P. virginiana subsp. praemorsa. It can be told from the latter by its longer (0.13 to 0.2 mm) trichomes; those of ssp. praemorsa are rarely even 0.1 mm long--use 60x lens.


3.P. virginianaLAMIACEAE PHYSOSTEGIA virginiana (L.) Benth. subsp. praemorsa (Shinners) Cantino Late Lionsheart. Stems to 1.8 m tall; rhizome short, usually strictly vertical and the perennating bud on the rootstock or at the tip of the rhizome, only infrequently with horizontal rhizomes to 20 cm long. Lower and midstem leaves sessile but not clasping, or the lower 1 to 7 pairs petiolate, but petiolate leaves usually early-deciduous, midstem leaves 2 to 18 cm long, elliptic to oblanceolate, obovate, ovate, spatulate, lanceolate, or linear, 0.2 to 3(4.4) cm broad, margins sharply serrate, apex usually acute to attenuate, occasionally obtuse, upper stem leaves slightly to strongly reduced, those of the second pair below the terminal raceme 0.8 to 11 cm long and 1/3 to 4 times as long as the internode directly above. Racemes 1 to 8, axes usually pubescent with eglandular hairs mostly not longer than 0.1 mm (rarely as long as 0.13 mm); floral bractlets lanceolate to ovate, attenuate, usually 2 to 8 mm long and 1 to 2.5 m broad; sterile bractlets (up to 40 pair) commonly present below the flowers; flowers (13)14 to 17 mm long, usually densely arranged, the calyces overlapping 1/4 to 1/2 their length. Calyx at anthesis (3.5)4 to 8 mm long, in fruit 6 to 10(11) mm long; corolla red-violet or paler and lavender-white, usually marked inside with purple, variously puberulent to tomentulose or glabrous. Cocci (2.5)2.9 to 3.8 mm long, trigonous, with concave sides, smooth. In a wide variety of habitats, including prairies, limestone areas, calcareous outcrops, ditches, stream margins, and so on; in our area known from calcareous sandstone outcrops in Grimes Co. W. VA and Cen. NC, W. to IL, MO, OK, and TX; scattered in W. TX, SE. NM; one record each from SE. NE and SE. MN. July-early Oct., rarely as late as mid-Dec. The latest-flowering of all our species. [P. virginiana (L.) Benth. forma candida Benke and var. reducta Boivin; P. praemorsa Shinners; P. variegata (Vent.) Benth.; P. serotina Shinners; D. variegatum Vent.]

Most easily confused with P. angustifolia but much later flowering where the ranges overlap and with shorter hairs on the raceme axis.

The species as a whole occurs from Que. to FL, W. to Man. and TX, with some scattered populations located in MT, NM, and N. Mex. Several cultivated varieties are known, but all known free-living populations in our area are wild-type subsp. praemorsa.


4.P. intermediaLAMIACEAE PHYSOSTEGIA intermedia (Nutt.) Engelm. & Gray Primary rhizome branched, yielding 1 to many elongate horizontal rhizomes to 40 cm long, these sometimes branched; stems to 1.2 m tall, often conspicuously enlarged or swollen at the base. Lowermost 3 to 8 pairs of leaves with petioles to 6 cm long, these usually deciduous prior to anthesis; blades of lower and midstem leaves 3 to 14 cm long and 0.3 to 1.5 cm wide, all lanceolate or some oblanceolate to narrowly elliptic, bases attenuate to cuneate on the lower leaves, becoming rounded-auriculate upwards, with at least a few clasping, apically acute to attenuate, margins entire to repand or with a few widely spaced blunt teeth, upper leaves little to only moderately smaller than the midstem leaves (those of the second pair below the terminal raceme 1.7 to 9(12) cm long and 1/3 as long to nearly as long as the internode immediately above, lanceolate, attenuate, auriculate, clasping, widest at or near the base. Racemes 1 to 5(10), axes densely puberulent to pubescent throughout or only sparsely so or glabrate near the base, hairs up to 0.15 mm long, rarely to 0.2 mm; floral bracts lanceolate or less often ovate, attenuate, (1.5)2 to 5(6) mm long, 1 to 1.5(2.5) mm wide; flowers generally 9 to 19 mm long (shorter if anthers aborted, which is common in our material), adjacent calyces at anthesis not overlapping. Calyx not conspicuously glandular-punctate and without stalked glands, tube at anthesis (2.2)2 to 4 mm long, lobes acute (rarely a few cuspidate), (0.7)1 to 2(2.8) mm long, calyx in fruit (3)4 to 7 mm long; corolla lavender, streaked and spotted inside with purple, puberulent to tomentulose. Cocci 2 to 2.5 (2.9) mm long, trigonous, the sides slightly convex and smooth. Swamps, ditches, marshes, etc. SE. MO, S. to OK, AR, LA, and TX; also reported from KY and IL, perhaps mistakenly. Late Mar.-late July. [D. intermedium Nutt.; P. micrantha Lundell].

See NOTE #2 at P. pulchella, below.


5.P. pulchellaLAMIACEAE PHYSOSTEGIA pulchella Lundell Rhizome strictly vertical, to 6 cm long, unbranched or with a few branches; stems to 14 dm tall, with 7 to 10(12) nodes below the inflorescence. Lowermost 1 to 4 pairs of leaves with petioles to 6.5 cm long, some of these usually present at anthesis, blades 2.5 to 11 cm long and 0.6 to 1.7 cm wide, narrowly elliptic to lanceolate or oblanceolate, basally cuneate to attenuate, apically obtuse, entire to repand or remotely crenate or dentate, sessile leaves of lower and midstem often with 1 to 3 pairs of weak primary veins (in addition to the midvein) arising from the base of the leaf, 3 to 15 cm long, 0.5 to 2 cm wide, lanceolate to oblanceolate or spatulate, bases usually auriculate and clasping, apex acute to obtuse, margins usually sharply serrate, upper leaves scarcely smaller to much smaller than the midstem leaves (those of the second pair below the terminal raceme 1.8 to 6.5(10) cm long and 1/3 as long to equalling the internode immediately above), usually with 1 to 3 pairs of weak veins arising from the base, lanceolate or less commonly elliptic or oblanceolate, bases auriculate and clasping, apices attenuate, margins sharply serrate, uppermost pair of leaves sometimes serrate only at the base or even entire. Racemes 1 to 3(7), the axes densely pubescent at the apex and sparsely pubescent to glabrous at the base, some (usually many) hairs 0.13 to 0.2 mm long and often to 0.25 mm long; floral bracts lance-ovate, attenuate, 2 to 5(6) mm long, 1 to 2(3) mm wide; flowers 16 to 30 mm long, loosely or closely spaced. Calyx not conspicuously glandular-punctate and without stalked glands, tube at anthesis (3)4 to 6 mm long, lobes acute to acuminate (sometimes a few cuspidate), 1 to 2.2(2.5) mm long, calyx in fruit 6 to 9 mm long; corolla deep lavender to red-violet (very rarely paler), the interior spotted or streaked with purple, puberulent, occasionally tomentulose or glabrous. Cocci 2.2 to 3 mm long, trigonous, the sides concave to slightly convex, smooth. Moist river bottoms, ditches, meadows. E. TX from OK to the Gulf. Early Apr.-early June (rarely to late June.)


NOTE 1: Similar to and easily confused with P. angustifolia; most readily distinguished by the deeper corolla color. Also, P. pulchella has fewer nodes below the inflorescence and tends to be ending its blooming season when P. angustifolia is beginning (mid-May.) P. pulchella has a tendency to retain its lower petiolate leaves longer--they are often seen on herbarium sheets.

NOTE 2: Species of Physostegia are generally reproductively isolated by blooming period (among other factors) and seldom hybridize in the wild. However, Dr. Cantino has identified a plant from our area that seems to be a cross between P. pulchella and P. angustifolia.



LAMIACEAE LAMIUM9. LAMIUM L. Dead-nettle, Henbit


Annual, winter annual, or perennial herbs from short taproots. Stems usually branched at the base and rooting at the lower nodes, decumbent at the base, the flowering portions erect. Leaves roughly cordate, lobed, toothed, or incised or merely crenate to serrate, petiolate. Flowers in cymules in the axils of leafy bracts, the clusters distinct on the lower stem and congested in the upper region. Calyx tubular to campanulate, 5-nerved and -toothed, more or less regular, teeth equal or the upper ones longer. Corolla bilabiate, the tube longer than the calyx, dilated at the throat, upper lip galeate, erect, entire to emarginate, ovate to oblong, lower lip 3-lobed, the middle lobe constricted at the base and emarginate or 2-lobed, the lateral lobes reduced or represented by convex undulations of the margin of the throat. Stamens 4, didynamous, ascending under the upper lip, the outermost pair seated lower in the corolla and longer than the other two; anther sacs divergent, pubescent. Style equally 2-lobed. Ovary 4-parted. Cocci trigonous, smooth or tuberculate.

About 40 species native to the Old World, with several naturalized in N. America; 2 in TX; 1 here. L. purpureum is found just to our east and has reddish flowers--it may someday be found here.

The common name refers to the foliage, which is nettle-like but is "dead," ie., does not sting.


1. L. amplexicaule L. Henbit, Dead-nettle. Annual, winter annual, or biennial; stems to 45 cm tall, branched at the base or from the lower axils, decumbent to erect or flexuous above, internodes reduced upwards, sparsely pubescent or appressed-hirsute. Leaf blades broadly ovate to reniform or orbicular, sometimes shallowly 3-lobed, 5 to 35 mm long and nearly as wide, basally truncate to cordate or widely cuneate, apically obtuse to rounded, crenate, variously pubescent, lower leaves with petioles to 3.5 cm long, petioles reduced upwards. Flowers (3)6 to 10 in cymules borne in the axils of foliose bracts; bracts similar to the leaves though seldom lobed, sessile and clasping or connate-perfoliate, ascending to spreading; verticils of flowers distinct or the uppermost crowded. Calyx densely pilose or villous, tubular, 4 to 7 mm long, teeth narrowly triangular, about equalling the tube; corolla red-violet, pubescent to pilose externally, especially the upper lip, glabrous within, 10 to 20 mm long, the tube slender and straight, 10 to 14 mm long, upper lip 3 to 5 mm long, entire to slightly emarginate, lower lip with the large median lobe obcordate, spotted, 1.5 to 2.5 mm long and the lateral lobes very small; cleistogamous flowers sometimes produced, white-pubescent externally. Cocci 1.5 to 2.4 mm long, trigonous, clavate to obovate, light brown to olivaceous, smooth and lustrous, sometimes white-mottled. Lawns, cultivated areas, roadsides, waste places, etc. Native to Eur. and naturalized nearly throughout TX and most of N. America. Flowering throughout the year, ours mostly late winter to early spring, primarily Feb.-Mar., one of the earliest flowers each year.

Though not as aromatic as some other mints, the leaves, stems, and flowers are tasty and can be added to salads, stews, and soups (Tull 1987).



LAMIACEAE TEUCRIUM10. TEUCRIUM L. Germander, Wood Sage


Annual or perennial herbs to ca. 15 dm tall. Stems simple or with opposite branches. Leaves serrate, crenate or pinnatifid or lobed, short-petiolate or nearly sessile. Flowers in terminal bracteate spikes or in the axils of reduced upper leaves or leafy bracts. Calyx campanulate to saccate, bilabiate to regular, with 5 teeth and 10 nerves. Corolla bilabiate but often appearing to be one-lipped, with all 5 lobes pulled to the bottom, the tube shorter than the calyx, upper lip very short and deeply notched, the lobes about equalling the lateral lobes of the lower lip, lower lip spreading, 3-lobed, the medial lobe much the largest. Stamens 4, exserted from the notch between the lobes of the upper lip; anther sacs divergent, filaments pubescent near the base. Style 2-cleft, the branches equal. Cocci ovoid, rugose, glabrous to sparsely pubescent.

About 300 species in tropical and temperate regions of both the Old and New Worlds; 3 species in TX; 1 here.


1. T. canadense L. var. canadense American Germander, Wood Sage. Perennial herbs from creeping rhizomes, sometimes also with tubers; stems simple or branched near the inflorescence, 3 to 10(15) dm tall, pubescent with the hairs retrorsely curled or villous, eglandular. Basal leaves absent, leaf blades elliptic or elliptic-lanceolate to oval, ovate, or lanceolate, 3 to 12(16) cm long and 1 to 4(6) cm wide, basally cuneate to rounded or truncate, apically acute to acuminate or occasionally obtuse, serrate (or less often crenate or only obscurely toothed), upper leaf surface glabrous to pubescent or hirsute, lower surfaces with minute curled, eglandular hairs; longest petioles 0.3 to 2(2.5) cm long. Flowers in (2-)4- to 6-flowered verticils arranged in a spiciform inflorescence 6 to 30 cm long; peduncles 0 to 1 mm long; flowers subsessile, pedicels 1 to 3(4) mm long; bracts of inflorescence equal to or longer than the calyx, lance-subulate, only the lowest leaflike or occasionally the entire inflorescence with leafy bracts or leaves. Calyx often purplish, lightly eglandular (rarely sessile-glandular), appressed-pubescent or canescent, sometimes with scattered longer hairs, bilabiate, 5 to 9 mm long, the tube 4 to 7 mm long, glabrous to lightly hirsute within, enlarged and cuplike but not inflated at maturity, teeth broadly ovate to triangular, usually obtuse, the upper 3 more or less united, 1 to 2 mm long, the lower 3 slightly longer; corolla 10 to 18 mm long, pink to lavender or purple, rarely white, commonly spotted and fading cream to yellowish, pubescent and with sessile or stipitate glands externally, tube 4 to 8 mm long, about equalling the calyx, the 2 lobes of the upper "lip" 1.5 to 3.5 mm long, erect, deltoid or narrowly deltoid, appearing lateral on the lower lip because of the deep, wide notch between them, lateral lobes of the lower lip similar but smaller and spreading, median lobe of the lower lip short-clawed, reflexed, broadly ovate, 6 to 10 mm long; stamens ascending between the lobes of the upper lip, then curved downward. Cocci ellipsoid-obovoid, 1.5 to 2.4 mm long, rugose, glabrous, yellow-brown to light reddish-brown. Wet areas--ditches, near streams, lakes, canals, marshes, etc. Nearly throughout TX; N. S., W. to MN and NE, S. to FL and TX. (The species as a whole from Que. to B.C., S. to Cuba and Mex.) May-Sept., ours primarily Jun.-July. [T. canadense L. var. virginicum (L.) Eat., T. virginicum L.]



LAMIACEAE MARRUBIUM11. MARRUBIUM L. Horehound


Native to the Old World; of the 45 species, we have the one naturalized throughout North America.


1. M. vulgare L. Common Horehound, Marrubio. Rhizomatous perennial herb; stems 4-angled, erect to ascending, often forming clumps to about 1 m high, white wooly-tomentose, especially below, often also with stellate hairs; herbage aromatic and with bitter sap. Petioles 0.5 to 4 cm long, reduced upwards; blades ovate to broadly ovate or nearly orbicular, 1.5 to 5 cm long, 1 to 4 cm wide, crenate (less often serrate), the teeth often alternating small and large, apically acutish to obtuse or rounded, basally cuneate to nearly truncate or rounded, wrinkly-veined, the upper surfaces green or gray-green and sparsely tomentose, the lower leaf surfaces wooly or tomentose, whitish, often also with stellate hairs. Flowers in cymules of 7 or more, appearing verticillate, the clusters borne on the upper 1/2 to 3/4 of the stem in the axils of foliaceous bracts; bracts reduced upwards; each flower subtended by a villous-pubescent, linear bracteole about as long as the flower, usually spinose and sometimes hooked. Calyx cylindric, the tube 3 to 6 mm long, faintly 10-nerved, villous-tomentose and with stellate pubescence, teeth 10 (rarely more), ca. 2 mm long, acicular, more or less uncinate (hooked), sometimes alternating larger and smaller, a tuft of hairs often present in the throat; corolla 4 to 6 mm long, white to creamy, tube only slightly longer than the calyx, bilabiate, upper lip entire, emarginate, or with 2 lobes, pubescent externally, lower lip 3-lobed, the middle lobe the broadest, obovate, often emarginate, and glabrous, the lateral lobes rounded and pubescent externally; stamens 4, included within the corolla tube, didynamous, the anterior pair the longer, anther sacs divergent; style included, stigma unequally 2-lobed. Cocci ovoid to oblong or elliptic, 1.2 to 2.3 mm long, smooth to warty (often obscurely so), dark brown. Waste places, roadsides and near pens and corrals. Scattered throughout most of TX; naturalized from Eurasia. Flowering throughout the year.

The leaves and flowering parts are used for flavoring candies and so forth. The plant also have medicinal properties; one favorite use is in cough drops and sore throat lozenges (Mabberley 1987).



LAMIACEAE PYCNANTHEMUM12. PYCNANTHEMUM Michx. Mountain Mint, Basil


Perennial herbs from long, slender rhizomes. Herbage aromatic. Stems well-branched, often corymbosely so and mostly in the region of the inflorescence. Leaves sessile to short-petiolate, linear to lanceolate or elliptic, entire to serrate; floral leaves often whitened or whitish. Flowers in corymbs with the branches evident or in terminal and solitary glomerules, these often dense and headlike, though sometimes loose and open. Bracts and bracteoles present. Calyx regular or somewhat bilabiate, glabrous within, tubular, 10- to 13-nerved, 5-toothed, the teeth shorter than the tube. Corolla whitish to purplish, much longer than the calyx, slightly bilabiate, the tube slightly enlarged upwards, the two lips usually dotted with purple, about equal in length, the upper lip entire or slightly notched, the lower reflexed and with 3 ovate to oblong, obtuse lobes, the middle lobe the largest. Stamens 4, exserted or not, the lower pair slightly longer than the upper; anther sacs parallel. Stigma lobes unequal. Cocci 3-sided, obovoid or ellipsoid.

About 21 species, primarily of the E. U.S.; 4 in TX and 2 here. The relationships between some species are obscure.

In spite of the common name, this is not the Basil of Italian and Mediterranean cooking--that is Ocimum.


1. Leaves mostly more than 5 mm broad, ovate to narrowly lanceolate; branches of

inflorescence easily visible; lower leaf surfaces light gray-green ...1.P.albescens

1. Leaves less than 5 mm broad, linear; flowers in dense, headlike clusters arranged in corymbs; lower leaf surfaces the same green as the upper leaf surfaces ..............................

...2.P.tenuifolium


1.P. albescensLAMIACEAE PYCNANTHEMUM albescens T. & G. ex Gray White Mountain Mint. Stems ca. 4 to 15 dm tall, branched above, branches ascending or somewhat spreading, pubescent below and whitish to canescent in the upper portions. Petioles 2 to 12 mm long, very narrowly winged; blades ovate to ovate-lanceolate or elliptic, 2.5 to 7 cm long, 1 to 2.5 cm wide, green and glabrous to pubescent above, all but the lowest much whitened-canescent and punctate below, apices acute, bases cuneate, margins subentire to remotely serrate. Inflorescences loosely cymose, all the branches visible, canescent, overall 1.5 to 4 cm wide, terminal and solitary (occasionally 2) or on short axillary branches, usually subtended by a pair of small leaves or leaf-like bracts, these much whitened; flowers and cymules within the cyme subtended by linear-lanceolate to subulate bracteoles, these to 4 mm long. Calyx canescent with minute, appressed hairs, bilabiate, 3.5 to 5 mm long, upper lip with 3 deltoid, obtuse teeth united for 1/2 or more their length, lower lip with 2 free, ovate to oblong, obtuse teeth; corolla white to lavender, usually spotted on the lower lip, pubescent and more or less sessile-glandular externally, sparsely pubescent within, 5 to 7.5 mm long, tube 3 to 4.5 mm long, gradually widened above, upper lip 2 to 4 mm tall. Cocci 1.1 to 1.4 mm long, weakly trigonous, dark brown, puberulent on the apex. Open woods and savannahs, thickets, and often along streams. E. TX; FL to TX, N. to MO and OK. Jun.-Nov.


2.P. tenuifoliumLAMIACEAE PYCNANTHEMUM tenuifolium Schrad. Slender-leaved Mountain Mint. Plant often forming colonies by horizontal rootstocks; stems 4 to 12 dm tall, glabrous or the rounded angles of the stem sometimes puberulent, usually with many sterile simple or forked axillary branches. Leaves sessile, linear to linear-lanceolate or linear-elliptic, 1.5 to 6 cm long, 1 to 5 mm wide, acute apically, basally attenuate, entire, glabrous to sparsely pubescent, margins ciliate, sometimes revolute. Flowers in dense, headlike glomerules 4 to 9 mm wide, solitary and terminal at the ends of the branches of a compact corymb; bracts 2.5 to 4 mm long, ovate to ovate-lanceolate, pubescent or canescent, ciliate, with the midvein thick and ending in a spiny tip; bracteoles similar to bracts but shorter and not as stiff; individual cymules and flowers sessile. Calyx regular, 3.7 to 5 mm long, tube canescent or only lightly so, teeth ca. 1 to 1.5 mm long, glabrous or glabrescent, lanceolate to lance-deltoid, apices apiculate or somewhat spiny; corolla 5 to 7 mm long, white to pale lavender or pinkish (usually spotted with purple), sessile-glandular and canescent outside, lightly pubescent within, tube 3 to 4.5 mm long, upper lip 2 to 2.5 mm long. Cocci 0.7 to 1 mm long, usually smooth, dark brown. Moist, grassy open woods, bogs, savannahs, old fields, meadows, and marshes. E. 1/4 of TX; GA and TX N. to N. Eng., Ont., NY, and OH; W. to MI, WI, MN. May-Oct, ours primarily (May)June(July.) [P. flexuosum B.S.P. (or (Walt.) B.S.P.)].





LAMIACEAE MENTHA13. MENTHA L. Mint


Perennial herbs, often with creeping rhizomes. Stems erect to decumbent, simple or branched. Herbage usually aromatic. Leaves sessile to petiolate, commonly punctate, ovate or lanceolate to elliptic, marginally serrate or less commonly crenate. Flowers in verticils at most of the nodes, towards the ends of the inflorescence the whorls crowded and the inflorescence resembling a dense or interrupted spike. Bracts of inflorescence foliaceous or reduced and similar to the bracteoles subtending individual flowers. Calyx campanulate to cylindrical or tubular, 10-(13-) nerved, 5-lobed, regular to weakly bilabiate. Corolla generally pale purple, pink, or whitish, slightly bilabiate, the upper lip entire or emarginate and the lower lip 3-lobed, the lobes rounded or obtuse, OR the corolla nearly regular, 4-lobed, with one lobe slightly larger. Stamens 4, nearly equal, erect, exserted (or included and nonfunctional in plants of hybrid and cultivated origin), of 2 kinds as to fertility in some species, filaments glabrous, anther sacs parallel. Stigma lobes equal. Cocci ovoid to ellipsoid, usually smooth, brownish, not produced by some cultivated species.

About 25 species and hybrids in temperate regions of the world; 4 in TX (1 native and 3 naturalized); the 3 naturalized species are possible around old homesites. (Some species are aggressive and invasive and will persist for years where planted.)

This genus includes the familiar garden mints--peppermint, spearmint, applemint, etc. Many have culinary or medicinal uses.


1. Leaves more or less tomentose on both surfaces; upper stem leaves sessile .......................

...1.M.xrotundifolia

1. Leaves glabrous to slightly pubescent; upper stem leaves petiolate or subsessile ...............2


2(1) Leaves all distinctly petiolate, mostly cuneate at the base; calyx longer than ca. 1.5 mm ......

...2.M.xpiperita

2. Leaves short-petiolate to nearly sessile, mostly rounded at the base; calyx generally shorter than 1.5 mm ...3.M.spicata


1.M. x rotundifoliaLAMIACEAE MENTHA rotundifolia (L.) Huds. Applemint, Roundleaf Mint. Stems primarily erect, simple or branched, ca. 3 to 15 dm tall, reflexed-pubescent to lanate or tomentose; herbage more or less tomentose and often viscid. Leaves sessile to short-petioled, elliptic to oblong or ovate, 3 to 7 cm long and 2 to 3.5 cm wide, obtuse to acute apically and rounded to cordate basally, the margins crenate to serrate, pubescent on both surfaces and the lower surface somewhat reticulate-rugose. Inflorescence to 1 dm in fruit and to ca. 1 cm in diameter, spikelike in appearance, nearly continuous, the lowermost whorls of flowers somewhat distinct from one another; bracts not foliaceous, ovate-lanceolate to lanceolate, acuminate, generally shorter than the flowers. Calyx campanulate, velvety, the tube 0.7 to 1 mm long, teeth about the same length and connivent (drawing together) in fruit; corolla white or pink, sometimes spotted with purple, ca. 4 mm long, puberulent. Cocci 0.5 to 9.7 mm long, broadly oblong, reddish brown and reticulate. Old fields, homesites, thickets, and waste places. Cen. and W. TX; naturalized form Eur. in many parts of N. Amer. May-Sept.

A natural hybrid between M. longifolia (L.) L. and M. suaveolens Ehrh.


2.M. x piperitaLAMIACEAE MENTHA piperita L. Peppermint. Plant from spreading rhizomes; stems erect to somewhat decumbent, branched, ca. 6 to 16 dm tall, often purplish. Leaves petiolate, ovate to lanceolate, basally cuneate to rounded, apically acute, strongly serrate, dark green, glabrous or lightly pubescent on the veins below. Inflorescence spikelike, continuous or interrupted, ca. 3 to 10 cm long, to 12 cm at maturity, 12 to 20 mm broad; bracts not longer than the calyces, narrowly lanceolate and acuminate, resembling the bracteoles, glabrous to only sparsely ciliate; pedicels relatively long. Calyx tube 1.7 to 2 mm long, glabrous, the teeth equalling the tube or shorter, 0.8 to 1.4 mm long, pubescent or hirsute, sometimes sparsely so, ciliate-margined; corolla rose-purple to white or lavender, glabrous; stamens apparently nonfunctional and cocci not produced. Damp or wet areas, mostly in W.-Cen. TX; naturalized from Europe in much of N.A. June-Oct. in TX, probably flowering sporadically until frost.

This is the mint used to flavor candies, gum, toothpaste, teas, ice cream, liqueurs, juleps, etc. Peppermint tea has anitflatulent properties and is also useful in soothing upset stomachs and cramps. It is thought to be a naturally occurring hybrid between M. spicata L. and M. aquatica L. (Mabberley 1987). Very similar to spearmint, M. spicata L. (see NOTE below).


3.M. spicataLAMIACEAE MENTHA spicata L. Spearmint. Plants from rhizomes; stems glabrous or lightly pubescent at the nodes, erect to ascending, 3 to 12 dm tall, usually branched. Leaves sessile or quite short-petiolate, ovate to oblong-lanceolate or ovate-lanceolate, the larger ones 3 to 9 cm long and 0.7 to 3 cm wide, bases rounded to sometimes subcordate, rarely cuneate, apically acute to acuminate, the margins serrate (sometimes unevenly so), glabrous or sparsely pubescent on the veins beneath, generally punctate below. Inflorescence spikelike, the floral whorls crowded toward the branch tips but the inflorescence commonly interrupted, especially below, overall ca. 4 to 8 cm long at maturity, 6 to 9 mm in diameter; bracts not leafy, but equal to or longer than the calyx, subulate-lanceolate or linear-lanceolate, 3 to 7 mm long and 0.5 to 1 mm wide, ciliate or glabrous. Calyx campanulate, sessile-glandular, ca. 1.2 to 1.5 mm long, the tube 0.7 to 0.9 mm long, teeth triangular, 1/3 to 1/2 as long as the tube to about equal, ciliate-margined; corolla white to lavender, 1.7 to 3 mm long, glabrous; stamens often included within the corolla throat and nonfunctional. Cocci not seen, apparently not produced. Moist meadows and fields, usually near habitation. Cen. and W. TX; naturalized from Europe in many parts of N. Amer. Jun.-Oct.

Used for flavoring candies, ice cream, gum, tea, etc., and in various herbal remedies.

NOTE: Very similar to M. x piperita; best distinguished by the different leaf shape, smaller calyx, and longer floral bracts.



LAMIACEAE STACHYS14. STACHYS L. Hedge nettle


Annual, biennial, or perennial herbs, the perennials spreading by rhizomes. Herbage generally pubescent or hispid. Leaves petiolate to sessile (ours usually with at least the lower leaves petiolate), entire to crenate or serrate. Flowers in verticils or axillary clusters, the inflorescence appearing to be a compact or interrupted terminal spike or raceme; flowers sometimes also in the axils of the upper leaves. Lower bracts of the inflorescence large and leaflike, upper ones much reduced; bracteoles present beneath each flower or absent. Calyx regular or nearly so, usually campanulate, 5- to 10-(15-) nerved, 5-toothed, the teeth equal and more or less spreading, often spiny-tipped. Corolla generally reddish to purplish or pink, bilabiate, the tube slender and about as long as or slightly longer than the calyx, upper lip erect, straight or somewhat arched or galeate, entire or shallowly emarginate, lower lip spreading or deflexed, longer than the upper lip, 3-lobed, the middle lobe much larger than the later ones and itself sometimes 2-lobed. Stamens 4, didynamous, ascending and arched under the upper lip, anther sometimes contiguous in pairs, the sacs of each anther divergent. Style equally 2-cleft, the lobes subulate. Cocci ovoid or oblong, obtuse, sometimes weakly 3-ribbed, usually smooth.

About 300 species in the temperate and warm regions of the world, with several scattered also in S. Afr. and S. Amer.; 6 in TX; only 1 found here.

Some European and Eurasian species are somewhat shrubby, some have medicinal value (Betony), some have edible tubers, and some are grown as ornamentals (Lamb's Ears) (Mabberley 1987).


1. S. crenata Raf. Hedge-nettle, Shade Betony. Hirsute annual or biennial; stems to ca. 3 dm tall, branched at the base, decumbent to erect, internodes often much longer than the leaves. Lowermost leaves with petioles about equalling the blades, the upper most sessile; blades ovate to oblong, to ca. 4 cm long, obtuse, basally truncate or cordate, decidedly crenate. Flower clusters few-flowered, pedicels 1 to 2 mm long; bracts ovate to nearly orbicular, subulate-tipped to mucronate. Calyx 3 to 5 mm long, the lobes generally shorter than the tube, ovate-lanceolate to lanceolate, glabrous above the middle, with spiny subulate tips; corolla pinkish lavender, or pink to blue or rarely white, upper lip bearded and the rest glabrous, lower lip 3-lobed, the lobes rounded or the middle one very slightly emarginate. Cocci granular, ca. 1 mm long. In shaded areas, preferring rocky or gravelly soils; woods, ravines, and prairies, also banks and open grounds, sometimes weedy. E., Cen., and S. TX; also N. Mex. Feb.-May, occasionally collected again in Sept. [S. agraria of authors, but not S. agraria Cham. & Schlecht., which is a separate species. Many old specimens from our area, however, were originally identified as S. agraria Cham. & Schlecht.]

Many specimens collected locally in the 1940's were misidentified as Mentha arvensis L.; incorrectly identified plants can still be found in some herbaria.



LAMIACEAE SALVIA15. SALVIA L. Sage


Annual or perennial herbs, some (but not ours) shrubs. Stems square, herbage often aromatic. Leaves cauline, or in one species largely basal, generally toothed or lobed. Flowers in few-flowered whorls forming contracted or interrupted spike-like inflorescences. Bracteoles present or absent. Calyx bilabiate, campanulate, 10- to 15-nerved, the upper lip entire or 3-cuspidate or 3-toothed, straight or somewhat arched, lower lip 2-lobed or -toothed. Corolla zygomorphic, 2-lipped, rather tubular, pubescent externally and glabrous within, usually some shade of red, blue, or white, upper lip straight and emarginate to galeate and entire, lower lip about twice as long as the upper, 3-lobed. Stamens only two, exserted from the corolla tube, ascending under the upper lip and exserted beyond the limb or included within the galea; connective well-developed and articulating with the filament, each filament with only one fertile anther sac on the upper branch, lower branch sterile or with a rudimentary half-anther, rarely a filament with 2 normal sacs. Stigma unequally 2-lobed, style included or slightly exserted. Cocci weakly trigonous, ovoid, smooth or minutely tuberculate.

About 700 species worldwide, primarily in temperate and tropical regions, especially S. Amer.; 22 species known from TX; 3 here.

Many Salvias are grown as ornamentals. Some of the TX species are among the finest native plants for landscaping. S. officinalis L. is the common garden sage used in cooking. S. sclarea L., Clary Sage, is used in medicines, perfumes, and flavorings (Mabberley 1987).


1. Leaves mainly in a basal rosette, lyrate; both sacs of each anther fertile; floral bracts minute; corolla whitish, light blue, or violet ...1.S.lyrata

1. Leaves cauline and basal, at most crenate to dentate; one sac of each anther

rudimentary; floral bracts larger; corolla red or blue (rarely white) ........................................2


2(1) Corolla bright red; leaves rhombic-ovate, widest at the ...2.S.coccinea

2. Corolla blue, rarely white; leaves lanceolate to linear, with cuneate bases ...3.S.azurea


NOTE: Reports of S. leptophylla Benth. from our area are based on misidentified specimens of white-flowered S. azurea. Specimens of "S. farinacea" from our area prove to be S. azurea or else taken from cultivation. It is possible S. farinacea might persist. It can be distinguished by its cylindrical, truncate calyx which is densely white-tomentose and usually tinged with violet.


1.S. lyrataLAMIACEAE SALVIA lyrata L. Cancer-weed, Lyre-leaf Sage. Perennial from a somewhat woody taproot; stem scapose to subscapose, simple or rarely branched, 3 to 8 dm tall; herbage more or less pilose-hirsute. Leaves mostly basal, basal leaves 5 to 15 or numerous, petiolate, obovate to oblanceolate or spatulate, spring leaves repand to sinuate to commonly lyrate-pinnatifid, the lobes somewhat dentate, to 3 dm long, summer and fall leaves usually nearly entire, 6 to 15 cm long, 2 to 7 cm wide, usually sparsely strigose-hirsute to pilose, apices rounded or acute, bases attenuate to rounded, often purple-tinted, petioles 2 to 7 cm long, cauline leaves if present 1 or rarely 2 pairs, elliptic-lanceolate or obovate, either petiolate and lyrate-pinnatifid or sessile and entire. Verticils of flowers 3 to 10, forming an interrupted, spike-like inflorescence, each small cymule in each whorl 2- to 5-flowered; floral bracts linear to oblong, 4 to 7 mm long, generally shorter than the calyx; bracteoles absent; pedicels 1 to 4 mm long. Calyx often purple-tinged, campanulate, 7 to 12 mm long at maturity, the tube hirsute, upper lip about equalling the tube, straight or slightly reflexed, rather truncate, with 3 cuspidate bristles/teeth, lower lip somewhat longer than the upper, with 2 narrowly-triangular to lanceolate, cuspidate, pointed teeth; corolla light blue to whitish or violet, sometimes with darker markings, roughly funnelform, 18 to 30 mm long, tube 12 to 20 mm long, upper lip straight, emarginate, much shorter than the lower lip; both arms of the anther connective often with a fertile cell. Cocci ca. 2 mm long, obovoid, dark brown, minutely tuberculate. Sandy open woods, clearings, meadows, lawns, etc. E 1/4 of TX; FL to TX, N. to N. Eng., NW. to IL, MO, and OK. Dec.-May (ours in bloom mostly Mar.-Apr.)


2.S. coccineaLAMIACEAE SALVIA coccinea P.J. Buchoz ex Etlinger Tropical Sage, Mirto, Mejorana. Perennial, herbaceous or somewhat suffrutescent; stems well-branched, 3 to 10 dm tall, canescent to glabrous or hirsute toward the base with long, spreading hairs. Leaves all cauline, petiolate, the petioles 0.5 to 4 cm long; blades veiny, ovate to deltoid-ovate, 1 to 7 cm long and 0.8 to 5 cm wide, acute to obtuse, basally truncate to cordate, margins crenate, mostly softly tomentulose below. Raceme virgate, flower clusters somewhat distant, each with few to several flowers; pedicels 3 to 6 mm long. Calyx 6 to 10 mm long, crisp-hirtellous, usually tinged with red, bilabiate, the upper lip entire, lower lip with 2 acute teeth; corolla bright or scarlet red, 2.5 to 3 cm long, externally pubescent to puberulent, tube 13 to 17 mm long, moderately enlarged above, upper lip 3.5 to 5 mm long, narrow, lower lip twice as long as upper and much broader; stamens exserted from beneath upper lip; fertile and infertile cells of each anther side by side, anthers and the usually glabrous style bright reddish. Cocci dark brown, 2.3 to 2.5 mm long, smooth. Sandy soils in thickets, brush, open woods. E., SE. TX; SC to FL, GA, AL, MS, LA, and TX; also Mex. Feb.-Nov. in TX, ours seen mostly in May and Oct. [Author sometimes given as L.].


3.S. azureaLAMIACEAE SALVIA azurea Lam. Blue Sage, Pitcher Sage. Perennial from a stout rootstock; stems erect, simple or sparingly branched in the upper portion, 5 to 15 dm tall, cinereous, the hairs short and recurved, sometimes scattered longer hairs present, rarely glabrate. Leaves all cauline, petioles 0.5 to 1.5 cm long; lower leaves often early-deciduous, blades 3 to 7 cm long, 1 to 2.5 cm wide, narrowly lanceolate, elliptic, obovate, oblong or oblanceolate, acute to obtuse, dentate to serrate, middle and upper leaves elliptic, lance-linear, lanceolate, or oblanceolate, 3 to 7 cm long, 0.5 to 2 cm wide, variously pubescent, acute, basally attenuate, sessile or with petioles to 1 cm long, serrate to dentate to entire. Whorls each with 2 to 8 flowers, arranged in a spike-like inflorescence, rather crowded above; bracts subulate to linear-lanceolate, 2 to 8 mm long, somewhat persistent; bracteoles similar to bracts but reduced; pedicels 1 to 3(5) mm long. Calyx oblong-campanulate, 4.5 to 10 mm long, pubescent, often bluish around the opening, somewhat bilabiate (cleft for about 1/4 of its length), upper lip 1.5 to 2.2 mm long, entire, lower lip about as long, with 2 triangular-ovate, acute but not aristate teeth; corolla deep blue, rarely white, 10 to 25 mm long, pubescent externally, tube 5 to 12 mm long, well-exserted from the calyx, upper lip galeate, entire, densely pubescent, lower lip 3-lobed, emarginate; sterile and fertile cells of each anther side by side; style densely hirsute. Cocci 2 to 2.8 mm long, ellipsoid-ovoid, brown, smooth or resinous-glandular. Dry prairies, oak-cedar woods, flats and openings, etc. Most of TX; SC and FL, W. to NE, CO, AR, TX and S. to Mex. May-Nov., ours primarily Sept.-Nov.

If varieties are recognized, most of the TX material belongs to var. grandiflora Benth. [S. pitcheri Nutt.] with recurved stem hairs rather than the spreading ones of var. azurea.



LAMIACEAE RHODODON16. RHODODON Epl.


One (or two) species endemic to Texas. B. L. Turner (1995) recognized a second species formerly in Stachydeoma and once considered part of this species.


1. R. ciliatus (Benth.) Epl. Annual, rather pleasantly aromatic (even in old herbarium sheets retaining the minty-antiseptic smell); stems 10 to 30 cm tall, erect, freely branched, commonly villous at least above, with coarse, decurved hairs, merely pubescent below. Leaves subsessile, 15 to 20 mm long, elliptic to oblong, glandular-dotted as seen with a lens, spreading or ascending, entire, apically obtuse and bases rounded to truncate, upper and lower leaf surfaces glabrous but the margins densely long-ciliate with hairs 1 to 3 mm long. Flowers in cymules of 3 to 7, these crowded in the axils of the uppermost leaves and bracts, forming compact clusters to ca. 4(6) cm long and 1.5 cm wide; floral bracts similar to the leaves, some becoming nearly oblanceolate; pedicels about half the length of the calyx, densely retrorsely pubescent. Calyx slightly distended on the lower side, 15-nerved, the base densely setulose with coarse hairs 2 mm long, tube tapered, 4.5 to 5 mm long, teeth parallel or connivent, deltoid-lanceolate or subulate, acute, subequal, about equalling the tube, often purple-tinged, sinuses membranous; corolla pale pink to purple, 5 to ca. 6 mm long, tube with a villous ring within, equal to or shorter than the lower lip, upper lip 2-cleft, the lower lip with 3 rounded lobes, the middle one spotted within; stamens 2, attached at or near the middle of the corolla tube and included completely within it, filaments very short, anthers connate, anther sacs widely divergent; style hirtellous to pubescent. Sandy soils, localized. Smith and Robertson Cos., S. to Refugio Co.; endemic. May-July, ours apparently mostly May-June. [Hedeoma texana Cory]

This little plant, with its bushy habit, purple flowers, and sprightly scent, would make a nice addition to the garden.



LAMIACEAE HEDEOMA17. HEDEOMA Pers. Mock Pennyroyal, False Pennyroyal


Aromatic herb or herbaceous perennial, sometimes somewhat suffrutescent, commonly forming clumps, tufts, or mats. Stems from a woody taproot or rootstock, erect or ascending, decumbent or trailing, some species (but not ours) rooting at the nodes, stems variously pubescent. Leaves petiolate to sessile, sometimes coriaceous, mostly linear, elliptic, lanceolate or ovate to oblong (occasionally rhombic), punctate on one or both surfaces, entire to serrate, acute to obtuse, variously pubescent. Flowers solitary or 2 to 15(31, but not ours) in whorls, usually well-spaced, but occasionally the inflorescence spike-like. Foliaceous bracts usually subtending flowers; bracteoles lanceolate to subulate, minute. Pedicels usually shorter than the calyx. Calyx slightly zygomorphic, tubular or tubular above and gibbous below, in some species tapering upwards, in any case 13-nerved (in ours), 5-toothed, upper and lower lips equal or the lower longer, teeth of lower lip typically longer than those of the upper, teeth in some species convergent and closing the calyx in fruit, in other species teeth spreading, usually a dense ring of hairs (an annulus) present inside near the summit of the tube, this more or less sealing the orifice. Corolla white to blue, lavender, or pink (some red, but not ours), bilabiate, glabrous to variously pubescent within, tube slender or funnelform, expanded upward, upper lip ligulate (or obcordate), straight or concave and subgaleate, apically emarginate, lower lip spreading (in some species, but not ours, erect), with 3 equal lobes or the median lobe larger. Stamens 4, the lower pair with functional anthers, ascending under the upper corolla lip and exserted from beneath it, anther sacs divergent to nearly parallel, connective well-developed, upper pair of stamens staminodial and (almost always) consisting of naked filaments. Style unequally 2-lobed, about as long as the fertile stamens. Cocci oblong, ovate, or orbicular, about twice as long as wide, brown to black, areolate to smooth, usually glaucous, usually mucilaginous when moistened.

About 38 species, primarily of the American SW. and adjacent Mex.; including 5 in S. Amer.; 2 the E. and Midwest U.S.; 10 in TX; 3 here. This treatment follows that of Irving (1980).

Some species are used in herbal teas, but as at least one species, H. pulegioides, has been used to induce abortions, they should be used with extreme caution and never by pregnant women.


1. Calyx teeth at maturity not convergent and closing the orifice, instead at least some teeth reflexed or spreading; annulus at the summit of the calyx tube; plants annual; corolla glabrous within ...1.H.hispida

1. Calyx teeth at maturity convergent and closing the orifice, not reflexed or spreading; annulus below the summit of the calyx tube; corolla pubescent within. ...............................2

2(1) Plants annual or herbaceous perennials (sometimes woody near the base), usually less than 25 cm tall; leaves green, more than 3 times longer than wide; calyx weakly hirsute, teeth in fruit strongly connivent; corolla 7 to 11 mm long; plants smelling of peppermint ......

...2.H.drummondii

2. Plants suffrutescent, usually more than 25 cm tall; leaves gray or dark green, usually less than 3 times longer than wide; calyx coarsely hirsute or white-villous, teeth in fruit

incompletely closing the orifice; corolla 8 to 15 mm long; plants smelling of lemon ..............

. ...3.H.reverchonii

var. reverchonii


1.H. hispidaLAMIACEAE HEDEOMA hispida Pursh Rough False Pennyroyal. Annual, 7 to 35(40) cm tall, from a slender taproot; stems quite square, usually solitary, erect or somewhat decumbent, simple or branching in the lower 1/3, puberulent below and densely retrorsely pubescent above. Leaves spreading, linear to linear-elliptic, usually more than 5 times longer than broad, 7 to 21 mm long, 1 to 3 mm broad, punctate (sometimes obscurely so), apically obtuse and basally attenuate and subsessile, margins entire and frequently revolute, glabrate to glabrous above, strigose below and on the margins, sometimes glabrous below. Verticils at nodes in the upper ca. 2/3 of the stem, sometimes more, well-spaced or sometimes congested in a spike-like inflorescence, each cymule with (3)5 to 8(15) flowers; peduncles less than 1 mm long; pedicels hirsute, 2 to 5 mm long; bracteoles stiff, lanceolate to subulate, exceeding the pedicels, hirsute-ciliate, (0.5)2.5 to 6.5 mm long. Calyx zygomorphic, 4.5 to 6 mm long, tube 2.8 to 3.2 mm long, gibbous or saccate on the lower side for about 3/4 its length, ca. 1.5 to 2 mm wide at the widest, hirsute with spreading hairs, upper teeth connate for ca. 1/2 their length, forming a lip 1.2 to 2 mm long, curved outward and wider than the tube, silky-pubescent on the inner face, teeth narrowly triangular, 0.5 to 0.9 mm long, hirtellous-ciliate, lower calyx teeth subulate, 1.5 to 2.5 mm long, slightly recurved upwards but not closing the calyx, hirsute to ciliate; annulus in throat dense, ca. 0.5 mm wide, located at the orifice. Flowers dimorphic. Cleistogamous flowers blue or white, small in proportion to the calyx, ca. 5.3 mm long, scarcely exserted from the calyx, glabrous within, lobes of lower lip not spreading. Chasmogamous flowers blue, 6 to 7 mm long, exceeding the calyx, also glabrous within, upper lip subgaleate, lower lip spreading, the middle lobe slightly emarginate. Cocci yellow-brown or sometimes darker, ca. 6 mm wide and 1.3 mm long, narrowly ovoid, areolate and strongly glaucous (occasionally smooth or not glaucous.) Prairies and pastures. E. 1/2 of TX; VT, W. to Alta., S. to SC, TX, and E. FL, along the Gulf Coast. Apr.-Jun. [Often seen listed as H. hispidum; Includes forma simplex Lalonde].


2.H. drummondiiLAMIACEAE HEDEOMA drummondii Benth. Drummond False Pennyroyal. Aromatic annual or perennial, smelling of peppermint, ca. 10 to 45 cm tall; stems single from a taproot or many from a branched rootstock and clump-forming, simple to branched in the lower 1/2 to 2/3, branches spreading at a 45o angle, glabrate below, pubescent to puberulent above with retrorsely curling hairs. Leaves bright green, linear to elliptic-oblong, 5 to 12(20) mm long and 1.2 to 4 mm wide, punctate on both surfaces, apically obtuse, basally attenuate and short-petiolate to subsessile, entire, glabrous or spreading-pubescent above, densely pubescent beneath. Verticils more or less crowded in the upper 1/2 to 3/4 of the stems, flowers 2 to 7 per cymule; peduncles to ca. 1.5 mm long; pedicels pubescent, 1 to 3.5 mm long; bracteoles lanceolate to subulate, 1 to 2 mm long, about 1/2 as long as the pedicels. Calyx 5 to 7.5 mm long, finely hirsute, gibbous/saccate for about 2/3 its length, 1.2 to 1.3 mm wide at its widest, tube tapered to a slender neck 0.5 mm wide, orifice closed after anthesis, upper calyx teeth subulate or narrowly triangular, ca. 0.2 mm wide at the base, 1.0 to 1.5 mm long, slightly connate, erect, connivent with the lower teeth, lower calyx teeth subulate, recurved to arch over the upper teeth, 0.3 mm wide at the base, 1.5 to 2.0(2.3) mm long, ciliate; annulus of hairs in calyx throat dense, ca. 1.5 mm wide, located about 1.5 mm below the juncture of the upper and lower teeth; corolla blue (sometimes rosy-lavender), 7 to 9 mm long, sparsely pubescent in the throat with a 0.7-1.5 mm wide annulus of hairs lower down in the throat, tube ca. 5 to 8 mm long, 1.0 mm wide, upper lip ligulate, lower lip 2.5 to 3 mm long, 3-lobed, the middle lobe slightly the largest and somewhat emarginate. Cocci dark brown, usually glaucous, narrowly ovoid, ca. 1.2 mm long and 0.4 mm wide, slightly areolate. Limestone soils and outcrops nearly throughout TX except absent in far E.; ours from chalk and limestone outcrops; MT and ND, S. to Mex., W. to CA and NE, E. to OK and TX. May-Sept., ours sometimes as late as Oct. [Includes var. drummondii and var. crenulatum Irving].


3.H. reverchoniiLAMIACEAE HEDEOMA reverchonii Gray var. reverchonii Reverchon False Pennyroyal. Suffruticose perennial from a somewhat woody taproot; stems 25 to 60 cm tall, those of the previous season often persistent, branched mostly at the base, the branches then branched only in their upper 1/2, ascending or slightly decumbent, densely pubescent to canescent above, the hairs tightly retrorsely curled, glabrous below; herbage with the odor of lemon. Leaves gray-green, usually erect (not spreading), coriaceous, elliptic-oblong, ranging in size from 9 to 14 mm long and 3 to 5 mm wide, apically obtuse, basally attenuate and sub-sessile, sparsely pubescent above, densely pubescent below and on the entire margins. Verticils well-spaced in the upper 1/2 of the stems, cymules with 2 to 4 flowers (sometimes more); peduncles 1.0 to 1.5 mm long; pedicels 3 to 4 mm long, retrorsely-curling pubescent; bracteoles subulate, to ca. 2 mm long. Calyx 6 to 8 mm long, the 4.5 to 5.5 mm long tube gibbous/saccate below and tapering to a narrow neck, orifice only partially sealed after anthesis, externally rather densely hoary-villous, hairs antrorsely curled and to 1.5 mm long, upper calyx teeth not or only slightly connate, hirsute-ciliate, subulate, ca. 0.2 mm wide at base and ca. 1 mm long, widely spreading laterally, slightly reflexed but with their bases convergent with the lower teeth, lower calyx teeth subulate, 2.0 to 2.5 mm long, ciliate, slightly recurved upwards; annulus of hairs in throat quite dense, ca. 1.5 mm wide, located ca. 1.5 mm below the junction of the upper and lower teeth; corolla showy, 10 to 15 mm long, lavender or white, marked in the throat, sparsely pubescent in the throat, densely pubescent in the tube, forming a distinct annulus, tube strongly dilated, to ca. 5 mm wide, upper lip ligulate, subgaleate, 4 to 5 mm long and ca. 5 mm wide at base, lower lip ca. 5 mm long and 4 to 5 mm wide, 3-lobed, the middle lobe slightly larger than the laterals. Cocci yellow-brown or darker, narrowly ovoid, ca. 1.2 mm long and 0.6 mm wide, glaucous, areolate. Calcareous soils and outcrops. Generally along the Ed. Plat. and in N.Cen. TX, S. to the Coastal Plain, a few on scattered outcrops in E. TX; OK to TX, also AR and AL. May-Jul.(Aug.) [H. drummondii Benth. var. reverchonii A. Gray; H. latum Small].

The very similar var. serpyllifolium (Small) Irving occurs primarily to our west and may perhaps find its way into the extreme west of our region. Distinguishing characters are a camphor scent, stems branched in the lower 1/3, calyx only 5 to 6 mm long with the upper teeth erect or only slightly spreading and smaller (8 to 10 mm long), and corolla with tube only very slightly dilated. [H. drummondii Benth. var. serpyllifolium (Small) Irving; H. sanctum Small].




LAMIACEAE MONARDA18. MONARDA L. Horsemint, Bee-balm, Monarda


Plants variously taprooted annuals, perennials from creeping rhizomes, or small shrubs (none of ours shrubby). Stems erect, simple or branched, variously pubescent. Herbage aromatic, punctate or glandular nearly throughout. Leaves ovate to narrowly elliptic, lanceolate, or linear, acute to acuminate or sometimes obtuse, toothed or entire, petiolate to subsessile. Flowers sessile to subsessile in 1 to several dense, semi-globular, headlike glomerules that are more or less evenly spaced along a spike-like inflorescence or else terminal at the ends of the branches, each glomerule subtended by foliaceous, often color-tinged outer bracts and more modified inner bracts, and each flower usually subtended by a linear or subulate bracteole. Calyx tubular, regular, 13- to 15-nerved, with 5 deltoid to aristate teeth shorter than to equalling the tube, throat hairy internally or not. Corolla bilabiate, throat slightly expanded, upper lip erect, entire, and linear or else falcate and sickle-shaped, lower lip broader, spreading or deflexed, 3-lobed, the lateral lobes ovate and obtuse, median lobes narrower and slightly notched, or lower lip with just one tooth. Stamens 2, inserted on the throat of the corolla and ascending under the upper lip, exserted or not so, anther sacs divergent. Stigma unequally 2-lobed, style exserted. Cocci obovoid-oblong, apically rounded and somewhat pointed at the base, smooth.

Probably about 12 species (the exact number depends on the treatment) from N. Amer., Can. to Mex; 11 in Texas and 5 here. This treatment follows that of Scora (1967), who made an exhaustive monograph. B. L. Turner (1994b) had a different interpretation of Texas species.

Many species are cultivated for herbal seasonings and teas. M. didyma (Oswego Tea, Bee-balm, Bergamot) is especially popular and is the flavoring for Earl Grey tea (Mabberley 1987). Plains Indians chewed the leaves of several species, ate the foliage as greens, and used the plants in herbal remedies (Kindscher 1987, 1992). M. citriodora has the essential oil citronellol used in perfumes and pomades; it also is reported to have insect repellent properties. Other species are good for potpourris or have medicinal uses (Tull 1987).


1. Flower clusters solitary and terminal (rarely two per stem); upper corolla lip usually erect and a continuation of the tube; expanded throat of corolla equal to or longer than the cylindrical tube; stamens exserted; leaves ovate to ovate-lanceolate, basally abruptly narrowed or subcordate ...........................................................................................................2

1. Flower clusters two to several, forming an interrupted spike-like raceme; upper corolla lip falcate/ sickle-shaped in side view; corolla throat usually shorter than the cylindrical tube; stamens usually not exserted; leaves elliptic to oblong or linear, generally gradually

tapered to the base ...................................................................................................................3






2(1) Corolla creamy white; glomerules lax and open so that the constituent branches are visible, pedicels 2 to 5 mm long; nodes and lower part of stem often long-pilose with spreading hairs ...1.M.lindheimeri

2. Corolla lavender-purple or yellow, rarely white: glomerules dense, pedicels 1 to 2 mm long; nodes (at least those of the upper portion) with downcurved hairs, usually glabrous below, occasionally with some spreading hairs ...2.M.fistulosa


3(1) Calyx teeth deltoid to narrowly triangular-lanceolate, not aristate; floral bracts not aristate ...

. ...3.M.punctata

3. Calyx teeth aristate; floral bracts aristate ................................................................................4


4(3) Floral bractlets all similar, oblong, generally reflexed to reveal the purple, rather densely- pubescent inner surface ...4.M.citriodora

var. citriodora

4. Floral bractlets not similar; outer ones broad, becoming progressively narrower inward, seldom reflexed, essentially glabrous except perhaps for the major veins ..............................

...5.M.clinopodioides

1.M. lindheimeriLAMIACEAE MONARDA lindheimeri Engelm. & Gray ex Gray Rhizomatous perennial 3 to 6 dm tall; stems simple or branched, glabrous to finely pubescent or rarely with dense, stiffish, spreading hairs, rounded angles of stems generally purplish or darker than the sides. Petioles of major leaves generally less than 5 mm long; blades all ovate or broadly ovate-lanceolate, from 1/2 as long to 2 times as long as the internodes immediately above them, 3 to 5 cm long, 1.5 to 2.5(4) cm broad, widest just above the base, glabrous, minutely puberulent or rarely with dense, spreading hairs, serrate or occasionally subentire (especially the uppermost pairs). Glomerules solitary at the ends of the branches, 2 to 5 cm broad (excluding the corollas), component branches of a cluster visible, at least on older specimens; outer bracts more or less foliose, usually reflexed. Calyx 9 to 12 mm long, covered with short, spreading hairs and sometimes with stiff hairs at the base of the teeth; corolla cream-white, 1.7 to 3 cm long, pubescent with short, curled hairs, upper lip comose, erect, 1 to 2 mm broad at apex, corolla tube 11 to 19 mm long, gradually expanded upwards; stamens attached 1 to 2 mm within the tube. Sandy soils at wood edges, in meadows, on slopes, etc. E. 1/4 of TX; also LA. Apr.-Aug.


2.M. fistulosaLAMIACEAE MONARDA fistulosa L. Wild Bergamot, Long-flowered Horsemint. Perennial from slender rhizomes; stems 3 to 15 dm tall, simple or branched above, pubescent in the upper portion with retrorsely curling hairs (at least on the angles), sometimes with longer hairs also, rarely glabrous, lower portion of the plant glabrous. Petioles distinct, 2 to 20(25) mm long; blades ovate to ovate-lanceolate or lanceolate, 3 to 10(11) cm long and 1 to 3.5 (5) cm wide, punctate, acute to acuminate, basally rounded to cuneate or sometimes truncate, margins coarsely serrate, serrate-dentate, or subentire, sparsely pubescent to glabrous above, spreading-hirsute or villous below, primarily on the veins, sometimes glabrate. Glomerules solitary, terminating the branches, 1.5 to 3 cm broad (excluding the corollas), dense, the individual branches within not visible; floral bracts few, usually reflexed, tinged pink or white or lighter green; outer bracts narrowly ovate, 1 to 2.5 cm long and 0.7 to 1.5 cm broad, acute to acuminate, entire, inner bracts generally narrower; bracteoles present or absent, linear, 6 to 10 mm long, ciliate-hirsute. Calyx 7 to 12 mm long, puberulent or pubescent with short, spreading hairs, tube 5.5 to 11 mm long, throat hirsute within and with dense, stiff hairs at the orifice, sometimes similar hairs at the base of the acuminate, spinose teeth which measure ca. 0.6 to 2 mm long; corolla pale to dark lavender, sometimes pinkish, rarely white, 2 to 3.5 cm long, externally puberulent to weakly pilose with soft curly hairs, tube slender, 15 to 25 mm long, gradually expanded upwards, upper lip shorter than the tube, comose or bearded at the apex, linear, ca. 1 to 2 mm broad, erect, usually forming a straight continuation of the throat; lower lip spreading or deflexed, the median lobe notched; stamens positioned ca. 1 mm within the tube, exserted from under the upper lip. Cocci oblong, 1.5 to 2 mm long, brownish or blackish, shiny. Dry open woods and wood edges, old fields, roadsides, wet meadows, and marshes. E. 1/4 of TX; Ont., Que. and N. Eng., W. to MT, S. to GA and Cen. TX, W. again to NE; also Mex. May-July, ours primarily June.

If subspecific taxa are recognized, our plants are subsp. fistulosa var. mollis with stems usually branched above, petioles (8)10 to 25 mm long on the longer leaves, and the corolla lavender or rarely white. [M. mollis L.].

The Plains Indians used this plant for many things--as a meat seasoning, in medicinal teas, and as a source of perfume for hair pomades (Kindscher 1987).


3.M. punctataLAMIACEAE MONARDA punctata L. Spotted Bee-balm, Horsemint. Annual, biennial, or perennial; stems to ca. 1 m tall, sparingly branched at the base or else branched before the inflorescence; herbage puberulent to pubescent with curled or spreading bristle-like hairs. Leaves primarily petiolate, sometimes to subsessile, lanceolate to oblong-lanceolate or oblong (or nearly linear in one or two varieties), varying greatly in size from 1.5 to 9.5 cm long, serrate to subentire. Glomerules (1)2 to several per stem, 1.5 to 2.5 cm broad excluding the corollas; floral bracts spreading to reflexed, yellowish to whitish or rarely purplish, suborbicular, oblong, or lanceolate, entire to remotely serrate, usually ciliate along the margins and at the base, the outermost bracts foliose. Calyx 4.5 to 9 mm long, generally finely pubescent, teeth lanceolate to narrowly triangular to deltoid, acute to acuminate or somewhat spinose, but never aristate; corolla white, yellowish, or pinkish, with or without purple or maroon spots, upper lip usually falcate and concealing the stamens. Generally on dry or sandy soils in roughly the E. 2/3 of TX; E. 1/2 U.S.

There are two subspecies and several varieties (see Scora 1967; Kartesz 1998); 6 in TX; and apparently only 4 here, all belonging to subsp. punctata. It is difficult to determine varieties from dry specimens; fresh material is preferable. For problematic plants, consult Scora's monograph (Scora 1967).


1. Leaves essentially glabrous or with minute curled hairs on both surfaces ..................................................................2

1. Leaves pubescent with longer, straight or spreading hairs throughout or primarily on the midvein below ...................3


2(1) Corolla yellowish with large maroon spots on the upper lip; leaves 5 to 7 cm long and 1 to 1.5 cm broad, equalling or longer than the internodes immediately above them; annual to short-lived perennial; primarily on prairies E. of the Brazos R. .............................................................................................1. var. intermedia (McCl. & Epl.) Waterfall

2. Corolla pale yellow, upper lip slightly stippled with small maroon dots; leaves 5 to 6 cm long and 7 to 13 mm broad, usually shorter than the internodes immediately above; primarily on the Rio Grande Plains ............. .........................................

. ........................................................................................................2. var. coryi (McCl. & Epl.) Cory ex Shinners


3(1) Corolla whitish to cream, upper lip with pale maroon spots; stem with few bristles, these not spreading ....................

.............................................................................................................................................3. var. lasiodonta Gray

3. Corolla deep yellow, upper lip with deep, dark maroon spots; stems with spreading bristles (usually from AR. and NE. TX but known from Leon Co.) ........................................................4. var. arkansana (McCl. & Epl.) Shinners


NOTE: Two somewhat shrubby, narrow-leaved (less than 8 mm broad) types, subsp. immaculata Penn. and subsp. punctata var. fruticulosa (Epl.) Scora [M. fruticulosa Epl.], are found to our south, the former in S. Cen. TX (centered in Bastrop, Gonzales, and Guadalupe Cos.) and the latter in the Rio Grande Plains. It is possible they may eventually show up here.


4.M. citriodoraLAMIACEAE MONARDA citriodora Cerv. ex Lag. subsp. citriodora var. citriodora Lemon Bee-balm, Lemon-mint, Horsemint. Annual or sometimes biennial; stems 3 to 8 dm tall, branched mostly in the inflorescence, seldom at the base, retrorsely pubescent. Petioles 2 to 15(30) mm long, narrowly winged; leaves lanceolate to elliptic-lanceolate or oblong, larger leaves 2.5 to 6 cm long and 8 to 12 mm broad, mostly longer than the internodes immediately above them, punctate, acute, basally cuneate to attenuate, serrate to subentire, ciliate, overall thinly pubescent or glabrate. Glomerules (1)2 to 6 per branch, forming an interrupted, spike-like inflorescence, each cluster 15 to 35 mm broad exclusive of the corollas; outer floral bracts foliose and spreading, less commonly reflexed, sometimes serrulate near the apex, inner bracts oblong to lanceolate, 1.5 to 3, cm long, usually less than 4 mm wide, abruptly acuminate and ending in a spinose bristle 2 to 5 mm long, generally reflexed only above the middle to reveal the densely canescent, often purplish inner surface, outer surface pubescent, margins more or less ciliate, at least below the middle; bracteoles primarily linear, 6 to 10 mm long, ciliate-hirsute. Calyx ca. 1 to 1.5 cm long, tube 7 to 14 mm long, externally pubescent with minute upwardly-curling hairs, densely hirsute at orifice, teeth aristate, (2.5)4 to 7 mm long, ciliate; corolla 1.5 to 2.5 cm long, white to pink or pale lavender, often purple-dotted on the lower lip, tube 7 to 19 mm long, slender at the base and abruptly expanded and funnelform at the throat, sparsely hirsute near the base of the lower lip, upper lip galeate and strongly falcate, shorter than the tube, the apical 1 to 2 mm pilose, emarginate, lower lip about equalling the upper, median lobe entire; stamens exserted from under the upper lip. Cocci oblong, 1.4 to 2 mm long. Widespread in sandy loams and rocky soils of slopes, hills, prairies, meadows, and savannahs in most of TX; MO & KS, S. to AR & TX, W. to SE. NM and Mex.; adventive elsewhere. Apr.-Oct.; ours seem to be primarily May-June. [M. dispersa Small].


5.M. clinopodioidesLAMIACEAE MONARDA clinopodioides Gray Basil Bee-balm. Annual; stems simple or sparingly branched at the base, rarely branched above, 2 to 5.5 dm tall, retrorsely pubescent, sometimes longer bristles present. Petioles 2 to 25 mm long; blades oblong, lanceolate or elliptic-lanceolate, 2 to 5.5 cm long, 4 to 15(20) mm wide, broadest above the middle, longer or shorter than the internodes directly above, punctate, acute, basally attenuate to cuneate, remotely serrate to subentire, ciliate, overall glabrous to pubescent. Glomerules 1 to 3 cm broad excluding the corollas, (1)2 to 7 per stem, forming an interrupted spike-like inflorescence; bracts usually purplish, outermost bracts ovate-lanceolate to lanceolate 1 to 1.8 cm long, 3 to 5 mm broad, glabrous above and glabrous to sparsely pubescent below, usually reflexed, abruptly acuminate to a bristle, entire or becoming serrate near apex, inner bracts similar but narrower, narrowly ovate to elliptic, glabrous on the inner surface, the outer surface puberulent, with the midvein and usually 2 (4 to 6) lateral veins well-developed, margins pectinate-ciliate; bracteoles 5 to 10 mm long, hirsute-ciliate, usually linear. Calyx tube 6 to 10 mm long, minutely pubescent with antrorsely curled hairs, throat and orifice densely hirsute, teeth usually reddish-purple, somewhat subulate, aristate, (3)4 to 6 mm long, hirsute-ciliate and antrorsely pubescent; corolla white to pink or purplish, 1.5 to 2(2.5) cm long overall, tube slender at base, 11 to 15 mm long, abruptly expanded to a funnelform throat 4 to 5 mm long, sparsely hirsute near the base of the lip, upper lip galeate, falcate, shorter than the tube, apex emarginate and hirsute, lower lip about equal to or shorter than the upper; stamens included or barely exserted from beneath the upper lip. Cocci 1.3 to 1.8 mm long, brownish. Sandy or caliche soils or granites in meadows, flats, fields, and open areas. E. 1/2 of TX; KS through OK to Atascosa Co. Apr.-June.

Eaten as a green and used as a seasoning by the Plains Indians (Kindscher 1987).



LAMIACEAE LYCOPUS19. LYCOPUS L. Water Horehound, Bugle-weed


Perennial herbs from rhizomes or slender stolons, tubers produced or absent. Stems simple or branched, erect. Leaves linear to ovate-lanceolate, petiolate, sessile, or with a sub-petiolar base, entire to deeply pinnatifid, punctate, pubescence various. Flowers in dense verticils at nearly every node; bracteoles present. Calyx tubular to campanulate (sometimes rotate in fruit), glabrous in the throat, 4- to 5-toothed, the teeth equal or subequal, obtuse to acute, subulate or cuspidate, shorter than to longer than the cocci. Corolla generally white, often marked with purple, usually regular, tubular to funnelform or campanulate, pubescent in the throat, 4- or 5-lobed, the lobes spreading or erect. Functional stamens 2 (the 2nd or upper pair present but reduced to staminodes), included within the corolla or slightly exserted, anther sacs divergent. Stigma equally bilobed, style included or exserted, ovary 4-lobed. Cocci obovoid, somewhat trigonous or else flat on one side and the other side convex, glabrous, often sessile-glandular or with resin globules, apex with a crest which is entire to irregularly toothed.

About 12 species in N. temperate zones; 4 in TX; 2 found here.


1. Mature cocci longer than the broadly triangular, acute to obtuse calyx teeth, largely obscuring them at maturity ...1.L.virginicus

1. Mature cocci much shorter than the subulate or acuminate calyx teeth, not obscuring them at maturity ...2.L.rubellus


1.L. virginicusLAMIACEAE LYCOPUS virginicus L. Virginia Bugle-weed. Plant from slender stolons, tubers mostly absent, base of stem not conspicuously thickened; stems simple or branched, 3 to 9(12) dm tall, puberulent to pubescent with curved hairs, angles obtuse and sides of the stem flat or shallowly grooved. Fresh leaves dark green or occasionally purple-tinged, to 15 cm long and 5 cm broad, ovate to ovate-oblong, elliptic, or lance-ovate, acuminate and apex and narrowing basally to a margined petiole, this taper with a "dip" toward the midrib and not straight, margins coarsely toothed to serrate, sometimes just in the distal portion, glabrous or sparsely pubescent on the major veins. Verticils dense, 6- to 20-flowered, 8 to 15 mm broad at maturity; bracteoles 0.5 to 1 mm long, linear, pubescent. Calyx tubular to campanulate, 4(5)-toothed, 1 to 2 mm long, usually glabrous, shorter than the cocci, teeth broadly triangular, acute to obtuse; corolla whitish, nearly regular, 4-lobed, the lobes not reflexed, 1.8 to 2.2 mm long, puberulent and sessile-glandular; stamens included with the corolla, style included or exserted. Cocci brownish, 1.6 to 1.9 mm long, much exceeding the calyx lobes and obscuring them at maturity, asymmetrical, the crest deeply and unevenly toothed. Roadside ditches, bogs, stream banks, etc. in SE. TX; N. Eng., W. to NE, S. to FL and E. TX. Aug.-Dec. in TX; ours collected nearly all in Oct.


2.L. rubellusLAMIACEAE LYCOPUS rubellus Moench Water Horehound. Plants from rhizomes and slender, branched stolons, often with tubers at the tips; stems simple or branched, 4 to 12 dm tall, glabrous to densely pubescent, the angles rounded and faces grooved. Leaves to 15 cm long and 5 cm broad, elliptic to elliptic-lanceolate or ovate-lanceolate, acuminate, basally narrowed into a margined petiole, the taper straight or with a "dip" and thus concave, margins sharply serrate or with low teeth, lower leaf surface glabrous to pubescent. Verticils dense, 6- to 20-flowered, ca. 1 cm broad; bracteoles 1 to 2 mm long, linear, pubescent. Calyx tubular to campanulate, with (5)4 equal teeth, 1.8 to 2.5 mm long, longer than the cocci, pubescent, the teeth narrowly triangular, acuminate and sharp-pointed, straight or recurved; corolla slightly bilabiate, whitish, often spotted with purple, 2.5 to 5 mm long (about twice as long as the calyx), 5-lobed, lobes subequal and spreading, sessile-glandular and pubescent; stamens and style slightly exserted but sterile filaments mostly included. Cocci brownish with a lighter band, overall 1 to 1.6 mm long and ca. 1 mm broad, at least partially hidden by the calyx at maturity, the dorsal angle rounded, lateral edges undulate, crest with 4 to 6 blunt tubercles or undulations or one of the points sharp and tooth-like. Bogs, marshes, ditches, seeps and shallow water, etc. E. TX; ME to MO and KS, S. to FL and TX. Aug.-Dec. in TX; ours collected generally Sept.-Nov.





CALLITRICHACEAECALLITRICHACEAE

Water-starwort Family


Small, slender-stemmed annual or perennial herbs, aquatic or terrestrial on muddy, damp, or wet soil. Leaves opposite, entire estipulate, in some species quite polymorphic: narrower when submersed and broader upwards and when floating or emersed. Plants monoecious or dioecious; flowers 1 to 3 in the axils of the leaves, sometimes subtended by a pair of minute, white, inflated bractlets. Perianth none; staminate flower consisting of a single stamen with a slender filament, anther 4-locular. Pistillate flower of a single pistil, carpels 2, locules 4, each with one ovule; styles 2, slender, often longer than the fruit, persistent or caducous. Fruit plump to somewhat flattened, splitting at maturity into 4 small, nutlike mericarps, these commonly winged along the margins, sometimes wingless, surface smooth, papillose, or reticulate.

A monogeneric family.


CALLITRICHACEAE CALLITRICHE1. CALLITRICHE L. Water-starwort, Water Chickweed

Characters as given for the family. Identification to species requires careful examination of mature fruit under magnification. Vegetative characters are less reliable as environment largely dictates habit. Apomixis occurs in some taxa, with any variation fixed in the resulting populations. This adds another degree of variability. The illustrations presented by Correll and Correll (1975) and Godfrey and Wooten (1979) are very helpful.

About 17 to 25 species nearly worldwide; 5 in TX, all of which should occur in our area.

Some taxa are cultivated in aquaria (Mabberley 1987).


1. Fruit as long as wide or longer; flowers subtended by 2 small, white, inflated bractlets; leaves usually dimorphic, the various types on the same or different plants; plants usually aquatic or forming a mat on mud on the margin of bodies of water ......................................2

1. Fruit wider than long; flowers without bractlets; leaves more or less uniform; plants mostly terrestrial ...................................................................................................................................3


2(1) Fruit ca. 0.2 mm longer than wide, widest above the middle but sides of mericarps curved outward so the fruit is thickest at the base; commissural grooves shallow ...1.C.verna

2. Fruit about as long as wide or only ca. 0.1 mm longer, widest near the middle and thickest just below the middle; commissural grooves narrow ...2.C.heterophylla


3(1) Fruit 0.3 to 0.8 mm wide, more or less gibbous at the base ...3.C.peploides

3. Fruit 0.5 to 1.2 mm wide, not gibbous at the base ..................................................................4


4(3) Fruit pedicellate; margin with a thin wing turned at right angles to the fruit surface or the wing revolute and appearing as a mere thickening of the margin ...4.C.nuttallii

4. Fruit sessile or with pedicel only to 0.2 (rarely 0.6) mm long; margin under high

magnification with a definite thin wing ...5.C.terrestris


1.C. vernaCALLITRICHACEAE CALLITRICHE verna L. (emend. Kütz) Common Water-starwort. Perennial from slender rhizomes, submersed or sometimes terrestrial on the margins of streams or ponds; stems to ca. 5 dm long, leafy throughout; herbage pale green. Leaves very variable, the lower, submersed ones linear, 4 to 11 mm long, 0.3 to 0.1 mm wide, 1-nerved, apically shallowly notched, transition to floating or emersed leaves usually gradual, upper leaves commonly broader, ultimate leaves often in a floating rosette, petiolate and with obovate to spatulate blades 7 to 13 mm long and 4 to 7 mm wide, apically rounded, 3-nerved from near the base, base attenuate and connate. Flowers subtended by minute, white, inflated bracts. Fruits sessile, oval to suborbicular in outline, 0.8 to 1.7 mm long, 0.6 to 1.4 mm broad, the length always more than the width, usually by ca. 0.2 mm, widest just above the middle, thickest at the base, margins widely spreading, each with a scarious wing broadest at the summit and narrower down the sides, surface sharply reticulate with the reticulae in vertical rows, commissural grooves shallow, styles usually caducous. Still,

shallow water or sometimes stranded on mud. E. and Cen. TX; Greenl. to AK, S. to WV, OH, IL, IA, NE, TX, NM, AZ, and CA; also Mex., Eur., and Asia. Mar.-June. Treated by Kartesz (1998) as C. palustris L. [C. palustris L. and var. verna (L.) Fenley ex Jepson].

NOTE: Care should be taken to observe mature fruits,as the immature fruits can widen when pressed so that they resemble fruits of C. heterophylla, below.


2.C. heterophyllaCALLITRICHACEAE CALLITRICHE heterophylla Pursh (emend. Darby) Larger Water-Starwort. Perennial from slender rhizomes, aquatic or sometimes stranded on mud; stems leafy throughout; herbage rather dark green. Leaves quite variable, the lower submersed ones linear to linear-lanceolate, 1 to 2 cm long, 0.5 to 1.2 mm wide, 1-nerved, apically shallowly notched, transition to floating or emersed leaves usually gradual, upper leaves broader, floating leaves usually in a tight rosette, obovate to spatulate, 6 to 15 mm long, 3 to 7 mm broad, 3-nerved from near the base, apically rounded, basally attenuate and connate. Flowers subtended by two inflated, falcate bractlets. Fruit oblong to cordate in outline, 0.6 to 1.2 mm wide and about as long, or to only ca. 0.1 mm longer or shorter, carpels more strongly rounded apically than basally, fruit widest about the middle, thickest slightly below the middle, margins wingless or with very narrow wings, at least apically, surface reticulate but the reticulae not in vertical rows, commissural groove very narrow, styles 1 to 6 mm long, spreading or erect, caducous or persistent. In still water and on mud. Throughout TX; N. S., W. to MN, S. to FL and TX; also W. MT and WA, S. to CA. and Guat. Feb.-July.


3.C. peploidesCALLITRICHACEAE CALLITRICHE peploides Nutt. Mat Water-starwort. Small, inconspicuous plant; stems creeping and rooting at the lower nodes, erect branches 1 to 5 cm tall. Leaves more or less uniform, slightly crowded at the branch tips, cuneate to spatulate-obovate, 2 to 5 mm long, apically rounded, with 1 to 3 very faint nerves. Flowers without bractlets. Fruits more or less sessile, black at maturity, wider than long, 0.5 to 0.8 mm wide, the mericarps elongating and pushing against each other so that the bases are bent outward at an angle or are gibbous, surface reticulate, commissural grooves narrow only at the top, stigmas ca. 0.2 mm long, reflexed, often persistent. On wet sand or mud and in low places. E. 1/2 TX; TN and SC to FL and SE. AR, W. to TX; also E. Mex. to Costa Rica. Feb.-May.


4.C. nuttalliiCALLITRICHACEAE CALLITRICHE nuttallii Torr. Nuttall Water-starwort. Tiny annual; stems prostrate, rooting at the lower nodes, simple or well-branched. Leaves mostly uniform, oblanceolate-obovate or spatulate, 3 to 4 mm long, 0.5 to 1.5 mm wide, apically rounded, very weakly 3-nerved. Flowers without bractlets, pedicellate, often buried in the mud at maturity. Filaments in male flowers 0.2 to 0.5 mm long, anthers 0.2 to 0.3 mm wide. Fruit wider than long, ca. 1 mm broad, 0.6 to 0.8 mm long, 0.3 to 0.5 mm thick, the mericarps flat-faced, the margins with a thin, definite wing at right angles to the fruit face or curled toward the face (revolute) and appearing as a thickened margin, stigmas ca. 0.8 mm long, slightly ascending to somewhat reflexed, sometimes persistent. Damp or wet soil of flat places, fallow fields, and other low places. E. TX; Gulf Coast from AL to TX, inland to AR and KY; also Mex. and Cen. Amer. Mar.-May. Treated by Kartesz (1998) as C. pedunculosa Nutt.


5.C. terrestrisCALLITRICHACEAE CALLITRICHE terrestris Raf. (emend. Torr.) Annual Water-starwort. Tiny annual; stems slender, simple or well-branched, to 6 cm long, leafy throughout. Leaves more or less uniform, obovate-oblanceolate to spatulate, 2 to 3.5 mm long, 0.5 to 1 mm broad, very weakly 3-nerved, apically obtuse to rounded, basally usually attenuate. Flowers not subtended by bractlets, sessile or nearly so, pedicel usually only to 0.2 mm long (rarely to 0.6 mm). Filaments of male flowers 0.1 to 0.2 mm long, anthers 0.1 to 0.2 mm wide. Fruit wider than long, 0.6 to 0.9 mm wide, 0.4 to 0.7 mm long, mericarps flat-faced and with rounded tops so fruit occasionally weakly heart-shaped, margins under high magnification with very narrow but definite wings, commissural groove present between mericarps except at the apex, style reflexed and usually persistent, 0.2 to 0.4 mm long. Wet, low, open spots of lawns, slopes, pastures, paths, fallow fields, etc. E. TX; ME, W. to E. KS, MO, IL, and OK, S. to VA, GA, LA, and E. TX. Mar.-June. [C. deflexa A. Br. var. austinii (Engelm.) Hegelm.; C. austinii Engelm.].





PLANTAGINACEAE</STRONG>PLANTAGINACEAE

Plantain Family


Annual, biennial, or perennial herbs from fibrous roots, a taproot, and/or a rootstock, acaulescent or with only a short stem. Leaves all basal or basal and alternate (opposite in some, but not ours). Flowers usually small, perfect or unisexual, regular or the calyx slightly irregular, borne in a bracted spike. Calyx and corolla both 4-merous, corolla sympetalous, commonly persistent and scarious or the lobes with scarious margins. Stamens 4 or (1)2(3), epipetalous and alternate with the corolla lobes. Ovary superior, bicarpellate, in Plantago 2-celled with axile placentation, or 4-celled due to the intrusion of the placentae (ovary 1-celled in other genera); style terminal, stigma 2-lobed. Fruit in Plantago a circumscissile capsule with 1 to 40 ovules per placenta, an achene or nutlet in other genera; endosperm well-developed.

3 genera and 255 species worldwide; 1 genus and 13 species in TX; 9 species known or likely here. Recent research indicates the family may belong within the Scrophulariaceae.



PLANTAGINACEAE PLANTAGO1. PLANTAGO L. Plantain


Characters of the family. Spikes with a bract beneath each flower. Sepals 4, the 2 nearest to the bract sometimes different from the other 2. Corolla salverform, the lobes usually scarious and persistent, erect and connivent or else spreading-reflexed after anthesis. Capsule circumscissile at or near the middle.

More than 250 species worldwide; 13 in TX; 9 known or likely in our area.

Many species of Plantago are weeds of lawns and fields. The seeds of some are mucilaginous when wet. This soluble fiber is used medicinally as a laxative (Mabberley 1987). The leaves of some species are edible fresh or cooked (Tull 1987).


1. Bracts primarily scarious, non-scarious only at the base and in the center, abruptly narrowed into a pointed, scarious tip; 2 sepals nearest the bract united into one unit with 2 midveins ...1.P.lanceolata

1. Bracts wholly herbaceous or herbaceous with scarious margins, the tips not usually scarious; all 4 sepals separate .................................................................................................2


2(1) Sepals and bracts glabrous; spikes ca. 2 to 4 mm broad in flower; leaves mostly less than ca. 3 mm broad ........................................................................................................................3

2. Sepals and bracts pubescent to villous; spikes ca. 3.5 to 12 mm broad in flower; leaves often more than 3 mm broad ....................................................................................................4


3(2) Corolla lobes erect in age; capsule about twice as long as calyx; seeds 10 to 30 per capsule, rather asymmetrical ...2.P.heterophylla

3. Corolla lobes reflexed in age; capsule only slightly longer than the calyx; seeds (2)4(8) per capsule, symmetrical ...3.P.elongata


4(2) Hairs of upper portion of scape (at least most of the longer ones) spreading at right angles ........................................................................................................................................5

4. Hairs of upper portion of scape (at least most of them) ascending or appressed ................6


5(4) Bracts 1 to 2.5 mm long; corolla lobes 0.8 to 2.3 mm long; mature seeds yellow-brown to black, 1.25 to 1.75 mm long, with a deep ventral groove, the margins not transparent .........

. ...4.P.virginica

5. Bracts 3 to 4.5 mm long; corolla lobes 2 to 3 mm long; mature seeds bright red to reddish-black, shallowly concave ventrally, the margins definitely transparent ......................

...5.P.rhodosperma


6(4) Bracts triangular-ovate, with broad scarious margins, each 1/4 to 1/3 the width of the bract ..........................................................................................................................................7

6. Bracts narrowly triangular-lanceolate, lance-linear, or aristate, with very narrow scarious margins only at the base ..........................................................................................................8


7(6) Leaves densely to sparsely woolly, floccose, or pubescent on both surfaces ........................

. ...6.P.hookeriana

7. Leaves glabrous above, thinly pubescent beneath ...7.P.wrightiana


8(6) Upper leaf surface glabrous; bracts in the middle of the spike 3 to 6 (or more) times longer than the calyces ...8.P.aristata

8. Upper leaf surface more or less densely pubescent; bracts in the middle of the spike shorter than to ca. 4 times longer than the calyces ...9.P.patagonica



NOTE: P. major L., introduced from Eurasia, is a common weed in N. Amer. No specimens have been seen from our area by the author, but should the plant show up, it is easily recognized. It has large, ovate to elliptic leaves to 30 cm long and glabrous bracts and sepals. P. rugellii Dcne. is similar, but has capsules dehiscent below the middle and narrowly-lanceolate bracts rather than the broadly ovate bracts of P. major. It is a plains species first reported for Texas in 1978--it may eventually be found here.


1.P. lanceolataPLANTAGINACEAE PLANTAGO lanceolata L. English Plantain, Buckhorn Plantain, Ribwort, Ribgrass, Ripplegrass. Perennial, sometimes flowering the first year, from an erect, often branched caudex 4 to 6 cm long, usually also with fibrous roots and often also with a taproot, generally tan-woolly at the crown; herbage villous to pubescent or glabrate. Leaves narrowly elliptic, lanceolate, or oblanceolate, (5)10 to 40 cm long, (0.5)1 to 4 cm broad, acute, long-tapered to the base, 3- to several-nerved, entire or often remotely denticulate. Scapes 1 to several, striate-sulcate, commonly strigose, 15 to 60 cm tall; spikes quite dense, ovoid-conic when young, becoming cylindric and 1.5 to 5(8) cm long and to 1 cm in diameter at maturity; bracts thin, more or less ovate, 1.5 to 2.2 mm long, the small midrib green and glabrous, extending only halfway up the bract, the remainder scarious-hyaline, the tip acuminate and hyaline, surpassing the calyx above; flowers perfect, staminate, or pistillate. Sepals ovate, 2 to 2.5 mm long, scarious, the midrib green and villous-ciliate, at least at the tip, the 2 sepals next to the bract fused into one more or less truncate structure with 2 midribs and 2 short tips, or sometimes entire; corolla lobes 2 to 2.5 mm long, ovate-lanceolate, spreading or reflexed; stamens 4, conspicuously exserted, anthers basally cordate. Capsule ellipsoid, 3 to 4 mm long, dehiscent near the base; seeds (1)2, dark brown or usually black, lustrous, more or less ellipsoid, concave on the adaxial face. Roadsides, lawns, fields, and waste areas; native of Eurasia; introduced and weedy throughout much of the U.S. and N. Amer. Spring-fall. [Includes var. sphaerostachya Mert. & Koch].

Reported from our area, and as a cosmopolitan weed, quite likely. However specimens seen by the author have all proved to be misidentified and were mostly the chasmogamous form of P. virginica.

Khaki or yellow-green dyes can be made from the leaves (Tull 1987). This plant was used by European settlers as a potherb, and is said to have been brought from Europe on purpose for that use (Tull 1987).


2.P. heterophyllaPLANTAGINACEAE PLANTAGO heterophylla Nutt. Simplespike Plantain. Rather small annual with a taproot and fibrous roots. Leaves linear to linear-subulate, few to many, 5 to 10(15) cm long, less than 5 (usually less than 3) mm broad, obtuse to acute and callous-tipped, glabrous or with a few short hairs. Scapes slender, shorter than to longer than the leaves, sparsely pubescent; spikes slender, to 13 cm long, less than 5 mm broad, rather loosely-flowered so that the axis is visible, lower flowers often rather remote; bracts glabrous, to ca. 2.5 mm long, about equalling to twice as long as the calyces, longer in the upper half of the spike, acute, the hyaline margins about as broad as the midrib, base more or less saccate. Sepals glabrous, ovate to oblong-ovate, scarious-hyaline with a green midrib, the 2 nearest the bract markedly asymmetrical and apically rounded, the 2 away from the bract asymmetrical only at the base and apically obtuse; corolla lobes triangular-ovate, less than 0.5 mm long, erect and connivent after anthesis; stamens only 2. Capsule ovoid, about twice as long as the calyx, tapered to a rounded apex, dehiscent about the middle; seeds 10 to 30 per capsule, dark brown to blackish, asymmetrically angled, ca. 0.8 mm long, pitted. Damp or wet sandy soils and rocky areas, fallow fields, woods, mudflats, and sometimes salt marshes. E. 1/3 TX; NJ and PA to MO, S. to FL and TX, W. to CA, S. to Argen. Mar.-May. [P. hybrida Bart. is the name used for this plant by several regional floras (e.g., Correll and Johnston 1970; Godfrey and Wooten 1981), but P. hybrida Bart. is more properly a synonym of P. pusilla Nutt., a separate species not found in TX. Some sources list P. elongata Nutt. as a synonym, but not to be confused with P. elongata Pursh, below.].


3.P. elongataPLANTAGINACEAE PLANTAGO elongata Pursh Slender Plantain. Small, slender, taprooted annual, 3 to 20 cm tall; herbage puberulent or sometimes glabrous, sometimes somewhat woolly at the crown. Leaves linear to narrowly linear, 2 to 10 cm long, 0.3 to 1.5(3) mm wide, somewhat succulent, blunt to acute, somewhat callous-tipped. Scapes usually longer than the leaves; spikes 2 to 10 cm long, totally glabrous, less than 5 mm broad, loosely to densely flowered but the axis at least partly exposed; bracts 1 to 2 mm long, shorter than to slightly longer than the calyces, ovate to triangular-ovate, the scarious margins about as wide as the herbaceous midrib, the anterior 2 asymmetrical, the posterior 2 similar but folded and strongly keeled; corolla lobes 0.5 to 1 mm long, usually reflexed in age; stamens 2. Capsule 1.5 to 3.5 mm long, slightly longer than the calyx, ovoid, dehiscent slightly below the middle; seeds (3)4 (or more), ca. 1.5 mm long if many, to 2.5 mm long if only 4, symmetrical, oblong to lanceolate, reticulate and commonly more or less pitted. Damp or dry sandy soils, commonly over rock; sometimes on salt or alkaline flats. Rare in E., S., and N. Cen. TX; quite probable in our area; Man. to B.C., S. to TX, CO, UT, and CA. Spring. [P. pusilla Nutt. is listed as a synonym by Correll and Johnston (1970) and Godfrey and Wooten (1981), but Kartesz (1998) lists P. pusilla as a valid species.].

Some sources (e.g., Cronquist, et al. 1984) recognize varieties or subspecies . If subspecific taxa are recognized, our plants are subsp. elongata, with (3)4(5) seeds.


4.P. virginicaPLANTAGINACEAE PLANTAGO virginica L. Pale-seeded Plantain, Hoary Plantain. Usually short-lived annual or winter annual, with a weak taproot and a very short, compact stem; herbage glabrate to villous, the hairs septate and usually not straight. Leaves mostly in a basal rosette, spreading or ascending, oblanceolate to obovate or spatulate, (2)5 to 10(20) cm long and to as much as 4 cm broad, obtuse to nearly acute, base long-cuneate or with a distinct petiole, usually with 3 major veins, entire or slightly toothed or repand. Inflorescences usually much longer than the leaves, scapes to 30 cm long, short-hirsute with spreading, septate hairs; spikes usually dense, sometimes interrupted, (2)3 to 25 cm long and ca. 8 mm broad; bracts lanceolate to linear-lanceolate or ovate-lanceolate, 1.5 to 2.5 mm long, usually shorter than the calyces, hirsutulous, mostly without scarious margins, ones at the base of the spike keeled and somewhat clasping. Sepals 1.5 to 2.7 mm long, oblong-ovate or -obovate, apically acute or obtuse, midrib or keel brownish and hirsutulous, margins broad and scarious. Flowers typically cleistogamous, with aborted styles and stamens, corolla lobes erect and connivent both before and after flowering, 0.8 to 2.3 mm long, sharply acute, scarious and pale orange; occasional flowers or sometimes a whole spike or an entire plant chasmogamous, the corolla lobes spreading or reflexed after anthesis and with styles well developed, stamens well-exserted, bracts and sepals more rounded apically than in cleistogamous flowers. Capsule ovoid or rhombic-ovoid, 2 to 2.5 mm long, about as long as the calyx, dehiscent about the middle; seeds 2, dull yellow-brown to nearly black, 1.2 to 2.1 mm long, without a hyaline margin. Very common on roadsides, lawns, open woods, thickets, salt marshes, dunes, waste areas, etc., often on rocky ground. E. 1/3 TX; MA and NY, W. to WI, IA, and KS, S. to FL and TX; introduced further west. Mar.-Jun. [Includes var. viridescens Fern.].

Most manuals have keys for and describe only the cleistogamous flowers, with the result that many specimens with chasmogamous flowers have been misidentified. In our area, many have been labeled P. lanceolata or P. media, the latter a species not found in TX.


5.P. rhodospermaPLANTAGINACEAE PLANTAGO rhodosperma Dcne. Redseed Plantain. Annual from a slender taproot. Leaves in a basal rosette, sometimes densely pubescent at the crown; blades oblanceolate to elliptic or elliptic-lanceolate, (1)6 to 20(35) cm long, to about 5 cm broad, apically acute to obtuse or rounded, basally long-cuneate or with an evident petiole to 7 cm long, pubescent on both surfaces, sometimes with a grayish cast, entire to remotely and coarsely dentate or pectinate, strongly 3-nerved from the base or only the midvein conspicuous. Scapes 1 to several, shorter than to much longer than the leaves, spreading-hirsute; spikes dense, (3)6 to 15(25) cm long, ca. 1 cm broad; bracts lanceolate, 3 to 4.5 mm long, those at the bottom the spike strongly keeled, midvein hirsute, margins scarious, narrow to broad, apex blunt to apiculate. Sepals ovate-lanceolate, slightly keeled, acute to acuminate, (2)2.5 to 3.2 mm long, unequal, midrib hirsute, margins broad and scarious; corolla lobes lanceolate to ovate-lanceolate, zygomorphic, but this scarcely discernible as they are erect and connivent both before and after flowering; chasmogamous flowers apparently rare or none. Capsule more or less ovoid, (2)3(4) mm long, dehiscent below the middle; seeds 2, bright red to reddish-black, 2 to 2.8(3) mm long, almost flat on both sides, usually with a thin, pale-hyaline margin. Roadsides or usually on gravel bars in washes or streams, and on rocky soils of slopes or brushlands; sometimes on sandy soils. Throughout much of TX but most common W. of the Blackland Prairies; MO and TN, W. to KS, TX, and AZ; adventive farther W. Mar.-May.


6.P. hookerianaPLANTAGINACEAE PLANTAGO hookeriana Fisch. & C. A. Mey. Hooker Plantain, Tallow Weed. Taprooted annual with a very short, compact stem. Leaves nearly rosulate, rather grass-like, narrowly oblanceolate to linear, to 30 cm long and to 2 cm broad, usually shorter and more slender, apex acute to obtuse, callous-tipped, margins entire or with a few small, scattered teeth, usually woolly or floccose on both surfaces, sometimes glabrate. Scapes erect, shorter than to greatly surpassing the leaves, glabrate to pubescent, most of the longer hairs of the upper scape ascending or appressed, though a few may be loose; spikes dense, capitate to cylindric and rather blunt-tipped in full flower, to 12 cm long and 8 m wide, or rarely much reduced and with only 2 flowers; bracts shorter than to equalling the calyces, glabrate or more commonly villous, broad-based, triangular-ovate, midrib herbaceous, margins ca. 1/4 to 1/3 the width of the bract, broad and scarious at the base and for 1/3 to the full length of the bract. Sepals oblong, ca. 3 mm long, rounded to blunt, with a darker midrib and broad, scarious margins; corolla lobes 1 to 4 mm long, suborbicular-ovate, apically obtuse and the base slightly lobed-cordate, whitish-scarious with a slightly thicker midrib, with a brown area at the base of each or with brown stripes running the entire length. Capsule ovoid, ca. 4 mm long; seeds 2, boat-shaped, ca. 3 mm long, and 1.3 mm broad, dull brown, finely pitted. Sandy, gravelly, or rocky soils of open woods, roadsides, pastures, dunes, and savannahs; also clay flats; in our immediate area primarily in sandy soils of the Carrizo formation in Leon and Robertson Cos.; E. TX, on the coast to the Rio Grande Valley, and W. to the Trans Pecos; possibly endemic to TX. Mar.-June.


7.P. wrightianaPLANTAGINACEAE PLANTAGO wrightiana Dcne. Wright Plantain. Annual or perennial, sometimes flowering the first season, from a short, erect, sometimes branched caudex and a taproot; stem none or rarely to 6 cm long. Leaves many, erect, alternate but crowded toward the base, linear to linear-oblanceolate, 3 to 12(20) cm long, 2 to 7(10) mm broad, tips obtuse to acute, callous or sometimes involute, with 3 to 5 inconspicuous ribs, upper surface glabrous, lower surface sparsely pubescent, herbage often darkening upon drying. Peduncles 1 to many, usually longer than the leaves but sometimes not so, (5)10 to 15(25) cm long, pubescence of upper scape appressed; spikes 1 to 10 mm long, to 9 mm broad, dense, usually many-flowered and cylindrical, but sometimes with only a few flowers, axis villous; bracts about equalling or shorter than the sepals, 2 to 3 mm long,ca. 1 mm broad, rigid, lanceolate to triangular-ovate, obtuse, with an herbaceous midrib and broad scarious margins at least at the base, villous to glabrate. Sepals narrowly obovate to oblong, obtuse to rounded, 2.7 to 3.2 mm long, herbaceous, dark green or brown, pilose, margins narrowly scarious; corolla lobes pale-scarious, ovate, obtuse and more or less cordate at the base, with a brown area at the base and/or somewhat of a thickening along the midrib. Capsule ovoid, apically rounded, 3.5 to 4.2 mm long, ca. 2 mm broad, dehiscent just below the middle; seeds 2, brown, a little constricted below the middle, finely pitted. Sandy, rocky, or gravelly soils of prairies, hillsides, open woods, stream valleys, etc. Primarily on or near the Ed. Plat; in our area known at least from Brazos Co. but infrequent; OK, TX, NM, and AZ. Apr.-July.


8.P. aristataPLANTAGINACEAE PLANTAGO aristata Michx. Bottlebrush Plantain, Bracted Plantain, Buckhorn. Taprooted annual; stem very short or obsolete, 2 to 3(6) cm long in the largest specimens; herbage dark green, often darkening on drying. Leaves crowded-alternate or rosulate, linear or narrowly oblanceolate, (3)6 to 17(20) cm long, 2 to 7(10) mm wide, more or less villous beneath, glabrous above, entire, margins with slender cilia. Peduncles 1 to several, erect, usually surpassing the leaves, 3 to 20 dm long, pilose to villous with spreading or appressed-ascending hairs; spikes cylindric, (2)7 to 15 cm long, 6 to 8(10) cm broad exclusive of the bracts, occasionally reduced and with only a few flowers; bracts aristate to linear, the longest at the base of the spike and 1 to 3 cm long, gradually reduced upwards, usually hirsute, scarious margin very narrow and usually noticeable only at the base. Sepals narrowly oblong-obovate, apically rounded, 2 to 2.5 mm long, pilose, only very narrowly scarious margined; corolla more or less zygomorphic, petals 1.8 to 2.2 mm long, suborbicular-ovate, rounded, spreading-reflexed after anthesis, base more or less cordate, margin involute; stamens 4. Capsule ovoid, 2.8 to 3.5 mm long, ca. 1.3 mm wide, dehiscent just below the middle; seeds 2, brownish or reddish, 2.3 to 3 mm long, outer face convex, inner face concave, the depression surrounded by a paler stripe, hilum tiny and shaped like an "8". Dry, infertile or sandy soils of roadsides, vacant lots, etc. E. 1/4 TX; IL and NE, S. to LA and TX, but naturalized in much of the U.S. and Can.: ME to MI, ND, MT, and OR, S. to N. Mex. May-Nov. [Includes var. nuttallii (Rapin) Morris; P. patagonica Jacq. var. aristata (Michx.) Gray].

See NOTE at P. patagonica, below.


9.P. patagonicaPLANTAGINACEAE PLANTAGO patagonica Jacq. Bristle-bracted Plantain, Patagonian Plantain. Annual, winter annual, biennial, or short-lived perennial (?) with short caudex or crown and a taproot, crown sometimes branched, each branch producing a rosette; stem usually obsolete or 1 to 4 cm long; herbage usually woolly-villous or floccose. Overwintering rosette leaves short, 0.5 to 3 cm long, narrowly oblanceolate to nearly obovate, major leaves linear to very narrowly oblanceolate, 2 to 15(20) cm long, 0.5 to 7(15) mm broad, acute to acuminate, usually callous-tipped and apiculate, with 1 to 3 veins, entire, lightly pubescent to woolly on both sides. Inflorescences 1 to several, shorter than to much longer than the leaves, 2 to 26 cm long, appressed-pubescent; spikes dense, (2)5 to 10(15) cm long, to ca. 8 mm broad, with many flowers or sometimes only a few, cylindrical if well-developed; bracts linear to linear-subulate, triangular-lanceolate or lance-linear, with only a very narrow scarious margin near the base, shorter than to noticeably longer than the calyces, 1 to 10(14) mm long, longest at the base of the spike and reduced upwards. Sepals narrowly obovate, 1.4 to 2.5 mm long, scarious margined, blunt, pilose; corolla lobes suborbicular to ovate-lanceolate, 1 to 2 mm long, spreading after anthesis, pale-scarious, with a brown patch at the base and a thickened median line, margins sometimes involute; stamens 4, included to slightly exserted. Capsule 3 to 3.5 mm long, dehiscent at or just below the middle; seeds 2, reddish-tan, 2.5 to 3 mm long. Sandy, gravelly, or rocky soils of prairies, plains, hillsides, pastures, and mountains; in our area the very pubescent type common on sandy uplands of the Carrizo formation, above bogs in Leon Co. Throughout TX except the extreme E.; B.C. to Sask., S. to TX, AZ, and CA; S. to S. Amer. Mar.-Jun. [Includes var. patagonica, var. spinulosa (Dcne.) Gray, var. gnaphalioides (Nutt.) Gray, and var. breviscapa (Shinners) Shinners; P. purshii R. & S. and var. spinulosa (Dcne.) Shinners, var. breviscapa Shinners, var. picta Pilger, and var. oblonga (Morris) Shinners; P. spinulosa Dcne.; P. picta Morris; P. wyomingensis Gandog.].

NOTE: Several varieties have been described, based on bract length, pubescence, and spike-vs.-peduncle ratios, but all intergrade. In addition, some longer-bracted forms are extremely close to P. aristata and may represent hybrids or products of introgression.






OLEACEAEOLEACEAE

Olive Family


Ours shrubs and trees. Leaves usually opposite, rarely alternate, simple to odd-pinnately compound, estipulate. Flowers basically solitary or cymose, grouped into clusters or panicles, usually from the axils of the previous season's leaves or sometimes on current season's growth, perfect or unisexual, plants dioecious, polygamodioecious, or polygamous. Sepals usually 4(5 to 16), united into a cup or free, sometimes lacking. Corolla usually sympetalous, 4-(6- to 12-)lobed, sometimes absent. Stamens 2(3 to 6), epipetalous, alternate with the corolla lobes. Nectary disk sometimes present. Ovary superior, bicarpellate; style 1, terminal, simple or with a 2-lobed stigma, sometimes absent; ovules pendulous-axile, 2 per locule. Fruit a samara, drupe, or capsule.

24 genera and about 900 species nearly worldwide, especially abundant in Asia; 5 genera and 24 species in TX; 4 genera and 8 species confirmed from our area.

The common olive, Olea europaea, belongs to this family. There are many ornamentals in the family as well, including Syringa (Lilac), Forsythia, Ligustrum (Privet), Osmanthus (Sweet Olive), and Jasminum (Jasmine). Some taxa provide timber, notably Fraxinus (Ash) and Olea (Mabberley 1987).


1. Leaves pinnately compound; fruit a samara .........................................................1. Fraxinus

1. Leaves simple; fruit a drupe or drupe-like ...............................................................................2


2(1) Flowers in sessile or subsessile axillary racemes or clusters, imperfect; corolla absent .......

..............................................................................................................................2. Forestiera

2. Flowers in terminal or axillary panicles, perfect or imperfect; corolla well-developed ..........3


3(2) Panicles axillary, drooping; petals linear, 1 to 3 cm long ..............................3. Chionanthus

3. Panicles terminal, not drooping; petals shorter, not linear .................................4. Ligustrum


NOTE: Specimens of Jasminum occasionally persist around old homesites, but do not escape cultivation in our area.



OLEACEAE FRAXINUS1. FRAXINUS L. Ash


Usually trees, sometimes shrubby or flowering when still small, deciduous. Branchlets terete (as in ours) or quadrangular; terminal buds with 1 pair of valvate scales or 2 or 3 pairs of 4-ranked scales, apices of outer pairs sometimes obscurely pinnate and somewhat foliose; lateral buds with 2 scales; leaf scars slightly raised, crescent-shaped to semi-circular or nearly round, vascular bundles many, in an arc. Leaves opposite, odd-pinnately compound (rarely reduced to 1 leaflet, and not in ours); petiole flat or grooved on the upper surface; leaflets subsessile to petiolulate, entire to serrate or undulate, usually paler beneath. Plants dioecious or with occasional perfect flowers. Flowers small, commonly anemophilous, in crowded racemes or panicles, appearing before or with the leaves. Calyx small, the 4 sepals free or fused, sometimes absent or caducous. Petals 2 to 6, usually 4, rarely connate or absent. Stamens 2(3), absent or abortive in pistillate flowers. Pistil 2-celled; style slender to broad, stigma lobes 2. Fruit a quickly-maturing 1-seeded samara or winged nutlet, the body terete to flattened, wing terminal or decurrent on the body.

About 65 species of the N. temperate region; 9 in TX; 2 confirmed from our area.

The wood of Ash trees is often tough and is often used for tools, sporting equipment, and so on. Some, including F. americana and F. velutina, are grown as shade trees (Mabberley 1987). The pollen can be allergenic (GPFA 1986).


1. Wing of samara decurrent less than 1/3 the length of the fruit body; leaf scars (except on the most recent year's growth) usually crescent-shaped, the upper margin concave; terminal bud wider than long, usually blunt ...1.F.americana

1. Wing of samara decurrent to the midpoint of the fruit body or below; leaf scars mostly semicircular, the upper margin flat or only slightly concave; terminal bud longer than wide, acute to acuminate. ...2.F.pennsylvanica


NOTE: Two species not indigenous to our area deserve mention. F. texensis (Gray) Sarg. is similar to F. americana but usually has only 5(7) leaflets that are rounded to abruptly acute and are usually 8 cm long or shorter; its samaras are 1.5 to 3 cm long. It occurs in N. Cen. and Cen. TX and may be present on some limestone areas. It has been reported from our area, but herbarium sheets seen by the author from this region all belonged to F. americana or F. pennsylvanica. F. velutina Torr., Velvet or Arizona Ash, is native to the Trans-Pecos. It is frequently planted in our area as a shade tree and can be recognized by samaras with a wing shorter than the body and decurrent scarcely to the middle. As far as is known, it does not escape cultivation here and collections taken in parks and "vacant" lots are usually either definitely cultivated or belong instead to F. pennsylvanica.


1.F. americanaOLEACEAE FRAXINUS americana L. White Ash, Fresno. Tree to 20(40) m tall; trunk to 0.8(1) m in diameter; mature bark gray-brown to dark gray, deeply furrowed. Branchlets terete, usually glabrous, dark green to gray- or orange-brown; terminal bud broader than long, blunt, usually flanked by a pair of lateral buds; leaf scars deeply crescent- or U-shaped except sometimes on the most recent year's growth. Petiole flat or grooved above; blades (10)18 to 40 cm long, 10 to 15(20) cm broad, leaflets 5 to 9, usually 7, usually conspicuously whitish below, with petiolules 4 to 10(15) mm long, leaflet blades ovate to ovate-lanceolate or elliptic-lanceolate, 5 to 15 cm long, 2 to 7(8) cm broad, usually acute to acuminate (sometimes blunt), base cuneate to rounded, sometimes oblique, margin entire or slightly dentate or undulate apically, dark green above, glaucous beneath and sparsely to moderately puberulent (often densely so on the veins) or glabrous. Flowers usually unisexual; calyx minute, usually cup-like and cleft on one side, toothed 1/4 to 1/3 its depth, persistent; anthers linear to oblong, 2.5 to 4 mm long, apiculate, filaments 0.1 to 1.3 mm long. Samaras 3 to 5 cm long at maturity, slenderly lanceolate or narrowly spatulate, 4 to 7 mm broad, apically rounded to acute, emarginate, or apiculate, wing decurrent 1/3 or less the length of the fruit body; fruit body terete or essentially so, slenderly elliptic, (2)2.6 to 3.3 mm broad, comprising ca. 1/2 to 2/5 the length of the fruit. Along watercourses and in deciduous woods. E. 1/3 TX; SE. Can. and ME to E. MN, S. to TX, LA, and FL. Flowering Feb.-Mar.(Apr.). [Includes var. biltmoreana (Beadle) J. Wright ex Fern., var. crassifolia Sarg., var. curtissii (Vasey) Small, var. juglandifolia (Lam.) Rehd., and var. microcarpa Gray; F. biltmoreana Beadle.]

This is an important timber tree in the U.S., the wood being used for cabinetry, tools, toys, and so on. The seeds are eaten by some birds, while deer, rabbits, and porcupines browse the foliage (Vines 1960).


2.F. pennsylvanicaOLEACEAE FRAXINUS pennsylvanica Marsh. Green Ash, Red Ash. Tree to ca. 20 m tall; trunk to 0.5 m in diameter, usually less; mature bark dark gray to brown, shallowly furrowed; branchlets terete, glabrous to densely pubescent; terminal bud obviously longer than wide, pointed, usually flanked by a pair of lateral buds; leaf scars semicircular, the upper margin straight or shallowly concave. Petiole flat or grooved above, greenish to brown or gray; lateral petiolules 2 to 7 mm long, slightly winged; leaves 12 to 30 cm long, 8 to 18(20) cm broad, leaflets 5 to 7(9), paler beneath but not whitish, ovate to elliptic-oblong or lanceolate, (3)6 to 15 cm long, 1.3 to 5(7) cm broad, rounded to acuminate or rarely emarginate, base cuneate to rounded, commonly oblique, margin entire to serrate (sometimes only vaguely so) bright green above, paler below and pubescent to glabrous beneath or pubescent only on the veins. Flowers unisexual, panicles usually dense. Calyx minute, cup-like, often split on 1 side, nearly entire to irregularly toothed, persistent; anthers linear to narrowly oblong, 2.5 to 4 mm long, filaments 0.1 to 0.6 mm long. Samara (2)2.5 to 4.5(7.5) cm long, 0.4 to 0.6(0.8) mm long, narrowly oblanceolate, acute to rounded or emarginate, the wing decurrent 1/2 or more the length of the fruit body; body more or less terete, linear to slightly oblong, 0.8 to 2(2.2) mm broad, making up 2/5 to 1/2 the length of the fruit. Along watercourses, in alluvial woods, in swamps, in depressions in savannahs, along fencerows, and in upland woods. Roughly the E. 1/3 to 1/2 TX except the extreme SE; S. Can. and ME to Man. and MT, S. to FL, LA, and TX. Flowering Feb.-Apr. [Includes var. integerrima (Vahl) Fern., var. lanceolata (Buckl.) Sarg., var. campestris (Britt.) F. C. Gates, var. subintegerrima (Vahl) Fern., and var. austinii Fern.; F. lanceolata Borkh.; F. darlingtonii Britt.; F. smallii Britt. The specific epithet is often misspelled "pensylvanica".]

The wood is of lesser quality than that of F. americana but is used for many of the same purposes. Some species of birds eat the fruit; the foliage is browsed by rabbits and deer (Vines 1960).



OLEACEAE FORESTIERA2. FORESTIERA Poir. Forestiera


Shrubs or small trees. Branchlets terete; terminal buds with 4 to 7 pairs of imbricate, 4-ranked scales, the scales not foliar or pinnatifid (cf. some Fraxinus), lateral buds commonly similar to the terminal but sometimes superposed; leaf scar semicircular, slightly raised, with 1 bundle scar. Leaves opposite or subopposite, arrangement sometimes made obscure by shortening of the internodes and the presence of spur shoots; blades simple, entire to serrate, estipulate, epidermis usually porulose (with tiny pits); petiole grooved on upper surface. Flowers unisexual or sometimes a few perfect, borne in small axillary clusters, appearing before or with the leaves. Staminate flowers: sepals 4(5), small and caducous or else absent, corolla none, functional stamens 2 to 6, rudimentary pistil often present. Pistillate flowers: sepals 4(5), small, abortive stamens 0 to 4(5), pistil 1, unilocular, style slender, stigma capitate to clavate. Fruit a slow-ripening drupe, endocarp thin, longitudinally ribbed before maturity, mesocarp becoming juicy only just before ripening, exocarp ripening through green to purple or nearly black, often with a whitish-glaucous bloom. Seeds 1(2), narrow, elliptic-oblong; endosperm plentiful.

15 species of the Americas, especially SW. N. Amer.; 5 in TX; 2 here, with another perhaps to be looked for.

Some species are cultivated for ornament or for wood suitable for lathework (Mabberley 1987).


1. Leaves usually more than 3.5 cm long, acute to acuminate, usually glabrous; petiole usually 5 to 11 mm long; plants of low, wet woods; flowers appearing before leaves are well-developed ...1.F.acuminata

1. Leaves usually less than 3.5 cm long, acute to rounded, minutely pubescent along the major veins above and soft pubescent below; flowers appearing after leaves are well- developed ...2.F.ligustrina


NOTE: F. pubescens Nutt. is found in the Trans-Pecos, Ed. Plat., Plains Country, and N. Cen. TX. Its leaf blades are variously shaped and either totally glabrous or with some pubescence on both surfaces; the flowers appear before the leaves; habitat preference is for open pastures and brushy

areas. It is not supposed to occur in our area, but the author has seen one leafless flowering specimen labeled "F. pubescens" from Robertson Co. The label includes no habitat information which would distinguish F. pubescens from F. acuminata.


1.F. acuminataOLEACEAE FORESTIERA acuminata (Michx.) Poir. Swamp Privet, Texas Adelia. Shrub or small tree to 6 m; main stems usually spreading or leaning; bark gray-brown; twigs gray-brown to blackish, often pubescent at first, becoming glabrate, lateral branches sometimes spine-like; terminal bud ovate, 1.5 to 3 mm long, gray-brown. Leaves deciduous; petioles 7 to 14(20) mm long; blades ovate to elliptic or rhombic, acuminate at both ends, (3)4 to 10 cm long, (1)1.5 to 3(5) cm broad, usually serrate or crenate (sometimes obscurely so), at least in the upper 1/2, sometimes some leaves entire, usually glabrous at maturity (sometimes pubescent when young) or with sparse pubescence on the midrib above. Flowers unisexual or sometimes some perfect, on separate plants, appearing before the leaves in the axils of the previous year's leaf scars. Staminate flowers in dense fascicles of 4 to 8, each cluster surrounded by 3 or 4 obovate bracts 4 to 5 mm long, inflorescence axis glabrous; sepals 4 and minute or else absent; stamens (1)2 to 6, anthers oblong, 0.7 to 1.1 mm long, filaments 3 to 4.5 mm long, sometimes a style-less, rudimentary pistil present. Pistillate flowers about 18 to 32 in a dense panicle surrounded by a few obovate to spatulate, deciduous bracts, inflorescence axis glabrous; sepals none or 0.1 to 0.2 mm long and caducous; pistil 0.8 to 1.2 mm long, style 0.1 to 0.2 mm long. Drupe 7 to 18 mm long, elliptic to oblong, usually 1-seeded, dark blue-black with a glaucous "bloom", commonly curved or wrinkled. Infrequent in wet or swampy woods. E., SE., and N. Cent. TX, W. to Victoria and Refugio Cos.; SC to IN, TN, and MO, S. to FL, LA, and TX. Flowering in spring. [Includes forma vestita Palmer; Adelia acuminata Michx.].

The fruit is eaten by wild ducks (Vines 1960).


2.F. ligustrinaOLEACEAE FORESTIERA ligustrina (Michx.) Poir. Privet Forestiera. Shrub to 4 m; bark gray-brown; twigs gray-brown, some often shortened and spur-like, pubescent initially, becoming glabrous. Leaves deciduous, petioles relatively short, to ca. 1 cm long, channeled on the upper surface and pubescent; blades rather thick, elliptic or elliptic-ovate, 2.5 to 4(6)cm long, apex obtuse, rounded or acute, base cuneate and very slightly decurrent on the petiole, margin serrulate, upper surface pubescent along the major veins, softly pubescent over all below. Flowers appearing after the leaves are well developed. Staminate flowers in dense clusters commonly crowded on spur shoots, clusters subtended by ovate to obovate, sometimes ciliate, bracts or bud scales; anthers short-oblong to suborbicular. Pistillate flowers in fascicles or reduced racemes, usually glabrous or sometimes pubescent, subtended by elliptic, ciliate bracts; calyx minute and often with 2 unequal lobes; abortive stamens usually present; ovary bilocular, 2 ovules per cell, style slender. Drupe blue-black, glaucous, ovoid-subglobose, (6)7 to 8 mm long. Sandy soils of woods; not uncommon but not abundant in our area. E. and SE. TX; AL, GA, FL, TN, and TX. Flowering summer-fall. [F. wrightiana Lundell; F. autumnalis (Michx.) Poir.; Adelia ligustrina Michx.].



OLEACEAE CHIONANTHUS3. CHIONANTHUS L. Fringetree


About 120 species (including taxa formerly in Liociera) of the tropics and subtropics of E. Asia and E. N. Amer.; 1 species in TX.


1. C. virginicus L. White Fringetree, Old-man's Beard, Flowering Ash. Polygamodioecious shrub or small tree with crooked branches, to ca. 10 m tall; twigs pubescent when young, becoming glabrate. Leaves simple, opposite to alternate, deciduous (sometimes persistent in coastal regions of TX); petioles to 25 mm long, pubescent; blades oval to oblong or oblanceolate, to 20 cm long and 6 cm broad but usually smaller than this extreme, base cuneate, apex acute to acuminate (sometimes obtuse), glabrous and deep green above, paler and sometimes pubescent below, margin entire. Flowering with or before the leaves at the ends of previous year's branchlets, inflorescences delicate, pendulous panicles to 20 cm long, the bases usually with leafy bracts similar to the foliage leaves but decidedly smaller; pedicels slender, pubescent. Flowers fragrant, perfect or imperfect but usually functionally unisexual, white to greenish white. Calyx minute, with 4 triangular lobes, persistent; corolla lobes 4 to 6, 1.5 to 3 cm long, ca. 2 mm broad, narrowly linear and ribbon-like, acute, sometimes with purple spots near the base; stamens 2, filaments very short and the anthers subsessile in the base of the corolla; ovary ovoid, 2-celled, style short, 2-lobed. Fruit usually 1-celled and drupe-like, ovoid-ellipsoid, to 18 mm long, purple with a glaucous bloom; seeds 1 to 3, ovoid, slightly reticulate. Damp woods, thickets, bluffs, etc., often on the edge of woods. E. TX; FL to TX, N. to NJ, PA, WV, OH, MO, and OK; cultivated and sometimes spreading from cultivation farther NW. Flowering Mar.-Apr. Fall color, if any, bright yellow. [The specific epithet is frequently misspelled "virginica"].

Some sources (e.g., Correll and Johnston (1970), and Vines (1960)) recognized 2 varieties. Var. maritimus Pursh is supposed to have more densely pubescent leaves and panicles than the typical var. virginicus.

Commonly cultivated (where conditions allow) for its unusual flowers. The bark has medicinal properties and has been used as a diuretic and a febrifuge (Vines 1960).




OLEACEAE LIGUSTRUM4. LIGUSTRUM L. Privet


Shrubs or small trees, evergreen or deciduous; twigs usually with conspicuous lenticels. Leaves opposite, simple, entire, usually relatively short-petiolate. Flowers in terminal panicles, perfect, regular, usually white or cream-colored. Calyx campanulate, with 4 small teeth. Corolla salverform or funnelform, tube shorter than to longer than the 4 spreading lobes. Stamens 2, epipetalous, included or exserted. Ovary 2-celled, ovules 2 per cell. Fruit a drupe-like berry, black or blue-black, 2-seeded or sometimes 1-seeded by abortion.

About 50 species of Eur., N. Afr., and E. and SE. Asia to Aust.; 3 escaped or naturalized in TX.

Several species are widely cultivated for hedge plants or small trees. The fruits of some are eaten by birds. The flowers of most are strongly scented, the aroma perceived by some people as unpleasant (Mabberley, 1987).


1. Leaves coriaceous, evergreen, 5 to 15 cm long ...1.L.lucidum

1. Leaves not coriaceous, deciduous or sometimes persistent, 1.2 to 5 cm long .....................2


2(1) Flowers pedicellate; leaves ovate to elliptic or oblong ...2.L.sinense

2. Flowers sessile or subsessile; leaves elliptic or narrowly oblong ...3.L.quihoui


NOTE: L. japonicum Thunb. is cultivated extensively in our area. Not usually escaping, it does occasionally persist in old landscapes. It is an evergreen shrub to ca. 3 m, with ovate, obtuse to acute leaves to about 8 cm--smaller, less pointed, and flatter than the leaves of L. lucidum.


1.L. lucidumOLEACEAE LIGUSTRUM lucidum Ait. f. Glossy Privet, Chinese Privet, Nepal Privet. Shrub or small tree to ca. 10 m; trunks commonly several from the base; bark smooth and light gray-brown. Leaves coriaceous, dark glossy green above with a paler midrib, paler below, oval to ovate or oblong, ca. 7.5 to 15 cm long, acute or more commonly acuminate, base rounded to widely cuneate, margin somewhat revolute, the whole leaf commonly more or less conduplicate (folded lengthwise); petioles 5 to 12(20) mm long, grooved on the upper surface. Panicles, terminal or axillary, erect, 5 to 25 cm long; flowers short-pedicelled to sessile. Calyx ca. 1.5 mm long, truncate or with 4 tiny lobes; corolla white, salverform to funnelform, the tube ca. 3 mm long, the 4 lobes spreading-reflexed, short-oblong, obtuse to acute, about as long as the tube; stamens exserted, filaments about as long as the corolla tube, anthers oblong, yellow; pistil usually scarcely exserted. Drupe subglobose to oblong or slenderly obovoid, sometimes curved, blue-black, ca. 5 to 8 mm long; seed linear to oblong. Native to China, Korea, and Japan; cultivated as a shrub or tree, occasionally persisting or escaping. Flowering in spring; fruit maturing fall. Several cultivars exist (Bailey, et al. 1976). Free-living material in our area tends to be the plain variety with the normal upright habit and dark green rather than variegated leaves.


2.L. sinenseOLEACEAE LIGUSTRUM sinense Lour. Chinese Privet, Trueno de Seto. Shrub (only rarely tree-like) to ca. 4 m tall; bark gray-brown. Leaves deciduous or remaining in mild winters, but scarcely coriaceous and not lustrous; blades ovate, rhombic, or elliptic to oblong-elliptic, 10 to 30 cm long on flowering branches, rarely to 7 cm, broadly obtuse to acute or minutely notched apically, base cuneate, glabrous above or sometimes minutely puberulent on the veins, often pubescent on the midrib below; petioles short, ca. 3 to 12 mm long, pubescent. Panicles terminal, generally loose but narrow and elongate, to 10 cm long, pubescent; flowers with definite, pubescent pedicels to 3 or 4 mm long . Corolla white, to 1 cm broad, salverform-tubular, the tube equalling the limb, limb with 4 spreading, oblong-ovate, acute lobes; stamens exserted, filaments about as long as the corolla tube, anthers oblong; pistil scarcely exserted at anthesis. Drupe subglobose to oblong or narrowly obovoid, ca. 4 mm in diameter, blue-black. Vacant lots, fencerows, woods, parks, and roadsides, especially common within communities. Native to China; cultivated and widely naturalized in our area. Flowering spring and summer; fruit maturing in the fall.

Sometimes confused with L. amurense Carr., which is distinguished by having the corolla tube ca. 3 times longer than the limb. Many old specimens of L. sinense can be found which are identified as L. amurense.


3.L. quihouiOLEACEAE LIGUSTRUM quihoui Carr. Waxleaf Ligustrum. Shrub to ca. 2 m, usually deciduous in our area, the leaves perhaps remaining in a mild winter; bark gray-brown; branches commonly horizontal. Leaves thin, not coriaceous nor especially lustrous, petioles glabrous or puberulent, very short or absent; blades elliptic to narrowly obovate or oblanceolate, narrowly oblong or occasionally linear, (1.5)2 to 3(4) cm long, apex obtuse, sometimes slightly notched, base cuneate, glabrous above, duller green and glabrous below or slightly pubescent on the midrib. Inflorescences pubescent, narrow, short (to 1 to 2 cm) racemes, usually paired, grouped into panicles to ca. 10 cm long; flowers to ca. 4 mm long, sessile or extremely short-pedicellate. Calyx pubescent, with 4 small teeth; corolla tubular-salverform, tube about as long as limb, lobes of limb ovate, acute to obtuse; stamens long-exserted, anthers shortly oblong; pistil shorter than the stamens, scarcely if at all exserted. Drupe 5 to 7 mm long, subglobose or slightly flattened, blue-black, glaucous; seeds 1 or 2. Fencerows, thickets, and roadsides, especially near communities. Native of China; cultivated and escaping in our area. Flowering spring to early summer (ca. Apr.-Jun.); fruit maturing in the fall.






<CENTER>SCROPHULARIACEAE</STRONG>

SCROPHULARIACEAE

Figwort Family


Primarily herbs, some shrubs and a few trees (our native plants all herbaceous), some semiparasitic. Stems usually terete, sometimes angled or winged. Herbage often blackening on drying. Leaves simple, estipulate, most often opposite, sometimes alternate, whorled, or basal, entire to pinnately or palmately lobed or divided. Inflorescence a spike, raceme, panicle (usually of cymes), or the flowers axillary. Many species with flowers subtended by bracts or leaves; some with bracteoles as well. Flowers perfect, 5-merous or 4-merous by reduction, bilabiate to regular. Calyx of 4 or 5 parts, fused or free, lobes shorter than to longer than the tube. Corolla 4- or 5-parted, sympetalous, weakly to strongly zygomorphic, commonly bilabiate, sometimes spurred, lobes imbricate in bud. Stamens inserted on the corolla tube, fertile stamens 4 or 2, if 4, usually didynamous; sometimes 4 fertile and 1 sterile (as in Penstemon) or 5 fertile stamens (as in Verbascum) present. Ovary superior, bicarpellate, bilocular, placentation axile, ovules usually many; style 1, slender, stigma entire to bilobed or forked. Fruit a capsule, dehiscing septicidally, loculicidally, or both, or sometimes poricidal or opening by bursting. Seeds usually many, small; embryo small and endosperm well-developed.

About 222 genera and 4,450 species worldwide, but most common in the temperate zone and mountains of the tropics. There are 33 genera and 110 species in TX; 18 genera and 34 species in our area, some of which are seldom collected. Useful references include Pennell (1935) and Vincent (1982).

The family is of little economic importance, save for Digitalis (source of cardiac drugs) and a few ornamentals, including Antirrhinum (Snapdragon), Linaria (Toadflax), and Paulownia (Princess Tree) (Mabberley 1987). Castilleja is the beloved Indian Paintbrush of Texas' spring.



1. Fertile stamens 5; plants woolly ........................................................................1. Verbascum

1. Fertile stamens 2 or 4; plants variously pubescent .................................................................2


2(1) Stamens with functional anthers 2 ...........................................................................................3

2. Stamens with functional anthers 4 ...........................................................................................6


3(2) Calyx and corolla 4-lobed .........................................................................................................4

3. Calyx and corolla 5-lobed ........................................................................................................5


4(3) Leaves orbicular; capsule globose; sprawling or mat-forming herbs .......2. Micranthemum

4. Leaves not orbicular; capsule flattened; plants erect to ascending ....................3. Veronica


5(3) Sterile filaments 2, forming hairy ridges in the pleated throat of the corolla; pedicels without bracteoles; corolla blue-violet to lavender-white .....................................4. Lindernia

5. Sterile filaments absent or not evident; pedicels with two bracteoles at the summit, these similar to and adjacent to the sepals; corolla yellow, white, or purple-tinged ......5. Gratiola


6(2) Staminodium present in addition to the 4 normal stamens; staminodium often bearded and exserted from the corolla throat ................................................................6. Penstemon

6. Staminodium none, androecium of 4 normal stamens ..........................................................7


7(6) Flowers subtended by brightly-colored bracts which are often showier than the flowers themselves ............................................................................................................7. Castilleja

7. Flowers not subtended by showy bracts; bracts, if any, foliaceous ........................................8


8(7) Corolla strongly spurred ............................................................................................8. Linaria

8. Corolla not spurred ..................................................................................................................9


9(8) Leaves pinnately parted ...................................................................................9. Leucospora

9. Leaves entire or toothed ........................................................................................................10


10(9) Flowers 4-merous and essentially regular and plants erect .............................10. Scoparia

10. Flowers 5-merous OR irregular OR plants prostrate ............................................................11


11(10) Calyx of separate or nearly distinct sepals ............................................................................12

11. Calyx synsepalous, with a distinct tube and 4 or 5 lobes ......................................................13


12(11) Stems four-angled; flowers zygomorphic; leaves with pinnate venation; bracts present at the base of the pedicel ...................................................................................11. Mecardonia

12. Stems terete; flowers slightly if at all zygomorphic; leaves with major veins all arising from the base, or only one vein obvious; bracts, if present, at pedicel apex and similar to the calyx ........................................................................................................................12. Bacopa


13(11) Corolla yellow, 3 to 5 cm long ..........................................................................13. Aureolaria

13. Corolla some shade of blue, pink, purple, or white; usually less than 3 cm long ...............14


14(13) Plants to 20 cm tall; leaves primarily basal; flowers 7 to 10 mm long; calyx lobes about equalling tube ...........................................................................................................14. Mazus

14. Plants mostly more than 20 cm tall; leaves primarily cauline; flowers generally longer than 10 mm; calyx lobes shorter than the tube, or if equal to the tube then calyx zygomorphic .....

.................................................................................................................................................15


15(14) Calyx pleated; stem more or less winged. ..........................................................15. Mimulus

15. Calyx not pleated; stem wingless ...........................................................................................16


16(15) Inflorescence a dense terminal spike, the flowers overlapping; inflorescence viscid- pubescent ............................................................................................................16. Bellardia

16. Inflorescence looser, a raceme or spike, or flowers axillary; inflorescence not viscid- pubescent (though perhaps pubescent) ...............................................................................17


17(16) Leaves linear; plants glabrous to pubescent .......................................................17. Agalinis

17. Leaves broader, lance-elliptic; plants rough-hairy ............................................18. Buchnera



SCROPHULARIACEAE VERBASCUM1. VERBASCUM L. Mullein


Erect biennial or perennial herbs from taproots. Stems usually tall (often to 1 m or more), often coarse, simple or virgately branched. Herbage commonly hairy. Leaves cauline and/or basal, the basal leaves in a rosette and the cauline leaves alternate and sessile or decurrent. Inflorescence usually spicate or racemose (often dense) or sometimes a dense panicle. Floral bracts reduced. Flowers often ephemeral. Calyx synsepalous, deeply and equally 5-parted. Corolla yellow or sometimes white, only slightly zygomorphic with a short tube and rotate limb, the 2 upper lobes of the limb slightly shorter than the lower 3 and external to them in bud. Stamens 5, all fertile (unusual for the family), anthers 1-celled, the filaments lanose or villous or the lower pair glabrous to only sparsely hairy. Style apex flat, stigma capitate. Capsule globose to ellipsoid, thick-walled, septicidal. Seeds many, ribbed or reticulate.

About 360 species, primarily of N. temperate Eurasia; 3 in TX; 1 here.

Some species are grown for ornament or have various medicinal uses (Mabberley 1987).


1. V. thapsus L. Common Mullein, Flannel Mullein, Woolly Mullein, Flannel Plant, Hag-taper. Biennial with a basal rosette the first year and a simple (sometimes branched) stem 3 to 20 dm or more tall the second year; herbage densely golden-brown woolly-tomentose throughout, the hairs usually dendritically branched. Basal leaves obovate to oblanceolate, obtuse to acute, 8 to 50 cm long, 2.5 to 14 cm wide, tapered to a petiole or subpetiolar base, entire to crenate, leaves reduced in size up the stem, cauline leaves sessile, the bases decurrent on the stem as wings, the blades elliptic to oblanceolate, acute to obtuse, entire to crenate. Inflorescence a dense spike or spikelike panicle to ca. 5 dm (or more) long; flowers sessile or on very short pedicels; floral bracts ovate to lanceolate, 12 to 18 mm long, acuminate. Calyx segments (5)8 to 12 mm long, lanceolate; corolla yellow, 12 to 20(35) mm broad, pubescent with stellate hairs; upper 3 filaments woolly-villous, lower 2 glabrous or only sparsely villous; flowers fragrant. Capsule stoutly ovoid, 7 to 10 mm long, with dense dendritic-branched or stellate hairs; style persistent and glabrous; seeds 0.7 to 0.8 mm long. Fields rocky or gravelly areas in open woods, sometimes in boggy places. Naturalized throughout much of N. Amer., native to Eurasia. Mar.-Nov.

The leaves have some medicinal properties and are sometimes included in cigarettes formulated for asthma sufferers (Mabberley 1987); the first year leaves are used in teas for coughs and sore throats (Tull 1987). There is a popular folk tale about the plant's degree of woolliness predicting the coming winter's harshness. Lightfast yellow dye can be made from the flowers and leaves (Tull 1987). Roman women made yellow hair dye from the flowers and Roman soldiers dipped the long stems in tallow and used them as torches (Tull 1987).



SCROPHULARIACEAE MICRANTHEMUM2. MICRANTHEMUM Michx.


Three species in the W. hemisphere; one found in the SE. U.S. and 2 endemic to FL and Cuba.


1. M. umbrosum (J. F. Gmel.) Blake Shade Mud-flower. Low, creeping, matted, glabrous herbs, often rooting at the nodes. Leaves opposite (rarely in 3's), sessile, rotund, 4 to 9 mm wide, entire, the main veins all arising at the base of the leaf. Flowers minute, best seen with a lens, solitary in the axils of the midstem leaves, commonly a flower in only 1 leaf at each node, or some nodes without flowers; pedicels short; bracts none. Calyx 1.5 to 2 mm long, cleft nearly to the base with 4 equal, oblanceolate segments; corolla white, bilaterally symmetrical, obliquely salverform, ca. 1.5 mm long, about equal to the calyx, tube short, overall nearly rotate, 4-lobed, the upper lobe larger than the lateral and lower lobes; stamens 2, curved or bent, the filaments appendaged near the base; style short, slender, stigmas short. Capsule globose, thin-walled, 1-celled at maturity, bursting irregularly rather than dehiscing neatly, several- to many-seeded. Mud and wet sand and along ponds, bogs, streams, and swamps. E. TX; FL to VA, MS, and LA, also tropical Amer. Known in our area from Leon Co.; perhaps occurring elsewhere but inconspicuous and easily overlooked. Apr.-Sept.



SCROPHULARIACEAE VERONICA3. VERONICA L. Speedwell


Annual, biennial, or perennial herbs, mostly low. Stems erect to decumbent or prostrate. True leaves opposite, all cauline (not to be confused with leafy floral bracts present in some species--these are sometimes alternate), entire to crenate, serrate, or lobed. Flowers in terminal or axillary racemes or solitary in the axils of foliaceous bracts. Sepals 4(5), distinct or untied only at the very base, usually all the sepals equal or in some species the lower pair the longer. Corolla with a very short tube and 4 lobes, the upper lobe representing the fusion of 2 lobes and hence the largest, the lower 3 lobes external in bud, corolla overall nearly rotate, blue, violet, or pink to white. Stamens 2, exserted, each anther with 2 more or less parallel cells. Style 1, stigma solitary and capitate. Ovary 2-locular. Capsule loculicidal, rarely septicidal at the apex as well, (in ours) emarginate or deeply notched into a cordate shape, strongly laterally compressed; in other species the capsule ovoid or orbicular and flattened to subterete. Seeds 4 to many, flattened, somewhat concave, smooth to rough.

About 180 species in the N. temperate zone, primarily Europe and Turkey; 6 in TX; 3 here.

A few are cultivated or used in herbal remedies. The common name (meaning goodbye) was given because the corolla usually falls as a unit as soon as the plant is picked (Mabberley 1987).


1. Flowers solitary in the axils of bracts similar to the cauline leaves but perhaps smaller; corolla generally more than 3 mm broad or long ...1.V.agrestis

1. Flowers in racemes, subtended by bracts which may pass gradually into the leaves, but the uppermost bracts at least very different from the stem leaves; corolla generally less than 3 mm broad or long .........................................................................................................2


2(1) Corolla violet-blue; floral bracts, or at least the upper ones, shorter than or about equalling the flowers ...2.V.arvensis

2. Corolla white; floral bracts all much longer than the flowers ...3.V.peregrina


1.V. agrestisSCROPHULARIACEAE VERONICA agrestis L. Wayside Speedwell. Annual; stems prostrate to reclining, to ca. 2 dm tall but often much less, pilose; herbage pubescent with small, thickened, at least somewhat-glandular hairs, especially on the petioles. Leaves obscurely palmately veined, ovate to oval, short-petiolate, to 1.5 cm long (often 1 cm or less), basally rounded-truncate, the margins rather deeply crenate-serrate; . Flowers in the axils of bracts resembling the leaves but smaller. Pedicels equalling or shorter than the leaves but longer than the calyx, ca. 5 to 12 mm long. Sepals ovate to elliptic, acute to obtuse, slightly exceeding or just equalling the capsule; corolla uniformly bright blue or sometimes with darker markings, 4 to 8 mm broad; style ca. 1 mm long. Capsule wider than long, notched-obcordate, strongly flattened, the apical lobes erect, with short eglandular and longer glandular hairs. Seeds ca. 9 to 12 per locule. Lawns, waste areas, roadsides, etc. E. and N. Cen. TX; naturalized from Eurasia in the E. U.S. and Mex. Early spring, more or less Mar.-Apr. [Treated as V. polita Fries. by many sources, including Correll and Johnston (1970); apparently, that is a separate species--see Kartesz (1998)].

Quite common in our area though not much collected--being so small it is easily overlooked, and its deciduous corollas defeat attempts to get good specimens.


2.V. arvensisSCROPHULARIACEAE VERONICA arvensis L. Common Speedwell, Corn Speedwell, Corn Sperry. Taprooted annual; stems erect to decumbent or ascending, sometimes procumbent, simple to well-branched, 0.5 to 2(3) dm long; herbage villous-hirsute to pilose or glandular-puberulent. Leaves sessile or the lowermost with very short petioles, rather conspicuously palmately veined, ovate to broadly ovate, 5 to 12(20) mm long and 3 to 13 mm broad, apically rounded to obtuse, basally subcordate to truncate or rounded, margins crenate to serrate. Inflorescence a terminal spike-like, bracteate raceme; bracts somewhat leaflike at the lower nodes, but becoming narrower and elliptic and entire up the stem, the uppermost not much resembling the stem leaves; pedicels short, 0.5 to 1(2) mm long. Calyx segments linear-lanceolate, acute, 3 to 6 mm long, the lower pair slightly longer than the upper; corolla blue to blue-violet, often with darker blue lines, inconspicuous, ca. 2 to 3 mm broad. Capsule 2.5 to 3.5 mm long, 3 to 4 mm wide, deeply notched-obcordate, the apical lobes spreading, strongly flattened, pubescent on the margins, the style 0.4 to 0.8(1) mm long, included between or slightly exceeding the lobes of the fruit. Seeds 0.7 to 1 mm long, flattened, ca. 8 to 12 per locule. Open woods, fields, sandy woods, lawns, and urban areas. E. and E. Cen. TX; naturalized from Europe throughout most of the U.S. and Can. Mar.-June.

Very similar to V. agrestis but distinguished by the non-leaflike upper bracts, shorter pedicels, and smaller corollas.


3.V. peregrinaSCROPHULARIACEAE VERONICA peregrina L. Purslane Speedwell, Necklace Weed. Annual from a short taproot; stems erect or ascending, simple or branched at the base (sometimes branched in the inflorescence in particularly robust plants), 0.5 to 2(3) dm tall; herbage sometimes somewhat succulent, glabrous or with gland-tipped hairs. Leaves sessile or the lowermost somewhat petiolate, narrowly oblong to oblanceolate, 0.5 to 2.6 cm long, 3 to 5(10) mm wide, entire to crenate or serrate, apically rounded-truncate to rounded-acute. Flowers in terminal, bracteate racemes; the bracts linear-oblanceolate, the lower somewhat similar to the stem leaves but becoming reduced and more entire up the stem; pedicels 0.5 to 2 mm long, generally shorter than the calyx. Calyx segments narrowly elliptic to lanceolate or oblanceolate, 3 to 6 mm long; corolla white, 2 to 2.5 mm broad; filaments short; style 0.1 to 0.4 mm long. Capsules roughly obcordate, flattened, 3 to 4(4.5) mm long, 3.5 to 4.5(5) mm wide, with a broad, shallow apical notch 0.2 to 0.5 mm deep. Seeds many, flattened, oblong, 0.5 to 0.8 mm long, ridged over the embryo. In and around tanks, streams, open woods, prairies, and dune areas; also common in lawns and waste areas. In most of TX; most of N. Amer. and introduced from there into many parts of the world. Feb.-Jun., our collections mostly Mar.-Apr.

There are two subspecies, both collected in our area.


subsp. peregrina Plants glabrous. [V. peregrina L. var. typica Penn.].


subsp. xalapensis (Kunth in H.B.K.) Penn. Plants with short, gland-tipped hairs, usually present even on fruit. [V. peregrina L. var. xalapensis (Kunth in H.B.K.) Penn.; V. xalapensis Kunth; V.sherwoodii M. E. Peck].


NOTE: Many of our plants don't fall neatly into either of these categories, showing some gland-tipped pubescence and having glabrous fruits.



SCROPHULARIACEAE LINDERNIA4. LINDERNIA All. False Pimpernel


Annual herbs. Leaves opposite, in TX material denticulate to subentire. Flowers solitary in the axils of leaves; bracteoles absent. Sepals 5, free or only weakly fused, commonly more or less linear. Corolla lavender blue to pale blue or blue-violet, zygomorphic, the limb shorter than the tube, upper lip with short, erect lobes and lower lip with 3 larger, spreading lobes. Fertile stamens 2, representing the shorter, upper pair, anther cells commonly divergent, filaments of lower stamen pair without anthers and forming 2 pubescent yellow ridges on the inside floor of the corolla, their ends projecting as sterile knobs. Stigmas distinct, platelike. Capsule ovoid to ellipsoid, septicidal, the septum persistent as a thin plate. Seeds many, minute, yellow to light tan, smooth or with fine transverse lines.

Roughly 50 to 80 species in warmer parts of the E. hemisphere, primarily Asia and Africa; 1 in Eur. Two species are found in TX, one variety of which is found here.


1. L. dubia (L.) Penn. var. anagallidea (Michx.) Cooperrider. Clasping False Pimpernel. Sstems 0.5 to 2 dm tall, 4-angled, slender and diffusely branched from near the base, sometimes rooting at the nodes; herbage glabrous. Stem leaves sessile, ovate to ovate-elliptic or ovate-lanceolate, 5 to 17 mm long, 2 to 8 mm broad, broadest at the rounded or clasping base (lowermost leaves sometimes somewhat narrowed), entire or serrulate. Pedicels solitary in the axils of the leaves, filiform, to ca. 25 mm long, usually 1 to 3 times the length of the subtending leaf. Calyx lobes linear, 2 to 4 mm long; corolla 7 to 9 mm long, white to pale lavender, all the flowers apparently chasmogamous (cf. var. dubia with many falling unopened). Capsule ellipsoid, 2.5 to 5 mm long, often exceeding the calyx, with persistent style ca. 1 mm long; seeds faintly reticulate to smooth. Sandy soils, margins of streams, ponds, and persistent puddles, and along ditches and other moist places. Nearly throughout TX excluding the Trans-Pecos; SE. Can. and N. Eng., W. to ND, S. to the Gulf states and Mex; also the Pacific NW. and CO. Apr.-Oct.; most of ours apparently from June or later. [L. anagallidea (Michx.) Penn.; Ilysanthes inequalis (Walt.) Penn.]

NOTE: Cooperrider (1976) combined the L. anagallidea and L. dubia. Our plants probably all belong to var. anagallidea, above. Var. dubia is distinguished by having shorter, stouter pedicels which are generally shorter than the subtending leaves, leaves rounded or narrowed at the base, and capsules generally equalled or exceeded by the calyx. Its late-season flowers are often cleistogamous, falling unopened.



SCROPHULARIACEAE GRATIOLA5. GRATIOLA L. Hedge-hyssop


Low herbs, usually perennial but some annual or biennial. Stems generally erect. Leaves opposite, sessile, palmately-veined, entire to toothed, sometimes obscurely glandular. Flowers solitary or paired in the axils of the leaves or leafy bracts, most species with 2 bracteoles at the apex of each pedicel, these often similar to the calyx segments. Flowers more or less bilabiate. Calyx deeply 5-lobed, the segments narrow and subequal to unequal. Corolla tube quadrangular, tubular, or narrowly campanulate, the limb basically 5-lobed, the upper lip 2-cleft or entire, external in bud, pubescent at the base within, lower lip 3-cleft, the lobes rounded to emarginate. Style dilated or bilabiate apically. Fertile stamens 2--the upper pair, with anther cells approximately parallel, commonly exceeded by a membranous extension of the connective, lower filaments vestigial or absent. Capsule ovoid to globose, glabrous, septicidal and usually also loculicidal (4-valved), 2-celled, membranous or coriaceous. Seeds many, small, reticulate.

About 20 species of temperate areas and tropical mountains; 2 in Eur. TX has 5; we have 4 of these.


1. Plants villous-hirsute; flowers subsessile; calyx lobes quite unequal; corolla only slightly longer than the calyx ...1.G.pilosa

1. Plants glabrous to puberulent; flowers usually pedicellate; calyx lobes subequal; corolla much longer than the calyx. .....................................................................................................2


2(1) Corolla bright yellow; stem less than 10 cm tall, thin, not fleshy; plants of sandy, often dry, soils; capsules almost twice the length of the calyx lobes ...2.G.flava

2. Corolla white (at lest the limb); stems usually taller than 10 cm, rather thick and/or fleshy; plants of wet or very muddy soils; capsule equal to the corolla lobes or only slightly longer ..

...................................................................................................................................................3


3(2) Pedicels stout, generally much shorter than 1 cm; capsule globose, usually slightly longer than the calyx; hairs on the upper side of the corolla throat filiform ...3.G.virginiana

3. Pedicels slender, less than 1 cm long; capsule globose-ovoid, about equalling the calyx; hairs on the upper side of the corolla throat clavate ...4.G.neglecta


1.G. pilosaSCROPHULARIACEAE GRATIOLA pilosa Michx. Hairy Hedge-hyssop. Perennial; stems firm, 1 to several from the base, stiffly erect, 1 to 7.5 dm tall, from a hard, subwoody caudex; herbage pilose or villous-hirsute throughout, stems sometimes purplish-green above. Leaves ovate to ovate-lanceolate (the upper leaves smaller and narrower), 1.2 to 2 cm long, 5 to 11 mm wide, sessile, broadly rounded and clasping at the base, apically acute to obtuse (the lower leaves sometimes blunter), crenate to serrate with a few teeth or entire, somewhat pebbled in texture above in addition to the pubescence, glandular punctate below. Flowers sessile or very short-pedicellate in the axils of the leaves; bracteoles 2, linear-lanceolate, ca. 9 mm long, at the apex of the pedicel, just below and slightly larger than the sepals. Calyx segments pubescent, markedly unequal, linear to linear-lanceolate, the long ones 5 to 7 mm long; corolla white or purple-tinged, sometimes blue-purple veined or sometimes slightly yellowish, 5 to 9 mm long, not much longer than the calyx, limb more or less bilabiate, pubescent within on the lower side; anther cells contiguous, vertical. Ovary slenderly conical or pyramidal, 4 to 5 mm long; seeds irregularly 4-sided, obscurely lined, brownish-yellow. Bogs, swamps, moist areas, often sandy woods, sometimes in water. E. 1/3 TX; FL to TN, N. to NJ, MD, NC, KY, TN, AR, and OK. Jun.-Aug. [Tragiola pilosa (Michx.) Small & Penn., Sophoranthe pilosa (Michx.) Small].


2.G. flavaSCROPHULARIACEAE GRATIOLA flava Leavenw. Golden Hedge-hyssop. Glabrous annual; stems less than 10 cm tall, clustered near the base, erect-ascending, slender, sometimes purple-tinged near the base. Leaves relatively few, sessile, clasping, linear-oblanceolate, to 15 mm long and 5 mm wide, often much less than this, entire or with a few reduced serrations, apically obtuse. Bracteole at apex of pedicel only 1, linear-spatulate, about as long as the sepals. Calyx lobes 3 to 5 mm long, linear to linear-lanceolate, apically obtuse; corolla to ca. 12 mm long, often no more than half that, tubular-funnelform in overall outline, usually described as having an orange-yellow tube and golden-yellow limb, but ours when fresh usually bright lemon yellow, drying golden. Capsule about twice as long as the sepals at maturity, ovoid-pyramidal, brown; seeds brown, coarsely reticulate. Sandy soils of prairies and fields, often under fairly dry conditions. Restricted to S. and Cen. TX and parts of LA. Feb.-Apr., ours apparently mostly Mar. [G. pusilla Torr.].

A charming plant. Though small, it is often found in large enough numbers to be striking. It is on the same scale and a nice counterpoint to our Bluets (Houstonia spp.), with which it is often found--at least one sheet in the TAMU herbarium is a mixed collection!


3.G. virginianaSCROPHULARIACEAE GRATIOLA virginiana L. Annual or perhaps the base sometimes biennial or weakly perennial (?), dark green, with a long hank of fibrous roots; stems, at least at the base, somewhat thickened or fleshy, usually in pressed specimens very flat and more than 1 cm broad, simple or with a few ascending branches, sometimes the lower branches rooting at the nodes, erect to decumbent, 1 to 4(5) dm tall; herbage glabrous to subglabrous below, somewhat glandular-pubescent above. Leaves 3- to 5-veined, elliptic-lanceolate to oblanceolate or oblong-obovate, (1.5)2 to 5(7) cm long, (5)7 to 25 mm wide, somewhat glandular-punctate, entire to sharply serrate, sessile or the lower ones short-petiolate or sometimes clasping, apically rounded to broadly acute. Pedicels stout, 1 to 4(12) mm long; bractlets at apex similar to the calyx segments but a little longer, 1.5 to 5.5(9) mm long. Calyx segments linear or lanceolate, acuminate, 3.5 to 4.5 mm long in flower and to 7.5 mm long in fruit; corolla milk-white to sometimes slightly honey-colored, often pink- or lavender-tinged or with purple lines, sometimes the tube greenish, flowers often drying lavender, 8 to 13(14) mm long, the early-season flowers largest, pubescent inside the throat with filiform hairs; stamens included. Capsule globose, (3.5)4 to 7 mm in diameter; seeds yellow or brown-yellow, slenderly cylindric, 0.7 to 0.9 mm long, finely reticulate with the longitudinal ribs the more prominent. Usually in water: streams, lakes, ponds, ditches, boggy areas, or occasionally in open, flat forest areas. E. 1/3 TX; FL and TX N. to NJ, MD, WV, IA, IN, IL, OH, KS, and MO. Mar.-May. Some sources recognize varieties. [G. sphaerocarpa Ell.].


4.G. neglectaSCROPHULARIACEAE GRATIOLA neglecta Torr. Annual; stems erect or diffuse, simple or branched, 0.5 to 2(4) dm tall, sometimes slightly succulent or woody at the base, upper internodes, pedicels, and expanding leaves rather clammy-puberulent. Leaves elliptic to rhombic-lanceolate, serrate (especially apically), tapered to base and apex or some leaves clasping at the base. Pedicels filiform, conspicuous, 8 to 20(25) mm long, glandular-pubescent; bractlets linear, similar to the calyx segments and as long or longer. Calyx segments subequal, narrowly ovate to lanceolate, acute to obtuse or rarely rounded, 3.2 to 5 mm long in flower, to 7 mm long in fruit; corolla (7)8 to 12 mm long, cream or honey-color to white, the tube commonly yellow (rarely pale lavender), pubescent inside the throat with clavate hairs, lobes of upper lip almost completely fused, lobes of the lower lip broad, emarginate; sterile stamens minute or absent, fertile stamens included, the connective between the anther cells ca. 0.7 mm long, generally longer and wider than the pair of anther cells. Capsule ovoid to globose, acuminate, 3 to 5 mm long; seeds basically cylindric, 0.5 to 0.6 mm long, yellowish, finely reticulate with the longitudinal ridges more prominent. Wet or muddy soils around ponds and in woods. E. 1/3 of TX; Can. from Que. to B.C., S. throughout most of the U.S. as far as TX, NM, AZ, AL, MS, and GA. Mar.-Jun. [G. gracilis Benth.; G. torreyi Small].



SCROPHULARIACEAE PENSTEMON6. PENSTEMON Mitch. Beard-tongue


Perennial herbs, some (but not ours) shrubby. Stems decumbent to erect, 1 to many from a taproot and herbaceous or woody crown. Herbage sometimes with a fetid odor, stems glabrous or pubescent; if pubescent, often glandular in the inflorescence. Leaves opposite, entire to toothed, subpetiolate or more commonly with well-developed petioles, basal leaves absent or present, if present sometimes tufted, stem leaves filiform to lanceolate or broadly ovate, sessile, often clasping. Inflorescence usually a terminal panicle made of cymes or verticils; bracts varying from reduced to prominent. Flowers varying from nearly regular to strongly bilabiate. Calyx of 5 equal to subequal sepals, free or nearly so, entire to erose, sometimes scarious-margined. Corolla often pink, purple, blue, white, or in some species red, tubular to salverform, funnelform, or tubular-funnelform, slightly to strongly pleated, occasionally gibbous, the upper lip with 2 lobes, the lower lip 3-lobed or -cleft, lower inner surface (floor) of throat and tube often with lines representing nectar guides, variously glabrous to pubescent within and without. Stamens 4, didynamous, epipetalous at or near the base of the corolla, filaments arching, anthers commonly opening across both the divaricate sacs and the connective. Staminodium filament-like, inserted on the corolla tube at about the same height as the ovary, often arched against the upper surface of the corolla, included to strongly exserted, often flattened and bearded at the apex. Stigma lobes united, capitate, style elongate and slender. Capsule ovoid to ovoid-pyramidal, septicidal or 4-valved, the valves firm to flexible. Seeds many, usually angled, sometimes slightly rounded, reticulate or rough.

About 250 to 300 species of N. Amer., especially the W.; 1 species in NE. Asia; 22 species in TX; apparently only 3 here. An old but useful reference is the work of Keck (1932).

Many species are grown as ornamentals, often as annuals. [Also spelled in old sources Pentastemon and Pentstemon.].


1. Corolla bright red; leaves thick, fleshy, and glaucous; upper leaves connate-perfoliate ........

. ...1.P.murrayanus

1. Corolla white to pale lavender or pink; leaves not thick, fleshy, or glaucous; upper leaves not connate-perfoliate ...............................................................................................................2


2(1) Corolla 3.5 to 6 cm long, widely inflated; floor of corolla throat not pleated ...2.P.cobaea

2. Corolla 2 to 3 cm long, narrow, little expanded; floor of corolla throat pleated .......................

. ...3.P.australis

var. laxiflorus


1.P. murrayanusSCROPHULARIACEAE PENSTEMON murrayanus Hook. Scarlet Penstemon, Cupleaf Penstemon. Stems 5 to 10(11) dm tall; herbage glabrous. Leaves entire, thick or somewhat fleshy, glaucous, the lowermost oblanceolate, long-tapered to a subpetiolar base, midstem leaves 5 to 10 cm long, 2.5 to 5 cm wide, ovate to oblong, upper leaves connate-perfoliate, suborbicular. Inflorescence glabrous, of several whorls, each with a few flowers. Sepals 5 to 7 mm long, lanceolate, acute to acuminate, with minute scarious margins; corolla bright red, even old herbarium sheets retaining some color, ca. 3(4) cm long, gradually-tapered funnelform, lobes of limb gently spreading; staminode glabrous, usually shorter than the fertile stamens. Capsules ovoid, 14 to 18 mm long including a beak ca. 5 mm long; seeds 3 to 4 mm long. Sandy soils in the E. part of TX; also OK, LA, AR. Mostly Apr.-May but sometimes as late as June. Pollinated by hummingbirds.


2.P. cobaeaSCROPHULARIACEAE PENSTEMON cobaea Nutt. Cobaea Penstemon, Fox-glove. (Not to be confused with Digitalis, the true Fox-glove.) Robust perennial; stems 1 to 3 (4) from a creeping, woody, often massive rootstock, erect or ascending, (1.5)2 to 6.5(10) dm tall, retrorsely puberulent below and glandular pubescent near the inflorescence. Leaves entire to serrate, glabrous to puberulent or pubescent, basal leaves often absent or withered by anthesis, if present lanceolate or oblanceolate to spatulate, 3.5 to 18(26.4) cm long, 0.8 to 5.5 (7.6) cm wide, subsessile to petiolate, the petioles often winged, apically acute to rounded, stem leaves sessile, passing to cordate-clasping above, lanceolate to ovate, 3.5 to 12(15) cm long, 1 to 4.5(5.4) cm wide, apically acute. Inflorescence 10 to 30(52) cm long, with 3 to 6(8) whorls, each cyme with 2 to 6 flowers; pedicels erect or appressed to the axis; bracts ovate, acute, glabrous to glandular-pubescent. Calyx viscid, glandular-pubescent, the segments lanceolate to lance-ovate, (8)10 to 16 mm long, 2.5 to 4.5 mm wide, acute, sometimes with narrow scarious margins at the base; corolla bilabiate, 3.5 to 5.5 cm long, white or pink to pale purple shades, externally glandular pubescent and viscid, throat abruptly inflated, 15 to 18(25) mm long, the orifice slightly constricted, the interior with dark pink or purple nectar guidelines, glandular pubescent, the upper lip arched, the lower lip spreading to reflexed; staminode included or slightly exserted, lightly golden-yellow bearded nearly its full length, the tip flattened and recurved, with wavy hairs to 3.5 mm long, the other hairs shorter and retrorse; anther cells divaricate. Capsule 13 to 18 mm long; seeds 2 to 3.5 mm long, black, angular. Loamy soil, prairies, and seeming to prefer calcareous soils. From the Rolling Plains to the Blackland Prairie, S. to the Gulf; NE. to MO, AR, OK, and TX. Listed as flowering May to June, but ours often in April. Some of our best colonies occur on the calcareous outcrops of Grimes and Washington Cos.


3.P. australisSCROPHULARIACEAE PENSTEMON australis Small subsp. laxiflorus (Penn.) Bennett Stems 1 to 5 from a simple or branched rootstock and taproot, erect or ascending, 2.5 to 6.5(7) dm tall, spreading- or retrorsely-puberulent below, glandular-pubescent near the inflorescence. Leaves nearly entire or more often serrate to dentate, glabrous to puberulent, the lower surface generally paler than the upper, basal leaves ovate or obovate to spatulate, 2.5 to 9 cm long, 0.5 to 2.5 cm wide, subsessile to petiolate, the petioles sometimes winged, apically acute to obtuse, stem leaves narrowly lanceolate to oblanceolate, (2)3 to 10.5(11) cm long, (2)5 to 22 mm wide, sessile, commonly clasping, acute to obtuse. Inflorescence glandular-pubescent, 5 to 26(32) cm long, with 3 to 7 well-spaced whorls, each cyme with 2 to 6 flowers; peduncles spreading to ascending, pedicels lax; bracts similar to the upper leaves but much smaller. Calyx glandular-pubescent, the lobes 2 to 5.5 mm long, 2 to 3 mm wide, lance-ovate to ovate, acuminate to acute, with slight or pronounced scarious margins; corolla white to tinged with pink or lavender, 2 to 2.8(3) cm long, bilabiate, tubular, little expanded, the lower side definitely pleated or ridged, the palate somewhat upward-arched and restricting the orifice, interior with red-purple nectar guidelines and white to pale yellow eglandular hairs, upper lip spreading, lower lip projecting beyond the upper; staminode well-exserted from the throat, flattened, sometimes slightly recurved at the tip, the last 8 to 10 mm with a dense golden beard of hairs to 1.5 mm long; anther sacs 1 to 1.3 mm long, divaricate; style glabrous. Capsule 8 to 10 mm long, pointed-ovoid; seeds brown to darker, 0.7 to 1.5 mm long, angular to rounded. Prefers acid soils, primarily on the Gulf coastal plain; common but not very abundant in E. TX; GA and FL, W. to OK, AR, LA, and TX. Mar.-June, ours mostly Apr. Treated by Kartesz (1998) as P. laxiflorus Penn. [Includes subsp. purpureus (Penn.) Penn., subsp. typicus Penn., and var. purpureus Penn.].

NOTE #1: Similar to P. tubiflorus Nutt., which differs in having broader, more-entire leaves and a trumpet-shaped corolla which is unpleated and without lines within. The staminode is only lightly bearded and not exserted. A plant primarily of the Ozarks, sometimes sporadic in NE. TX. Unfortunately, one local key gave it preeminence in the local flora and mentioned P. australis (as P. laxiflorus) as being reported from our area but not seen. This has led to numerous misidentifications of our plants as P. tubiflorus. Penstemon species have been known to expand their ranges dramatically over the years; it would not be impossible to one day encounter the true P. tubiflorus here.

NOTE #2: We are just to the S. of the range of P. digitalis Nutt. It is similar to P. australis subsp. laxiflorus and P. tubiflorus but has an unpleated, unlined corolla abruptly expanded above the base; corolla lobes more or less extended rather than spreading. It may someday make its way south into our area.



SCROPHULARIACEAE CASTILLEJA7. CASTILLEJA L. f. ex Mutis Indian Paintbrush, Paintedcup


Perennial or less commonly annual or biennial herbs, some species sometimes woody at the base, hemiparasitic. Stems decumbent to erect. Leaves alternate, sessile, entire to pinnately lobed, mostly cauline. Inflorescence a spike or spike-like raceme; bracts foliose below and becoming more brightly colored and conspicuous upward, usually brighter and showier than the flowers themselves. Flowers markedly bilabiate. Calyx tubular, the apex 4-cleft or -lobed or more commonly the segments fused and the calyx thus 2-lobed, these 2 primary lobes entire to emarginate or (slightly) 2-cleft. Corolla often inconspicuously colored, greenish or yellowish to cream, strongly zygomorphic, tubular, narrow, the upper lip galeate (hooded), straight and erect, its lobes fully fused and enclosing the anthers, lower lip very short with 3 rudimentary teeth or subpetaloid lobes, these external in bud. Stamens 4, didynamous, each anther with 2 unevenly-placed cells--one attached by its middle and the other, smaller, cell suspended by its apex. Stigma 1 (lobes united) or shortly bilabiate. Capsule ovoid, somewhat asymmetrical, bilocular, loculicidal. Seeds many, seed coat loose, alveolately-reticulate.

About 200 species, primarily in W. N. Amer. and a few in S. Amer., the W.I., and Eurasia. There are 9 species in TX; 2 known from our area.


1. Primary lobes of the calyx (10)13 to 22 mm long, the segments of each lobe linear to lanceolate, distinct for at least 1/3 the length of the primary lobes; plants perennial ..............

...1.C.purpurea

1. Primary lobes of the calyx 6 to 9 mm long, the segments wholly united, forming 2 broad, truncate (or at most emarginate) primary lobes; plants annual ...2.C.indivisa


1.C. purpureaSCROPHULARIACEAE CASTILLEJA purpurea (Nutt.) G. Don Perennial from a somewhat woody crown; stems clustered, 1 to several, 2 to 3(4) dm tall, erect to ascending, simple to sparingly branched, villous-lanate. Leaves villous-cinereous; bracts lanceolate to widely lanceolate, with 1 or 2 pairs of lateral lobes. Calyx (20)25 to 34 mm long, the primary lobes (10)13 to 22 mm long and their segments 7 to 16 mm long, linear-lanceolate. Bracts and calyces brightly colored on their distal ends (see the varieties below), the corolla inconspicuously colored; corolla 25 to 40 mm long, the tube 16 to 25 mm long, upper lip (galea) 9 to 13 mm long, the lower lip somewhat protruding, (1.5)2 to 7 mm long. Capsule to 1 cm or more long; reticulae of seeds enclosing areas that are nearly hexagonal. Most commonly on gravelly and sandy soils. Cen. TX and the Ed. Plat. to the N. Rio Grande Plains; S. KS and S. MO through OK to TX.

There are 3 varieties in TX, with one known from our area and another perhaps within a portion of our range.


var. purpurea Bracts and calyces distally purple to purple-red; lower lip of the corolla 1.5 to 3(4) mm long. Localized on limestone outcroppings. E. Cross Timbers and N. Blackland Prairies, common on the Grand Prairie, W. to W. Cross Timbers and S. to Central Mineral Region. SW. MO and E. OK, SW. through Cen. TX. Known from our area. Apr.-May. [Euchroma purpurea Nutt.].


var. lindheimeri (Gray) Shinners Bracts and calyces distally yellow-orange to red, occasionally brownish-orange; lower lip of corolla ca. 1.5 to 3(4) mm long. Calcareous clays and gravels or sandy soils or prairies and hills. S. Blackland Prairies and Grand Prairie to the SE. part of the Ed. Plat. and N. Rio Grande Plains. Endemic. Mar.-May. No specimens seen from our area but we do have pockets of its favored habitat in our region; it likely occurs here. It is similar in color to C. indivisa and perhaps it has been collected but misidentified. [C. lindheimeri Gray; C. mearnsii Penn.; C. williamsii Penn.].


NOTE: A variety with yellow-tipped bracts and calyces, var. citrina (Penn.) Shinners, occurs to the NW. of our area and is not expected here. [C. citrina Penn.; C. labiata Penn.]. Our yellow-flowered paintbrushes are forms of C. indivisa.


2.C. indivisaSCROPHULARIACEAE CASTILLEJA indivisa Engelm. Indian Paintbrush, Texas Paintbrush. Annual from a slender taproot, hemiparasitic, presumably on grasses; stems 2 to 4 dm tall, usually strict, pilose below, passing to villous in the inflorescence. Leaves linear to lanceolate, entire or with 1 to 2 pairs of short, spreading lobes, the blades puberulent and the nerves and margins pilose-ciliate, basal rosette scarcely developed, if present, not persisting until anthesis. Floral bracts rhombic to broadly oblanceolate, 7 to 13(18)mm broad, commonly with 2 short lateral lobes near the apex, the wide midlobe mucronate. Calyx 18 to 25(28) mm long, the primary lobes entire to truncate-rounded or at most emarginate, 6 to 9 mm long; bracts and calyces distally various shades of red, with orange-red the most common, occasionally blood-red or yellow forms found; corolla 20 to 28 mm long, the tube 15 to 20 mm long, the hooded upper lip 6 to 9 mm long, lower lip reduced but still projected forward. Capsules ca. 1 cm long or slightly longer. Reticulae of seeds enclosing slightly elongated areas. Sandy loam soils on the Gulf Coastal Plain and the E. 1/2 of TX, a few from lower elevations in the Grand Prairie; also SE. OK. Mar.-June, with sporadic summer and fall blooming in our area (at least.)

This is one of the plants encouraged by the TX Highway Department. It provides spectacular displays in early spring, especially in contrast with Blubebonnets and Evening Primrose. Often plentiful even where not planted. A greenish-yellow dye can be made from the stem, leaves, and flowers. American Indians used this and other species to obtain tan to rust colors (Tull 1987).



SCROPHULARIACEAE LINARIA8. LINARIA Mill. Toadflax


Glabrous annual or perennial herbs. Stems erect or spreading, usually simple, some (not ours) coarse. Leaves alternate or the basal leaves opposite (rarely whorled), all sessile, pinnately veined, generally entire. Inflorescence a terminal raceme or spike, with reduced, alternate bracts; bracteoles absent. Flowers often showy. Calyx deeply parted, with 5 subequal segments. Corolla yellow (but not in ours) or blue-violet to white, strongly zygomorphic and bilabiate, the tube ventrally spurred at the base, upper lip 2-lobed, external in bud, lower lip often arched up into a palate, partially closing the throat, its 3 lobes usually reflexed. Stamens didynamous, each anther with 2 divergent, glabrous cells. Stigma solitary and capitate. Capsule ovoid to cylindric or globose, thin-walled, irregularly loculicidal at the apex. Seeds usually many, angled, tuberculate, or with thin wings.

About 100(150?) species in N. temperate regions of the world, especially Mediterranean Europe; 2 species in TX; 1 here with 2 similar but distinguishable varieties. Our plants treated in Nuttallanthus by Kartesz (1998)

Some, especially L. vulgaris and L. maroccana, are cultivated for ornament. A few used locally in medicines (Mabberley 1987). The common name refers to the flax-like leaves and the irregular (toad-like?) shape of the corolla. The flowers, when very fresh, can be made to snap open and shut like miniature Snapdragons.


1. L. canadensis (L.) Dum. Two varieties as distinguished below. Some sources, e.g. Kartesz (1998), place this species in the genus Nuttallanthus.


1. Surfaces of seed smooth and the angles usually thin and sharp; corolla less than 1 mm long, exclusive of the spur, which is 2 to 6 mm long .............................................................................................................1a. var. canadensis

1. Surfaces and the rounded angles of the seeds densely tuberculate (use a strong lens); corolla more

than 1 cm long, exclusive of the spur, which is 5 to 9 mm long .........................................................1b. var. texana


1. var. canadensisOld-field Toadflax. Annual or winter annual; flowering stems slender, to 7 dm tall, erect to ascending, with short, spreading, prostrate branches often present at base (these with their small leaves sometimes mistaken for compound leaves), lowermost leaves opposite-approximate, oblanceolate to spatulate, 0.5 to 1 cm long, 2 to 3.5 mm broad, stem leaves alternate, sparse upwards, 1 to 2.5(3) cm long, 1 to 2(3) mm wide, narrowly linear. Raceme congested at first, elongating in fruit; pedicels 1 to 7 mm long. Calyx segments 2.8 to 3.4 mm long, broadly lanceolate; corolla (in ours) pale blue-violet with a lighter palate, 8 to 10 mm long, excluding the spur, which is 2 to 6 mm long. Capsule 2.8 to 3.5 mm long, splitting from the top downwards for 1/3 to 1/2 its length; seeds smooth or only sparsely tuberculate, the angles generally thin and sharp. Sandy-grassy areas in open woods, rare on caliche. E. 1/3 of TX; widespread from SE. Can. and ME to SD, S. to FL and E. TX. Mar.-May. Treated as Nuttallanthus canadensis (L.) D.A. Sutton by Kartesz (1998) [Reeves (1977) incorrectly listed the plant as L. canadense].


1b. var. texana (Scheele) Penn. Texas Toadflax. Annual or winter annual; flowering stems to 7 dm tall, erect, with short, spreading, prostrate branches often produced at the base (with all their narrow leaves, these sometimes look at first like compound leaves), leaves opposite to approximate, narrowly linear or at most oblong to spatulate, 0.5 to 3 cm long, 1 to 3.5 mm wide, somewhat sparse on the upper portion of the stem. Pedicels 1 to 7 mm long. Calyx segments broadly lanceolate, 2.8 to 3.4 mm long. Corolla (in ours) pale blue-violet, (7)10 to 14 mm long, exclusive of the spur, which is 5 to 9(11) mm long and curved downward. Capsule 2.5 to 3.5 mm long, rupturing at maturity from the top downward for 1/3 to 1/2 its length; seeds with densely tuberculate faces and angles, the angles rounded. Sandy soils in fields, open woods, grassy pine forests, and also roadsides, vacant lots, etc. Common. E. 2/3 TX, rare westward; widespread from B.C. to Mex., E. to the Atlantic Coast through Midwest and SE. states. Feb.-May. Treated as Nuttallanthus texanus (Scheele) D. A. Sutton by Kartesz (1998). [L. texana Scheele].



SCROPHULARIACEAE LEUCOSPORA9. LEUCOSPORA Nutt.


A monotypic genus.


1. L. multifida (Michx.) Nutt. Low, taprooted annual herb; stems erect, diffusely branched, plants to ca. 2 dm tall and about as wide; herbage puberulent to pubescent or hispid throughout. Leaves opposite, petiolate, 1 to 3 cm long, triangular-ovate in overall outline, pinnately (to bipinnately) parted with 3 to 7 linear-cuneate to oblanceolate, ascending divisions. Flowers relatively small, borne singly or in pairs in the axils of the leaves; pedicels 5 to 10 mm long, without bracteoles. Calyx deeply 5-parted, the equal, subulate to lanceolate or linear lobes united only at the base, 2.8 to 4 mm long at anthesis and to 5.5 mm long in fruit; corolla bilabiate, tubular, pink to pale lavender, the throat and tube greenish-yellow within and with a purplish ring at the mouth, 3 to 5 mm long, only slightly longer than the calyx, lips shorter than the tube, the lobes slightly spreading, upper lip 2-lobed, external in bud, lower lip 3-lobed, the lobes longer than the upper ones. Stamens 4, included, didynamous, the anthers approximate, anther cells more or less parallel; style 2-lobed; stigmas cuneiform. Capsule ovoid, acute, thin-walled, septicidal; seeds many, oval to oblong, pale green-yellow, ridged and grooved when seen under high magnification. In mud and sandy gravels of river and stream banks and shores. Primarily E. and Cen. TX; Ont. to OH to IA and KS, S. to GA, AL, MS, LA, and TX. June-Oct. [Conobea multifida (Michx.) Benth.].



SCROPHULARIACEAE SCOPARIA10. SCOPARIA L. Broomwort


About 20 species, primarily in tropical America. The one TX species has been found in our area.


1. S. dulcis L. Sweet-broom, Sweet Broomwort. Annual or perennial herb; stems erect, 2 to 10 dm tall, often much-branched, glabrous to pubescent, if branches slightly pubescent, especially so at nodes. Leaves opposite or in whorls of 3 or 4, short-petiolate, ovate-lanceolate to rhombic-ovate, coarsely serrate (at least in the distal 1/3 to 2/3) or the upper leaves nearly entire, apices acute, bases narrowed or cuneate, glandular punctate, lower leaves 2 to 4 cm long, ca. 6 to 16 mm broad. Flowers axillary, often in most of the axils, usually solitary; pedicels as long as or shorter than the petioles of the subtending leaves. Calyx lobes 4, oblong to obovate to elliptic, acute, sometimes ciliate, ca. 1.5 to 2 mm long, equaling or longer than the calyx tube; corolla white to light lavender or sometimes pinkish, essentially rotate and regular, 3 to 4 mm broad, 4-cleft, densely bearded-lanate within, the lobes about equal, obtuse, ca. 1 mm long; stamens 4, subequal, filaments filiform, anther sacs parallel or divergent; style clavate apically, stigma truncate or notched. Capsule subglobose to globose-ovoid or widely ellipsoidal, ca. 2 mm long or broad, slightly longer than the calyx lobes, septicidal, valves entire, membranaceous; seeds many, angular. Disturbed moist to dry soils, especially sands, often in open woods. E. TX; S. U.S. to S. Amer.; also Old World tropics. According to Pennell (1935), probably only adventive in the U.S. Apr.-Nov. Very few specimens seen from our area.

This pan-tropical weed is used to sweeten well-water. (Mabberley 1987).



SCROPHULARIACEAE MECARDONIA11. MECARDONIA R. & P.


Perennial herbs. Stems much-branched, diffuse or erect, 4-angled or -winged, often dull yellow-green and commonly turning black upon drying. Leaves opposite, tapered to a sessile base or shortly wing-petioled, more or less glandular-punctate, toothed, at least on the distal margin. Flowers solitary in the axils of leaves or foliaceous bracts, each slender pedicel with 2 bractlets at its base, these smaller than the bracts. Sepals 5, free or nearly so, rather unequal, the outer ones wider than the inner. Corolla bilabiate, the tube somewhat quadrangular and slightly enlarged upward, abruptly expanded into a slightly 2-lipped limb, the throat closed by a palate, upper lip 2-lobed, somewhat pubescent within, lower lip 3-lobed, all the lobes shorter than the tube, truncate and usually emarginate. Stamens 4, didynamous. Ovary cylindric to ovoid, acute, style short, stigma lip-like. Capsule glabrous, septicidal, slightly loculicidal at the apex. Seeds many, cylindric, reticulate.

About 15 species in the warmer parts of the Americas. We have both species found in TX.


1. Corolla yellow; leaves generally ovate to oval, abruptly cuneate, usually less than 2.5 cm long; 3 outer sepals broadly lanceolate to ovate, overlapping the shorter, narrower, inner ones ...1.M.procumbens

1. Corolla white, often veined purple; leaves generally oblanceolate, gradually tapered to base, usually 2.5 cm or more long; sepals lanceolate, subequal, the inner 3 only slightly narrower than the outer ...2.M.acuminata


1.M. procumbensSCROPHULARIACEAE MECARDONIA procumbens (P. Mill.) Small Prostrate Mecardonia. Plants variable in form, glabrous throughout; stems prostrate to decumbent or weakly erect, generally branched and wide-spreading, to ca. 1 to 4 dm long. Leaves sessile, oval to ovate or obovate or elliptic, obtuse to subacute, abruptly cuneate at the base, serrate to serrulate or crenate above the middle, 1 to 2.5(3) cm long. Pedicels slightly shorter than to 5 times longer than the subtending leaves, solitary in the axils; basal bracteoles lanceolate to oblanceolate, to ca. 5 mm long. Calyx 4 to 9 mm long, the outer 3 sepals ovate to broadly lanceolate, overlapping the 2 narrower inner ones; corolla 6 to 12 mm long, exceeding the calyx, bilabiate, (bright) yellow with darker lines on the more-or-less-united upper pair of lobes. Capsule ellipsoid, 4 to 5 mm long, shorter than to about as long as the calyx. Mud and water of ditches, streams, lagoons, ponds, and depressions. Cen., S., and W. TX; FL to LA, TX, AR, and Mex.; S. to S. America. Generally Mar.-Nov., some of ours as early as Feb. [M. vandellioides (Kunth in H.B.K.) Penn.; M. peduncularis (Benth.F) Small; M. viridis Small; M. montevidensis (Spreng.) Penn.; Pagesia peduncularis (Benth.) Penn.; P. vandellioides (H.B.K.) Penn.; some local material from the first half of this century is erroneously labeled Bacopa procumbens (Mill.) Greenm.].


2.M. acuminataSCROPHULARIACEAE MECARDONIA acuminata (Walt.) Small Purple Mecardonia, Sawtooth Water-hyssop. Glabrous perennial from a somewhat woody crown; stems erect or sometimes ascending or prostrate, simple to much-branched, 4-angled, 1 to 6.5(7) dm tall. Leaves sessile (sometimes obscurely petiolate), firm, oblanceolate to elliptic or oblong-lanceolate, tapered to a cuneate base, apically serrate and narrowly obtuse, overall (1)2 to 4(5) cm long, 5 to 12 mm wide. Flowers solitary in the axils of the leaves; pedicels slender, 7 to 35 mm long, generally longer than the subtending leaves and the 2 small, linear bractlets at their bases. Flowers bilabiate; sepals 6 to 9(10) mm long, subequal in length but the outer 3 oblong-lanceolate, wider than the 2 lanceolate inner ones; corolla white, commonly tinged or veined with purple or lavender and/or with a yellowish throat, 6 to 11 mm long, the upper lip noticeably bilobed, bearded within at the base, lower lip 3-lobed. Capsule 4 to 8 mm long, narrowly ovoid or ellipsoidal. Bogs, ditches, and low spots in fields and prairies, swamps, and pinelands. E. TX, known from our area, especially Robertson Co.; DE and MD, S. to MO, KS, TN, VA, LA and E. TX. Aug.-Oct. [Bacopa acuminata (Walt.) Robins.; Pagesia acuminata (Walt.) Penn.].

Some sources recognize varieties.



SCROPHULARIACEAE BACOPA12. BACOPA Aubl. Water-hyssop


Low herbs, mostly perennial, succulent or nearly so, some (but not ours) aromatic when crushed. Stems prostrate or creeping, sometimes floating, often mat-forming. Leaves opposite, sessile or nearly so, entire, glandular punctate (sometimes only obscurely so.) Flowers solitary (uncommonly in pairs) in the axils of the leaves, often only 1 leaf of each pair with a flower. Pedicels in some species (as in one of ours) with 2 linear bractlets subtending the calyx. Calyx of (4)5 sepals, nearly free, equal in length but of different widths (sometimes appearing to be in 2 series)--the uppermost the largest, the 2 lowest intermediate, and the 2 lateral the smallest, all sometimes enlarging in fruit. Corolla overall white to blue, tubular-campanulate, with (3,4)5 subequal, spreading lobes, upper lip entire to notched or 2-cleft and the lower lip 3-lobed. Stamens usually 4, didynamous (sometimes 3 or 2), the filaments glabrous. Ovary globose to ovoid or elliptic-cylindrical and septicidal, loculicidal, or both, usually equal to or shorter than the persistent calyx. Style with 2 terminal stigmas or stigma 1, dilated and nearly capitate. Seeds many, alveolate-reticulate.

About 50 to 100 species in warm regions of the world, especially the Americas; 3 species in TX; 2 expected here.

A few species are cultivated in aquaria.


1. Leaves basally cuneate, 1-nerved; pedicels with 2 bracteoles just below the calyx ..............

...1.B.monnieri

1. Leaves suborbicular to obovate, palmately many-nerved; pedicels without bractlets .............

...2.B.rotundifolia

1.B. monnieriSCROPHULARIACEAE BACOPA monnieri (L.) Penn. Coastal Water-hyssop. Plants glabrous, fleshy; stems with the potential to be highly-branched, main stems radiating outward, prostrate, rooting at the nodes, mat-forming; flowering branches decumbent to slightly ascending, 1 to 3 dm tall. Leaves obovate, spatulate or oblanceolate, cuneate at the base, succulent and glabrous, 5 to 17 mm long, 3 to 8 mm wide, obtuse to rounded apically, the margins entire or those of the upper leaves somewhat slightly wavy, each with only one conspicuous nerve; internodes commonly longer than the leaves. Flowers solitary in the axil of one leaf of some pairs; pedicel shorter than to slightly longer than the subtending leaf, elongating to 1 to 2.5 cm in fruit; bractlets at apex of pedicel linear, 2 to 3 mm long. Sepals unequal, the largest becoming 5 to 7 mm long and 3 to 4(5) mm broad, lanceolate to ovate; corolla campanulate to rotate, only slightly bilabiate, 8 to 10 mm long, lobes about equalling the tube, glabrous within, the tube greenish to pale yellow, the limb white to pale blue-violet, lavender or pink, with a darker ring within the tube at about 1/2 its length; stamens 4, subequal, but the longer pair slightly exserted. Capsule 4 to 7 mm long, more or less shorter than the calyx, narrowly conic to slightly ovoid; seeds olivaceous, short- or obliquely-oblong, ca. 0.5 mm long, alveolate-reticulate, with the longitudinal striations the more prominent and the openings between striations longer than wide. Mud and sand in low spots, among dunes, around ponds, streams, ditches, etc. E., Cen., and S. TX; FL to TX, N. to VA; also found in the tropics and subtropics outside the U.S. Apr.-Sept. [Includes var. cuniefolia Fern.; Bramia monnieri (L.) Penn.]


2.B. rotundifoliaSCROPHULARIACEAE BACOPA rotundifolia (Michx.) Wettst. Disc Water-hyssop. Plants aquatic or semi-aquatic; stems to 6 dm long, soft, spongy, prostrate, creeping, or floating, rooting at the nodes, pilose to hispid in the younger, terminal portions, the tips ascending or floating. Leaves sessile, entire, orbicular to broadly ovate-cuneate, 12 to 27(40) mm long, (8)12 to 25 mm wide, with several conspicuous palmate veins, clasping or nearly so, bases rounded or sometimes broadly subcuneate, apically broadly rounded. Flowers 1 to 4 in the axils of most of the upper leaves; pedicels usually shorter than the subtending leaves, 6 to 17(23) mm long, without bractlets at the apex. Calyx pubescent, 1/2 to 1/3 as long as the pedicel, segments 3 to 4.5 mm long at anthesis and elongating to ca. 8 mm in fruit, the outer ovate to nearly round and the inner 2 lanceolate; corolla white with a yellow throat, exserted from the calyx, campanulate, 6 to 10 mm long, the limb about as broad, the 5 lobes subequal, slightly shorter than the tube; stamens 4, somewhat exserted, anther cells parallel. Capsule globose to ovoid, about as long as the sepals, to ca. 5.5 mm long; seeds many, tan, ellipsoid or falcate-cylindrical, minutely tailed at either end, alveolate-reticulate, ca. 0.5 mm long. Mud and water around lakes, pools, ditches, ponds, etc. Throughout TX; IN and IL to MN, MT, and ND, S. to MS, LA, and TX; also ID and CA. May-Nov. [Macuilliama rotundifolia (Michx.) Raf.; Bramia rotundifolia (Michx.) Britt.; Hydranthelium rotundifolium (Michx.) Penn.].



SCROPHULARIACEAE AUREOLARIA13. AUREOLARIA Raf. False Fox-glove, Oakleech


Annual or perennial herbs, said to be parasitic on Quercus (Fagaceae in general, sometimes Ericaceae), usually turning black upon drying. Stems branched or rarely simple. Lower leaves opposite, entire to deeply pinnately lobed, becoming smaller and irregularly alternate up the stem. Flowers showy, solitary and pedicellate in the axils of the uppermost leaves or bracts. Calyx 5-parted, campanulate to cup-shaped, lobes shorter than to longer than the tube, triangular to linear, often unequal. Corolla yellow, zygomorphic but weakly bilabiate, the tube campanulate, somewhat oblique, the limb 5-lobed, the lobes shorter than the tube, distinct, equal, rounded, spreading, the lower 3 external in bud. Stamens included, 4, didynamous, the lower pair the longer, filaments flattened, slender, at least the lower pair villous or lanate; anther sacs parallel, basally awned, pubescent. Style slender, stigma 1, capitate, exserted. Capsule ovoid to ellipsoid, loculicidal, seeds few to several, winged or not so.

About 10 species of the SE. U.S., with 1 in Mex.; 4 in TX; only 1 here.

Some species are cultivated for ornament.


1. A. grandiflora (Benth.) Penn. Perennial herb to 15 dm tall, widely branched, the foliage cinereous-pubescent. Leaves opposite, the lowermost ovate in overall outline, more or less pinnatifid, the divisions themselves often toothed or lobed, leaves progressively reduced and less pinnate upward, the very uppermost lanceolate to narrowly ovate, serrate or even entire. Flowers opposite or subopposite in the axils of bracteal leaves, these entire to serrate or laciniate; pedicels stout, upcurved, 5 to 14 mm long at anthesis, densely and minutely pubescent; flowers large, showy, weakly bilabiate. Calyx with pubescence similar to that of the pedicels, synsepalous, campanulate or tubular to hemispheric, overall 10 to 15(18) mm long, 5-lobed, the lobes entire to dentate, lanceolate or triangular, usually shorter than the tube, often unequal, the longer ones ca. 5 to 11 mm long; corolla yellow, campanulate, 38 to 50 mm long, throat open, lobes broadly rounded, spreading, ciliolate. Capsule glabrous, ovoid, acute to

acuminate, 1.5 to 2 cm long; seeds few to several, winged or wingless. Dry open woods and along streams, mostly in the forests of E. TX; IN to MN and IA, S. and W. to LA and TX. June-Oct. [Gerardia grandiflora Benth.].

There are 3 varieties in TX as follows. We definitely have the first, and the other two are mentioned for completeness--they may yet be found in our area.


var. grandiflora Bigflower Oakleech. Petioles mostly 5 to 6 mm long; floral bracts coarsely toothed; calyx lobes broadly lanceolate, decidedly dentate. All of our material seems to be of this variety.


var. serrata (Torr. ex Benth.) Penn. Petioles generally 10 to 15 mm long; upper leaves similar to the serrate to entire bracts, abruptly smaller than the lower leaves; pedicels somewhat slender, 7 to 13 mm long in fruit; calyx lobes linear to lanceolate, entire to slightly dentate.


var. cinerea Penn. Petioles generally 10 to 15 mm long; upper leaves only gradually smaller than the lower; bracts entire to shallowly serrate, lanceolate and acuminate-attenuate; pedicels relatively stout, 5 to 9 mm long in fruit; calyx lobes linear to lanceolate, entire to slightly dentate. [Gerardia grandiflora Benth. var. cinerea (Penn.) Cory.].



SCROPHULARIACEAE MAZUS14. MAZUS Lour.


About 20 to 30 species, primarily from Australasia, a few naturalized elsewhere. We have one of the two species naturalized in the U.S. (The other is M. miguelii Makino, rare in N.C.)


1. M. pumilus (Burm. f.) Steenis Plants erect, slender; stems to ca. 2 dm tall; herbage loosely pubescent to minutely puberulent. Leaves opposite, mostly crowded at or near the base of the plant, obovate to cuneate-oblong, abruptly tapered to a narrow subpetiolar base, coarsely dentate with few teeth, basal leaves 2 to 5 cm long. Flowers alternate, to 7 or more in loose, terminal, somewhat secund racemes; floral bracts minute or obsolete; bracteoles minute or absent; pedicels ca. 5 to 7 mm long, elongating in fruit. Calyx with 5 lanceolate to ovate-lanceolate lobes, about equal to or a little longer than the campanulate tube; corolla bilabiate, lavender-blue to pale blue, the palate yellow or whitish and bordered in reddish-brown, 7 to 10(11) mm long, barely twice as long as the calyx, pubescent within, upper lip slightly bifid, erect, lower lip 3-lobed and much larger than the upper, with 2 swellings in the throat; stamens 4, included, anther cells divaricate. Capsule 3 to 4 mm long, roughly globose, shorter than the calyx, with 2 entire valves. Grasslands and roadsides, lawns, etc. E. TX; PA to MO, KS, LA, and TX, also OR and WA; naturalized from E. Asia. Mar.-Oct. [M. japonicus (Thunb.) O. Ktze.].

When Pennell wrote his treatment of the Scrophulariaceae in 1935, this plant was found in the vicinity of a few cities. It has since spread rather widely.

The flowers of this small plant are charming, somewhat reminiscent of Viola or some Salvia species. It makes a nice addition to the garden as a rock garden plant or ground cover.


SCROPHULARIACEAE MIMULUS15. MIMULUS L. Monkey-flower


Annual or (as in ours) perennial herbs, the perennial species with rhizomes or stolons. Stems erect or ascending, sometimes prostrate, branched or unbranched. Herbage often glandular-pubescent to viscid or pilose, sometimes glabrous. Leaves opposite, pinnately or palmately veined, petiolate or sessile, entire to toothed or laciniate, often basically lanceolate. Flowers usually solitary in the axil of each leaf or in a bracted raceme; bracts leaf-like and progressively smaller upwards, bracteoles absent. Pedicels often well-developed. Flowers slightly to strongly bilabiate. Calyx synsepalous, tubular to campanulate or often inflated, angled or prismatic, essentially pleated, equally or unequally 5-lobed, the lobes markedly shorter than the tube. Corolla yellow to red, purple, or bluish, bilabiate, campanulate, the upper lip with 2 erect to reflexed lobes external in the bud, the lower lip 3-lobed, spreading or deflexed, 2 ridges present in the throat to form a palate sometimes partially or wholly closing the orifice, interior usually bearded. Stamens 4, didynamous, inserted near the middle of the corolla tube, anthers approximate in pairs, the sacs of each anther divergent. Style elongate, with 2 platelike stigmas which close together after being touched by an insect. Capsule enclosed by the persistent calyx, thin-walled to coriaceous, cylindrical, loculicidal, sometimes also splitting down the septum dividing the placentae, glabrous. Seeds many, generally yellowish, oblong to oval of fusiform, reticulate with the "cells" in rows or varying to nearly smooth.

About 150 species, primarily of the W. U.S., also S. Afr. and Asia; 5 species are foundin TX; 1 here.

Some are cultivated for ornament, especially M. x hybridus Hort. ex Sieb. & Voss, which comes in a wide array of colors and grows best in moist, shady places (Mabberley 1987).


1. M. alatus Ait. Sharpwing Monkey-flower. Glabrous, stoloniferous perennial; stems erect to ascending, 4-angled, the angles more or less winged, 3 to 7 dm tall, simple or branched above. Leaves opposite, petiolate, broadly ovate to lanceolate, 5 to 8(15) cm long, ca. 2 to 6 cm wide, acute, tapered below to a narrow-margined petiole to 25 mm long, margins serrate, sometimes scabrous-ciliolate. Pedicels somewhat stout, shorter than the calyx, 2 to 8 mm long in flower and to 14 mm long in fruit. Calyx 11 to 18 mm long, not inflated, sharply angled-pleated, puberulent within, the lobes essentially equal, broad, each 0.8 to 2.6 mm long, with an extension of the midnerve forming an awn on each, the margins sometimes scabrous-ciliolate; corolla blue to violet, sometimes tinged pink or lavender, rarely white, 2 to 2.8 cm long, the tube slightly exserted from the calyx and the throat nearly closed by the palate, lobes of the upper lip erect and markedly reflexed, the longer lower lip spreading, pubescent inside and out; stamens and style included, anthers 0.8 to 1.3 mm long. Capsule ovoid, obtuse, 8 to 11 mm long, loculicidal at the apex; seeds oblong, ca. 0.3 mm long, light brown, papillate or finely reticulate. Wet areas, especially along woodland streams, etc. E. TX, W. to the Blackland Prairie; S. Ont., NY, and CT to NE, S. to FL and TX; known in our area at least from Leon Co. June-Nov.



SCROPHULARIACEAE BELLARDIA16. BELLARDIA All.


A monotypic genus adventive from the Mediterranean region.


1. B. trixago (L.) All. Annual from taproot and equally large secondary roots; stems erect, in plants of our area ca. 20 to 50 cm tall, simple or branched near the inflorescence, roughly quadrangular with the angles rounded, hairs reflexed; herbage viscid-pubescent with short, pale hairs. Leaves opposite, sessile, simple, lanceolate, the margins with widely-spaced, spreading-ascending acute teeth or small lobes, to ca. 7 cm long and 1 cm broad (excluding the teeth), cinereous viscid-pubescent, the hairs ascending and a small tuft present at the base of each leaf, margins (at least in dried plants) slightly revolute, the midvein and secondary veins prominent beneath. Inflorescences dense spikelike racemes, terminating the main stem and each of the branches in the upper part of the plant, the main spike to ca. 7 to 15 cm long and 2 cm broad, excluding the bracts; lowermost floral bracts similar to the leaves, lanceolate-acuminate, viscid-pubescent, bracts becoming progressively ovate upwards, ca. 1 to 1.5 cm long and 1 cm broad, and increasingly covered with stalked glands. Calyx ca. 1 cm long, densely covered with stalked glands, overall campanulate-urceolate, with 4 lobes or teeth, teeth less than half the length of the tube, unequal and the sinuses between unequal; corolla white-lavender, markedly zygomorphic, ca. 2 to 3 times longer than the calyx, upper lip slightly galeate, of 2 fused lobes, entire, stalked-glandular pubescent externally, the lower lip 3-lobed, the lobes spreading-reflexed, median lobe ovate-oblong, the apex bluntly acute, the ear-like lateral lobes overlapping the median, their lower margins elongate and rounded, upper margins straight to slightly curved, ending in a blunt, acute tooth (overall lobe shape approximately half-reniform), somewhat stalked-glandular pubescent within and on the margins; stamens 4, didynamous, anther cells parallel, anthers bearded below and at the apex; style slender, to 1.5 cm long, gradually expanded into the stigma. Capsule densely pubescent, ovoid; seeds oblong, with longitudinal lines. Native to the Mediterranean area, including Europe; also known from S. Afr. Colonies exist in the Mideast countries, Ethiopia, Africa, Australia, and parts of N. and S. Amer. Formerly known only from old fields and grassy areas around San Francisco Bay. Discovered in TX in 1980 in Navarro Co. on the Blackland Prairie; known in our area and apparently increasing in range (Do, et al. 1996). April (to June.) For illustration and more information, see Lipscomb and Ajilvsgi (1982). A population of very short, depauperate plants exists along FM 2818 in Bryan, Brazos Co.

Large stands of the plant are undoubtedly ornamental and would do well in a garden.




SCROPHULARIACEAE AGALINIS17. AGALINIS Raf. Gerardia


Annuals (including all TX species) or perennial parasitic or hemiparasitic herbs, most turning black on drying. Stems erect, 4-angled by decurrent lines from the leaf bases or else subterete, branched, glabrous or with varying degrees of ascending scabrous pubescence. Leaves opposite, linear to filiform, sometimes lanceolate, in some species also clustered on short axillary shoots or in axillary fascicles, leaves usually entire or one species with some of the largest lower leaves 3-cleft, upper surface commonly scabrous to scabridulous, the lower surface various, often glabrous, margins sometimes revolute. Inflorescence a terminal, bracted raceme or in 1 species the racemes grouped and more or less paniculate; bracts leaf-like below and reduced upwards, often becoming alternate, often only one of a pair of flowers present. Pedicels short and clavate to long and filiform, often enlarged just below the flower. Flowers relatively large and showy, slightly to strongly irregular. Calyx synsepalous, campanulate to hemispheric, the tube shorter than to usually longer than the 5 lobes or teeth, lobes or teeth triangular to lanceolate or subulate, acute to acuminate, sometimes the posterior one longer than the other 4. Corolla sympetalous, somewhat inflated, pink-purple or (rarely) white (almost never so in local material), commonly with 2 yellow lines and purple or reddish spots within the open throat, tube more or less campanulate, often distended on the lower side, lobes 5, often marginally ciliate, truncate to emarginate or rounded, subequal or the upper 2 smaller, upper lobes spreading to reflexed or in some species projecting, the lower 3 spreading, external in bud. Stamens 4, didynamous, the lower pair the longer, filaments pubescent at least basally; anther cells 2 per stamen, obtuse to cuspidate or caudate basally, villous. Stigma solitary, rather elongate, flattened. Capsule globose to cylindric or ovoid or ellipsoidal, loculicidal. Seeds many, triangular or with rounded edges, reticulate to varying degrees.

About 40 to 60 species in temperate America, depending upon what one allows the genus to include; 16 in TX; 7 possible here. Formerly Gerardia, a genus which included plants now in several genera. Plants formerly in Tomanthera have been returned to Agalinis by some authors--this would bring the TX total to 18 species.

A. purpurea is sometimes cultivated for ornament; all our local species have some ornamental value.

Agalinis, at least locally, is a confusing genus. Because plants do not make good herbarium specimens (most turn black) and because Reeves (1977) included only two species (neither of them common here), there exists a wealth of misidentified specimens. Without fresh material, some of these puzzles are virtually irresolvable.


1. Pedicels mostly less than 1 cm long, stout to slender, equal to or shorter than the calyx at anthesis (sometimes a few longer) ..........................................................................................2

1. Pedicels (or at least some) more than 1 cm long, generally filiform, always longer than the calyx at anthesis .......................................................................................................................4


2(1) Calyx lobes nearly as long as or longer than the tube (generally 3 to 6 mm), sinuses between the lobes narrow; largest leaves linear, 2 to 6 mm broad ...1.A.heterophylla

2. Calyx lobes much shorter than the tube, generally less than 3 mm long, sinuses broad and shallow; leaf blades narrowly linear to subulate, at most 2 to 4 mm broad ...................3






3(2) Axillary fascicles well-developed; primary leaves usually less than 2.5 cm long, erect- ascending or arcuate-ascending, somewhat appressed; stem subterete, scabridulous; stigma 3 to 4 mm long ...2.A.fasciculata

3. Axillary fascicles poorly-developed or absent; primary leaves to 4 cm long, spreading or arcuate-ascending; stems 4-angled, smooth to scabridulous; stigma 2 to 3 mm long ..........

...3.A.purpurea


4(1) Upper 2 corolla lobes arched forward or held flat over the stamens .....................................5

4. Upper 2 corolla lobes spreading or reflexed ...........................................................................6


5(4) Upper corolla lobes less than half as long as the lower, flattened and projecting forward; axillary fascicles well-developed; stem ridges and sides conspicuously scabrous-

scabrellous ...4.A.homalantha

5. Upper corolla lobes more than half the length of the lower, concave-arched; axillary fascicles absent or few; stem glabrous or nearly so ...5.A.tenuifolia

subsp. leucanthera


6(4) Largest leaves narrowly to broadly linear, flat; calyx ribbed on the major veins; corolla strongly upcurved; inflorescence racemose, often secund; leaves of flowering branchlets generally appressed ...6.A.strictifolia

6. Largest leaves filiform, grooved on the upper surface; calyx not ribbed; corolla slightly upcurved; inflorescence often racemose-paniculate; leaves of flowering branchlets spreading-reflexed ...7.A.navasotensis


1.A. heterophyllaSCROPHULARIACEAE AGALINIS heterophylla (Nutt.) Small ex Britt. Prairie Agalinis. Plants dark green, often tinged with purple, turning black on drying (except sometimes in very dry years); stems to 1 m tall, glabrous to scabridulous, the upper branches stiff and paniculately spreading or virgate. Leaves generally linear-lanceolate to lanceolate, somewhat erect, thickish, scabrous above and on the margins, the larger leaves 1.5 to 3(4.5) cm long, 2 to 6(8) mm wide, attenuate to mucronate-acute, the lowermost sometimes 3-cleft or laciniate (but these, if produced, often absent at anthesis), leaves of upper branches short and somewhat subulate. Pedicels short, 1 to 3 mm long, ascending, not lengthening much in fruit. Calyx campanulate, (5)6 to 10 mm long, the tube 3.5 to 6.5 mm long, lobes triangular-lanceolate, acute to attenuate, somewhat spreading in age, 3 to 6 mm long, usually equal to or obviously longer than the tube on many of a plant's flowers (this character alone is enough to distinguish this species in our area most of the time); corolla some shade of pink-purple, purple-spotted within and with 2 yellow lines in the throat, 2.5 to 3 cm long, tube somewhat gibbous on the lower side, the lobes 5 to 9 mm long, broadly rounded, spreading; anthers 2.8 to 3.5 mm long. Capsule subglobose or slightly longer than wide, 5 to 8 mm long; seeds dark brown to black, ellipsoid or trapezoidal, 0.8 to 11 mm long, reticulations deep and irregular. Prairies, along streams, roadsides, vacant lots, open woods, etc. In TX, E. of a line from Grayson to Cameron Cos.; S. MO and E. KS, S. to LA, W. AL, and E. TX. June-Oct. (Nov.) [G. heterophylla Nutt., G. crustata Greene].

Far and away our most common Agalinis. Unfortunately Reeves (1977) omitted this species, so many older specimens of this plant are misidentified as A. purpurea, a species he did include.


2.A. fasciculataSCROPHULARIACEAE AGALINIS fasciculata (Ell.) Raf. Stout dark green herb, darkening on drying; stems 4 to 7(10) dm tall, scabrous-pubescent, main stem nearly terete to weakly 4-angled, branches angled; axillary fascicles usually well-developed. Leaves erect-ascending or curled or arched, sometimes closely arcuate-ascending, scabrous, entire, linear, 1 to 3.5 cm long, 1 to 2 mm wide, leaves of the fascicles smaller, bracteal leaves in the inflorescence also smaller, shorter than the flowers. Racemes elongate, usually 12- to 30-flowered; pedicels 2 to 4(4.5) mm long at anthesis, shorter than the calyx, spreading-ascending. Calyx broadly campanulate to hemispheric, tube 3 to 5(5.5) mm long, lobes acuminate, 0.4 to 2 mm long, sinuses subquadrate to broadly rounded; corolla rose-pink, (16)20 to 35 mm long, lobes 5.5 to 10 mm long, spreading, rounded to truncate; anthers 2.5 to 3.5 mm long, with acute to cuspidate bases. Capsule globose-ovoid, 4.5 to 6(7) mm long; seeds dark brown to blackish, trapezoidal to triangular, 0.7 to 1.1 mm long, reticulae elongate and irregular. Moist or dry soils in savannahs, open wooded areas, and dune hollows and tidal marshes. E. and N. Cen. TX; FL to TX, N. to VA, SE. KS, MD, MO, and AR. Sept.-Oct. [G. fasciculata Ell. and forma albiflora E. J. Palm.].


3.A. purpureaSCROPHULARIACEAE AGALINIS purpurea (L.) Penn. Dark green herb, blackening on drying; stems (1.5)4 to 12 dm tall, usually in the shorter end of the range, glabrous to sparsely scabridulous, commonly 4-angled; branches numerous above, virgate and wide-spreading, 4-angled; some axillary fascicles often present. Leaves wide-spreading or arcuate-ascending, somewhat scabrous to smooth or merely with scabrous margins, entire, narrowly linear, acute, 1 to 4(5.5) cm long, 0.8 to 2(3) mm wide, leaves of axillary fascicles smaller. Pedicels spreading, shorter than or equalling the calyx, 1 to 5 mm long. Calyx hemispherical, 4 to 5.5(6) mm long, the tube 2 to 4 mm long, lobes triangular-lanceolate, acuminate to acute, 1.0 to 2.2 mm long, generally shorter than the tube, sinuses broadly v-shaped to rounded; corolla rose-pink to pink (sometimes purplish), (17)20 to 35 mm long, the spreading lobes 5 to 9 mm long; anthers 2.5 to 3.2(4) mm long. Capsule globular to subglobose, 4 to 6 mm long; seeds dark brown to black, trapezoidal to triangular, 0.7 to 0.2 mm long, the reticulae irregularly elongate. Moist sandy soils in bogs, seeps, moist prairies, pinelands, and barrens and along pond and lake shores and moist roadsides. E. TX, probably present in the E. part of our area, but usually seen in deep E. TX; ME to MN and NE, S. to the Gulf, FL to TX. Sept.-Oct. [G. purpurea L.].


4.A. homalanthaSCROPHULARIACEAE AGALINIS homalantha Penn. Plants darkening on drying; stems 4 to 7 dm tall, slender; branches many, ascending-spreading, slightly scabridulous. Leaves spreading, linear and acuminate, main cauline leaves 2.5 to 3 cm long and 1 to 1.5 cm wide, becoming smaller upwards, somewhat scabrous on the upper surface; axillary shoots or fascicles well-developed, though shorter than the subtending leaves. Racemes 4- to 10-flowered, elongated; pedicels ascending-spreading, somewhat scabridulous, slender and slightly clavate, 1 to 2 cm long at anthesis and to 3 cm long in fruit, 1.5 to 2.5 times as long as the bracts. Calyx tube 3 to 3.5 mm long, the apex truncate, lobes 1 to 1.2 mm long (rarely much longer), lanceolate, acuminate, the apex of the calyx tube and the interior of the lobes densely puberulent; corolla lavender to pinkish, 2.3 to 2.7 cm long, pubescent externally, the tube straight, 1.5 to 1.8 cm long, upper lobes 3 to 4 mm long, less than half as long as the lower lobes, flat-projecting over the stamens and style, lower lobes 8 to 9 mm long, spreading, all lobes ciliate, corolla pubescent within in a narrow line below the posterior sinus. Capsule ca. 6 mm long, subglobose. Sandy soil of post oak woods, cut-over fields. E. and N. Cen. TX, in the cross-timbers; also S. OK. None seen from our area, but according to Pennell (1935), known from the Brazos Valley in the southernmost part of our area; probably present but uncollected. Aug.-Oct. [G. homalantha (Penn.) Penn.].


5.A. tenuifoliaSCROPHULARIACEAE AGALINIS tenuifolia (Vahl) Raf. var. leucanthera (Raf.) Penn. Plants often darkening on drying, but not always; stems to about 5 dm tall, usually shorter, smooth and glabrous to subglabrous, much-branched. Leaves finely scabridulous to glabrous, spreading to arched-ascending, mostly linear, acuminate, the largest 3 to 7 cm long, 1 to 2 mm wide, about as long as the lower pedicels but generally shorter than the upper; axillary fascicles present or absent. Pedicels widely divergent, filiform, exceeding the calyces, 7 to 20 mm long. Calyx hemispherical, 3.5 to 5.5 mm long, tube 2 to 4 mm long, the lobes broadly triangular to subulate, apiculate, generally less than 1 mm long in this subspecies, sinuses broadly rounded; corolla of this subsp. 15 to 23 mm long, pink to mallow-purple (sometimes paler), glabrous except for cilia edging the subequal lobes, upper lobes concave-arched, projecting over the anthers and stigma; anthers 1.5 to 2.2 mm long. Capsules 3 to 7 mm long, globose; seeds dark brown to blackish, bluntly trapezoidal or triangular, 0.7 to 0.9 mm long, the reticulae irregularly elongate. Moist areas along streams, near ponds, meadows, fields, low woods, etc. Primarily NE. TX; ME to Que, W. to S. Man., E. ND, and E. WY, S. to

FL, E. TX, OK, and E. CO. Sept.-Nov. [G. tenuifolia Vahl. subsp. leucanthera (Raf.) Penn.].


NOTE: A. tenuifolia is a highly variable species with a wide range of corolla sizes; it may also have externally pubescent corollas.


6.A. strictifoliaSCROPHULARIACEAE AGALINIS strictifolia (Benth.) Penn. Plants darkening on drying; stems 5 to 8 cm tall; branches many, rigidly ascending or ascending-spreading, more or less 4-angled and striate, glabrous. Leave scabrous to scabridulous or nearly glabrate on the upper surface, those of stem and main branches spreading to reflexed, 2 to 3.5 cm long, 1 to 3 mm wide, linear and acuminate, leaves of flowering branchlets strongly ascending, more or less appressed to branchlets, linear-subulate, to ca. 1.3 cm long. Racemes slightly elongate, 4- to 10-flowered, often only 1 of each pair of flowers developed; pedicels ascending-spreading, glabrous, slender but expanded just below the receptacle, 1 to 2.5 cm long in flower, 1.5 to 3 cm long in fruit. Calyx tube campanulate or hemispheric, evidently or obscurely rib-veined, apex little if at all truncate, lobes 1 to 2 mm long, triangular-lanceolate to subulate, acuminate, apex of calyx tube and interior of lobes minutely pubescent; corolla 2 to 2.5 cm long, pink-purple to lavender, tube 1.2 to 1.5 cm long, strongly upcurved, lobes 8 to 10 mm long, spreading, truncate to erose, the margins ciliate, externally glabrous except for a line of hairs below the posterior sinus. Capsule about as broad as long, somewhat quadrangular, irregular, to ca. 6 mm long; seeds dark, areas between the reticulations horizontal. Old fields and open areas in scrub and in mesquite plains. Austin and Travis Cos. S. and W. to Rio Grande; also neighboring Mex. Pennell gives records N and E of our area as well as further S. along the Brazos. Not seen but perhaps someday to be collected from our area. Sept.-Nov. and May. [G. strictifolia Benth.].


7.A. navasotensisSCROPHULARIACEAE AGALINIS navasotensis Dubrule & Canne-Hilliker Navasota False-foxglove. Plants medium- to dark-green, often tinged with purple, maroon, or bronze, darkening little if at all if promptly pressed; stems 2.8 to 9 dm tall, minutely scabridulous below and sparsely to moderately so above, branches divaricate. Leaves filiform, spreading to ascending or often recurved, narrowly U-shaped in cross-section, 1.2 to 3(4) cm long, 0.5 to 1(1.2) mm broad, acute to acuminate, margin silicified and scabrous. Inflorescence racemose or commonly a panicle with racemose branches, racemes or branches with up to 8 floriferous nodes, terminated by leaves or bracts and no flowers appearing terminal, when flowers paired, usually not blooming concurrently; pedicels slender, spreading or ascending, (6)8 to 21 mm long at anthesis, always longer than the calyx, elongating to up to 2.5 cm in fruit. Calyx at anthesis obconic to somewhat campanulate or funnelform, straight-sided, apically more or less truncate and with minute teeth, tube 2.2 to 3.7 mm long, 3 to 4.4 mm broad, unribbed, interior with a narrow band of hairs below the sinuses, sinuses broad and straight to slightly concave; corolla lavender- to rose-purple, (13)16 to 25(27) mm long, tube 2 to 3.5 mm long, throat slightly upcurved, lower side slightly gibbous, pilose externally and within in a short, narrow, longitudinal band below the sinuses of the adaxial lobes and in a narrow horizontal band among the filament bases, lobes ca. 6 to 9 mm long, all spreading or spreading-reflexed, subequal or the lower ones longer by 0.5 to 2.5 mm, glabrous except for the densely ciliate margin; anthers densely villous, the halves of the adaxial pair placed slightly one above the other; style to 1.5 cm long, stigma 2 to 4.5 mm long, yellow-papillose. Capsule ovoid or obovoid-oblong, 4 to 7 mm long, 4 to 4.5 mm broad, longer than the calyx; seeds 0.8 to 2.3 mm long, irregularly trapezoidal, testa reticulate the cells irregular polygons. Known at present only from a single calcareous sandstone outcrop in Grimes Co. and from sandy roadsides a few miles distant. Probably a very restricted endemic. Sept.-Oct. First described from our area by Canne-Hilliker and Dubrule (1993).



SCROPHULARIACEAE BUCHNERA18. BUCHNERA L. Bluehearts


Perennial herbs, rough-hairy or pilose-hispid, blackening on drying, apparently root-parasitic. Stems generally simple. Leaves opposite or the uppermost alternate, sessile, elliptic to ovate-lanceolate. Flowers opposite, subtended by bracts in an open terminal spike, each subtended by 2 additional bractlets. Calyx tubular-cylindrical, obscurely nerved. Corolla tube straight or curved, the 5 lobes shorter than the tube, nearly equal, oblong to cuneate-obovate, flat. Stamens 4, included, anthers 1-celled. Style clavate, entire. Capsule many-seeded, ovoid-pyriform, 2-valved and loculicidal.

About 100 species, mostly of the Old World tropics and subtropics; 2 in TX; 1 here.


1. B. americana L. Stems to ca. 6 dm tall, slightly pilose to glabrate, root commonly thickened. Leaves rough-pubescent, 3-veined or not so, the larger ones elliptic to oblanceolate , ovate-lanceolate, or lanceolate, to 10 cm long and 18 mm broad, repand- or sinuate-dentate to entire. Bracts 2 to 3 mm long, spreading. Calyx lobes triangular, acute, to ca. 1 mm long, the anterior sinus the deepest; corolla violet to purple to (rarely) white, tube 7 to 10(12) mm long, lobes triangular-ovate, 3 to 5(8) mm long. Capsule ca. 5(7) mm long, largely enclosed in the persistent calyx. Sandy or gravelly soils in the S. 1/2 of TX; FL to TX, N. to NC. Quite uncommon here. Apr.-Nov. [B. floridana Gand.].





OROBANCHACEAEOROBANCHACEAE

Broomrape Family


Perennial herbs, usually root-parasites of other plants and lacking chlorophyll. Stems fleshy, yellow to brown, white, or purple, single or clustered, commonly with alternate scales rather than leaves. Flowers solitary in the axils of the scales or (as in ours) in simple or bracted spikes. Sepals 2 to 5, united below, equal or unequal, calyx sometimes spathe-like. Corolla tubular, more or less bilabiate, upper lip entire or with 2 lobes, lower lip 3-lobed, corolla withering and persistent. Stamens 4, didynamous, epipetalous on the corolla tube; anthers white, caudate, persistent. Ovary superior, bicarpellate, unilocular. Fruit a 1-celled capsule with 2 valves, each valve with 1 or 2 placentae, each bearing numerous tiny, wind-dispersed seeds.

A family of 17 genera and 230 species of the N. Hemis., especially common in the temperate and subtropical Old World; 3 genera and 7 species in TX; 1 species known from our area.



OROBANCHACEAE OROBANCHE1. OROBANCHE L. Broomrape


Plants as described for the family, usually more or less glandular or pubescent throughout. Stems fleshy, from an underground caudex, ca. 3 to 30 cm tall, purple to brownish or white. Flowers solitary or in spikes. Calyx lobes nearly equal. Corolla purplish to yellowish or reddish, tube slightly curved, limb bilabiate, upper lip commonly nearly erect and the lower recurved. Capsule with 4 placentae.

About 150 species of temperate and warm regions; 5 in TX; 1 collected in our region.


1. O. multiflora Nutt. Plants usually stout, 0.5 to 3 dm tall, the underground portion glabrous, base of stem commonly enlarged; herbage viscid-pubescent, usually grayish. Cauline scale-leaves lance-ovate, obtuse, 5 to 12 mm long. Inflorescence dense, spike-like or thyrse-like, rarely branched and making up about 1/2 to 1/3 the height of the plant, the axis often purple; flowers sessile or the lower with pedicels to 5 mm long, each flower subtended by (1)2 lance-linear, usually opposite bracts adnate to or adjacent to the calyx. Calyx 8 to 17 mm long, lobes erect, lance-linear, attenuate, externally glabrous or viscid-pubescent, often purple or viscid internally; corolla 15 to 35 mm long, tube purple to yellow or whitish, lips 4 to 12 mm long, rose or pale purple to yellow, upper lip erect, lobes 3 to 5 mm long, rounded to mucronate; anthers glabrous or woolly, positioned at about the throat or lower; style 5 to 9 mm long, about as long as the filaments, elongating in fruit, stigma peltate. Capsule equalling or shorter than the calyx, topped by the style, longitudinally dehiscent; seeds angular, dark. Parasitic on members of the Asteraceae: Hymenopappus, Artemisia, Heterotheca, etc.; in sandy soils of beaches, roadsides, prairies, etc. CO to WY and WA, S. to CA, TX, and Mex.

There are two varieties, both treated by Kartesz (1998) as O. ludoviciana Nutt. subsp. multiflora (Nutt.) Collins, comb. nov. ined. (The name is likely to be accepted and appear in later works.)


var. multiflora Largeflower Broomrape. Calyx 12 to 17 mm long; corolla 25 to 35 mm long, upper lip 9 to 12 mm long, purple; anthers commonly pubescent. The collections from Brazos Co. are believed to be this variety.


var. pringlei Munz. Calyx 8 to 10 mm long; corolla 20 to 50 mm long, lips 5 to 7 mm long, rose-purple or lighter. This plant occurs in the same range as the typical variety, so perhaps also present here.






ACANTHACEAEACANTHACEAE

Acanthus Family


Ours annual or perennial herbs. Vegetative parts usually with cystoliths which appear as minute lines. Leaves opposite, simple, estipulate. Flowers perfect, zygomorphic to nearly regular, hypogynous, solitary or in crowded, few-flowered spikes or cymes. Calyx equally 5-parted, persistent. Corolla bilabiate or 5-lobed. Stamens 4, didynamous, or else stamens 2 and 2 staminodia present; anthers (1-)2-celled, longitudinally dehiscent, cells often unevenly placed. Ovary bicarpellate, bilocular, with 2 to 10 ovules per locule; style single, slender, stigmas 2. Fruit capsular, commonly flattened, clavate or fusiform, dehiscent longitudinally by 2 valves, often explosively dehiscent. Seeds borne on small papilliform or usually hook-shaped funicles; seed coat smooth to rough, often mucilaginous when wet.

About 350 genera and 4,300 species, primarily of the tropics; 13 genera and 48 species in TX; 5 genera and 8 species confirmed from our area.

The family includes many cultivated ornamentals, including the common pot or conservatory plants Aphelandra (Zebra Plant), Fittonia, Crossandra, and Justicia (including Shrimp Plant). Thunbergia includes several very showy climbing vines (Mabberley 1987).


1. Corolla 2.5 cm long or longer, 5-lobed and nearly regular; stamens .....................1. Ruellia

1. Corolla less than 2.5 cm long, at least somewhat bilabiate; stamens 2 or 4 .......................2


2(1) Flowers in dense or loose spikes ............................................................................2. Justicia

2. Flowers in axillary cymes or solitary or clustered in the axils .................................................3


3(2) Flowers yellow; leaves lanceolate.. ...................................................................3. Hygrophila

3. Flowers pale pink-purple to white; leaves elliptic to ovate, obovate, or oblanceolate...........4


4(3) Stamens 2; leaves broadly elliptic to ovate; perennial ........................................4. Dicliptera

4. Stamens 4; leaves narrowly obovate to oblanceolate; annual ......................5. Dyschoriste



ACANTHACEAE RUELLIA1. RUELLIA L. Ruellia, Wild Petunia


Ours perennial herbs from a short, sometimes branched caudex, some also with short rhizomes. Leaves sessile or petiolate, entire to remotely crenate. Flowers usually large (2.5 cm or more long) and showy, solitary or in few-flowered cymes or glomerules in the axils or in terminal cymose panicles, in some species few-flowered lateral panicles of cleistogamous flowers also produced. Calyx segments 5, usually narrow. Corolla in ours some shade of purple or blue-purple (in other taxa also white, red, or yellow), funnelform to salverform, the tube narrow and usually longer than the campanulate throat, limb slightly zygomorphic, with 5 subequal, usually obtuse, spreading lobes. Stamens 4, didynamous, anther sacs basally rounded. Stigma lobes unequal. Capsule oblong-clavate or fusiform, with a constricted, stipe-like base. Seeds mucilaginous when wet, in our species round and flat.

About 150 species of tropical to temperate N. Amer.; 15 species listed for TX (Hatch, et al., 1990); 3 entities locally (but see note after R. humilis).

Some species are cultivated for their showy flowers. R. brittoniana Leonard is commonly grown in our area and may persist but escapes infrequently, if at all. It is a tall plant, to 1 m, with petiolate, linear to lanceolate leaves to 27 cm long, and short flowering branches in the axils of the upper leaves; its corollas are 2.5 to 4.5 cm long.

NOTE: The treatment of Ruellia in Texas has varied over the years--with no two treatments alike. The following is based upon the work of Tharp and Barkley (1949), Wasshausen (1966), Correll and Johnston (1970), Hatch, et al. (1990), B. L. Turner (1991), and Kartesz (1998).


1. Flowers in terminal panicles; lateral panicles of cleistogamous flowers often produced, but these often absent by anthesis of chasmogamous flowers ...1.R.nudiflora

1. Flowers in axillary glomerules or cymes, or solitary in the axils .............................................2


2(1) Calyx lobes narrowly lanceolate, acuminate, flat to the tip, 2 to 4 mm broad; leaves definitely petiolate; some forms with only cleistogamous flowers ...2.R.strepens

2. Calyx lobes narrowly linear, with slender, bristle-like tips; leaves sessile to short-petiolate; cleistogamous flowers infrequent ...3.R.humilis


1.R. nudifloraACANTHACEAE RUELLIA nudiflora (Engelm. & Gray) Urban Violet Ruellia. Perennial from a slightly woody base; stems ultimately to 70 cm tall and more or less glabrous, erect to ascending. Spring leaves oblong to narrowly ovate-oblong, to 6 cm long and 3 cm wide, lower surface strongly reticulate, slightly grayish and moderately short-pubescent, margins distinctly crisped or undulate, base decurrent on a petiole to 2.5 cm long, summer leaves to 12 cm long, commonly smaller, reduced above. Spring flowers cleistogamous, in clusters or reduced panicles in the axils; summer and fall flowers chasmogamous, produced in a terminal panicle. Sepals narrow, subulate-attenuate, hispid to slightly scabrous, with or without gland-tipped hairs, but not viscid, 11 to 15 mm long at anthesis, elongating slightly in fruit; corolla purple-blue, usually noticeably curved, ca. 5.5 cm long, 3.5 to 4 cm wide, opening in the early morning and falling in the afternoon, tube 1.5 to 2 cm long, throat markedly broader. Capsules of cleistogamous flowers clavate, ca. 1.5 cm long, 3.5 to 4 mm broad, minutely but densely retrorse puberulent, seeds 4 or 5 per locule, ca. 3.5 mm long, 3 mm broad, brown; capsules of chasmogamous flowers 1.6 to 2 cm long, 4 to 4.5 mm broad, densely finely pubescent, seeds usually 18 or fewer, ca. 3 mm wide and about as broad, apically rounded or apiculate, dark brown. Widespread in roadsides, fields, thickets, etc., commonly in sandy soil. Chiefly in the S. 1/2 TX, N. to N. Cen. TX; also NE. Mex. Flowering throughout the year; local collections primarily Jun.-Nov. [As treated here, includes var. hispidula Shinners and var. glabrata Leonard; Dipteracanthus nudiflorus Engelm. & Gray].

NOTE: Kartesz (1998) followed the recommendation of B. L. Turner (1991) and reduced the plants formerly known as R. runyonii Tharp and Barkl. to varietal status under R. nudiflora (Engelm. & Gray) Urban var. runyonii (Tharp & Barkl.) B. L. Turner. This taxon is not present in our area, but if this combination is accepted, our plants then become R. nudiflora (Engelm. & Gray) Urban var. nudiflora.


2.R. strepensACANTHACEAE RUELLIA strepens L. Limestone Ruellia, Smooth Ruellia. Perennial from a short, branched caudex and sometimes short rhizomes; stem erect, simple or with a few ascending branches, 3 to 10 dm tall, slightly quadrangular, glabrous to pilose-hirsute. Leaves ovate to sometimes ovate-lanceolate or -elliptic, 5 to 13(18) cm long, (1.5)2.5 to 5(9) cm broad, usually 1.5 to 2 times longer than the internode above, thin-textured, acute to short acuminate (sometimes obtuse), base attenuate to cuneate or rounded, margin entire to very slightly undulate or scarcely crenulate, commonly ciliate, surfaces glabrous to short-strigillose or sparsely hirsute; petioles to 2 cm long, commonly with a narrow wing. Cleistogamous flowers borne in the axils, in compact, subsessile cymes subtended by 2 lanceolate, glabrous to sparingly hirsute bracts 10 to 15 m long, 1.5 to 2.5 mm broad. Chasmogamous flowers borne usually at 1 to 3 of the midstem nodes, 1 to 3 per cyme on peduncles to 6(9) cm long, subtended by 2 broad, leafy bracts, flowers each subtended by 2 minute, obovate bractlets. Calyx segments lanceolate to linear-lanceolate, flat all the way to the apex, 9 to 20 cm long, (1.5)2 to 4 mm broad, villous-ciliate to hirsute, commonly with glandular hairs, shorter than to longer than the capsules, calyx tube 1 to 3 mm long; corolla lavender to light blue-violet or dark violet (rarely white), externally glabrous or pubescent on the veins, (3)4 to 7 cm long, the tube about half this length, throat widely funnelform, limb 2 to 4 cm broad, ciliate or eciliate; filaments 2 to 3.5 cm long, anthers 2.5 to 3 mm long; style 2 to 4.5 cm long, usually not persisting after corolla falls. Capsules glabrous, 10 to 15(20) mm long; seeds round and flat, ca. 2.8 mm broad, brown or reddish, sometimes slightly hirsute. Low woods, stream courses, and talus slopes; often in calcareous areas and/or where shady. E. and N. Cen. TX; PA to IA, NE, and KS, S. to SC and TX. Flowering Apr.-May, collected later in fruit. [Includes var. cleistantha Gray and var. micrantha (Engelm. & Gray) Britt.; Dipteracanthus strepens (L.) Nees; D. micranthus Engelm. & Gray].




3.R. humilisACANTHACEAE RUELLIA humilis Nutt. Rhizomes, if any, knobby; stem erect to decumbent, from short, , simple to well-branched, to 8 dm tall, sometimes strongly quadrangular, glabrous to white-villous-hirsute. Leaves commonly somewhat coriaceous, ovate to lanceolate or elliptic, less commonly obovate, 2 to 5(8) cm long, 1 to 3.5(4.5) cm wide, apically generally acute, but sometimes obtuse or rounded, base truncate to widely cuneate, sessile or with a short petiole to 3 mm long, surface hirsute to villous, margin entire to slightly undulate, usually pilose-ciliate. Floral bracts lanceolate to linear-lanceolate, elliptic, or oblanceolate; flowers usually 1 to few in the axils of the middle and upper leaves, very short-pedunculate. Calyx hirsute or villous-hirsute, tube 2 to 3 mm long, lobes linear-attenuate, the tips long and bristle-like, 15 to 30 mm long, much longer than the capsules, only 0.7 to 1.5 mm broad; corolla lavender or pale bluish, 3 to 8 cm long, tube usually obviously longer than the throat and limb, usually pilose outside, to 45 mm long, limb 2 to 4 cm wide (in rare, fall-produced cleistogamous flowers, corolla tubular and closed); filaments 2 to 5 cm long; style 2.5 to 6 cm long, commonly persisting after the corolla falls, at least for a period, longer stigma lobe usually recurved. Capsule tan or brown, 10 to 16 mm long, glabrous; seeds brown or reddish brown, round and flat, ca. 3 mm broad, sometimes marginally hirsute. Open woods, old fields, savannahs, prairies, outcrops, etc. Mostly in the E. 1/4 TX, rare in the Ed. Plat. and Panhandle; PA and WV to MI, NE, and KS; S. to FL and Tx. Apr.-Oct., ours usually June or later.

Many varieties have been described (see, e.g., Wasshausen (1966) and Correll and Johnston (1970)). However, most of them intergrade, and it seems logical to combine them, an approach taken by GPFA (1986), B. L. Turner (1991); and Kartesz (1998). If combined, synonyms thus include var. depauperata Tharp & Barkl., var. expansa Fern., var. frondosa Fern., and var. longiflora (Gray) Fern.; R. ciliosa var. longiflora Gray.

NOTE: Low Ruellia, the plants treated as R. humilis Nutt. var. humilis by Hatch, et al. (1990) may be distinct. Kartesz (1998) lists them under R. caroliniensis (J. F. Gmel.) Steud. subsp. ciliosa (Pursh) R. W. Long var. cinerascens (Fern.) Kartesz & Gandhi. I have been unable to find a satisfactory discussion of this in the literature. R. caroliniensis as presented by Wasshausen (1966), Correll and Johnston (1970), Hatch, et al. (1990), and B. L. Turner (1991) is not present in our area.



ACANTHACEAE JUSTICIA2. JUSTICIA L. Waterwillow, Justicia


Perennial, rhizomatous herbs (as ours) or shrubs. Stems more or less 4-angled at midstem; cystoliths often evident. Leaves petiolate, entire. Flowers in axillary spikes (as in ours) or panicles, or sometimes solitary; floral bracts small, varying from linear to triangular or subulate, usually not concealing the calyces. Calyx (4-)5-parted, the lobes narrow and equal or subequal. Corolla white to pink or purple (or red), sometimes with darker markings, with a distinct tube and a bilabiate limb, upper lip 2-lobed, usually reflexed to some degree, lower lip 3-lobed. Stamens 2, shortly exserted from the corolla throat but usually shorter than the lips; anther cells 2, connective slender to broad, lobes parallel or oblique, one or both cells tailed or apiculate. Capsule clavate or linear-clavate, slender to broad, sometimes constricted below and often tapered above the seed-bearing portion. Seeds 4, round and flat.

About 240 species of tropical, subtropical, and temperate parts of N. Amer.; 5 in TX; 2 here.

Some species are cultivated for ornament, including the familiar Shrimp Plant, J. brandegeeana (Mabberley 1987).


1. Flower spikes dense, head-like ...1.J.americana

1. Flower spikes loose, commonly slender and 1-sided ...2.J.ovata

var. lanceolata


1.J. americanaACANTHACEAE JUSTICIA americana (L.) Vahl American Waterwillow, American Dianthera. Perennial forming colonies by slender, creeping rhizomes; stems 3 to 10 dm tall, erect or decumbent below and ascending above, simple or branched, often rooting at the nodes, lower internodes sometimes swollen. Leaves linear to narrowly linear-elliptic or narrowly-lanceolate or -oblong, (4)6 to 15(20) cm long, (0.5)0.8 to 2.5(3) cm broad, gently tapered to an acutish or sometimes obtuse apex, base cuneate to attenuate, sessile or with a very short petiole to 1 cm long, margin entire to rather obscurely crenate. Peduncles axillary, and perhaps also terminal, 4 to 17 cm long, rather stiffly erect, shorter than to slightly longer than the subtending leaves; flowers opposite, crowded, sessile or essentially so, each subtended by 3 short-ovate or triangular bracts. Calyx lobes subequal, lanceolate to linear-subulate, acute, 4 to 6.5 mm long, ciliolate, tube ca. 1 mm long; corolla (6)9 to 12(14) mm long overall, sometimes pubescent internally, violet to nearly white, upper lip emarginate and reflexed, often darker, lower lip palate-like at the base, with 2 spreading lateral lobes and a deflexed central lobe, usually marked with darker dots; stamens 2, filaments attached to the apex of the corolla tube, 6 mm long, anther cells dark violet or mottled violet and white, separate and more or less unequal, the terminal one transverse and the lower lateral; stigmas unequal. Capsule 11 to 18 mm long, clavate, with a sterile lower part about equalling the seed-bearing part and the calyx; seeds densely warty, reniform-orbicular, 2 to 3.5 mm long. In shallow water or on mud near streams, rivers, ditches, ponds, etc., usually in the sun. E., SE., and N. Cen. TX and the Ed. Plat; Que. and VT. to MI, WI, KS, and OK, S. to GA and TX. Apr.-Oct. [Includes var. subcoriacea Fern.; Dianthera americana L. and var. subcoriacea (Fern.) Shinners].


2.J. ovataACANTHACEAE JUSTICIA ovata (Walt.) Lindau var. lanceolata (Chapm.) R. W. Long Lance-leaved Waterwillow. Perennial, sometimes solitary, but commonly rhizomatous and colonial; stems 1 to 5 dm tall, simple or branched, erect or spreading, angled. Leaves membranous, linear to linear-elliptic or elliptic-lanceolate, to ca. 10 cm long, 0.5 to 3(4) cm broad, sessile or narrowed to a short subpetiolar base, acuminate, entire or with a few shallow undulations, surfaces glabrous or minutely puberulent. Spikes terminal or terminal and axillary, sometimes branched, usually longer than the leaves, slender and loose, commonly 1-sided, ca. 3 to 10 cm long; floral bracts small, short-ovate to triangular or subulate, minutely pubescent marginally. Calyx lobes narrowly linear, (4)5 to 76(10) mm long, less than 1 mm wide, short-pubescent marginally and sometimes on the midribs; corolla purple to white, sometimes marked with purple or white, in this variety ca. 10 mm long, the tube slightly saccate basally, the 2 lips about as long as the tube, upper lip truncate or retuse lower lip 3-lobes, the lateral lobes obtuse and the center 1 lobe truncate to retuse; stamens slightly exserted, filaments ca. 3 to 4 mm long, attached about the middle of the corolla tube; anther cells unequal, 1 terminal and transverse, ca. 0.8 mm long, and the other lower, lateral, ca. 1 mm long, and apiculate, connective broad; ovary 1.5 mm long, style ca. 1 cm long. Capsule 1 to 1.5 cm long, the sterile base about as long as the body; seeds 4, ca. 2 mm wide, smooth to slightly papillose, margin thick. Swamps, wet woods and prairies, and around ponds, depressions, and rivers. SE. TX; SE. VA, S. to FL, W. to MO, KY, TN, OK, and TX. Mar.-Jun. [J. lanceolata (Chapm.) Small].



ACANTHACEAE HYGROPHILA3. HYGROPHILA R. Br.

A genus with worldwide distribution. Many species have been published; about 25 are still recognized. We have the one species found in TX.


1. H. lacustris (Schlecht. & Cham.) Nees Lake Acanthus. Perennial, colonial from rhizomes; stems decumbent or erect, to ca. 1 m long, bluntly quadrangular, branched. Leaves opposite, lanceolate, 5-13 cm long, 5 to 25 mm broad, entire, acute, tapered to the base, upper surface with cystoliths visible as short lines under magnification, a few bristles present between the bases of each pair. Flowers clustered in the axils of the leaves, sessile; bracts linear. Calyx segments lanceolate, ca. 5 mm long, narrowly white-margined and minutely pilose; corolla yellow, puberulent, bilabiate, the upper lip 2.5 mm long and bidentate, the lower lip with 3 lobes; stamens 4 and didynamous or else 2 of the stamens represented by staminodia, filaments of each pair united by a membrane at the base, anthers 2-celled, minutely mucronate; posterior stigma lobe reduced. Capsules narrowly oblong, 8 to 12 mm long; seeds 16 to 18. Wet areas along streams, swamps, or ponds, etc. SE. TX, known from Madison Co.; FL and GA to TX. June-Sept.



ACANTHACEAE DICLIPTERA4. DICLIPTERA Juss.


Herbs, stems often hexagonal in cross-section. Leaves petiolate, generally ovate, entire to undulate. Flowers 1 to several in panicles or spikes composed of contracted cymes, subtended by pairs of conspicuous bracts that usually conceal the calyces. Calyx 5-parted, regular, sometimes hyaline. Corolla bilabiate, the tube narrow, lips entire or slightly toothed. Stamens 2, inserted at the top of the corolla tube, the anther sacs separate, one terminal and one lateral. Capsule elliptic, ovoid, of subglobose, elastically dehiscent; seeds 2 to 4.

About 150 species of tropic and temperate regions; 2 in TX; 1 here.


1. D. brachiata (Pursh) Spreng. Annual herb; stems 3 to 7(8) dm tall, usually widely branching, sometimes straggling, sometimes rooted at the lowest nodes, glabrous to pilose-pubescent or occasionally villous or hirsute, angled, nodes often swollen. Leaves elliptic to ovate, oblong-ovate, or ovate-lanceolate, (2)5 to 15 cm long, (1.5)2 to 7 cm broad, often smaller, usually acuminate, sometimes acute or obtuse, base rounded or narrowed, somewhat decurrent on a petiole (0.5)2 to 7 cm long, margin entire to slightly toothed, ciliate, surfaces glabrous to pilose-pubescent or sometimes hirsute on the veins below, upper surface usually with cystoliths visible as lines. Flowers in axillary cymes, sessile or subsessile, or at the branch tips the inflorescence appearing somewhat spike-like where the leaves are small or absent; subtending bracts obovate to spatulate, 4 to 7 mm long and 2 to 5 mm wide, acute to acuminate or cuspidate, ciliate, usually concealing the calyces. Calyx membranous or sub-hyaline, 2.5 to 4 mm long, tube only 1 to 1.8 mm long, lobes subulate, 1.5 to 2.5 mm long, sometimes ciliate; corolla 11 to 15(20) mm long, pale pink, purple, lavender, blue-purple, or white, commonly with darker spots, especially on the lower lip and throat, tube slender, 3 to 6 mm long, upper lip erect, entire or shallowly notched, lower lip usually entire, externally pubescent, sometimes with glandular hairs; stamens extending up under and enclosed by the upper lip, reaching its tip, filaments sparsely hirsute toward the base or glabrous, anther cells ca. 1.7 mm long, ellipsoid; style 10 to 15 mm long, stigma with 2 equal lobes. Reduced, cleistogamous flowers often produced. Capsule ovoid, obovoid, or subglobose, with a sterile basal part ca. 1 mm long and an apiculate and emarginate tip, the tip also ciliate, the body pilose; seeds 2 to 4, dark reddish-brown to black, nearly round, flattened, ca. 2.5 mm broad, muricate, glabrous to papillose-pubescent. Moist, shady areas. S. and Cen. TX; VA to IN, MO, and KS, S. to FL and TX. July-Oct. [Includes var. glandulosa (Scheele) Fern. and var. ruthii Fern.; Diapedium brachiatum (Pursh) O. Ktze.].



ACANTHACEAE DYSCHORISTE5. DYSCHORISTE Nees. Dyschoriste


Perennial herbs. Stems prostrate to erect, variously pubescent or glabrous. Leaves sessile or petiolate, commonly entire. Flowers in cymes, spikes, or capitate clusters, terminal or axillary, subtended by bractlets and leafy bracts. Calyx lobes 5, usually subulate-setaceous and ciliate. Corolla slightly to strongly bilabiate, 5-lobed. Stamens 4, didynamous, filament bases united, epipetalous on the base of the corolla tube, pubescent; anther cells basally mucronate, parallel or slightly divergent. Ovary glabrous, ovules (1)2 per cell; 1 lobe of stigma reduced, the other oblong and flattened. Capsule shorter than the persistent calyx, oblong-linear, somewhat reluctantly 2-valved. Seeds 2 to 4, suborbicular, flattened, mucilaginous when moistened.

About 65 species of tropical and subtropical regions; 3 species in TX; 1 infrequent here.


1. D. linearis (T. & G.) O. Ktze. Narrowleaf Dyschoriste. Plants from a short, branched caudex, commonly tufted but sometimes rhizomatous; stems strongly erect, stiff, (10)18 to 25(42) cm tall, usually branched below, usually pubescent to hispid, sometimes sparsely pubescent to glabrous. Leaves linear-oblanceolate to oblong-spatulate or sometimes narrowly obovate, (1)1.5 to 4.5(6.5) cm long, 2 to 7(11) mm broad, rounded to mucronulate, entire or very slightly undulate, ciliate, rather rigid, pubescent on the veins and the midrib, sessile or essentially so. Flowers 1(2 to 3) in compact, axillary cymes, subtended by bractlets and by leafy, oblanceolate bracts which can be as large as the leaves. Calyx ca. 15 to 22 mm long overall, lobes united for (1/4)1/3 to 1/2 their length, to 1 mm wide, subulate-setaceous, hispid on the veins, margins long-ciliate; corolla pale pink-purple or whitish, 16 to 23(27) mm long, more or less bilabiate, pubescent externally, upper lip 2-lobed, lower lip 3-lobed, lobes rounded, tube 4 to 7 mm long, shorter than the abruptly-spreading limb; filaments pubescent, the shorter pair 10 to 15 mm long, longer pair 13 to 18 mm long, anther cells oblong; style about as long as the stamens, pubescent. Capsule included in the calyx, 4-seeded, oblong-linear, glabrous. Widespread in TX except the Pineywoods, but most common on rocky-grassy slopes, on caprock, and on silty flats in the W. 2/3 TX; very infrequent in our area; OK to TX, NM, and N. Mex. Apr.-Aug.






PEDALIACEAE(including Martyniaceae)
PEDALIACEAE

Sesame Family


Plants of the Western Hemisphere annual or perennial herbs. Herbage usually with glandular pubescence, the hairs tipped with 4 or more mucilage-filled cells, often strongly-scented. Leaves usually opposite, sometimes alternate above, simple, estipulate. Flowers perfect, usually showy, in terminal racemes, subtended by bracts which often conceal nectaries. Calyx of 5 unequal sepals or sepals united into a 5-lobed tube. Corolla sympetalous, somewhat bilabiate, 5-lobed. Functional stamens 4 and didynamous, with a fifth stamen rudimentary or functional stamens 2 and the other 3 represented by staminodia; anthers gland-tipped. Ovary commonly subtended by a nectary disk, in our taxa superior, bicarpellate, bilocular; style 1, stigma with 2 flat lobes. Fruit a drupe-like capsule with a somewhat fleshy exocarp and a woody endocarp, 2-valved, loculicidally dehiscent or indehiscent, imperfectly 4- or 5-celled due to intrusion of the parietal placentae, commonly crested and with a terminal, curved, 2-horned beak. Seeds 4 to many, the testa variously sculptured; endosperm none.

There are 18 genera and 95 species of tropical and warm regions, especially common in coastal areas; 1 genus and 5 species in TX; 1 species in our area. New World plants were once separated as Martyniaceae and were presented as such by Correll and Johnston (1970). Old World Pedaliaceae have axile placentation and axillary flowers.

Sesamum (sesame) and Proboscidea include some oilseed plants. There are also a few taxa cultivated for ornament (Mabberley 1987).



PEDALIACEAE PROBOSCIDEA1. PROBOSCIDEA Schm. Devil's Claw, Unicorn-plant


Taprooted annuals or perennials from tuberous roots. Stems erect to decumbent. Herbage usually odiferous. Leaves opposite or alternate, entire to pinnately or palmately lobed. Flowers in terminal racemes (racemes sometimes appearing axillary if the lower portion of the plant keeps growing), the floral axis elongating; pedicels at first erect to spreading but elongating and becoming recurved; bracts beneath pedicels small, linear to oblanceolate, usually deciduous or sometimes absent; paired bractlets (sometimes rather fleshy) present below the calyx. Calyx spathe-like, more or less tubular and unequally 5-lobed, split almost to the base on the lower side, deciduous. Corolla in TX material red-purple, pink, cream, white, or yellow to orange, slightly bilabiate, 5-lobed, tubular-cylindric to campanulate or funnelform, commonly somewhat gibbous. Fertile stamens 4, didynamous (sometimes 2), adherent to one another by their edges; anther sacs diverging by ca. 180o. Ovary at first unilocular, the 2 parietal placentae intruding and bifurcating, the fruit ultimately 4-locular, ovules in 1 row along the edge of each of the 4 placental flanges; exocarp fleshy, endocarp woody, reticulate and sculptured, lower part of the fruit crested ventrally and sometimes also dorsally, upper portion tapered into a beak 1.25 to 3.5 times as long as the body, this beak ultimately curved and splitting lengthwise into 2 curling horns. Seeds many, angled, seed coat corky, usually black, sometimes white.

About 9 species of warm parts of the Americas; 5 in TX; 1 here.

Some species are oilseed plants; others have fruits edible when young. Mature fruits can injure grazing animals (Mabberley 1987).


1. P. louisianica (P. Mill.) Thell. subsp. louisianica Common Devil's Claw, Unicorn-plant. Taprooted annual; stems and branches erect or sometimes decumbent, 1.5 to 6(10) dm tall and plants spreading up to 1 m; herbage viscid-pubescent and with a strong odor. Leaves opposite or the upper alternate, petioles 3 to 25(30) cm long, densely pubescent; blades reniform-orbicular to widely ovate, 3 to 20(30) cm long, to 30 cm broad, about as wide as long or a little wider, apex rounded, obtuse or broadly acute, basally cordate, margin entire or sinuate, venation palmate. Flowers (4)8 to 20(26) in a loose terminal raceme; pedicels 2 to 4 cm long at anthesis thickening and elongating in fruit to ca. 4.5 cm long; pedicel bracts linear to linear-lanceolate, 5 to 11 mm long, deciduous; calyx bracts oblong-falcate to ovate, to ca. 10(15) mm long, generally less than 2 mm broad. Flowers zygomorphic, calyx thick and greenish or thinner and somewhat yellow, 10 to 22 mm long, lobes about half as long, acute to obtuse; corolla to ca. 55 mm long, base tubular-cylindric for 2 to 5 mm long, broadly campanulate and gibbous above for 15 to 25 mm, lobes 1.5 to 2 cm long, 1.5 to 3 cm broad, dingy white to purplish or pinkish, spotted or mottled with yellow or reddish purple, the inner tube spotted with red-purple and the lower portion with yellow stripes, occasionally nearly clear red-violet; filaments glabrous to villous or tomentose basally, glandular on the curved portion; flowers protandrous; style longer than the stamens. Body of fruit plump, to 10 cm long and 3 cm thick, the curving horns 1.5 to 3 times longer, endocarp woody and reticulate; seeds black, narrowly ovoid, with 1 convex and 2 flat sides. Cultivated fields, waste places, meadows, stream banks, and playas. Throughout much of TX, primarily the Cen. and NE. parts; native to the SE. U.S. and Mexico, but scattered/introduced N. to ME, MN, SD, and CO. [Martynia louisianica P. Mill; P. jussieui Scmid.].

The other taxon in this species, subsp. fragrans (Lindl.) Brett. is the plant treated as P. fragrans (Lindl.) Dcne. by Correll and Johnston (1970). Bretting (1983) pointed out the similarities and combined the two.

The unusual fruits have various uses. Native Americans of the U.S. and Mexico used strips of the pods in basketweaving, while European settlers sometimes cultivated the plant for its flowers. The immature fruits (less than 3 cm long) are edible steamed or pickled (similar to and substitutable for okra) and have even been grown commercially for canning (Tull 1987). The seeds are eaten in Mexico and are served fresh, toasted, or ground into meal. Various tribes of Native Americans also made food use of the roots or leaves (Tull 1987).






BIGNONIACEAEBIGNONIACEAE

Catalpa or Bignonia Family


Trees, shrubs, or woody vines. Leaves opposite or the uppermost sometimes alternate, simple or pinnately compound, entire to toothed or lobed, estipulate. Flowers perfect, hypogynous, in terminal panicles or racemes or paired or whorled at the nodes or in the axils, bracts and bracteoles commonly present. Calyx 4- or 5-lobed, sometimes bilabiate. Corolla from nearly regular and tubular or campanulate to strongly bilabiate, 5-lobed, the lobes usually imbricate. Stamens 2 or 4, epipetalous, alternate with the corolla lobes, the 5th or 3rd, 4th, and 5th stamen(s) sometimes present as staminodia; anther cells often divergent. Pistil 1, bicarpellate, bilocular, with 2 axile placentae per locule (as in ours) or else unilocular with 2 or 4 parietal placentae; ovules commonly many. Fruit capsular. Seeds many, usually flattened and winged, sometimes comose.

112 genera and 725 species, primarily of the tropics and especially common in S. Amer.; 6 genera and 7 species in TX; 2 genera and 2 species here.

Some of the tropical taxa are timber sources. The family includes many ornamentals including Jacaranda, Paulownia, Catalpa, and Campsis (Mabberley 1987).



1. Plants vines or viney shrubs; flowers red to orange .............................................1. Campsis

1. Plants trees; flowers white to purple .......................................................................2. Catalpa



BIGNONIACEAE CAMPSIS1. CAMPSIS Lour. Trumpet Creeper


Two species, 1 in the U.S. and 1 in E. Asia, both with ornamental qualities (Mabberley 1987).


1. C. radicans (L.) Seem. ex Bureau Trumpet Creeper, Cow-itch Vine, Trumpet-honeysuckle. Deciduous viney shrub with aerial rootlets, sprawling over vegetation or climbing trees and structures to 10 m or more. Leaves opposite, odd-pinnately compound, to 30 cm long or more, leaflets 5 to 13, petiolulate, blades ovate to ovate-oblong or elliptic-lanceolate, acuminate, basally rounded to cuneate, to 9 cm long and 4 cm broad, but usually smaller, serrate, glabrous above, commonly somewhat pubescent below on the veins and/or with some hairs present on the rachis between the petiolule bases. Flowers in terminal panicles or corymbs of ca. 8 to 15; pedicels ca. 1 cm long, stout. Calyx tubular-campanulate, 2 to 2.5 cm long, the tube much longer than the triangular lobes, leathery, reddish-brown; corolla tubular-funnelform, red-orange or orange, the tube sometimes paler within and with darker markings, ca. 6 to 9 cm long, lobes rounded; functional stamens 4, didynamous, appressed to the upper side of the corolla tube, with divergent anthers ca. 4 mm long, dehiscing downward, 5th stamen non-functional or sometimes vestigial; ovary subtended by a nectariferous ring, bilocular, style exserted beyond the anthers at maturity, stigma lobes 2, flat and rather large. Capsule stipitate, cylindric-oblong, 8 to 12(20) cm long, leathery, beaked, keeled along the sutures, loculicidally dehiscent, the interior divided by a thin partition that separates from the valves; seeds many, flat, ovoid, with 2 large, trapezoidal, papery wings. Woods, roadsides, floodplains, hillsides, etc. E. 1/2 TX; NJ to ND, IA, KS, and OK, S. to FL, and TX. May-Oct., most of our collections from late summer and fall. [Tecoma radicans (L.) Juss.].

This plant is sometimes cultivated for ornament, but it is very aggressive, capable of pulling down structures and often escaping. Orange- and yellow-flowered cultivars exist (Bailey, et al. 1976). The flowers are attractive to hummingbirds.



BIGNONIACEAE CATALPA2. CATALPA Scop. Cigar-tree, Catalpa


Deciduous trees. Twigs without terminal buds, winter buds with several outer scales. Leaves opposite (sometimes whorled), simple, entire or very shallowly lobed, with 3 to 5 veins from the base, usually with purple nectary spots on the undersides in the axils of the major veins. Flowers few to many in terminal racemes or panicles. Calyx bilabiate or splitting irregularly. Corolla irregular, with 2 upper and 3 lower lobes. Fertile stamens 2. Pistil bilocular, style slightly longer than the stamens. Capsules terete in cross-section, long and linear, dehiscent by 2 valves. Seeds many, oblong, 2-winged, each wing with a terminal tuft of hairs.

11 species of E. Asia and N. Amer.; 2 species in TX, both probably only adventive from the SE. U.S. (Elias 1980; Duncan and Duncan 1988); 1 species found in our area.


1. C. speciosa (Barney) Warder ex Engelm. Northern Catalpa. Fast-growing tree to 30 m; trunk ultimately to 1 m in diameter; crown narrow to broad; bark red-brown, with shallow grooves and thick scales; twigs initially green and pubescent, later orange to reddish-brown and glabrous. Leaves ovate to ovate-oblong, 15 to 30 cm long, 10 to 20 cm wide, long-acuminate, base truncate to broadly cordate, entire or very rarely with 1 to 3 small teeth, green and glabrous above, densely pubescent beneath, without an odor; petiole to 15 cm long. Panicle 15 to 20 cm long, with fairly few flowers, glabrous; floral bracts linear, pubescent, irregularly spaced along the axis. Calyx lobes ovate to obovate, 6 to 12 mm long, apiculate; corolla to ca. 5 cm long and 4 cm broad, gibbous on the lower side, white, with 2 longitudinal yellow stripes and a few purple-brown spots within, the lobes undulate-ruffled, 2 upper, 2 more or less lateral, and 1 larger, notched lobe on the bottom; filaments epipetalous on the lower corolla lobe, 2 cm long, strongly curved near the anthers, anthers ca. 4 mm long, 3 vestigial stamens attached to the upper corolla lobes; ovary cylindrical, style white, 2 cm long, stigma lobes ovate, flat, 2 mm long. Capsule 20 to 45 cm long, ca. 1 to 1.5 cm thick, with irregular shallow ridges, dark brown, thick-walled, the valves remaining more or less rounded on the back after dehiscence; seeds flat, 4 to 5(8) mm broad, with a wing at either end, overall 3 to 4 cm long, each wing with a terminal tuft of straight hairs. Mostly in damp woods and on the edges of streams and swamps in E. TX; in our area commonly cultivated and frequently escaping to vacant lots and roadsides. More or less IN to IL, MN, and ND, S. to TN and TX; escaped or naturalized in much of the SE. U.S., N. to N. Eng.

Often cultivated for its ornamental flowers or as an attractant for caterpillars used as fishing bait (Duncan and Duncan 1988). The wood is used for fenceposts, telephone poles, and cheap furniture, but is soft and rather weak, leaving the trees susceptible to storm damage (Elias 1980). In our area the foliage is often yellow and unsightly by August.

NOTE: Southern Catalpa, C. bignonioides Walt., is occasionally cultivated in our area, but is not known to be naturalized here and probably only rarely (if ever) escapes. It is distinguished by having leaves scarcely or only abruptly short acuminate, 10 to 25 cm long, with an unpleasant musky odor when crushed. The panicles have more flowers, the blossoms are smaller, 2.5 to 4 cm long, liberally spotted, and the lower lobe not notched. The capsules are 15 to 40 cm long and only about 1 cm wide; the valves flatten after dehiscence. The seeds are 2.5 to 4.5 mm broad, and the hairs of the terminal tufts are connivent into a point at each end. Native from FL to GA and AL; naturalized from S. N. Eng. to E. TX and the Great Plains.






LENTIBULARIACEAELENTIBULARIACEAE

Bladderwort Family


Annual or perennial herbs; plants terrestrial, aquatic, or amphibious, carnivorous by various means: in TX Pinguicula with glandular-viscid leaves and Utricularia with bladder-like traps. Leaves alternate, cauline or in a rosette, varying from entire, broad and flat to finely dissected with thread-like segments. Flowers 1 to several on a scape or in a scapose raceme, zygomorphic, perfect; floral bracts commonly present. Calyx bilabiate, with 2 or 5 lobes. Corolla strongly bilabiate, lower lip more or less 3-lobed and with a definite palate, the base spurred. Stamens 2, epipetalous on the base of the corolla tube, included. Ovary superior, bicarpellate, unilocular, with free-central placentation; style short or absent, stigmas 2. Fruit ovoid to globose, with 2 or 4 valves but often splitting irregularly or dehiscent via a pore or slit(s). Seeds tiny, the shape and ornamentation diagnostic.

4 genera and about 245 species nearly worldwide; 2 genera and 11 species in TX; 1 genus and 4 species here.



LENTIBULARIACEAE UTRICULARIA1. UTRICULARIA L. Bladderwort


Ours aquatic, wet-terrestrial, or amphibious herbs, without true roots but with rhizoids commonly present. Turions or winter buds present on the stolons of some species; a few have tubers. Stems/stolons elongate and creeping or floating, commonly with slender, often finely dissected, leaf-like structures possibly fully of stem origin (Godfrey and Wooten 1979; Taylor 1989). Some or all of the vegetative parts with stalked (or sessile) ovoid or pyriform bladder-like traps, these usually with hairs or flagellae around the orifice and with 2 minute 2-armed or 4-armed glands within; traps triggered by the presence of small organisms to open, the lower pressure within actively sucking the prey inside. Flowers 1 to many on a scape held above the water or substrate, bracts present at the base of the pedicels, sometimes flanked by or concealing bracteoles, sterile bract-like scales sometimes present on the peduncle. Calyx and corolla both bilabiate; calyx 2-parted nearly to the base, the upper part usually larger than the lower. Corolla spurred, yellow (as in ours), white, pink, or purple. Stamens 2, anthers convergent. Cleistogamous flowers produced in some species. Fruit usually a 2-valved capsule, but often dehiscent via pores or slits or else circumscissile. Seeds variable and diagnostic but best viewed by SEM.

214 species nearly worldwide, according to the most recent monograph (Taylor 1989); 10 species in TX; 4 here.

The genus includes plants with diverse habits and habitats, some resembling orchids and epiphytic, others growing in the tanks of bromeliads, in bogs, in savannahs, and so on. Traditional pressing methods tend to produce poor specimens. Our smaller species are best floated onto mounting paper to preserve the delicate anatomy of stems and traps; preservation in fluid is also an option. Some species are cultivated as oddities (Taylor 1989).


1. Peduncle with a whorl of 4 to 7 cylindrical (flat in pressed specimens) floats; lower lip of corolla more or less deeply 3-lobed, about equalling the upper ...1.U.radiata

1. Peduncle without a whorl of floats (perhaps with a whorl of leaves); lower lip of corolla entire, or if 3-lobed then much longer than the upper ............................................................2


2(1) Corolla 15 mm or more long from the sharp tip of the curved spur to the top of the upper lip; spur ca. 10 mm or more long; leaves linear, undivided ...2.U.cornuta

2. Corolla usually less than 15 mm long; spur usually less than 10 mm long, the tip blunt or bifid; leaves entire or diffusely divided .....................................................................................3


3(2) Bract at the base of the pedicel peltate; lower lip much longer than the upper; leaves undivided ...3.U.subulata

3. Bract at the base of the pedicel attached by its base; lower lip equalling or smaller than the lower; leaves dichotomously divided ...4.U.gibba


1.U. radiataLENTIBULARIACEAE UTRICULARIA radiata Small Whorled Bladderwort. Free-floating annual or sometimes perennial, rhizoids none; stolons to 0.5 m long (sometimes longer), branched; internodes 1 to 3 cm long. Leaves 1 to 10 cm long, divided from the base into 2 roughly equal, filiform segments, these many times dichotomously divided, the ultimate segments filiform-capillary, terete, with minutely setulose apices. Traps many, positioned laterally on the next-to-last divisions of the leaves, 0.7 to 2 mm long, ovoid, mouth lateral, with 2 long, bristle-like, branched appendages and a few smaller, simple bristles. Inflorescences relatively few and widely spaced, erect, emergent, 7 to 25 cm long; peduncle terete and spongy, bearing 3 to 10 cm below the apex a whorl of 4 to 7 floats, these spongy, cylindrical, little if at all united basally, 2 to 4 mm broad, the sides parallel for most of their length, sharply tapered to the base and slightly so to the apex, the distal ends with a number of leaf-like segments with thickened bases and sometimes a few traps; floral bracts attached by their bases, transversely oblong, broader than long, 2 to 3 mm long, more or less clasping, apex more or less 3-lobed, with the middle lobe more distinct and more acute; bracteoles none; flowers 1 to 7, usually relatively closely spaced; pedicels erect at anthesis, usually ascending or spreading in fruit, 0.2 to 1.8 cm long at anthesis. Calyx lobes subequal, suborbicular, 2.5 to 3.5 mm long, glandular, upper lobe rounded, lower lobe rounded to retuse; corolla 0.9 to 2 cm long, yellow with brown markings and nerves, upper lip about as long as the lower, rounded, lower lip with a bilobed swelling at the base and the margin 3-lobed, usually more or less deeply, spur narrow-conic, shorter than the lower lip, tip rounded to subacute, rarely briefly bifid; ovary globose, lower lip of stigma suborbicular and ciliate, upper lip rudimentary. Capsule globose, 3 to 5 mm in diameter, topped by a style ca. 1 mm long, circumscissile; seeds flattened-globose, ca. 1 mm broad, studded with many blunt projections ca. 0.2 to 0.3 mm long. Still water of ponds, lakes, ditches, swamps, and slow-moving streams. E. TX; N. S. and ME, S. to FL, W. to TN, IN, and TX. Mar.-June. [U. inflata Walt. var. minor Chapm. and var. radiata (Small) Stone].


2.U. cornutaLENTIBULARIACEAE UTRICULARIA cornuta Michx. Horned Bladderwort. Plants terrestrial or amphibious, probably perennial; rhizoids many, 1 to 2 cm long, well-branched, glandular; stolons many, slender, to several cm long, branched. Leaves numerous from the stolons, undivided, narrowly linear or oblanceolate, to 4 cm or more long, to 0.5 mm broad, acute, 1-nerved, petioles none or indistinct. Traps many on the rhizoids, stolons, and leaves, ovoid, 0.3 to 0.8 mm long, mouth lateral and without bristles but with a dorsal lip. Inflorescence solitary, scapose, 4 to 40 cm long; peduncle straight, glabrous above and with a few glands below, with numerous scales (sterile bracts); bracts below pedicels attached basally, widely ovate-deltoid, acute, ca. 2 mm long; bracteoles narrowly lanceolate, acute to acuminate, slightly longer than the bracts, 1-nerved and keeled; flowers 1 to 6, usually crowded; pedicels 1 to 2 mm long, erect. Calyx lobes unequal, the upper ovate, 4.5 to 7 mm long, acute, glabrous, plicate and ridged along the nerves, lower lip shorter, ovate-deltoid, more or less flat and only inconspicuously ridged, apex bifid with 2 sharp points; corolla yellow, sometimes marked with copper, 1.5 to 2 cm long, upper lip shorter than the lower, ovate-oblong, retuse, base narrowed, lower lip convex-galeate, more or less orbicular in outline, entire, the palate with a marginal rim, spur slender-conic, curved forward, very pointed, slightly longer than the lower lip, ca. 7 to 12 mm long; ovary ovoid, lower lip of stigma suborbicular, upper similar but smaller. Capsule ovoid, ca. 4 mm long, dehiscent by 1 ventral, longitudinal slit with thickened margins; seeds ca. 0.25 mm long, ellipsoid-ovoid, reticulate-rugose. Marshes, swamps, bogs, pools, and muddy shores, quite showy where numerous; E. 1/2 TX;

Newf. to N. Ont. and MN, S. to FL, W. to Alta., OH, MI, AR, and TX; also Caribbean. Apr.-Sept.

This species is superficially similar to U. juncea Vahl, which is found in E. TX and which may some day be found here. U. juncea has a looser raceme with more and smaller (to 1.5 cm long) flowers whose spurs have a bifid tip.


3.U. subulataLENTIBULARIACEAE UTRICULARIA subulata L. Zig-zag Bladderwort. Annual, usually delicate and often diminutive, terrestrial or in wet places; rhizoids in groups of ca. 6, usually less than 1 cm long, with short branches; stolons many, filiform, branched, several cm long. Leaves usually many, undivided, petiolate, narrowly linear to narrowly oblanceolate, subacute, to 2 cm long and 1 mm broad. Traps numerous on stolons and leaves, ovate, 0.2 to 0.7 mm long, mouth lateral, with 2 subulate, bristle-like appendages. Inflorescences solitary, simple or sometimes branched, often zig-zagging from node to node, (2)10 to 50 cm tall; peduncle filiform, glabrous or commonly papillose to short-hispid below; scales many, peltate more or less lanceolate to fusiform, the end acute, sometimes ciliate; bracts subtending pedicels peltate, shield-like, circular to transversely elliptic, ends rounded, 1 to 2 mm long; bracteoles none; flowers 1 to 25, well-spaced, pedicels ascending, very slender, 0.2 to 1.5 cm long. Calyx lobes about equal, broadly ovate to orbicular, rounded to subtruncate, 1 to 1.5 mm long; chasmogamous flowers: corolla yellow, 0.5 to 1 cm long, upper lip much shorter than the lower, broadly ovate, rounded, lower lip rhombic to widely cuneate, with a prominent rounded swelling at the base, margin deeply 3-lobed, palate pubescent, spur narrowly cylindric to subulate, slightly shorter than to slightly longer than the lower lip, often inconspicuous in pressed specimens, tip rounded or with 2 to 4 very small teeth; cleistogamous flowers white or reddish. Ovary globose, style very short, lower lip of stigma semicircular, upper lip reduced. Capsule globose, 1 to 1.5 mm in diameter, dehiscent by an ovate ventral pore or flap; seeds obovoid, ca. 0.23 mm long, 0.16 mm broad. In peat and sand of bogs or on seepage slopes or pond shores. E. TX (also Winkler Co. in W. TX); nearly pantropical; in N. Amer. from N. S. and N. Eng. S. to FL, W. to TN, AR, and TX; also Mex., Caribb., Cen. and S. Amer. Mar.-June. [Includes var. cleistogama Gray, U. cleistogama (Gray) Britt; many other synonyms exist, but none frequently encountered in modern treatments for our region. Erroneously omitted from the checklist of Hatch, et al. (1990).].


4.U. gibbaLENTIBULARIACEAE UTRICULARIA gibba L. Cone-spur Bladderwort, Swollen-spurred Bladderwort. Plants annual or perennial, usually floating, but sometimes anchored in floating debris or on mud; rhizoids absent or else a few cm long and with a few very short dichotomous branches; stolons usually many, well-branched and forming mats, to 20 cm long or more. Leaves many, 0.5 to 1.5 cm long, usually with 2 primary segments (sometimes only 1), sparingly dichotomously branched with up to (4)8 capillary ultimate segments or sometimes 1 branch of each pair replaced by a trap. Traps relatively few, lateral on the primary to penultimate leaf divisions, ovoid, 1 to 2.5 mm long, mouth lateral with 2 dorsal, bristle-like, branched appendages and lateral and ventral bristles. Inflorescences emergent, erect (sometimes reduced and submersed, bearing cleistogamous flowers), 1 to 20(30) cm long, solitary or commonly 2 together or produced at a short interval; peduncle straight (not zig-zag); scales usually 1(2) at about the middle or often absent, similar to the bracts; bracts attached by their bases, semi-clasping, more or less semicircular, apically rounded to truncate or slightly toothed, ca. 1 mm long; bracteoles none; flowers (1)2 to 6(12), crowded or well-spaced; pedicels erect or spreading, 0.2 to 3 cm long. Calyx lobes subequal, broadly ovate to orbicular, rounded, 1 to 3 mm long; corolla yellow, often with rusty nerves, 4 to 25 mm long, the 2 lips nearly equal or the upper larger (in pressed specimens the flowers often horizontal, in which case the lower lip is usually the one with the spur), upper lip broadly ovate to suborbicular, usually slightly to markedly 3-lobed, lower lip orbicular to transversely elliptic, base with a prominent rounded swelling, margin rounded, palate pubescent, spur slenderly conical or narrowly cylindric, slightly shorter to markedly longer than the lower lip, tip obtuse or obscurely bifid; ovary globose, lower lip of stigma transversely elliptic, upper lip obsolete. Capsule globose, 2 to 3 mm in diameter, tardily dehiscent by 2 lateral valves; seeds imbricate on the placenta, plumply lenticular, roughly circular in outline, 0.8 to 1 mm in diameter, with a broad, thick, encircling wing which has a rounded lobe on the portion nearest the apex of the capsule, the remainder of the margin minutely and irregularly dentate, ventral surface of the body smooth or sometimes with bluntly-conical projections. Mud or shallow water of bogs or seepage slopes; if in deeper water usually not flowering or flowering only if supported by floating debris or submerged vegetation. E. 1/2 TX; pantropical; in N. Amer. extending to E. and W. U.S. and Can. (absent from the Great Plains and Rocky Mts., N. to N. S. and B.C.; also in Cen. and S. Amer., Europe, Asia, Afr., and Oceania. Jun.-Oct. [U. biflora Lam., U. pumila Walt., U. obtusa Sw.].

According to Taylor (1989), U. gibba is very variable and includes plants formerly separated as U. biflora and having larger flowers with relatively longer, more slender spurs. Taylor points out that there is a continuum of variation in corolla size and shape and that there is no correlation with variance in vegetative characters.






CAMPANULACEAE(including Sphenocleaceae)
CAMPANULACEAE

Bluebell Family


Texas material annual or perennial herbs (elsewhere some shrubs or stem-succulent trees). Leaves alternate, usually spirally arranged, estipulate, usually simple and always so in ours. Inflorescence racemose, spicate, or cymose; flowers usually perfectly 5-merous except the gynoecium. Calyx in ours usually 5-parted, usually free above the ovary or sometimes briefly united. Corolla in ours of 5 united petals, regular or irregular (as in Lobelia). Stamens in ours epipetalous or attached to a nectary disk, alternate with the corolla lobes, free or connivent (as in Campanula) or more or less connate by the anthers and filaments (as in Lobelia). Ovary in ours at least partly inferior, of 2 to 5 united carpels; style 1, usually with a tuft of pollen-collecting hairs below the (1-)2- to 5-lobed stigma. Fruit a capsule (as in ours) or berrylike, usually with as many locules as carpels and axile placentation, but occasionally unilocular with 2 parietal placentae; seeds many.

Including Sphenoclea, 88 genera and about 1,950 species distributed world wide; 5 genera and 19 species in TX; 3 genera and 6 species in our area.

Many taxa are cultivated for ornament, especially species of Campanula--Bellflower, Platycodon--Balloon Flower, and Lobelia. Others have medicinal properties (Mabberley 1987).



1. Flowers in dense, leafless terminal spikes; flowers ca. 2.5 mm long, white; capsule circumscissile ....................................................................................................1. Sphenoclea

1. Flowers in more open, leafy-bracted spikes or racemes; flowers more than 2.5 mm long, usually red or blue; capsule loculicidally dehiscent or with a lateral pore .............................2


2(1) Corolla zygomorphic; anthers united around the style ...........................................2. Lobelia

2. Corolla regular; anthers free ................................................................................3. Triodanis



CAMPANULACEAE SPHENOCLEA1. SPHENOCLEA Gaertn.


An Old World genus with 1 or 2 species, 1 widely introduced or naturalized in warm areas of the New World. This genus is sometimes segregated as a separate family, the Sphenocleaceae. It differs from the rest of the Campanulaceae by having rounded anthers, truly epipetalous filaments, and a more or less unlobed style, and it lacks the typical brush of hairs on the style. Its taxonomic affinities are unclear, though Cosner, et al. (1994) found that DNA analysis place it closest to the Solanales.


1. S. zeylanica Gaertn. Chicken Spike, Piefruit. Annual; stem to 1 m or more tall, branched from the axils, when growing in water becoming thick and spongy or corky basally, hollow, and often with roots at the nodes. Petioles to 2 cm long; blades elliptic, entire, 6 to 12 cm long, 2 to 3.5 cm broad, tapered to both ends. Inflorescences terminal on the branches, dense, cylindrical spikes 2.5 to 8 cm long, to 10 mm broad, tapered to the tip; peduncle 1.5 to 3.5(10) cm long; each flower subtended by a tiny spatulate bract 2 to 3 mm long and 1 to 1.3 mm broad, and by 2 smaller bractlets ca. 1 mm broad; pedicels none. Calyx lobes triangular to suborbicular, 1.5 to 2 mm broad, a little wider than long, tips rounded, erose and scarious, at maturity flattened over and nearly concealing the ovary, deciduous with the fruit; corolla white, ca. 2.5 mm long, slightly longer than the calyx, lobes triangular, obtuse, ca. 1 mm long, tube 1 to 1.5 mm long; stamens inserted below the middle of the corolla tube, anthers attached dorsally, stamens deciduous with the corolla after anthesis; ovary fully inferior, style erect, 0.5 mm long. Capsule ca. 2.5 to 4 mm broad, bilocular, the septum perpendicular to the axis of the spike, circumscissily dehiscent, the calyx falling with the top portion; seeds many, ca. 0.4 to 0.5 mm long, pale yellow-brown, minutely pitted and somewhat shiny. Wet lowlands and flats, also common in rice fields. Not common in our area, but known from Washington Co. (McVaugh 1951); E. and S. TX; native of the Old World tropics and widely naturalized; in N. Amer. from SC to FL, W. to TX and AR.

This plant is a common weed of rice fields. In some parts of the world, it is eaten as a leaf vegetable (Mabberley 1987).



CAMPANULACEAE LOBELIA2. LOBELIA L. Lobelia


Ours annual, biennial, or perennial herbs; sap sometimes milky and/or acrid. Stems simple or branched. Leaves alternate. Inflorescence usually racemose, with leafy bracts, sometimes the racemes panicled; flowers inverted (resupinate) at anthesis, the pedicel twisted. Calyx regular, 5-parted, in some taxa the lobes with basal auriculate appendages. Corolla red, purple, or blue, occasionally white or pink, the upper lip with 2 lobes and the lower lip with 3 usually shorter and broader lobes, tube cleft nearly to the base between the 2 upper lobes, in some species the tube also incompletely separated from the bottom up on either side, forming "windows" and the tube then said to be fenestrate. Stamens 5, anthers united around the style, the 2 lower ones smaller and with tufts of pale hairs apically, the 3 larger ones pubescent dorsally or at the tip, filaments united above for ca. 1/3 to 1/2 their length. Ovary inferior (occasionally more or less free of the hypanthium), bilocular; style elongating and pushing pollen out of the stamen tube, stigma 2-lobed, with a ring of hairs below the tip. Fruit in ours a 2-valved capsule loculicidal at the apex. Seeds many, small, pitted-reticulate.

About 365 species, primarily of tropical and warmer parts of the world, a few temperate in distribution; 9 species listed for TX (Hatch, et al. 1990); 3 rather variable species present in our area.

Many species are cultivated for their flowers, especially L. cardinalis (with red flowers) and L. erinus (with blue or white flowers), commonly used as a bedding plant and in hanging baskets. The genus has an alkaloid chemistry; some taxa, especially L. inflata, have medicinal uses. Some species in Africa are tree-like stem-succulents (Mabberley 1987).


1. Flowers red; corolla 3.5 cm long or more; filaments 20 mm long or more ...1.L.cardinalis

1. Flowers blue, purple, lavender, or white; corolla usually not more than 2.5 cm long; filaments 15 mm long or less ...................................................................................................2


2(1) Filaments more than 5 mm long; corolla tube 6.5 mm long or more; flowering Apr.-June ....

...2.L.puberula

2. Filaments less than 5 mm long; corolla tube 6 mm long or less; flowering Aug.-Sept. ........

...3.L.appendiculata


1.L. cardinalisCAMPANULACEAE LOBELIA cardinalis L. Cardinal Flower. Perennial, reproducing by basal offshoots; stems usually simple, 40 cm to 2 m tall, 2 to 15 mm in diameter, glabrous to pubescent, sometimes reddish-purple below; herbage glabrous to hirtellous. Petioles 0.5 to 1.5 cm long or the upper leaves sessile or nearly so; blades ovate to lance-ovate, linear, or oblanceolate (especially below), 2.5 to 20 cm long, 0.5 to 5 cm broad, largest usually at midstem, acute to acuminate or the lower sometimes nearly obtuse, tapered to the base, margins denticulate to irregularly serrate. Inflorescence a terminal raceme 1 to 7 dm long, with few to 100 flowers, 1 per node, often somewhat secund (1-sided); floral bracts leaf-like below, linear or the lower slightly broader, reduced upwards, and at the top of the inflorescence shorter than the flowers; pedicels bristly-pubescent, ca. 3 to 10 mm long, shorter than the bracts, each with 2 bracteoles at the base. Flowers 3 to 5 cm long; calyx body campanulate to conic, 5 to 15 mm long, strongly ribbed, glabrous to sparsely hispid, teeth herbaceous, slenderly triangular to subulate, with short, deltoid bases, acuminate, 8 to 16(20) mm long, sometimes with minute basal auricles, glabrous or the tips ciliate, calyx enlarging and the lobes elongating after anthesis; corolla typically bright red (rare albino or pink forms occur), glabrous to slightly puberulent, especially within, tube generally 15 to 20 mm long, fenestrate as well as split down one side, the lobes of the lower lip spreading-deflexed, ovate to narrowly elliptic, from ca. 1/3 to 1/2 to nearly as long as the tube, upper 2 lobes erect and linear or linear-lanceolate, shorter than the lower lobes; filament tube 15 to 30 mm long, longer than the corolla, reddish, the filaments united for more than 1/2 their length, usually ciliate basally, anther tube bluish-gray, ca. 4 to 5.5 mm long, the 2 smaller anthers with a tuft of white hairs, the 3 larger anthers glabrous to lightly pubescent. Capsule about 1/2 inferior, 5 to 8(10) mm long, ca. 6 to 10 mm broad, cup-shaped; seeds ovoid to ellipsoid, ca. 0.6 to 1.2 mm long, areolate and commonly tuberculate to some degree. Wet to moist soil of meadows, streambanks, pond margins, swamps, and open woods, mostly where shade is not too heavy. Throughout most of TX where habitat is suitable, except the far S. portion; N. B., S. to FL, W. to S. Ont., MI, WI, NV, and CA; also Mex. May-Dec.; TAMU collections primarily Sept.-Oct.

This is a variable species. The traditional approach has been to recognize 2 subspecies, one in the east and another in the west, the eastern subspecies with 2 varieties and the western one with 3--our plants were then subsp. cardinalis var. cardinalis. However, a recent study of variation in the species throughout its range (Wysocki and Lammers 1995) revealed a continuum of variation with no justifiable categories at the infraspecific level. All subspecific and varietal names are thus rendered synonyms.

The plant is frequently cultivated for its showy flowers, which are attractive to hummingbirds. The roots and flowers were used in love charms by several Native American tribes, and the Cherokee cultivated it as a medicinal plant (Kindscher 1992).


2.L. puberulaCAMPANULACEAE LOBELIA puberula Michx. Downy Lobelia, Purple Dewdrop. Perennial from an erect caudex; stem erect, usually simple, 3 to 16 dm tall, commonly more or less tinged with purple, usually densely short-hirsute, sometimes pilose or rarely glabrate. Leaves variable in number, sessile or the lower tapered to the base, the lower larger, 3 to 12 cm long, to 4 cm broad, usually 2 to 4 times longer than wide, oblanceolate to obovate, oblong, elliptic, or lanceolate, mid-cauline leaves largest, reduced upwards, uppermost more or less grading into the floral bracts, all acute to obtuse, both surfaces more or less pubescent, margins subentire to serrate or dentate, the teeth callous-tipped or represented by small callosities, upper leaves usually more sharply and deeply toothed, often with small teeth alternating with the larger. Racemes (2)15 to 30(5) cm long, dense or interrupted, with up to 75(140) flowers; floral bracts usually more or less foliaceous, lanceolate, toothed, reduced upwards, mid-raceme bracts generally 1 to 1.5(2.5) cm long, 3 to 5(8) mm broad, acute; pedicels stout, 3 to 5 mm long in fruit, hirtellous; bracteoles 2, filiform, green, at the base of the pedicel or slightly above, 1 to 1.5 mm long. Flowers ca. 15 to 24 mm long; calyx tube/hypanthium more or less shallow at anthesis, becoming hemispheric with age, broadest above, with a flaring rim, strongly ribbed, somewhat angular, 5 to 9 mm wide, sometimes densely pubescent, calyx lobes lanceolate to lance-subulate, with broad bases and long-tapering tips, 5 to 12(15) mm long, usually 2 to 4 mm broad at the base, margins usually ciliate, entire to callose-denticulate or occasionally with definite teeth, auricles small, triangular, declined; corolla purple or blue with a white "eye" (varying to pink or occasionally white), tube 5 to 8 mm long, fenestrate in addition to the lateral split, usually pubescent externally, interior of lip smooth to hirsute, the 3 lobes of the lower lip oblong to ovate, usually at least slightly longer than the tube, acute to obtuse, 2 upper lobes lanceolate, erect; filament tube 6 to 15 mm long, pale blue-gray, basally pubescent, commonly united 1/3 its length, anther tube 3 to 6 mm long, blue-gray, the 2 small anthers with terminal tufts and the 3 larger dorsally pubescent. Capsule ca. 1/2 inferior, oblong to hemispheric, 4 to 7 mm long, broader than long, to ca. 10 mm broad; seeds oblong-linear or ellipsoid, 0.5 to 0.7(1) mm long, lustrous, alveolate-reticulate. Usually in moist areas--bogs, moist woods, wet fields, thickets, roadsides, etc., often in acid soils. E. TX; S. NJ. to IL, S. to N. FL, E. TX, and OK. Aug.-Sept.

This species is wide-ranging and quite variable. Several intergrading varieties or subspecies have been described, though some modern sources (e.g., Godfrey and Wooten 1979) chose not to recognize them. In our area, we tend to have 2 types of plants. The first kind are rather robust, puberulent to glabrous plants with many flowers, filament tube 6 to 7(9) mm long, calyx not densely pilose basally, and white eye of corolla continuous. These are found throughout the area but usually not in bogs. The second kind are slender, rather hirsute plants with very few flowers, filament tube 8 to 11 mm long, calyx tube densely pilose, and white eye of corolla divided into 2 lens-shaped patches. These plants are usually found in acid soils of bogs. The first type were treated by Correll and Johnston (1970) as L. puberula Michx., and by both Kartesz (1998) and McVaugh (1951) as L. puberula Michx. var. puberula. The few-flowered bog plants were treated by Correll and Johnston as L. reverchonii B. L. Turner and by Kartesz and McVaugh as L. puberula Michx. var. pauciflora Bush [= subsp. pauciflora (Bush) Bowden]. It may be true that these bog plants represent merely a collection of characters distributed throughout the range of variation in the species (McVaugh 1951) and are not worthy of separate taxonomic treatment. In our area, however, the two forms are certainly distinct, and there are few, if any, intermediates.


3.L. appendiculataCAMPANULACEAE LOBELIA appendiculata A. DC. Earflower Lobelia. Annual or biennial from fibrous roots; stem erect, simple or with a few branches, 1.5 to 9 dm tall, glabrous or sparsely chaffy-pubescent with short hairs, especially on the angles below. Basal leaves commonly with blades lanceolate to oblanceolate or elliptic, ca. 2 to 6 cm long, rounded apically, petioles to 1 cm long, shorter than the blades and margined, but often these basal leaves absent by flowering time, stem leaves thin, glabrous or hirtellous at the base on the upper side and on the margins, more or less sessile and often clasping, blades oblong below, passing to oblong, elliptic, or ovate above, the larger, lower ones to ca. 8 cm long and 1 to 2(3) cm broad, cuneate to rounded basally and apically rounded, margins more or less denticulate with minute callosities, upper leaves smaller, more pointed, and generally sessile, the last ca. 4 to 10 cm below the flowers. Raceme loosely-flowered, 6 to 18(30) cm long, sometimes 1-sided, single, or the plant with a few weak racemes from the upper axils; floral bracts markedly different from the stem leaves, 8 to 18 mm long, longer than the pedicels, linear to narrowly lanceolate or subulate, margins callous-denticulate; pedicels 4 to 8 mm long, with 2 minute (0.5 to 0.7 mm long), rough-hairy bracteoles basally. Flowers 10 to 15 mm long; calyx tube in anthesis ca. 1 mm long, conic, long-campanulate in fruit and 5 to 6 mm long, somewhat inflated, to 3.5 to 6 mm broad, lobes linear-subulate to linear-lanceolate, ultimately 4 to 8 mm long, bristly ciliate, at maturity with basal auricles 1 to 3(5) mm long, drying blue or purplish, ciliate-margined, usually at least partly obscuring the calyx tube; corolla lilac to violet, pubescent at the base of the lip within and with a pair of tubercles, tube 4 to 5 mm long, split down 1 side but not also fenestrate, the 3 lobes of the lower lip broadly ovate, about as long as the tube, the 2 upper lobes lanceolate and shorter; filament tube 2 to 4 mm long, glabrous and slightly deflexed, pale, filaments united about 1/2 their length, anther tube 1.9 to 2.5 mm long, blue-gray, the 2 small anthers tufted and the larger usually dorsally pubescent. Capsule 2/3 or more inferior, horizontal or nodding, nearly hemispheric, about twice as long as wide; seeds 0.4 to 0.6 mm long, irregularly oblong, amber, lustrous, alveolate-reticulate. Open sandy areas such as pinelands, prairies, old fields, and roadsides, often where moist. E. TX; AL to TX, AR, OK, KS, MO, and IL. Apr.-June. [If plants treated by some as L. gattingeri Gray are submersed in this species, they become var. gattingeri (Gray) McVaugh and our plants become var. appendiculata.

This plant is very similar to L. spicata Lam., which differs (theoretically) in having small triangular auricles which do not obscure the calyx, stem more or less pubescent all the way around (at least below), calyx lobes ciliate, and leaves tapered to the base and not clasping. However, at least in E. TX, it is possible to find plants intermediate in both floral and vegetative characters, and the 2 extremes growing side by side. It is the author's suspicion that the two species may represent a single variable entity. In our immediate area, however, our plants are nearly all of the definitely "appendiculata" type. The epithet spicata has priority should the two species ever be merged.



CAMPANULACEAE TRIODANIS3. TRIODANIS Raf. ex Greene Venus' Looking-glass


Taprooted annuals. Stems simple or branched below (rarely to midstem), commonly 5-angled. Leaves alternate, passing into the floral bracts above, commonly glabrous above and with some form of pubescence below, the decurrent bases merging into the stem angles. Flowers usually blue or purple, more or less sessile, in groups of 1 to 3 in the axils, the lower usually reduced and cleistogamous, later flowers chasmogamous. Calyx with 3 to 5 usually herbaceous, equal lobes, these usually entire save for 1 small glandular-callose tooth on each margin near the base. Corolla rotate to short-funnelform, with 5 equal, spreading lobes. Stamens 5, filaments linear above, broader and usually pubescent below; anthers linear, longer than the filaments. Style 1, pubescent distally, stigma with 3 to 5 lobes. Corolla, stamens, and style reduced in cleistogamous flowers. Capsule cylindric to prismatic, clavate, obovoid, or ellipsoid, topped by the persistent calyx lobes, 1- to 3-celled, dehiscent usually by as many lateral pores as cells, the pores linear to oval, opening from base to apex as a cartilaginous flap curves upward. Seeds many, most of them the product of the cleistogamous flowers (but not always fertile), brownish, usually less than 1 mm long, lenticular or quadrangular, placentation usually axile (parietal in some species, but not ours).

7 species of N. Amer. and 1 in the Mediterranean; 7 in TX; 2 here.


1. Seeds more or less quadrangular (use strong magnification), the surface with low

longitudinal ridges ...1.T.texana

1. Seeds lenticular or biconvex, smooth or rough, often lustrous ...2.T.perfoliata


1.T. texanaCAMPANULACEAE TRIODANIS texana McVaugh Plants slender; stems (10)25 to 50(75) cm tall, hirsute below, upper portions retrorsely hispid or scabrous. Leaves (5)15 to 30(40) mm long, 6 to 15 mm broad, generally 1.5 to 2 times longer than wide, rounded to obtuse, leaves sessile or the lower tapered to the base, lower blades oblong to obovate, midstem leaves elliptic, passing to ovate above, lower leaves hirtellous to hirsute below, upper leaves and bracts sparingly hirtellous to glabrous, all bracts and leaves marginally crenate and ciliate. Floral bracts broader, more ovate, somewhat clasping, obtuse to acute and/or mucronate, the median ones 4 to 22 mm long, 6 to 7 mm wide. Flowers 1 to 3 per node in 10 to 30(40) axils, the lower 1/3 to 2/3 cleistogamous and the upper chasmogamous, the terminal one cleistogamous or chasmogamous, usually only 1 flower per node maturing fruit. Calyx lobes triangular to triangular-ovate, acute and often mucronate or with a slender tip, glabrous to slightly ciliate. Cleistogamous flowers: calyx lobes 3 to 5, (1)1.5 to 3 mm long, 0.5 to 1.2 mm broad, sometimes dissimilar; corolla less than 0.5 mm long. Chasmogamous flowers: calyx lobes 3 to 6(7) mm long, 0.9 to 1.5(2) mm broad; flowers lavender to purple, 7 to 10(14) mm long, the tube 1.5 to 2.5 mm long, lobes 2 to 5 mm broad. Filaments 1.5 to 2.2 mm long, anthers (1.7)2.1 to 3 mm long; style 5 to 7.5 mm long, branches 1 mm long or less. Capsule glabrous to sparingly hirtellous, ellipsoid to ovoid or those of chasmogamous flowers truncate and clavate, 4 to 6(7) mm long, 1.5 to 2.5 mm broad, dehiscence pores usually 2 on cleistogamous fruits, oval, at about the middle of the capsule, the central cartilage narrow and the margins broad and scarious, curled after dehiscence; seeds dark brown, shiny, 0.4 to 0.6 mm long, 0.2 to 0.3 mm long, 4-angled with flat or concave faces and rounded angles, surface with low, narrow, anastomosing longitudinal ridges. Woods, prairies, pinelands, depressions, and open banks, generally in sandy soils. Endemic to E. TX. Apr.-May. [Specularia texana McVaugh].


2.T. perfoliataCAMPANULACEAE TRIODANIS perfoliata (L.) Nieuw. Taprooted annual, erect to reclining; stems (8 or less)15 to 60(80 or more) cm tall, usually simple and slender, basally pilose to hispid or hirsute, upper parts similarly pubescent or else merely retrorsely hispid or scabrous. Leaves sessile or the lowermost short-petiolate, 10 to 35 mm long, 6 to 25 mm broad, 1.5 to 2.5(3) times longer than wide, ovate-cordate to ovate or elliptic, or the lowermost (commonly absent at flowering time) obovate or elliptic, all blunt to rounded or obtuse, becoming more acute upwards, crenate and/or undulate-crisped, lower surface scabrous or with hairs like the stems (especially on the veins), varying to glabrous, bracts and leaves generally ciliate-margined; bracts generally broader and more ovate than the leaves, sometimes even reniform, acute, commonly mucronate, often more coarsely toothed or sometimes with enlarged basal teeth and nearly lobed, occasionally nearly glabrous, often not concealing the fruit, the median ones 25 mm long and 30 mm broad, (0.5)0.6 to 1.8(2.5) times longer than wide. Flowers 1 to 4(8) per node, usually all but one aborting, at 10 to 50 nodes, 1/3 to 2/3 or nearly all the nodes producing cleistogamous flowers, terminal node usually with a chasmogamous flower. Calyx lobes deltoid to lanceolate or deltoid-ovate, tip acute to subulate or sometimes with a capillary bristle, glabrous to hispid-scabrous, at least on the margins; corolla less than 0.5 mm long. Cleistogamous flowers: calyx lobes 1 to 5, 1 to 3(4) mm long, 0.3 to 2 mm broad. Chasmogamous flowers: calyx lobes (3.5)4 to 8.5 mm long, 0.7 to 2.5(3) mm broad; corolla lavender to purple or bluish (rarely white), tube usually bluish, 5 to 12(13) mm long, tube 1 to 3.5(5) mm long and lobes 2 to 4.5(5.5) mm wide. Filaments 1.3 to 2.5 mm long, anthers (1.4)1.6 to 3.3 mm long; style 3.5 to 7.5 mm long, branches less than 1 mm to ca. 1.5 mm long. Capsule glabrous to scarious or sparingly hispid, (2)4.5 to 7(9) mm long in cleistogamous flowers, to 10 mm long from chasmogamous flowers, obovoid, ovoid, ellipsoid, clavate, or oblong, dehiscence pores rounded to oval or elliptic, varying in position on the capsule; seeds ellipsoid, chestnut-brown, biconvex or slightly flattened, smooth to muriculate, faintly lustrous to highly polished, 0.5 to 0.65 mm long. Roadsides, waste places, prairies, moist banks, canyons, cultivated fields, etc., usually at low elevations. Common in TX except in the High Plains and avoiding desserts and permanently wet places. Mar.-July.

The above description includes characters found throughout the whole of the species. There are two varieties, both present in our area. These were formerly treated as separate species, but Bradley (1975) determined them to be part of the same taxon. Intermediates do occur, as do plants with characters of both this species and T. texana. Further investigation may elucidate the relationship.


var. perfoliata Clasping Venus' Looking-glass. Bracts as wide or wider than long, often with 1 or 2 pairs of veins visible below in addition to the midvein; usually several of the upper nodes with chasmogamous flowers; corolla of chasmogamous flowers 8 to 12(13) mm long; capsule 4 to 6(8) mm long from cleistogamous flowers and to 10 mm long from chasmogamous flowers; dehiscence pore well below the apex of the capsule, usually by 1 to 1.5 mm and at about the middle of the capsule, often near the base; seeds 0.5 to 0.6 mm long, usually not shiny but sometimes smooth or else muriculate to varying degrees. ME and Ont. to B.C., S. to FL, E. TX, KS, CO, MN, AZ, CA, and Mex. [Specularia perfoliata (L.) A. DC.].


var. biflora (R. & P.) Bradley Small Venus' Looking-glass. Bracts usually longer than wide, usually only the midvein prominent below or sometimes 1 additional pair obvious; only the terminal flower chasmogamous and the rest cleistogamous or sometimes scattered chasmogamous flowers present; corolla of chasmogamous flowers 5 to 9(11) mm long; capsule (2)4.5 to 7(9) mm long, the pore very near the apex, mostly only 0.3 to 0.5 mm below it; seeds 0.5 to 0.65 mm long, smooth and lustrous. VA to NE, S. to FL and TX, also S. OR to Baja California; also S. Amer.. [T. biflora (R. & P.) Greene; Specularia biflora (R. & P.) A. Gray].

Because this variety has a high percentage of cleistogamous flowers, it tends to preserve its suite of characters and remain fairly distinct from var. perfoliata.







RUBIACEAERUBIACEAE

Coffee or Madder Family


Herbs or shrubs (elsewhere, especially the tropics, also commonly trees or vines). Leaves opposite or whorled, simple and entire; stipules present, often interpetiolar and conspicuous, sometimes foliose and adding to the apparent number of leaves per node, sometimes deciduous. Flowers solitary or in cymes, panicles, or heads, perfect (as ours) or unisexual, regular, epigynous. Sepals in ours 4(6 or 8) or sometimes absent, crowning the ovary and persistent or sometimes circumscissily deciduous. Corolla in ours funnelform, salverform, or rotate, with (3)4(5) lobes valvate, imbricate, or contorted in bud. Stamens usually as many as the petals, inserted in the corolla throat, anthers introrse. Ovary inferior or half inferior, often topped with a nectary disk, (1-)2-(3-) celled, ovules 1 to many per cell; style 1, commonly divided above, flowers sometimes heterostylous. Fruit a capsule, berry, drupe, or often schizocarpic, separating into nutlet-like units or cocci.

630 genera and about 10,400 species worldwide, especially common in the tropics or subtropics; 16 genera and 59 species in TX; 11 genera and 23 here.

The family includes the economically important coffee (Coffea) and quinine (Cinchona). Some tropical species are timber trees. Ornamental genera include Gardenia, Ixora, Pentas, and so on (Mabberley 1987).



1. Plants shrubs or small trees .........................................................................1. Cephalanthus

1. Plants herbs ..............................................................................................................................2


2(1) Leaves apparently whorled ......................................................................................................3

2. Leaves apparently opposite .....................................................................................................4


3(2) Flowers 4 to 8 in terminal heads, subsessile; corolla pale purple .....................2. Sherardia

3. Flowers in axillary or terminal cymes or 1 to few on small branchlets, sessile to distinctly pedicelled; corolla white or greenish ........................................................................3. Galium


4(2) Plants evergreen, trailing herbs; fruit a red, berry-like drupe; flowers in pairs with their ovaries united .........................................................................................................4. Mitchella

4. Plants non-evergreen herbs, erect to prostrate; fruit dry; ovaries of separate flowers not united .........................................................................................................................................5


5(4) Fruit a circumscissile capsule ..........................................................................5. Mitracarpus

5. Fruit an apically-dehiscent capsule or else schizocarpic and splitting into single-seeded units ...........................................................................................................................................6


6(5) Ovules and seeds more than 1 in each cell, the 2-celled capsule dehiscent loculicidally across its apex ..........................................................................................................................7

6. Ovules and seeds only 1 in each cell; fruit schizocarpic .........................................................9


7(6) Flowers in terminal cymes .....................................................................................6. Hedyotis

7. Flowers axillary or else solitary and terminal ..........................................................................8


8(7) Flowers sessile, 1 to few in the axils of the leaves and/or in a dense terminal cluster; ovary completely inferior; seeds angled ...................................................................7. Oldenlandia

8. Flowers pedicellate, axillary or solitary and terminal; ovary partially superior; seeds globose with a ventral depression or else saucer-shaped ................................8. Houstonia


9(6) Flowers in terminal, bracted clusters; ovary and fruit with 2 to 4 (usually 3) cells; calyx falling as a unit before carpels separate .............................................................9. Richardia

9. Flowers axillary or both terminal and axillary; ovary and fruit with 2 cells; calyx teeth remaining on the carpels after they separate from one another ..........................................10


10(9) Flowers in dense axillary and terminal clusters; mature carpels separating from each other, one of them taking the septum and so closed, the other open; plants glabrous ..........

. ......................................................................................................................10. Spermacoce

10. Flowers 1 to 3 in the axils of the leaves; mature carpels separating but usually both remaining closed; plants wholly or partially pubescent or scabrous .....................11. Diodia






RUBIACEAE CEPHALANTHUS1. CEPHALANTHUS L. Buttonbush


Shrubs (most of ours) or small trees. Leaves opposite or sometimes whorled, petiolate. Flowers in dense spherical, pedunculate heads. Calyx tube obconic-obpyramidal, with 4 or 5 teeth. Corolla whitish or yellowish, tubular-funnelform, with 4 or 5 teeth imbricate in bud. Stamens 4; anthers with 2 basal cusps. Ovary 2-celled, each cell with 1(to 3) ovules; style filiform, long-exserted, stigma capitate. Individual fruits small, dehiscent from the base upwards into 2 to 4 closed, 1-seeded units.

6 species of N. America and the tropic; 2 in TX; 1 here.


1. C. occidentalis L. Common Buttonbush, Honey-balls, Globe-flowers. Shrub or small tree, to 15 m tall but usually shorter; trunk to 3 dm in diameter, usually smaller, stems usually several from the base; twigs brownish-gray, glabrous or puberulent. Leaves opposite or sometimes in 3's or 4's, petioles 0.5 to 3(5) cm long, glabrous to puberulent; blades ovate or ovate-oblong to lanceolate or broadly elliptic, (6)9 to 15(19) cm long, 3 to 5(9) cm broad, abruptly short- or long-acuminate, basally cuneate to subcordate, glabrous above, glabrous to pilose below; stipules 2 to 4(6) mm long, deltoid, acute to acuminate, usually with glandular-dentate margins, closer to the petiole bases than the sides of the stem, sometimes deciduous by late season. Peduncles terminal and from the upper axils, simple or commonly branched, to 10 cm long, glabrous or puberulent; heads 1.5 to 3 cm broad in flower, flowers many, subtended by filiform-clavate bractlets which are pilose above. Calyx ca. 1 mm long, with 4 to 5 shallow, rounded teeth, densely pubescent internally, usually glandular; corolla white or tinged with pink, tube 5 to 9(13) mm long, with 4 or 5 ovate or oval lobes, glabrous externally, sparingly pubescent within, each sinus of the limb with a small black gland; stamens included. Fruit schizocarpic, obconic, 4 to 8 mm long, splitting into indehiscent nutlets, each with 1 brown seed with a white aril. Wet woods, swamps, pond margins, and along streams. Nearly throughout TX; N. B. and Que., W. to MN and KS, S. to FL and TX; scattered CA, AZ, and NM; also Mex. Jun.-Sept. [Includes var. pubescens Raf. and var. californicus Benth.].

The seeds are eaten by waterfowl, especially ducks. Deer sometimes eat the leaves and shoots. Dense stands provide good cover for wildlife (Elias 1980).



RUBIACEAE SHERARDIA2. SHERARDIA L. Field Madder, Sherard

An Old World genus with 1 or 2 species; we have the 1 species introduced in TX.


1. S. arvensis L. Spurwort, Herb Sherard, Field-Madder. Annual from a slender, branched, reddish root; stems usually several to many, decumbent or prostrate, simple or branched, to 40 cm long, quadrangular, glabrous except the angles scabridulous-pilose. Stipules foliose so leaves apparently 4 to 6 per node, sessile, ovate-lanceolate to obovate or elliptic, acute to cuspidate, to ca. 18 mm long, ca. 5 m broad, usually longer on upper portions, glabrous except for the serrate-hispid, hyaline margins and often minute prickles on the midvein below. Flowers about 4 to 8 in terminal heads, subtended and exceeded by an involucre of ca. 8 leafy, lanceolate bracts. Sepals 4 to 6, triangular-lanceolate, persistent in fruit and usually accrescent; corolla pale lilac to nearly white or sometimes pinkish, funnelform, limb with 4 or 5 pointed lobes, ca. 4 to 5 mm broad, tube slender, about 2 or more times longer than the lobes; stamens 4 or 5; style bifid, stigmas capitate. Fruit ca. 4 mm long, capped by the persistent

sepals, splitting into 2 obovoid, appressed-bristly, indehiscent mericarps ca. 2 mm long and 1 mm wide, each with 1 seed. Roadsides, lawns, fields, and thickets. E. 1/3 to 1/2 TX; native to the Medit., Eur. and W. Asia; introduced in TX and much of E. N. Amer.; also the Pacific region of the U.S. and Can. Mar.-May.




RUBIACEAE GALIUM3. GALIUM L. Bedstraw, Cleavers


Annual or perennial herbs, sometimes rhizomatous, roots often reddish. Stems erect, spreading, ascending, or reclining on other plants, slender, 4-angled. Herbage glabrous to pilose-scabrous. Leaves apparently whorled, usually 4 or more per node, sessile or short-petiolate; stipules foliaceous and adding to the apparent number of leaves per node. Flowers usually in axillary or terminal cymes or panicles, or sometimes the inflorescences reduced and the flowers solitary in the axils or 1 to few on undivided branchlets; pedicel usually jointed to the calyx. Calyx teeth more or less obsolete. Corolla rotate, with 3 or 4 short, deltoid or ovoid lobes, lobes valvate in bud. Stamens (3)4, included. Styles 2, stigmas capitate. Ovary 2-celled, each cell with 1 ovule. Fruit dry, splitting into 2 globose, indehiscent, seedlike mericarps.

About 400 species worldwide; 19 in TX; 7 to be expected here (5 confirmed and 2 probable.)

The genus includes several species that are fragrant when dry and which were formerly used as mattress stuffing (hence the common name). Other species have been used medicinally in their native regions or have uses as dye plants (Mabberley 1987).


1. Fruits glabrous and smooth (occasionally tuberculate) .........................................................2

1. Fruits with bristles or hairs, these usually hooked ..................................................................3


2(1) Corolla 4-lobed, 2 to 3 mm broad; main stem leaves usually in 4's (sometimes 5's or 6's) ...

. ...1.G.obtusum

2. Corolla 3-lobed, less than 2 mm broad; main stem leaves usually in 5's or 6's (sometimes only 4's) ...2.G.tinctorium


3(2) Flowers and fruits sessile or with very short pedicels, in the axils or on the axis of the inflorescence ............................................................................................................................4

3. Flowers and fruits with definite pedicels usually more than 1 mm long, at the ends of the inflorescence branches ............................................................................................................5


4(3) Flowers solitary at the nodes of the (usually) unbranched main stem; leaves usually 10 mm long or less ...3.G.virgatum

4. Flowers spaced ca. 1 cm or more apart on a usually-zig-zagging axis often 5 cm or more long; leaves mostly 15 mm long or longer ...4.G.circaezens


5(3) Leaves usually in 4's, generally oval to elliptic ...5.G.pilosum

5. Leaves usually in 6's to 8's, usually oblanceolate to lance-elliptic .........................................6


6(5) Angles of stems and margins of leaves retrorsely hispid; leaves mostly oblanceolate; plants taprooted annuals ...6.G.aparine

6. Angles of stems smooth; leaf margins with ascending hairs; leaves mostly elliptic-

lanceolate; plants perennials from stolons ...7.G.triflorum


1.G. obtusumRUBIACEAE GALIUM obtusum Bigel. Bluntleaf Bedstraw. Perennial from very slender rhizomes; stems (1.5) 2 to 8 dm long or tall, branched from the base or throughout, rather stiff, nodes pubescent, angles smooth or sometimes hispidulous. Leaves of main stems usually in 4's, rarely in 5's or 6's, elliptic-oblong to lanceolate or broadly linear, ca. 4 to 7 times longer than wide, the larger ones 1.5 to 3 cm long, 1 to 8 mm broad, variable, obtuse, slightly spreading, margins and often midrib slightly scabrous or ciliate. Inflorescences cymose, terminal on the stems and branches; individual cymes with 2 to 4 flowers on straight, ascending pedicels; pedicels often divergent in fruit. Corolla white, 2 to 3(4) mm broad, lobes 1 to 1.3 mm long, acute. Fruit ca. 2.5 to 3.5 mm across, halves globose, smooth or somewhat tuberculate, sometimes only 1 carpel developing. Bottomland woods, swamps, and wet shores. E. TX; reported from our area but specimens seen by the author usually proving to be G. tinctorium; N. S. and Que. to MN and SD, S. to FL and TX, also to AZ. Mar.-July. [Includes var. obtususm and var. ramosum Gl.; G. trifidum L. var. latifolium Torr.; G. tinctorium Gray but not G. tinctorium L., below. Kartesz (1998) recognizes 3 subspecies.].


2.G. tinctoriumRUBIACEAE GALIUM tinctorium (L.) Scop. Dye Bedstraw, Stiff Marsh Bedstraw. Perennial from thin, reddish rhizomes and roots; stems slender, 1 to 6 dm tall or long, usually reclining or sprawling to ascending, commonly mat-forming, at least the younger portions with retrorsely scabrous angles. Leaves of main stems in 5's or 6's (rarely 4's), linear-elliptic to oblanceolate, 5 to 12(20) mm long, 1 to 2(3) mm broad, those at a node often unequal, apically rounded to obtuse, glabrous except more or less retrorsely scabrous on the midvein and margins. Inflorescences simple or branched; peduncles to ca. 2 cm long; flowers usually in ultimate groups of 3 (sometimes 1 or 2), on smooth, straight, spreading pedicels 3 to 8 mm long, each group with or without a subtending whorl of tiny bractlets. Corolla less than 2 mm broad, 3-lobed, white. Mature fruit ca. 2 to 3 mm across, the halves smooth, glabrous. Swamps, marshes, bogs, wet roadsides, meadows, ditches, etc. E. 1/4 TX; definitely present here; Newf. to Ont. and NE, S. to SC, KY, MO, and TX. Mar.-Aug. [Includes var. subbiflorum Rydb., var. floridanum Wieg., and subsp. floridanum (Wieg.) Puff; G. obtusum Bigel. var. floridanum (Wieg.) Fern.; G. subbiflorum Rydb.; G. claytonii Michx.; G. trifidum L. subsp. tinctorium (L.) Hara or var. tinctorium (L.) T. & G.; Asperula tinctoria L.; Often listed with the authority as L.].


3.G. virgatumRUBIACEAE GALIUM virgatum Nutt. Southwest Bedstraw. Annual, taproot and lower stem often reddish; stems erect, 1 to several from the base, usually simple or sometimes branched from near the base, 5 to 20(40) cm tall, angles retrorsely hispid or glabrate. Most leaves in 4's, elliptic, oblong, lance-oblong, or narrowly lanceolate, (2)4 to 7(10) mm long, usually shorter than the internodes, acute to obtuse, surfaces hispid and margins ciliate. Flowers solitary, sessile or subsessile in the leaf axils shorter than the leaves. Corolla white, the 4 lobes 0.5 mm long. Mature fruits deflexed, usually uncinate-hispid, 2 to 2.5 mm broad. Prairies, rocky woods, outcrops, glades, and barrens. In TX primarily on the Ed. Plat. and NW., known at least from Grimes and Robertson Cos.; LA and TX, N. to TN, MO, and KS. [G. texanum Scheele, but not G. texense Gray].

Probably more common in our area, but easily overlooked. Occasional plants have smooth fruits and glabrate foliage. These were once referred to var. leiocarpum T. & G., but they are almost always found with the typical form and are not really distinct.


4.G. circaezensRUBIACEAE GALIUM circaezens Michx. Woods Bedstraw, Cross Clover, Wild Licorice. Perennial from a short, branched crown; stems 1 to several, simple or branched from the base, erect or ascending, 2 to 4(6) dm tall, angles glabrate to hairy. Leaves in 4's, ovate-lanceolate, ovate-oblong, or elliptic, broadest near the middle, (1.5)2 to 5 cm long, 1 to 2.5 cm broad, 3- to 5-nerved, obtuse, surface glabrous to pubescent, especially beneath and on veins, lower surface sometimes with yellow translucent dots (but not punctate as in G. pilosum, below), margins ciliate. Inflorescences terminal and from the upper axils, simple or with 1 or 2 spreading forks, branches commonly zig-zagging, often 5 cm or more long; flowers solitary, generally 1 cm or more apart, usually closely subtended by tiny bracts, sessile or very short-pedicelled on the axes of the inflorescence (sometimes the most distal flower appearing at first glance pedicellate, but this "pedicel" actually the terminal segment of the axis). Corolla greenish-purple, lobes ca. 1 mm long, acute to acuminate, usually externally pubescent. Fruits ultimately deflexed, uncinate-hispid, ca. 3 mm broad. Dry or rich woods, primarily in N. Cen. and E. TX; present in our area but not common--many specimens bearing the name are actually G. pilosum. ME to MI and MN, S. to FL and TX. Apr.-Oct.

Larger, more pubescent plants are referred to var. hypomalacum Fern. var. circaezens includes less robust, more glabrous plants [Includes var. glabellum Britt., var. glabrum Britt., and var. typicum Fern.; G. bermudense L. (pro parte); G. rotundifolium L. var. circaezens (Michx.) Kuntze; G. boreale Walt. (but not G. boreale L.); G. brachiatum Muhl. (not G. brachiatum Pursh); G. circaeoides R. & S.].


5.G. pilosumRUBIACEAE GALIUM pilosum Ait. Hairy Bedstraw. Perennial from a short, branching caudex; stems 1 to 7(10) dm tall or long, erect to ascending, with a few branches near the base and otherwise unbranched to the inflorescence, sparsely to densely pilose on the angles below (rarely glabrous). Leaves in 4's, oval to elliptic or oblong, rarely suborbicular, broadest about the middle, 1 to 2.5 cm long, about 1/2 as broad, usually with 3 nerves, apex generally obtuse-apiculate, surfaces pilose, usually at least sparsely glandular-punctate on the underside, margins ciliate-hispid. Inflorescences terminal and from the upper axils, more or less paniculate, 2- or 3-forked, branches divaricate; pedicels obvious, flowers in groups of (1)2 or 3, often each group subtended by tiny bractlets. Corolla greenish white or purplish, lobes 1.2 to 2 mm long. Fruit with hooked bristles, 2.5 to 3.5 mm broad. Dry woods, prairies, thickets, or occasionally in clearings near water but where not very wet (e.g. at road-river intersections). Definitely present here. E. 1/3 TX; NH and S. Ont. to MI, S. to NC, TN, KS, and TX. May-Aug.

If varieties are recognized, our plants are mostly var. puncticulosum (Michx.) T. & G. with stem and leaves with short, upcurved hairs. This form is more common in the southern part of the range and on the Coastal Plain. [G. punctatum Pers., G. puncticulosum Michx.; G. purpureum Walt. (but not G. purpureum L.); G. walteri J. F. Gmel.].

Plants with stem and leaf hairs straight have been called var. pilosum. This form is more common in the northern part of the range, extending to LA and TX, with some introgression with var. puncticulosum. [G. bermudense L. (pro parte); G. puncticulosum Michx. var. pilosum (Ait.) DC.].


6.G. aparineRUBIACEAE GALIUM aparine L. Catchweed Bedstraw, Sticky Bedstraw, Goose-grass, Cleavers, (Velcro-Plant). Taprooted annual; stems weak, usually reclining or straggling over other foliage, often tangled, 1 dm to 1(2) m long, retrorsely hispid on the angles and pubescent at the nodes. Leaves usually in 6's or 8's, linear-oblanceolate, 1 to 8 cm long, shorter below and longer above, 1-nerved, cuspidate or rounded and apiculate, retrorsely hispid-scabrous on the margins and midvein beneath, often also rough appressed-pubescent on the surfaces. Peduncles axillary, usually longer than the leaves, usually with 3 to 5 flowers, or sometimes the flowers in (1 to)3's on short branches. Corolla white, 1 to 2 mm long. Fruits uncinate-bristly, (1.5)3 to 4(5) mm across. Lawns, roadsides, prairies, woods, thickets, seashores, and waste places; our most common species. E., Cen., and S. TX; Newf. to AK, S. to FL, TX, and CA; either native or introduced over much of the N. temperate region of N. Amer. and Eur. Mar.-May. [Includes var. echinospermum (Wallr.) Farw., var. intermedium (Merr.) Briq., var. minor Hook, and var. valliantii (DC.) Koch; G. valliantii DC.; G. spurium L. and varieties.].

Occasional plants of dry places have much smaller leaves (1 to 3(4) cm long) and fruit. These were referred to var. echinospermum or var. valliantii, but the differences seem to be due entirely to habitat (GPFA 1986).

Roasted, dried, mature fruits make a very good coffee substitute (GPFA 1986; Tull 1987). Tull (1987) gives a recipe for a tan dye made from all of the plant except the roots. Young leaves and stems can be eaten as a potherb or cooked like greens (Tull 1987), though the prickly hairs make them unpleasant to eat raw.


7.G. triflorumRUBIACEAE GALIUM triflorum Michx. Sweet-scented Bedstraw, Fragrant Bedstraw. Perennial from thin rhizomes; stems weak, scrambling on other vegetation or prostrate, simple or branched, 2 to 8(12) dm long, retrorsely scabrous or pubescent on the lower angles, varying to more or less glabrous. Leaves of main axes usually in 6's (sometimes 4's or 5's), narrowly elliptic to oblanceolate, cuspidate, 2 to 6 cm long, the upper ones slightly reduced, margins antrorsely ciliate or scabrous, midvein scabrous below. Inflorescences axillary, sometimes terminal; peduncles elongate, each 3-flowered or sometimes branched and with many flowers, flowers all pedicellate, 2 to 4 mm across. Corolla white or greenish white, the 4 lobes 1 to 1.5 mm long. Fruit uncinate-bristly, ca. 1.5 to 2.2 mm broad. Woods and thickets. E. and N. Cen. TX; Newf. to AK, S.to GA, MS, LA, TX, and CA; also Greenland and Mex.; more or less circumpolar; reported from the Blackland Prairies and probably present, but no specimens seen by the author from our area. Some sheets labeled G. triflorum proved to be misidentified. May-Sept.

The dried foliage is sweet-scented.



RUBIACEAE MITCHELLA4. MITCHELLA L. Partridge-berry


2 species, 1 of the SE. U.S. and the other from Japan.


1. M. repens L. Partridge-berry, Two-eyed Berry, Running Box. Evergreen herb; stems prostrate, creeping and rooting at the nodes, forming mats, pubescent to glabrate; herbage glabrous to minutely and sparsely pubescent. Leaves leathery, opposite, short-petiolate, blades ovate to cordate-ovate or suborbicular, 8 to 25 mm long, basally truncate to shallowly cordate, apically obtuse to rounded, glossy above, sometimes with whitish veins, secondary veins arched toward the apex and not reaching the margins, margins entire but sometimes uneven; stipules triangular-subulate, ca. 2 mm long, inconspicuous. Peduncles shorter than the subtending leaves, terminal on the shoots and branches, each bearing a pair of fragrant flowers united by their ovaries. Calyx 3 to 4 mm long, with minute lobes; corolla white, funnelform, usually with 4 lobes (occasionally with 3, 5, or 6), 10 to 14 mm long, the tube much longer than the ovate-oblong, spreading lobes, lobes densely pubescent within; flowers heterostylous--all flowers on a given plant with either short stamens and a long style or long stamens and a short style; style 1, with 4 linear stigma lobes. Fruits drupes, remaining in pairs or only 1 maturing, 4 to 7(10) mm in diameter, aromatic, bright red (or occasionally white in forma leucocarpa Bissel), topped with the minute calyx lobes, each drupe with 4 small, bony stones. Rich woods, moist or dry knolls, and stream banks. E. TX; N. S. to Ont. and MN, S. to FL and TX. May-July.

The fruits are edible, but are said to be rather bland (GPFA 1986).



RUBIACEAE MITRACARPUS5. MITRACARPUS Schult. & Schult.


Annual (as ours) or perennial herbs or small shrubs. Stems and branches usually quadrangular in cross-section; stipules sheath-like, with marginal bristles. Flowers in dense terminal and axillary heads. Calyx teeth 4(5), the lateral ones subulate and longer than the corolla, the other 2 smaller. Corolla white, salverform or funnelform, with 4 spreading, valvate lobes. Stamens 4. Ovary usually bilocular, with 1 ovule per cell; style branches slender. Capsules usually paired, fragile, circumscissile. Seed oblong or globose.

30 species of tropical America, with 1 introduced to the Old World; 2 in TX; 1 here.


1. M. hirtus (L.) DC. Taprooted annual; stems to ca. 4 dm tall, branches 4-angled; herbage hirtellous (or minutely so) throughout. Leaves sessile or with short petioles, oblong to elliptic, 2 to 6 cm long, acute to obtuse, tapered to the base; stipules 2 to 3 mm long, with marginal bristles about as long. Heads axillary and terminal, many-flowered, 6 to 12 mm broad. Lateral sepals ca. 2 mm long, linear-lanceolate and acuminate; corolla 2.5 to 3 mm long, tube longer than the lobes. Capsule shorter than the calyx, circumscissily dehiscent about the middle. Very rare in grasslands of the Coastal Prairies, inland usually to Bexar and Wilson Cos., known in our area at least from Brazos Co. (TAMU 007808, from 1946--perhaps no longer present); also W.I. and tropical America. May-July. [Mitracarpus villosus (Sw.) DC.; Mitracarpum hirtum (L.) DC.].



RUBIACEAE HEDYOTIS6. HEDYOTIS L. (s.s.) Bluets, Star Violet


Material of the continental U.S. annual or perennial herbs. Stems erect to spreading, ours well-branched. Leaves opposite, entire. Flowers in terminal cymes, 4-merous. Corolla salverform to funnelform, frequently internally pubescent. Stamens 4. Ovary bilocular, ovules several per cell. Capsule partially superior, protruding above the calyx. Seeds concavo-convex, with the hilum on a ridge on the ventral face.

Various numbers of species, depending upon interpretation; 28 listed for N. Amer. including Hawaii (Kartesz 1998); 5 in TX; 1 here.

For much of their recent history, our plants have been treated in Hedyotis (s.l.), as by Correll and Johnston (1970) and Hatch, et al. (1990). Recent work, such as that of Terrell (1996), supports the division of Hedyotis (s.l.) into segregate genera. The characters separating the segregate genera (pollen and seed morphology, chromosome number, etc.) are largely unobservable in the field or in the average classroom, and there are no correlating vegetative characters. It is only by chance that our Hedyotis (s.s.) can be separated from the other genera on the basis of inflorescence type.


1. H. nigricans (Lam.) Fosb. var. nigricans Prairie Bluets. Perennial from a stout, woody taproot or branched caudex; stems few to many from the base, sometimes clump-forming, 5 to 60 cm tall, usually branched, glabrous to minutely scabrous on the angles or occasionally short-pubescent below. Leaves opposite, essentially sessile or with petioles 1 to 3(5) mm long, linear-filiform to lanceolate (occasionally elliptic), 1 to 3(4) cm long, 0.5 to 4 mm broad, margins often revolute, margins and nerves of lower side usually minutely scabrous. Flowers in terminal leafy, pedunculate or sessile cymes, heterostylous. Hypanthium somewhat turbinate, 0.7 to 1.2 mm long; calyx lobes lanceolate to slenderly triangular, 1(2) mm long, sparingly ciliate; corolla white, pink, or pale purple, funnelform, tube (2)3 to 4(6) mm long, lobes about equalling or shorter than the tube, (1)2 to 4(5) mm long, pubescent internally. Capsule obovoid-turbinate or less often globose, 1/2 to 1/4 protruding above the calyx, 2.5 to 3 mm long; seeds 1 to 1.3 mm long, black, the ridge on the concave face inconspicuous, surface minutely papillate. Variable in vegetative characters and widespread throughout the state; in our area of roadsides, prairies, hillsides, rock outcrops, and even marshy areas. This variety FL to AZ, N. to GA, OH, MI, and NE; also Mex. Apr.-Nov. [Includes var. filifolia (Chapm.) Shinners, var. rigidiuscula (Gray) Shinners, and var. scabra (S. Wats.) Fosb.; Houstonia nigricans (Lam.) Fern.; etc.].


RUBIACEAE OLDENLANDIA7. OLDENLANDIA L. Bluets, Mille-graines


Annual or perennial herbs (as ours) or small shrubs. Stems erect to spreading, simple or branched. Leaves opposite, sessile or short-petiolate, entire; stipules short, fused to the leaf bases or petioles. Flowers solitary or few in the axils and/or in dense terminal clusters, sessile or very shortly pedicellate, homostylous. Calyx tube globose to ovoid. Corolla in ours rotate, white, with 4 lobes valvate in bud. Stamens 4. Ovary bilocular, ovules in ours many per cell; style 1, stigmas 2, short-linear. Capsule essentially fully inferior, loculicidally dehiscent across the apex. Seeds many, angular, without a ventral concavity.

Possibly 100 species worldwide, but most abundant in the Old World; 3 in TX; 2 here.

Our plants have, for much of their recent history, been treated in Hedyotis (s.l.), as by Correll and Johnston (1970) and Hatch, et al. (1990). Recent work, such as that of Terrell (1996), supports the division of Hedyotis (s.l.) into segregate genera. The characters separating the segregate genera (pollen and seed morphology, chromosome number, etc.) are largely unobservable in the field or in the average classroom, and there are no correlating vegetative characters. It is only by chance that our Oldenlandia can be separated from the other genera on the basis of inflorescence type.


1. Stems glabrous; leaves mostly linear; plants perennial ...1.O.boscii

1. Stems usually hirsute; leaves mostly ovate-elliptic; plants annual ...2.O.uniflora


1.O. bosciiRUBIACEAE OLDENLANDIA boscii (DC.) Chapm. Plants perennial; stems spreading or prostrate, usually branched and often clump-forming, 0.5 to 3 dm long, glabrous or occasionally hirtellous. Leaves sessile or nearly so, mostly linear or slenderly linear-elliptic, 10 to 25 mm long, 1 to 3 mm broad; stipules membranous, with a few long cilia. Flowers solitary or in small clusters in the axils. Calyx 2 to 3 mm long, with lanceolate, glabrous or roughened lobes about equalling the tube; corolla rotate, white or the lobes sometimes pink-tipped, ca. 1 mm long, shorter than the calyx lobes. Capsule to 2.5 mm broad, glabrous or short-ciliate. Pond margins, riverbanks, floodplains, and savannahs. E. TX, rare in S. Cen. TX; FL to TX, N. to SE. VA, TN, and SE. MO. May-Aug. [Hedyotis boscii DC.]


2.O. unifloraRUBIACEAE OLDENLANDIA uniflora L. Annual or summer annual; stems erect to spreading, 1 to 6 dm long, branched, occasionally glabrate but usually white-hirtellous, villous, or pilose, especially on the angles. Leaves essentially sessile or with petioles 1 to 2.5 mm long, ovate to ovate-elliptic or -lanceolate, the larger ones 1 to 2.5 cm long, 4 to 8 mm broad, acute to obtuse, midvein and margins usually white-hirsutulous; stipules membranous, long-triangular, with long marginal bristles, or sometimes divided.  Flowers solitary or in clusters in the axils, sometimes also in terminal, bracted clusters, subsessile. Calyx 2.5 to 4 mm long, pilose, lobes broadly lanceolate to lance-ovate, oblong, or ovate-oblong, more or less pilose, about as long as the calyx tube; corolla rotate, white, ca. 1 mm long, shorter than the calyx lobes. Capsule to 2 mm broad, usually white hirtellous; seeds ca. 0.2 mm long or wide, black. Wet streambanks and edges of bogs and ponds. E. TX; FL to TX; N. to S. NY and SE MO. Jun.-Sept.(Oct.) [Hedyotis uniflora (L.) Lam. and var. fasciculata (Bertol.) W. H. Lewis; H. fasciculata Bertol; O. fasciculata (Bertol.) Small].



RUBIACEAE HOUSTONIA8. HOUSTONIA L. Bluets


Ours smallish annual (elsewhere also perennial) herbs with fibrous roots (some other taxa rhizomatous). Stems erect, prostrate, or creeping. Leaves opposite, 1-nerved. Flowers terminal and solitary or in some taxa axillary, on pedicels 2 to 50(70) mm long, hetero- or homostylous, sometimes cleistogamous. Calyx 4-parted. Corolla salverform or slightly shorter, 4-lobed. Capsule (1/4)1/2(2/3) inferior. Seeds saucer-shaped or else nearly globose, ventral face concave or with a circular opening leading to a subglobose cavity, hilum often on a ridge in the depression or cavity, seed coat reticulate.

These plants have long been treated in Hedyotis (s.l.), as by Correll and Johnston (1970) and Hatch, et al. (1990). Recent work, such as that of Terrell (1996), supports the division of Hedyotis (s.l.) into segregate genera. The characters separating the segregate genera (pollen and seed morphology, chromosome number, etc.) are largely unobservable in the field or in the average classroom, and there are no correlating vegetative characters. It is only by chance that our Houstonia can be separated from the other genera on the basis of inflorescence type.

20 species of the U.S., Can., and Mex.; 11 in TX; 4 here.

A few spaces are cultivated for ornament, notably H. caerulea (Mabberley 1987).


1. Flowers usually 1 per node in the axils; middle and upper leaves linear ...1.H.subviscosa

1. Flowers terminal on the branches; middle and upper leaves proportionately broader .........2


2(1) Corolla white, 2 to 5.5 mm long, the tube about as long as the corolla lobes .........................

. ...2.H.micrantha

2. Corolla usually purple, blue-violet, lilac, or pink (occasionally white), 3.5 to 12 mm long, the tube mostly longer than the calyx lobes ............................................................................3


3(2) Corolla purple, blue-violet, lilac, or rarely pink or white, the tube and lobes each ca. 3 to 4 mm long; leaves primarily elliptic or narrowly elliptic ...3.H.pusilla

3. Corolla usually rose or pink (rarely white or pale violet), the tube and lobes each ca. 6 mm long; leaves mostly oblanceolate ...4.H.rosea

1.H. subviscosaRUBIACEAE HOUSTONIA subviscosa (C. Wright ex A. Gray) A. Gray Nodding Bluets. Taprooted winter or spring annual; stems 3 to 20(30) cm tall or long, erect to spreading or decumbent, well-branched, sparsely to densely pubescent or glabrate, sometimes with glandular areas. Leaves sessile and linear or the lower ones oblanceolate and tapered to a short petiole, cauline leaves 5 to 25 mm long, 0.2-1.5(3) mm broad, glabrous to hirtellous, obtuse to acute, margins revolute; stipules to 2 mm long and 3 mm broad, slightly sheathing, truncate to rounded, sometimes with a few teeth and/or glands. Flowers homostylous, solitary in the axils on pedicels 2 to 7 mm long, pedicels usually up-curved or recurved in fruit. Hypanthium densely pubescent; calyx lobes 0.4 to 2 mm long, 0.2 to 1.2 mm broad, triangular to narrowly lanceolate, acute to acuminate, pubescent; corolla salverform or short-funnelform, white to light pink, 1.5 to 3 mm long, tube 1 to 7 mm long, lobes 0.5 to 1.5 mm long, 0.5 to 0.8 mm long, minutely puberulent to glabrous internally; anthers included or exserted. Capsules (1/2)3/5 to 4/5 inferior, 1.8 to 3(3.5) mm long, 2.5 to 4(5) mm broad, wider than long, densely white-hirtellous, the apex retuse or rounded; seeds 0.7 to 1.1 mm long, 0.45 to 1.1 mm broad, thin and saucer-shaped, the ventral face with a ridge along the hilum. Sandy soil of roadsides, pastures, creek margins, and dunes. Cen. and S. TX and the Gulf Coast; known from Burleson Co.; probably also NE. Mex. Mar.-May. [Hedyotis subviscosa (Gray) Shinners; Oldenlandia subviscosa Wright ex A. Gray].


2.H. micranthaRUBIACEAE HOUSTONIA micrantha (Shinners) Terrell Southern Bluets. Fibrous-rooted annual or winter annual; stems 1-11 cm tall, erect to spreading, simple or branched, delicate. Basal leaves few or none, lowermost and midstem leaves tapered to a petiole equaling or longer than the blade, upper leaves short-petiolate to sessile, blade and petiole together 2 to 20 mm long, 0.5 to 5.5 mm broad, elliptic to narrowly elliptic or ovate, or the lower spatulate, apex rounded, obtuse, or acute, glabrous or minutely ciliate; stipules to 2.5 mm long and 3 mm broad, triangular-lanceolate, rounded-truncate, entire or toothed, with or without a few glands. Flowers homostylous, solitary and terminal on pedicels 2 to 25 mm long; buds erect or nodding. Hypanthium glabrous; calyx lobes 1.2 to 3 mm long, 0.5 to 1.5 mm wide, ovate to lanceolate, acute to obtuse; corolla white with a yellow eye, 2 to 5.5 mm long, short-salverform, tube 0.8 to 2.5 mm long, shorter than to little longer than the calyx lobes, lobes 0.8 to 3 mm long, ca. 0.5 to 2 mm wide, ovate, about equalling the corolla tube, glabrous; anthers included. Capsules 2 to 3.5 mm long, 3 to 5(5.7) mm broad, usually wider than long, subglabrous or slightly flattened, about 1/2 inferior, apically retuse; seeds brown or black, 0.5 to 1 mm broad, more or less subglobose, with a ventral opening, reticulate. Lawns, roadsides, pastures, and waste areas. E. and SE. TX; GA to TN, AR, and TX. Feb.-Apr. [Hedyotis australis H. Lewis and Moore; Hedyotis crassifolia Raf. var. micrantha Shinners].

Frequently found with H. rosea and H. pusilla, below.


3.H. pusillaRUBIACEAE HOUSTONIA pusilla Schoepf Tiny Bluets, Low Bluets, Star-violet, Small Bluets. Fibrous-rooted annual; stems 1.5 to 15(25) cm tall, erect or spreading, branched or simple, delicate, glabrous to puberulent or scabridulous. Basal laves few or none, lower and midstem leaves with a petiole shorter than to longer than the blades, the upper short-petiolate to sessile, blades and petioles together to 2.5 cm long, blades elliptic to narrowly elliptic, ovate, or spatulate, acute to obtuse, glabrous to puberulent, sometimes ciliate; stipules triangular-lanceolate, acute to rounded or caudate, 0.5 to 2 mm long, to 2 mm broad, entire or toothed, with or without glands. Flowers slightly fragrant, homostylous, terminal on erect pedicels 2 to 30(50) mm long; buds nodding or reflexed. Hypanthium glabrous to ciliolate; calyx lobes 0.5 to 5 mm long, 0.2 to 2 mm broad, lanceolate to ovate, obtuse to acute; corolla 3.5 to 10(12.5) mm long, salverform, usually purple, violet, blue-violet or lilac (occasionally pink or white), the base of each lobe commonly with small reddish or brownish blotches, the eye often brownish, yellow, or orange-ish, tube 2 to 5.5 mm long, usually longer than the calyx lobes, slightly widened around the anthers, lightly pubescent to glabrous within, lobes 1 to 5(7) mm long, 1 to 4 mm broad, ovate, glabrous; anthers included. Capsules 1.5 to 3.5(4) mm long, 2.5 to 5(6) mm broad, usually broader than long, subglobose or slightly flattened, about 1/2 inferior, apically retuse, glabrous; seeds 0.5 to 1 mm long, subglobose with a ventral opening, brown, reticulate. Lawns, roadsides, pastures, and fields. E. 1/3 TX; MD (introduced) to FL, W. to KY, IL, IA, KS, OK, and TX. Jan.-Apr. [Includes forma albiflora Standley, forma rosea Steyerm., and forma albiflora Wunderlin & Hopkins; Hedyotis crassifolia Raf.; Houstonia minima L. C. Beck; Hedyotis minima (L.C. Beck) Torrey & Gray; Houstonia caerulea L. var. minor (Michx.) T. & G.; etc.],


4.H. roseaRUBIACEAE HOUSTONIA rosea (Raf.) Terrell Rose Bluets. Fibrous rooted annual, usually very small in stature; stems 1 to 4(7) cm tall, delicate, simple or branched, glabrous to scabrous-puberulent. Basal leaves, if present, like the stem leaves but larger, stem leaves with petioles usually shorter than the blades, petioles and blades together 3 to 20 mm long, 0.3 to 3 mm broad, generally oblanceolate or narrowly spatulate, narrowly elliptic, or linear, glabrous to puberulent, margin sometimes ciliolate; stipules definitely sheathing, to 2 mm long and broad, rounded to truncate, with or without glands. Flowers slightly fragrant, homostylous, solitary and terminal on pedicels 6 to 12 mm long, relatively large compared to the size of the plant. Hypanthium glabrous; calyx lobes 1 to 2.3(3) mm long, 0.4 to 1(1.5) mm broad, shorter than the corolla tube, lanceolate, usually obtuse; corolla light pink, rose, white, or occasionally light lavender, usually with a yellow eye, 5 to 12 mm long, more or less salverform, tube 3.5 to 8 mm long, expanded around the anthers, throat densely pubescent within, lobes from equalling to 1/3 the length of the tube, (1.5)2 to 4(6) mm long, 1.2 to 3.3 mm broad, ovate; anthers included. Capsules 1.5 to 3.5 mm long, (2)2.5 to 4(5) mm broad, a little wider than long, subglobose or slightly flattened, 1/4 to 1/2 protruding from the calyx, apex emarginate; seeds (0.4)0.6 to 1 mm long, 0.35 to 0.7 mm broad, brown or black, more or less subglobose, with a ventral opening, reticulate-areolate. Lawns, roadsides, pastures, fallow fields, etc. Infrequent in E. 1/3 TX; AL to TX, N. to OK, AR, and extreme S. MO. Feb.-Apr. [Hedyotis rosea Raf.; Houstonia pygmaea Muller & Muller; Hedyotis taylorae Fosb. in Shinners; Hedyotis pygmaea R. & S.].



RUBIACEAE RICHARDIA9. RICHARDIA L. Mexican Clover


Tap- or fibrous-rooted annual or perennial herbs, generally prostrate to ascending and well-branched. Leaves subsessile to short-petiolate; stipules sheath-like, edged with bristles. Flowers in pedunculate terminal clusters of several to many, subtended by 2 to 4 bract-like leaves. Flowers (3)5 to 6(8)-merous. Calyx teeth lanceolate, united basally, circumscissily dehiscent and falling as a unit at or prior to the separation of the carpels. Corolla funnelform. Stamens (2)4 or 6, usually exserted. Style branched or entire, stigmas 2 to 4, linear to spatulate; carpels (2)3 to 4(6). Fruit at maturity splitting longitudinally into indehiscent or slightly open mericarps, each 1-seeded, leaving behind an inconspicuous central axis.

12 species, primarily of the S. U.S., S. to S. Amer.; a few adventive elsewhere; 4 species in TX; 2 here.

Reference: Lewis and Oliver (1974).


1. Leaves ovate to elliptic-lanceolate; calyx usually 6-parted; stems usually more than 15 cm long ...1.R.scabra

1. Leaves linear-lanceolate; calyx lobes (3)4(6); stems usually less than 15 cm long ...............

. ...2.R.tricocca

subsp. tricocca


1.R. scabraRUBIACEAE RICHARDIA scabra L. Rough Mexican-clover. Taprooted annual; stems to 8 dm long, decumbent or sprawling, pilose to hispid. Leaves with petioles to ca. 5 mm long; blades ovate to elliptic-lanceolate (rarely linear-lanceolate), 1 to 6.5 cm long, 3 to 13 mm broad, obtuse to acute, basally attenuate, scabrous on both surfaces, especially on the major veins below; stipules ca. 2 mm long, with bristles ca. 2 to 5 mm long. Flowers usually 20 or more per cluster, subtended by 2 or 4 bracteal leaves, these often without stipules, the second pair smaller. Calyx 2.5 to 3.5 mm long, the 6 lobes ca. twice as long as the tube, lanceolate, glabrous, margins ciliate; corolla funnelform to salverform, white or tinged with pink-lavender, tube 2 to 8 mm long, lobes 0.5 to 2.5 mm long; stamens 6, exserted or included; carpels usually 3. Fruit splitting into 3(or 6) mericarps, 2 to 3.5 mm long, ca. 1.5 to 2 mm broad, oblong to obovoid, the outer faces papillose and usually strigillose, the inner face closed and with a narrow groove, base somewhat indented. Roadsides, fields and waste places, commonly in sandy soil. FL to TX, N. to VA and AR, S. to Peru and Bolivia; also W.I.; present and presumed adventive in Africa and in the U.S. (Indiana). June-Oct.

Reportedly eaten by cattle, while the root has emetic properties (Correll and Johnston 1970; Mabberley 1987).


2.R. tricoccaRUBIACEAE RICHARDIA tricocca (T. & G.) Standl. subsp. tricocca Prairie Buttonweed. Perennial from a branched taproot; stems several, hispid, branched from the base, prostrate or spreading, the plant forming mats or clumps to 40 cm broad. Leaves coriaceous or stiff, sessile, linear-lanceolate, 4 to 25 mm long, 1 to 5 mm broad, acuminate to acute, sparsely hispid (especially on the veins below) to nearly glabrous, margins revolute in dried material; stipules sheath-like, 1 to 2.5 mm long, marginal bristles whitish or pale, to 6 mm long. Flower clusters terminal (very rarely subterminal or appearing axillary because of surpassing development of lateral branches), flowers ca. 3 to 10 per head, subtended by 4 bracteal leaves similar to the stem leaves. Calyx 2.5 to 3 mm long, puberulent to hirsute, the (3)4(6) lobes lanceolate, equal or nearly so; corolla white, funnelform, (3)4 to 6 mm long, with 3 or 4 lobes 1 to 2 mm long; ovary with (2)3(4) carpels, stigmas with 2 to 4 obscure lobes. Fruit splitting into (2)3(4) mericarps, 2(3)mm long, ca. 1 mm wide, usually hispid, rarely glabrous, sometimes reddish-purple apically. Sandy or sandy-clay soils in brushlands or open woods, also roadsides and fields. This subspecies from LA to TX, S. to Mex., inland to the Post Oak Savannah and Monterrey, Mex. (A second subspecies inhabits the interior of Mexico). Flowering Apr.-Jun.; collected with fruit into fall. [Diodia tricocca T. & G.; Crusea tricocca (T. & G.) Heller].



RUBIACEAE SPERMACOCE10. SPERMACOCE L. Buttonweed


Annual or perennial herbs, usually low in stature. Leaves opposite; stipules represented by a ciliate membrane uniting the leaf bases or petioles. Flowers in dense axillary or terminal clusters. Calyx tube short obovate to obpyramidal, lobes usually 4, triangular to lanceolate. Corolla whitish to blue-purple, salverform or funnelform, 4-lobed, the lobes valvate in bud. Stamens 4, epipetalous on the corolla tube. Ovary bilocular, ovules 1/locule; style 1, slender, stigma lobes 2. Fruit a sessile schizocarpic capsule, sometimes paired, usually leathery or hard, splitting at maturity into 2 carpels, 1 taking the septum and so closed, the other open on the inner face.

About 100 species of warm parts of the Americas; 4 in TX; 1 here.


1. S. glabra Michx. Smooth Buttonweed. Perennial from a branched, buried or surface-lying caudex; stems 1 to several, simple or branched, 1 to 6 dm long or tall, erect or usually spreading or procumbent, obscurely angled, glabrous. Leaves short petioled, blades elliptic to lanceolate, oblanceolate, or oblong-lanceolate, 2 to 8 cm long, acuminate to acute, tapered to the base, glabrous; stipular sheathing membrane with several filiform bristles 2 to 5 mm long. Flowers in clusters of 10 to 40 in the upper axils. Calyx tube turbinate, 2 to 2.5 mm long, lobes 1 to 3 mm long, lance-oblong or triangular-lanceolate, spreading; corolla white, 2 to 3 mm long, slightly longer than the calyx, densely white-hairy in the throat; filaments and styles short. Fruit 2 to 4 mm long, more or less turbinate or obpyramidal, topped by the persistent calyx lobes, eventually splitting into 2 parts, 1 with the septum. Lake shores, river banks, low woods and clearings. E. 1/3 TX; FL to TN, N. to OH, IN, IL, MO, and KS. May-Oct.; collections with fruit from Nov.


RUBIACEAE DIODIA11. DIODIA L. Buttonweed

Annual or perennial herbs. Stems prostrate to decumbent or ascending. Leaves opposite, generally sessile, entire, but commonly with bristly-scabrous margins; stipules sheath-like, with marginal bristles. Flowers sessile, in groups of 1 to several in the axils. Calyx teeth usually 2 to 4, equal or unequal. Corolla funnelform or salverform, with 3 or 4 lobes. Stamens usually 4, epipetalous on the corolla tube, usually exserted. Style filiform, simple or bifid, usually exserted; ovary bicarpellate. Fruit leathery or hard, topped by the persistent calyx lobes, maturing into 2, 1-seeded, indehiscent carpels, splitting apart or remaining together.

About 30 species of tropical and warm America and Africa; 2 in TX, both found here.


1. Fruit leathery, 6-ribbed, glabrous to villous, topped with 2(3) calyx teeth; carpels usually remaining together; corolla salverform, tube 7 to 10 mm long; style bifid ...1.D.virginiana

1. Fruit hard, unribbed, usually hispid, topped with 4 calyx teeth; carpels easily separated; corolla funnelform, tube 2 to 6 mm long; style simple ...2.D.teres


1.D. virginianaRUBIACEAE DIODIA virginiana L. Larger Buttonweed, Virginia Buttonweed. Perennial from a woody, branched root; stems spreading, procumbent, or erect, sometimes rooted at the lower nodes, 1 to 6 dm or more long, branched, glabrous to hirsute-villous. Leaves sessile but usually tapered to the base, elliptic-oblong to elliptic-oblanceolate or linear-lanceolate, acute to acuminate or obtuse, 2 to 7(9) cm long, 4 to 12(20) mm broad, glabrous to minutely scabrous, especially below, margins minutely bristly-scabrous; stipule bristles stiff and slightly flattened, stipules sometimes sparsely hirsute. Calyx teeth 2(3), lanceolate or linear-lanceolate, 4 to 6 mm long, more or less pubescent; corolla white, salverform, tube 7 to 10 mm long, the limb abruptly expanded into lobes 3 to 4 mm long, pubescent externally; style bifid. Fruit ellipsoid or oblong-ellipsoid, 5 to 9 mm long, to 5 mm broad, glabrous to villous, more or less hard, topped with the persistent calyx lobes, mericarps each with 3 prominent dorsal ribs, the carpels only rarely separating. Sandy soils of lake shores, savannahs, prairies, meadows, and stream banks. E. 1/3 TX; FL to TX, N. to NJ, IL, and MO. May-Oct. [D. hirsuta Pursh; D. tetragona Walt.].

If varieties are recognized, our plants are var. virginiana.



2.D. teresRUBIACEAE DIODIA teres Walt. Rough Buttonweed, Poor-joe. Taprooted annual herb (but the taproot sometimes woody); stems erect, branches prostrate to ascending, 2 to 8 dm long, glabrous to puberulent or hirsute on the angles. Leaves stiff, sessile, linear to lanceolate, 2 to 5 cm long, to 1 cm broad, acute to acuminate, base rounded to slightly clasping, minutely scabrous on both surfaces and margins ciliate-scabrous; stipular bristles abundant, filiform, commonly reddish-brown, often as long as the flowers and longer than the fruits. Flowers in axillary groups of 1 to 3, and so 2 to 6 per node. Calyx teeth lanceolate to ovate, sometimes unequal, 1.8 to 2.2 mm long, ciliate; corolla funnelform, whitish or tinged with pink-purple, 4 to 6 m long, the limb not widely flaring; style undivided. Fruit obovate-turbinate, unribbed, usually hispid or hispidulous, 4 to 5 mm long, topped by the persistent sepals, readily separating into 2 nutlets, often one of them taking 3 calyx teeth and the other only 1. Sandy soils of woods or open areas, pastures, wood edges, etc. E. 2/3 TX, rare on the Ed. Plat.; FL to TX, N. to VT, NY, PA, IL, IA, and KS. May-Nov. [Diodella teres (Walt.) Small].

Our plants are probably all var. teres. [Includes var. setifera Fern. & Grisc., with a slender bristle at the tip of each young leaf.].





CAPRIFOLIACEAECAPRIFOLIACEAE

Honeysuckle Family


Ours woody vines or shrubs, occasionally small trees (elsewhere also herbs). Leaves opposite, simple, usually estipulate or the stipules early deciduous. Flowers in various arrangements, perfect, regular or zygomorphic. Calyx 3- to 5-lobed, sometimes quite small. Corolla tubular to funnelform, rotate, or campanulate, sometimes rather bilabiate. Stamens (4)5, alternate with the corolla lobes, inserted on the corolla tube. Ovary with 2 to 5 carpels, ovules pendulous 1 or several per cell, sometimes only 1 or 2 cells maturing; style 1, obsolete to elongate, stigma capitate or 2- to 3-(5-) lobed. Fruit a drupe or berry (as in ours) or a capsule.

16 genera and about 365 species, primarily of N. temperate and tropical montane regions; 5 genera and 22 species listed for TX by Hatch, et al. (1990); 4 genera and 7 species here.

A traditional family treatment is presented here. Some recent phylogenetic studies support the removal of Sambucus and Viburnum to the Adoxaceae and/or merging the Caprifoliaceae with the Valerianaceae and Dipsacaceae. For a discussion of this, see Zomlefer (1994).

The family includes many ornamentals, especially in Lonicera, Viburnum, and Wiegela.


1. Leaves pinnately or bipinnately compound .......................................................1. Sambucus

1. Leaves simple ...........................................................................................................................2


2(1) Plants trees or large shrubs to 5 m or more; fruit a 1-seeded drupe; flowers in compound cymes ....................................................................................................................2. Viburnum

2. Plants small shrubs, climbing shrubs, or vines; fruit 2- to 4-celled and 2- to many-seeded; flowers solitary or in terminal and/or axillary clusters or whorls .............................................3


3(2) Plants vines and viney shrubs; corolla funnelform to tubular, more than 1 cm long; fruit red or black, with several seeds ............................................................................3. Lonicera

3. Plants small shrubs; corolla campanulate, less than 1 cm long; fruit mostly reddish-pink to purple, 2-seeded ......................................................................................4. Symphoricarpos



CAPRIFOLIACEAE SAMBUCUS1. SAMBUCUS L. Elder-berry


Shrubs or coarse herbs, rarely arborescent, often spreading by suckers. Stems pithy; bark with prominent lenticels. Herbage sometimes rank-smelling when crushed. Leaves odd-pinnately compound or rarely bipinnate, leaflets in TX material serrate, acuminate; stipules present or absent. Flowers in terminal compound cymes, regular, small, 5-merous. Calyx tiny or obsolete. Corolla rotate, the 5 lobes spreading, longer than the tube. Stamens 5. Style nearly rudimentary, stigma lobes 3 to 5; ovary 3- to 5-celled, with 1 ovule per cell. Fruit a berrylike drupe with 3(4 to 5) cartilaginous stones, often edible.

9 species of temperate and tropical regions; 2 in TX; 1 here. This treatment follows the work of Bolli (1994). Sambucus is placed in the Adoxaceae by Thorne (see Zomlefer 1994) or removed by some to its own family, the Sambucaceae.

Some species have edible fruit or are dyeplants (Mabberley 1987).


1. S. nigra L. subsp. canadensis (L.) R. Bolli Common Elder-berry, European Black Elder. Shrub, but the stems only slightly woody, to 4 m tall; branches glaucous and with thick white pith. Leaflets 5 to 11 or the lowermost pair sometimes divided again, ovate- or obovate-elliptic to lanceolate, 4 to 18 cm long, 2 to 5.5(7) cm broad, apically caudate to acuminate, base rounded to cuneate, sometimes oblique, margin serrate, upper surface glabrous or puberulent on the midvein, lower surface pubescent to glabrate; petioles 3 to 9 cm long, with a groove on the upper side; petiolules 2 to 7 mm long; stipules present but early deciduous. Cymes with 3 to 5 rays from the base, much-divided, to 35 cm broad in flower, flat-topped or concave, branches and pedicels glabrous or occasionally hispidulous; flowers fragrant. Corolla creamy-white, 4 to 5 mm broad, lobes elliptic. Fruit purple to purple-black, 4 to 6 mm broad, glabrous; stones yellowish, 2.5 to 3 mm long, 1.5 to 2 mm broad, obovoid, roughened. Usually in damp or wet soil of low areas, especially streambanks and swamp margins, occasionally in drier open woods. In TX, primarily in the E. and Cen. regions, rare in the Panhandle; N. S. and Que., W. to Man. and SD., S. to FL, TX, Mex., and W.I. Flowering May-July. [S. canadensis L. and vars. canadensis, laciniata Gray, and submollis Rehd.; S. cerulea Raf. var. mexicana (K. Presl. ex DC.) L. Benson; S. mexicana K. Presl. ex DC.; S. orbiculata Greene; S. simpsonii Rehd. ex Sarg.].

The fruits are edible, and can be used to make jelly or wine. Cooked or dry fruit can be added to muffins, pies, pancakes, etc., but raw fruits are reported to be unpalatable (Tull 1987). The flowers are also edible, used in pancakes, fritters, and wine (Medve and Medve 1990). The berries, leaves, and twigs can be used to produce dyes of edible colors (Tull 1987). Elder-flower water is used as a skin lotion (Medve and Medve 1990). Lampe (1985) notes that the whole plant is mildly poisonous, capable of causing diarrhea and vomiting, though the cooked fruits and flowers are deemed safe.




CAPRIFOLIACEAE VIBURNUM2. VIBURNUM L. Viburnum, Arrow-wood


Deciduous or evergreen shrubs or small trees. Leaves opposite (rarely whorled), simple, petiolate, estipulate (as ours) or the stipules adnate to the petioles or represented by glands; leaf buds naked or with paired scales. Inflorescences terminal, compound, umbel-like or panicle-like cymes; bracts and bractlets small and promptly deciduous. Flowers regular and perfect or the peripheral flowers in a cluster sometimes radiate, slightly irregular, and neuter (not usually so in ours). Calyx with 5 shallow lobes. Corolla rotate to open-campanulate, white to cream or pinkish, 5-lobed, the lobes imbricate in bud. Stamens 5, inserted on the base of the corolla, included or exserted, anthers introrse. Style none, stigmas 1 to 3 on the apex of the ovary, locules 3, 2 of them abortive and the 1 remaining with 1 pendulous ovule. Fruit a drupe, soft-fleshed, with 1 plump or flattened pyrene.

About 150 species of temperate and warm regions, especially in Asia and N. Am.; 8 species in TX (according to Hatch, et al. (1990); 2 of these combined by Kartesz 1998); 3 species here. This genus is removed to the Adoxaceae in Thorne's treatment (see Zomlefer 1994).

A number of species are cultivated for ornament, especially those with showy, sterile flowers. Some have edible fruit or useful wood (Mabberley, 1987).



1. Leaves entire or occasionally obscurely crenate ...1.V.nudum

1. Leaves serrulate .......................................................................................................................2


2(1) Margins serrulate; blades coriaceous............... ...2.V.rufidulum

2. Margins coarsely serrate, dentate, or crenate; blades sometimes membranaceous.............

.......................................................................................................................... 3. V. dentatum


1.V. nudumCAPRIFOLIACEAE VIBURNUM nudum L. Possumhaw Viburnum, Swamp-haw. Shrub or small tree to 5(6) m tall; bark gray to brown, more or less smooth; buds brown or gray-brown; twigs, petioles, and parts of lower leaf surfaces at least somewhat rusty-pubescent or with tiny rusty scales. Petioles to 1 cm long; blades coriaceous, elliptic to oblanceolate or obovate (rarely ovate), 5 to 15 cm long, 2.5 to 7.5 cm wide, rounded to abruptly acute (occasionally truncate and abruptly acute), base cuneate to rounded, margin entire to vaguely crenate, revolute, upper surface shiny and glabrous, lower surface paler. Inflorescences to 10(11) cm wide, on peduncles to 1.5 cm or more long; pedicels ca. 5 to 15 mm long. Calyx minute; corolla creamy white, ca. 5 mm broad, lobes rounded; filaments long-exserted, ca. 4 to 5 mm long. Drupe subglobose to plumply ellipsoid, pointed at both ends, blue or blue-black, glaucous, 7 to 10 mm in diameter, flesh bitter; stone globose-obovoid, smooth. Sandy swampland, bog margins, swamp forests, and wet pine woods. E. TX; FL to TX, N. to KY and AR. Flowering Mar.-June.; fruiting collections from fall (Oct.).

This species is closely related to plants treated by Correll and Johnston (1970) as V. nitidum Ait. Kartesz (1998) combines the two species under V. nudum. If the species are merged, our plants become var. nudum [V. nudum L. var. angustifolium T. & G., var. grandifolium Gray, var. ovale Wood, and var. serotinum Ravenal ex Chapm.; V. cassinoides L. var. angustifolium (T. & G.) Shinners]. If combined, the V. nitidum plants then become V. nudum L. var. cassinoides (L.) T. & G.


2.V. rufidulumCAPRIFOLIACEAE VIBURNUM rufidulum Raf. Southern Black-haw, Rusty Black-haw, Downy Viburnum, Rusty Nannyberry. Shrub or small tree to 10 m tall; trunk to 10 cm in diameter, usually less; bark blackish, breaking up into squares; winter buds, young branchlets, petioles, and parts of lower leaf surfaces sparsely to densely pubescent with rusty, scurfy hairs. Petioles slightly winged, to 2 cm long; blades elliptic to obovate or oblanceolate, 3.5 to 6.5(9) cm long, 2.5 to 4(5.5) cm broad, leathery, glossy above, obtuse to rounded (sometimes acutish), basally cuneate, margin serrulate. Inflorescences with 3 or 4 primary rays, branched several times, to 10 cm wide, branches more or less rusty-scurfy. Calyx with minute, rounded lobes; corolla creamy white, 4 to 8(10) mm broad, lobes rounded; stamens long-exserted, filaments 3 to 5 mm long. Drupe ellipsoid, dark blue or blue-black, glaucous, 1 to 1.5 cm long, pointed at both ends, flesh sweet; stone flattened. Upland woods, thickets, stream banks, fencerows, and along wood edges. E. and Cen. TX; FL to TX, N. to VA, OH, IN, IL, MO, and KS. Flowering Feb.-May, fruit well-developed by June -July. [Includes var. margarettiae Ashe; V. rufotomentosum Small; V. prunifolium L. var. ferrugineum T. & G.].

The fruit is edible raw or cooked in sauces or jelly (Correll and Johnston 1970; Tull 1987). The bark has been used in antispasmodic remedies, though its efficacy is doubtful (Vines 1960).


3.V. dentatumCAPRIFOLIACEAE VIBURNUM dentatum L. Southern Arrow-wood. Shrub or small tree to ca. 5 m tall; bark gray to brownish or reddish; branchlets very straight, young twigs commonly densely stellate-pubescent, becoming glabrous. Petioles to 15 mm long, stellate-tomentose to glabrous, slender; blades generally membranous but sometimes thicker, especially in plants of sunny, dry areas, ovate to ovate-lanceolate or nearly orbicular, to a maximum of 15 cm long and 10 cm broad, usually much smaller, apically acute to short-acuminate or rounded, basally rounded or subcordate, margin coarsely serrate to dentate or crenate, the teeth more or less triangular, upper surface glabrous or subglabrous, lower surface glabarous or with some stellate hairs, lateral veins straight. Inflorescence with 5 to 7 rays, 5 to 7 cm borad, on terminal or axillary peduncles 3 to 6 cm long. Hypanthium glabrous to glandular or slightly bristly; corolla 5 to 8 mm long, roatate; style short to nearly lacking, pubescent; stigmas 3. Drupe blue-black, ovoid to subglobose, 5 to 10 mm long, the flesh soft; stone plumply ellipsoid, deeply grooved on the ventral face. Woods and thickets. MA to FL, W. to AR, LA, and E. TX; in our area known at present only from far E. Grimes Co. Flowering Apr.-June; fruit maturing late summer-fall. [V. pubescens (Ait.) Pursh].

This species is variable in leaf shape and pubescence; 5 varieties are listed by Kartesz (1998) for North America; 2 are found in TX. More collections are needed to determine whether one or the other (or both) is present here.


var. dentatum has hypanthium glabrous to glandular and proportionately narrow leaves.


var. scabrellumT. & G. has corolla and hypanthium setose to hirsute and leaves that are proportionately broader. [Includes var. ashei (Bush) McAtee; V. scabrellum T. & G. ex Chapm. and var. dilutum McAtee.; V. ashii Bush].



CAPRIFOLIACEAE LONICERA3. LONICERA L. Honeysuckle


Woody vines or shrubs. Leaves opposite, sometimes leathery, entire or rarely with teeth or lobes, short petiolate or sometimes leaves connate-perfoliate. Flowers often fragrant, paired in the axils or in 3-flowered cymules or on axillary or terminal rachises. Calyx very small, 5-lobed. Corolla funnelform to tubular or slightly campanulate (not in ours), from nearly regular to strongly zygomorphic and bilabiate, with 4 lobes in the upper lip and 1 in the lower, tube sometimes gibbous or spurred at the base. Stamens 5, more or less equal, exserted, attached about the midpoint of the corolla tube. Style slender, stigma capitate. Ovary 2- to 3-carpellate, ovules pendulous, several per cell. Fruit a several-seeded berry with axile placentation.

180 species, mostly in the N. hemisphere; 4 in TX; 2 here.

Many species are cultivated for ornament and fragrance, but some, such as L. japonica, can become aggressive weeds (Mabberley 1987).


1. Flowers red; uppermost 1 or 2 pairs of leaves of each branch connate; berries red ..............

. ...1.L.sempervirens

1. Flowers white to yellow; leaves all distinct; berries ...2.L.japonica


1.L. sempervirensCAPRIFOLIACEAE LONICERA sempervirens L. Trumpet Honeysuckle, Coral Honeysuckle, Evergreen Honeysuckle. Viny shrub; stems twining or climbing, sometimes rooting at the nodes, to several m long; bark of older branches reddish-gray, shredding, young branches glabrous or essentially so. Leaves ovate to elliptic, obovate, or linear-elliptic, 3 to 7 cm long, 2 to 4 cm broad, acute to obtuse, base rounded to cuneate, uppermost 1 to few pairs one each branch connate-perfoliate, all more or less evergreen, leathery and glabrous on both surfaces or the lower surface minutely puberulent, dark green or olive above, paler and commonly glaucous below. Flowers 4 to 6(20 or more) in sessile whorls forming an interrupted spike; calyces subtended by small, blunt bracts. Calyx small, with 5 blunt, ovate-triangular teeth less than 1 mm long; corolla deep red, orange-red, or sometimes yellow, commonly tinged with yellow within, slenderly tubular-funnelform, 3 to 5.5 cm long, tube sometimes gibbous on the bottom near the base, usually expanded about the middle where stamens are attached, limb slightly zygomorphic, lobes rounded-ovate, the lowermost larger than the others and the upper 2 slightly overlapping at the base; stamens and style slightly exserted or included, anthers versatile. Berries red or red-orange, 6 to 10 mm in diameter, glabrous; seeds flattened-ovoid, yellow to brown, 4 to 5 mm long, 3 to 3.5 mm wide. Woods, thickets, roadsides, vacant lots, railroad embankments, etc. E. TX; FL to TX, N. to NY, OH, IA, and NE; escaped from cultivation elsewhere. Mar.-June, sometimes sparingly into fall. [Includes var. sempervirens, var. minor Ait., and var. hirsutula Rehd.; Phenianthus sempervirens (L.) Raf.].

Commonly cultivated for the showy flowers, which are attractive to hummingbirds. Several varieties exist in the nursery trade. This plant is not as aggressive as Japanese Honeysuckle (below), according to Odenwald and Turner (1990). The fruit is eaten by birds (Vines 1960), but should be considered inedible by humans (Tull 1987).


2.L. japonicaCAPRIFOLIACEAE LONICERA japonica Thunb. Japanese Honeysuckle. Viny shrubs; stems trailing, twining, or climbing, to several m long, sometimes rooting at the nodes; older stems red-brown, bark sometimes shredding, branchlets and young stems pubescent with tawny hairs. Leaves semi-evergreen, coriaceous, ovate to oblong or elliptic, 3 to 8 cm long, 1.5 to 3(4) cm broad, obtuse to acute and apiculate, base rounded to truncate or cuneate, margin ciliate, usually entire but on new growth sometimes toothed or lobed, upper surface pubescent at least on the midvein, lower surface pubescent to glabrate; petioles 2 to 8(11) mm long, pubescent, terminal leaves not connate; bracts below flower clusters similar to stem leaves but much smaller, usually more or less ovate. Flowers intensely fragrant, in pairs on short axillary peduncles, sometimes clustered at the branch tips where internodes are short; bractlets very tiny, roundish. Calyx lobes ca. 1 mm long, triangular to lanceolate, acuminate, glandular and pubescent; corolla bilabiate-funnelform, 2 to 4.5(5) cm long, white to cream, yellow, or pink-tinged, commonly darkening with age, tube slender, limb about 1/2 the length of the flower, 4 linear-oblong lobes in the upper lip and 1 below, usually recurved, corolla minutely and sparingly pubescent inside and out, the outside usually also glandular; stamens long-exserted, anthers versatile; style slender, elongate, stigma minutely 2-lobed. Fruits black, 5 to 8 mm in diameter; seeds ovoid, dark brown, 2 to 3.3 mm long and 2 mm broad, longitudinally ribbed and rugose. Roadsides, vacant lots, pastures, railroad embankments, wood edges and openings in woods. E. and Cen. TX; native of Asia, escaped from cultivation and naturalized from FL to TX, N. to NY, OH, and KS. Mar.-July and sporadically until frost. [Includes var. chinensis (P. W. Wats.) Baker; Nintooa japonica (Thunb.) Sweet].

Various cultivars exist in the nursery trade, some of them with purple-tinted foliage (Bailey, et al. 1976). This plant is a very aggressive weed in our area, out-competing native plants. Nonetheless, it is frequently planted for its fragrant flowers and it does make a good ground cover or screen. The nectar is sweet, honey-like, and edible (though some people are violently allergic to it). The fruits of U.S. plants are edible, though there are (dubious) reports of poisonings in Europe (Lampe 1985). Yellow and gray dyes can be made from the leaves, while the stems are useful in weaving baskets and wreaths (Tull 1987).



CAPRIFOLIACEAE SYMPHORICARPOS4. SYMPHORICARPOS Boehm. Snow-berry, Coral-berry, Wolf-berry


Shrubs, usually low and well-branched; bark of older stems exfoliating in strips or shreds. Leaves opposite, simple, usually entire (as ours) or sometimes toothed or lobed, short-petiolate, estipulate. Flowers in terminal or axillary clusters or reduced cymes or solitary in the axils of the upper leaves; tiny bracts commonly present. Calyx with (4)5 small teeth. Corolla pink to whitish or greenish, (4-)5-lobed, campanulate to funnelform or salverform, sometimes slightly gibbous at the base, often pubescent internally, with 1 to 5 nectaries at the base. Stamens (4)5. Style elongate, stigma capitate or vaguely 2-lobed; ovary 4-locular, 2 of the cells with several ovules each, but abortive, the 2 fertile cells each with 1 pendulous ovule. Fruit a globose to ovoid or ellipsoid berry-like drupe, white or red, with 2 more or less flattened stones.

17 species, 16 in N. Am. and 1 in China; 6 species in TX; 1 here.

Some species have ornamental uses (Mabberley 1987).


1. S. orbiculatus Moench Coral-berry, Indian-currant, Buckbrush. Low shrub to 1(1.5) m, often only half that, rhizomatous and sometimes colonial; branches erect or ascending, light brown to purplish, older branches with gray, shredding bark; twigs sparingly to densely pubescent or villous, becoming glabrate. Petioles 1 to 4 mm long, pubescent; blades elliptic to ovate or suborbicular, (1.5)2 to 4(5) cm long, 1 to 3.5 cm broad, obtuse to acute, base rounded to somewhat cuneate, glabrous to sparingly and minutely pilose above, lower surface often paler, somewhat glaucous, variously pubescent, margin entire or occasionally with a few irregular, blunt teeth. Flowers in short, dense, axillary spikes on current season's growth; bracts minute and pubescent. Calyx teeth 5, triangular, ciliate, less than 1 mm long, persistent; corolla broadly campanulate, slightly upturned and somewhat gibbous below, 2.5 to 3(4) mm long, 2 to 3 mm broad, pinkish or greenish, villous internally, glabrous externally, lobes spreading, rounded, about as long as the tube; stamens and style included, filaments shorter than the anthers, anthers ca. 1 mm long; style ca. 2 mm long, pilose. Fruit pink-purple to red or coral, ellipsoid to subglobose, 4 to 7 mm long, with a beak 1 mm long, glaucous; stones ovoid to ellipsoid, with 1 flat side, 2.5 to 3.5 mm long and 2 mm broad, ends blunt, smooth. Upland and low-lying woods, especially along streams, also pastures and ravines. E. 1/3 TX; NY to FL, W. to TN, NM, CO and SD. Flowering Apr.-July; fruit coloring in fall and commonly remaining over the winter. [S. vulgaris Michx.; S. spicatus Engelm.; S. symphoricarpos (L.) MacM.].

Sometimes cultivated for ornament, erosion control, or as a wildlife food. Many species of birds eat the fruit. Yellow-variegated and white-fruited forms are known (Vines 1960).





VALERIANACEAEVALERIANACEAE

Valerian Family


Annual (as ours), biennial, or perennial herbs. Leaves in a basal rosette and/or opposite, simple, estipulate, sometimes with sheathing bases. Flowers small, regular or irregular, perfect or unisexual, in cymes, thyrses, or heads, usually with bracts and bractlets. Calyx minute and/or developing tardily as the fruit matures, sometimes ring-like or toothed, sometimes inrolled and forming a pappus-like structure on the fruit. Corolla funnelform to rotate, salverform, or tubular, tube often saccate, gibbous, or spurred, the (3)4 or 5 lobes imbricate in bud. Stamens 1 to 4, epipetalous, alternate with the corolla lobes. Ovary inferior, 3-locular, with 1 fertile and 2 sterile cells or the 2 sterile cells rudimentary; style 1, stigma unbranched or with 2 or 3 branches or lobes. Fruit achene-like, with 1 seed in the 1 fertile cell.

17 genera and about 400 species worldwide, especially in the N. temperate zone and the Andes; 2 genera and 7 species in TX; 1 genus with 4 species in our area (one sometimes included within one of the others.) This family is placed by Thorne in a broadly-defined Caprifoliaceae (see Zomlefer 1994).

Members of the family often have an odor described as that of "wet dog". Some species have medicinal or perfume uses, principally species of Nardostachys (spikenard) and Valerian. Other members are edible or have ornamental uses (Mabberley 1987).



VALERIANACEAE VALERIANELLA1. VALERIANELLA P. Mill. Corn Salad, Lamb's Lettuce


Annual or biennial herbs, often smelling of "wet dog" when dry. Stems erect, dichotomously branched. Leaves slightly succulent, basal ones in a rosette, petiolate to subsessile, entire, cauline leaves sessile, sometimes connate basally, usually with a few teeth. Inflorescence a terminal headlike or dichotomously branched cyme, subtended by lanceolate to oblong, entire to glandular-fimbriate-serrulate bracts. Flowers very small, regular to slightly irregular, usually perfect (sometimes unisexual). Calyx more or less obsolete or represented by 5 minute teeth. Corolla white, bluish-white, or pale pink, funnelform or salverform, slightly saccate at the base, our species with a gibbosity at the base of the throat on the lower side, lobes 5, equal or unequal. Stamens (2)3, attached near the apex of the corolla tube, exserted, anthers 4-lobed. Style 3-lobed, exserted; ovary and fruit 3-celled, only 1 cell fertile. Fruit glabrous to variously pubescent, 2-celled or more commonly 3-celled, with only 1 fertile, a groove separating the 2 sterile cells, some species (but not ours) with the calyx inflated or forming a parachute or pappus.

About 50 species of N. temperate regions to N. Afr.; 5 in TX; 4 here.

Some species, especially V. locusta of Europe, are edible as potherbs (Mabberley 1987). Our plants are all edible as potherbs or in salads, but they taste better before they flower.

Species in our area are very similar in gross morphology and can vary greatly in stature. Mature fruit are needed for identification. The following treatment follows Eggers (1969), GPFA (1986), and Hatch, et al. (1990) in combining several very similar species and varieties which differed chiefly in fruit and corolla size.





1. Fertile cell of fruit only 2/3 to 1/2 the combined width of the 2 sterile cells when viewed from above; in dorsal view the fertile cell not nearly concealing the 2 sterile cells; fruit subglobose in outline ...1.V.woodsiana

1. Fertile cell of fruit slightly narrower to much wider than the combined width of the 2 sterile cells when viewed from above; in dorsal view the fertile cell nearly or fully concealing the 2 sterile cells; fruit mostly ovoid to elliptic in outline ................................................................2


2(1) Fruits white uncinate-pubescent; the 2 sterile cells ca. 1/2 as long as the fertile cell and projecting laterally ...2.V.amarella

2. Fruits pubescent with short hairs or glabrous; the 2 sterile cells about as long as the fertile cell .............................................................................................................................................3


3(2) Fruits ellipsoid-oblong, pubescent, the sterile cells parallel and the groove between them narrow and shallow; fertile cell with a prominent dorsal midnerve; herbage glabrous except perhaps for tufts of hair at the base of the leaves and some marginal cilia ................

. ...3.V.florifera

3. Fruits ovoid or ovoid-elliptic, glabrous or pubescent, the sterile cells usually divergent from one another and the groove between them wide or at least deep; fertile cell without a single prominent midnerve; herbage usually at least sparsely pubescent, often only on the lower stem and leaf margins, in addition to tufts near the leaf bases ...4.V.radiata


1.V. woodsianaVALERIANACEAE VALERIANELLA woodsiana (T. & G.) Walp. Woods Corn Salad. Annual; stem 1.5 to 5 dm tall, pubescent along the angles. Leaves pubescent on the margins and/or the midribs below, lower leaves entire and spatulate, slightly connate basally, upper cauline leaves oblong-ovate, not connate, with a few teeth basally. Bracts elliptic-lanceolate, acute to acuminate, glabrous but usually glandular-fimbriate-serrulate apically. Inflorescence open to rather dense. Corolla white, 1.5 mm long, funnelform, the tube much shorter than the limb; stamens and style exserted; stigma lobes short. Fruit yellowish, ca. 2 mm long, subglobose, glabrous to puberulent; fertile cell commonly with a short beak, slenderly oblong to oblong-lanceolate, dorsally weakly 1-nerved and flattened, only 1/2 to 2/3 as wide as the combined width of the 2 inflated, widely-divergent sterile cells, groove separating sterile cells wide. Moist clay or sandy soils of woods, prairies, and low areas. E. TX; uncommon in our area; also E. OK. Mar.-May. Included by Kartesz (1998) in V. radiata (L.) Dufr.


2.V. amarellaVALERIANACEAE VALERIANELLA amarella (Lindh. ex Engelm.) Krok Hairy Corn Salad. Annual; stem 1.5 to 3 dm tall, glabrous. Leaves glabrous except for small tufts of uncinate hairs on either side of the base at the node; lower leaves ovate- or obovate-spatulate, entire, upper leaves oblong-obovate, sessile. Inflorescence dense, bracts glabrous, ovate to lanceolate to elliptic-lanceolate, rounded to acute, usually with hyaline margins. Corolla white, 1.5 to 3 mm long, funnelform, the limb usually equalling the tube and throat together; stamens and style usually exserted, sometimes either one short or abortive; stigma lobes short. Fruit 1.5 to 2 mm long, brown, subglobose-ovoid, hirsute with whitish, uncinate hairs; fertile cell large and equalling or wider than the combined width of the 2 sterile cells; sterile cells about 1/2 as long as the fertile cell, projecting laterally, sometimes each with a shallow depression and the whole reminiscent of a "pig nose", groove between the 2 sterile cells very shallow or obscure. On calcareous or rocky soils of open woods, hills, prairies, low areas, and barrens. Primarily in Cen. TX; to be expected in the W. portion of our area; also OK and possibly E. KS. Mar.-May.

Unfortunately, almost all Valerianella keyed in Reeves (1977) came out to this species, resulting in a large number of misidentified specimens over the years.


3.V. floriferaVALERIANACEAE VALERIANELLA florifera Shinners Annual; stems 9 to 18 cm tall, glabrous. Leaves glabrous or the lower scabrous-ciliate, especially apically, entire, lower leaves oblong to oblong-spatulate, sessile, upper leaves oblong to oblong-lanceolate or slenderly deltoid-oblong, sessile. Bracts lanceolate to elliptic-lanceolate, acute, glandular-toothed or lacerate, glabrous. Inflorescence usually rather open, the clusters few but many-flowered. Corolla white, funnelform, the tube 1.4 to 1.5 mm long, the throat 1.3 to 1.5 mm long, lobes of limb 1.8 to 2 mm long; stamens and style exserted. Fruit ca. 1.7 mm long, 0.6 mm broad, narrowly elliptic in outline, about twice as long as wide, sparsely pubescent with straight, spreading hairs (apparently occasionally glabrous?); fertile cell about as wide as or commonly wider than the combined width of the sterile cells, slightly flattened dorsally and with a distinct mid-line (at least apically); sterile cells low, rounded, not divergent but parallel to one another. Sandy moist soil of post oak woods. Endemic to Cen. and S. Cen. TX. Mar.-May.


4.V. radiataVALERIANACEAE VALERIANELLA radiata (L.) Dufr. Beaked Corn Salad, Narrow-cell Corn Salad. Annual; stem 1 to 6 dm tall, usually at least sparsely pubescent along the angles. Leaves usually pubescent on the margins and midrib below, lower leaves spatulate or oblong-spatulate, entire and connate, upper leaves oblong-ovate to ovate, not connate, commonly with coarse teeth basally. Bracts lanceolate, the outer sparsely ciliate or sometimes weakly glandular-fimbriate-serrulate apically, inner bracts usually glabrous. Inflorescence loose. Corolla white, funnelform, 1.5 to 2(3) mm long, tube shorter than the limb; stamens and style exserted; stigma lobes short. Fruit usually yellowish, ovoid to ovoid-ellipsoid, glabrous to finely pubescent, fertile cell slightly narrower to wider than the combined width of the sterile cells, rounded or slightly flattened dorsally, only rarely with a weak median nerve; sterile cells usually divergent from one another and the groove between them narrow to wide, usually rather deep, a shallow groove present on each side between the sterile and fertile cells. Low, moist areas, vacant lots, roadsides, prairies, pastures, old fields, etc. E., SE., and Cen. TX; apparently our most common species; CT and PA, S. to FL, W. to MO, KS, AR, and TX. Mar.-May. [Includes var. radiata, var. fernaldii Dyal, and var. missouriensis Dyal; V. stenocarpa (Engelm. ex Gray) Krok and var. parviflora Dyal].

Eggers (1969) combined V. radiata (with ovoid fruits) with the very similar V. stenocarpa (with more elliptic fruits). This treatment certainly makes determinations easier. Kartesz (1998) maintains V. stenocarpa as a separate species, but does place its var. parviflora under V. radiata.





ASTERACEAE (COMPOSITAE)ASTERACEAE

Sunflower Family


Our plants mostly herbs--a few are vines or shrubs. Leaves alternate or opposite; stipules absent. Flowers 1 to many in heads or capitula on a common receptacle, each head subtended by involucral bracts or phyllaries; sometimes each flower also subtended in the inflorescence by a small bract or scale called a pale (plural: pales or chaff). Flowers variously perfect, staminate, pistillate and fertile, pistillate but sterile, or altogether neuter. Calyx reduced to a pappus of hairs, bristles, awns, etc. or absent. Corolla of 5 fused petals, regular or irregular, sometimes absent. Stamens 5, usually fused by their anthers around the style (if present). Gynoecium of 2 fused carpels, inferior, style branches two. Fruit an achene, sometimes called a cypsella.

For our purposes, flowers can be divided into 3 groups: A) DISK FLOWERS: Perfect flowers with regular, usually 5-toothed corollas; inflorescences having only disk flowers are called discoid. B) RAY FLOWERS: Flowers often pistillate and sterile, but sometimes fertile or altogether sterile; corolla zygomorphic with the limb all drawn to one side, forming a strap-shaped ligule which is usually 3-toothed; inflorescences having disk flowers in the center and ray flowers on the periphery are called radiate. C) LIGULATE FLOWERS: Perfect, fertile flowers with irregular corollas; as with ray flowers, the limb is a strap-shaped ligule, but usually 5-toothed; inflorescences having only ligulate flowers are called ligulate.

This is the largest dicot family, with 1,100 genera and roughly 20,000 species worldwide; 91 genera and 196 species can be found locally.

Composite taxonomy is difficult, and the family has been further broken down into about 13 tribes. (For the local flora, however, an artificial key is more practical than one based on relationships.) In recent years, there has been a great deal of progress in resolving intergeneric relationships, with the result that many traditional genera, e.g., Heterotheca, Aster, and Haplopappus, have been divided into smaller segregate genera. The following treatment attempts to present the most recent, widely-accepted generic concepts, with synonymies that include both older (and perhaps more-familiar) names and newer names that are not yet widely accepted.

Taxonomy in the Asteraceae seems to be especially fraught with subspecies and varieties, often with no two treatments of a particular group recognizing the same complement of species and subspecific taxa. The treatments presented here represent the consensus--if such exists--or the most recent treatments for which I have been able to find detailed literature support.

Proper identification usually requires careful dissection of the inflorescence and flowers under magnification (a black microscope stage is useful), with close attention to the structure of the phyllaries, pappus, and achene.

The family is of surprisingly little economic importance except for a few crops such as Sunflower and Jerusalem Artichoke (Helianthus). Some are ornamentals (Helianthus, Liatris, Coreopsis, Cosmos, etc.). Some have medicinal properties (Mabberley 1987).


1. Heads with only disc or with both disc and ray flowers; sap not milky ....................................2

1. Heads with only ligulate flowers; sap usually milky or colored ............................................96


2(1) Heads with only disk flowers, or at least all corollas regular or none of them with manifest ligules. (Includes Soliva, whose "ray" flowers have no corollas, Centaurea, whose outer disk florets are enlarged, and Artemisia, whose ligules are minute .......................................3

2. Heads with both ray and disk florets, though ligules not always large .................................36


3(2) Perfect flowers absent; plants monoecious or dioecious .......................................................4

3. Perfect flowers present, these fertile or infertile; imperfect flowers sometimes also present ..................................................................................................................................................8


4(3) Plants dioecious; stamens of staminate flowers united by their anthers ...............................5

4. Plants monoecious; stamens of staminate flowers distinct ....................................................6


5(4) Plants herbs; heads and leaves woolly .............................................................1. Antennaria

5. Plants shrubs; heads and leaves not woolly .......................................................2. Baccharis


6(4) Staminate and pistillate flowers in the same heads; phyllaries separate or united, but never completely enveloping the achenes ......................................................................3. Iva

6. Staminate and pistillate flowers in different heads; phyllaries of the pistillate involucre united and completely enveloping the achenes ......................................................................7


7(6) Bracts of staminate involucre separate; pistillate involucre covered with conspicuous, hooked spines .....................................................................................................4. Xanthium

7. Bracts of staminate involucre united; pistillate involucre smooth or with only a few tubercles or straight teeth .....................................................................................5. Ambrosia


8(3) Plants herbaceous perennials or subshrubs, aromatic, densely gray pubescent; corollas of peripheral of flowers ("ray" flowers") tubular, with a v-shaped notch on one side, well- developed ligules absent; pappus absent; receptacle naked ............................6. Artemisia

8. Plants without the above combination of characters ..............................................................9


9(8) Peripheral disk flowers expanded and somewhat ray-like .............................20. Centaurea

9. Peripheral disk flowers usually not expanded or ray-like .....................................................10


10(9) Plants densely white or gray woolly-tomentose or floccose, at least on the under-surfaces of the leaves; leaves entire ....................................................................................................11

10. Plants glabrous or pubescent, but not woolly-tomentose or floccose; leaf margins various; OR if leaves somewhat white- or gray-pubescent underneath, then leaves pinnately lobed or toothed ...............................................................................................................................14

11(10) Leaves decurrent on the stem, forming wings; white-tomentose pubescence primarily

confined to the undersurfaces of the leaves and decurrent wings ................7. Pterocaulon

11. Leaves not decurrent on the stem; white woolly pubescence covering more than just the undersurfaces of the leaves ..................................................................................................12


12(11) Pappus absent; receptacle chaffy; individual florets mostly obscured by pubescence in the inflorescence ..........................................................................................................8. Evax

12. Pappus present; receptacle naked or with no conspicuous chaff; individual florets

discernible upon dissection of the inflorescence ..................................................................13


13(12) Leaf apices retuse or truncate, often with a terminal bristle ....................................9. Facelis

13. Leaf apices acute, acuminate, or rounded; terminal bristle absent ......10. Gnaphalium s. l.


14(10) Heads with perfect flowers that are fertile or sterile and fertile pistillate flowers. (Examine both central and peripheral flowers for functional stamens and developing achenes.) .....15

14. Heads with only perfect, fertile flowers ..................................................................................20


15(14) Leaves pinnatifid or pinnately compound ..............................................................................16

15. Leaves entire or merely toothed ............................................................................................18


16(15) Plants more than 15 cm tall, branching above the base ...............................11. Parthenium

16. Plants less than 15 cm tall, acaulescent or branching at the base ......................................17


17(16) Achenes sessile; styles persisting on achenes as small spines .............................12. Soliva

17. Achenes stipitate on the receptacle; styles not spiny .............................................13. Cotula


18(15) Phyllaries usually dotted with small globules of resin; plants with a strong disagreeable odor ........................................................................................................................14. Pluchea

18. Phyllaries without resin globules; plants usually without a disagreeable odor ....................19

19(18) Phyllaries of 2 kinds: 1 row of outer, short, calyx-like bracts and 1 or 2 rows of inner, linear bracts; heads 1 cm or more long ....................................................................15. Erechtites

19. Phyllaries all alike; heads generally less than 1 cm long ....................................16. Conyza

20(14) Pappus mainly or entirely of capillary bristles, slender bristles, or slender awns, these bristles smooth, roughened, or plumose .............................................................................21

20. Pappus composed of stiff awns, coarse bristles, scales, or teeth; OR pappus absent ......31


21(20) Pappus bristles of two different types or lengths, usually an outer row of short, flat bristles and an inner row of longer bristles. (See Figure A in the illustration for couplet 20, above).. ............................................................................................................................17. Vernonia

21. Pappus bristles uniform or of random lengths; not of just two different types or lengths ...22


22(21) Heads few-flowered, arranged in congested clusters; each cluster in turn subtended by a whorl of involucre-like leaves .....................................................................18. Elephantopus

22. Heads few- or many-flowered; not arranged in congested, involucrate clusters ................23


23(22) Receptacle chaffy, bristly, or hairy ........................................................................................24

23. Receptacle naked ..................................................................................................................26


24(23) Leaves white-mottled and spiny ...........................................................................19. Silybum

24. Leaves not white-mottled, sometimes spiny .........................................................................25


25(24) Pappus bristles plumose, at least below ..............................................................21.Cirsium

25. Pappus bristles not plumose but sometimes barbellate--with the side hairs only as long as the width of the hair axis .......................................................................................22. Carduus


26(23) Pappus bristles plumose or strongly barbed. (See Figure B of drawing at couplet 20) .....27

26. Pappus bristles occasionally roughened, but not plumose or barbed .................................28


27(26) Corolla rose or purple; heads in cymes, racemes, or spikes .................................23. Liatris

27. Corolla whitish; heads solitary or in cymes or panicle-like inflorescences .......24. Brickellia


28(26) Stems twining .........................................................................................................25. Mikania

28. Stems not twining ...................................................................................................................29


29(28) Flowers bright yellow .........................................................................................26. Bigelowia

29. Flowers pale yellow or not yellow at all .................................................................................30


30(29) Leaves opposite; flowers white, bluish, purplish, or rosy .........................27. Eupatorium s. l.

30. Leaves alternate; flowers whitish or pale yellow ........................................28. Arnoglossum


31(20) Pappus composed mainly or wholly of 2 or 3 rigid awns above the angles of the achene, OR pappus absent. (See Figures C,D, and G at couplet 20) ..............................................32

31. Pappus composed of scales or teeth. (Figures E and F at couplet 20) ..............................33


32(31) Leaves finely dissected, the divisions filiform or nearly so .........................59. Thelesperma

32. Leaves entire or divided, if divided then divisions not filiform ................................60. Bidens


33(31) Receptacle chaffy .............................................................................................29. Marshallia

33. Receptacle naked ..................................................................................................................34


34(33) Phyllaries white and membrane-like, almost petaloid ...........................30. Hymenopappus

34. Phyllaries herbaceous, not white and membrane-like ..........................................................35


35(34) Flowers yellow .....................................................................................................53. Helenium

35. Flowers whitish, rose, pink, or purplish; never yellow ........................................31. Palafoxia


36(2) Peripheral flowers actually disk flowers with expanded, tubular, 5- toothed corollas .............

...........................................................................................................................20. Centaurea

36. Peripheral flowers either typical ray florets or else with very reduced ligules .....................37


37(36) Pappus of disc florets partly or wholly of (usually many) very fine round or capillary bristles (small scales or bristles may also be present) .....................................................................38

37. Pappus of disc florets of coarse bristles (if bristles slender, then flattened or just a few), scales, awns, teeth, etc. OR pappus absent. ........................................................................52


38(37) Disc corollas cream-colored; ligules of ray flowers poorly developed ................16. Conyza

38. Disc corollas yellow, bluish, or rose; ligules of ray flowers well-developed ........................39


39(38) Ligules of ray flowers white, blue, rose, etc; never yellow ....................................................40

39. Ligules yellow .........................................................................................................................43


40(39) Leaves, if any, small and often represented by spines ..............................33. Chloracantha

40. Leaves present, well-developed (perhaps narrow); spines absent .....................................41



41(40) Phyllaries usually in 1 to 3 series, not overlapping much vertically and usually not green at the tips; ray flowers ca. 25 to 300 per head ........................................................32. Erigeron

41. Phyllaries usually in 3 or more vertically-overlapping series, usually green at the tips; ray flowers ca. 5 to 60 per head ..................................................................................................42


42(41) Heads in a flat-topped, corymbose or umbelliform arrangement; pappus bristles in 3 series, the outermost very short; achene terete ..........................................34. Doellingeria

42. Heads variously arranged, but inflorescence not flat-topped; pappus bristles in one series; achene flattened .......................................................................35.Aster (Symphyotrichum)


43(39) Disc flowers with pappus bristles of 2 distinct types or of 2 different lengths, usually a short outer ring (sometimes scale-like) and a longer inner ring. (Do not confuse pubescence on the achene with a pappus series.) (See Figure B at the drawing for couplet 37) ................44

43. Disc flowers with pappus bristles uniform or of random lengths, but never of 2 distinct lengths or types. (Figures A at couplet 37) ...........................................................................46


44(43) Ray flowers without pappus; herbage aromatic and glandular ...................37. Heterotheca

44. Ray flowers with pappus; herbage little or not aromatic or glandular ..................................45


45(44) Lower stem leaves linear, grass-like, with parallel veins; herbage with long, soft, silvery hairs .....................................................................................................................38. Pityopsis

45. Lower stem leaves oblong-lanceolate to oblanceolate, veins not parallel; herbage pilose but not silvery ..................................................................................................39. Chrysopsis

46(43) Phyllaries of two very different sizes, the inner bracts in 1 or 2 series, equal, linear, erect, and not much vertically or horizontally overlapping; the outer series of bracts smaller, forming a calyx-like structure or this sometimes absent ......................................................47

46. Phyllaries in several regular, vertically and/ or horizontally overlapping series ...................48


47(46) Leaves markedly reduced up the stem, at least some lobed .............................41. Packera

47. Leaves not or only gradually reduced upwards, none lobed ..............................40. Senecio


48(46) Plants annual; heads more than ca. 5 to 6 mm wide ...........................................................49

48. Plants perennial, often rhizomatous; heads less than ca. 6 mm wide ................................50


49(48) Heads solitary; leaves toothed along entire margin ...................................42. Rayjacksonia

49. Heads arranged in inflorescences; leaves toothed only in the distal portion if at all ...............

.............................................................................................................................43. Croptilon


50(48) Receptacle chaffy ...............................................................................................44. Euthamia

50. Receptacle not chaffy (or sometimes chaffy only around the periphery) ............................51


51(50) Inflorescence corymbose, flat-topped or nearly so; phyllaries oblong or oblong-obovate, blunt ...............................................................................................................45. Oligoneuron

51. Inflorescence racemose or paniculate, pyramidal or cylindrical, not flat-topped; phyllaries usually linear-lanceolate to linear .......................................................................46. Solidago


52(37) Pappus of disc florets wholly or partly of several to many long, slender awns or bristles, these rigid, flattened, coarse, or clavate. (Figures C and D at couplet 37) .........................53

52. Pappus of disc florets composed of scales, strap-like appendages, teeth, a small crown, or just a few awns or bristles OR pappus absent. (Figures E-K at couplet 37) ...................57


53(52) Pappus of the ray flowers caducous, seldom intact on the achenes; margins of upper leaves with small, blunt, even teeth ....................................................................47. Grindelia

53. Pappus of the ray flowers persistent; margins of upper leaves various,

if toothed, the teeth sharp or uneven .....................................................................................54


54(53) Ray flowers bluish ..................................................................................................36. Eurybia

54. Ray flowers yellow ..................................................................................................................55


55(54) Phyllaries without scarious margins ................................................................48. Prionopsis

55. Phyllaries with scarious margins ............................................................................................56


56(55) Pappus members of two lengths: a shorter, outer set and a longer, inner set. (Figure D at couplet 37) ......................................................................................................49. Xanthisma

56. Pappus members all of one length ....................................................................43. Croptilon


57(52) Pappus of disk flowers of 5 (or more) white linear, strap-shaped appendages (See Figure K at couplet 37); pappus of ray flowers a small scaly crown ....................50. Amphiachyris

57. Pappus of disk and ray flowers not as above ........................................................................58


58(57) Main feature of the pappus of the fertile flowers (only disk flowers if both ray and disk flowers fertile) 2 to several scales, these awn-tipped in some genera, sometimes over the shoulders of the achene or sometimes forming a scaly crown. (Figures E, F, and G at couplet 37) .............................................................................................................................59

58. Main feature of the pappus of the fertile flowers (only disk flowers if both ray and disk flowers fertile) 1 to several (mostly few) rigid awns or bristles, usually over the angles of the achene; OR pappus reduced to a blunt or toothed crown or tiny projections over the shoulders of the achene OR pappus absent. (Figures H, I, and J at couplet 37) ...............72


59(58) Receptacle naked or at most somewhat pubescent ............................................................60

59. Receptacle chaffy or bristly ....................................................................................................65


60(59) Ligules white, rose, violet, or some shade of purple .............................................................61

60. Ligules yellow ..........................................................................................................................62


61(60) Ligules of ray flowers usually rose, violet, or purple (white only in rare albinos) ....................

.............................................................................................................................31. Palafoxia

61. Ligules of ray flowers white when fresh ..................................................51. Aphanostephus


62(60) Heads less than ca. 6 mm wide ......................................................................52. Gutierrezia

62. Heads more than ca. 6 mm wide ...........................................................................................63


63(62) Phyllaries reflexed at maturity ............................................................................53. Helenium

63. Phyllaries not reflexed at maturity ..........................................................................................64


64(63) Outer phyllaries united at the base to ca. 1/2 their length, noticeably thickened; inner phyllaries separate, in 1 series, leathery, slightly longer than the outer ...................................

........................................................................................................................54. Hymenoxys

64. Outer phyllaries separate and not thickened; inner phyllaries in 2 series, herbaceous with membranous margins ....................................................................................55. Tetraneuris


65(59) Leaves opposite ......................................................................................56. Tetragonotheca

65. Leaves alternate .....................................................................................................................66


66(65) Ligules white ....................................................................................................11. Parthenium

66. Ligules yellow or red, often with brown or darker red markings ...........................................67


67(66) Disk flower pappus scales linear, narrow at the base, sometimes somewhat bristle-like; ligules of ray flowers entire ....................................................................................................68

67. Disk flower pappus scales wider at the base, sometimes terminating in long awns; ligules of ray flowers toothed apically ................................................................................................70


68(67) Phyllaries free; pappus scales of 2 types: a long, lanceolate inner series alternating with a linear, subulate outer series (Figure A at couplet 67, above); phyllaries not dotted with glands ................................................................................................................49. Xanthisma

68. Phyllaries connate at the base (in ours); pappus scales linear and all alike (Figure B at couplet 67); phyllaries dotted with dark or orange glands ....................................................69


69(68) Leaves or lobes of leaves linear-filiform; phyllaries strongly connate for 2/3 or more their length and their margins free usually less than 1/2 their length; receptacle essentially naked ...........................................................................................................57. Thymophylla

69. Leaves simple, linear or broader; phyllaries connate 1/3 to 2/3 their length but separating at maturity, their margins free to the base; receptacle with fringe, bristles, or scaly ridges ....

. ......................................................................................................................58. Dysodiopsis


70(67) Ray flowers 4 to 5, ligules with 2 apical teeth ..............................................59. Lindheimera

70. Ray flowers more than 5, ligules with 3 to 5 apical teeth or lobes .......................................71


71(70) Ligules deeply 3- to 5- toothed; pappus most conspicuous on disk flowers, composed of long-awned scales. (Figure C at couplet 67) ....................................................60. Gaillardia

71. Ligules minutely 3-toothed; pappus most conspicuous on ray flowers, composed of 2 awnless scales, one over each shoulder of the achene. (Figure D at couplet 67) ................

. .....................................................................................................................61. Engelmannia


72(58) Receptacle naked ..................................................................................................................73

72. Receptacle chaffy, the chaff adnate to the achenes or free ................................................77


73(72) Ligules yellow ..........................................................................................................................74

73. Ligules white or tinged with blue, purple, or rose, never yellow ...........................................75


74(73) Margins of phyllaries scarious ...........................................................................62. Bradburia

74. Margins of phyllaries not scarious ......................................................................47. Grindelia


75(73) Disk achenes (at least) prismatic, 5-ribbed ................................................63. Chaetopappa

75. Disk achenes (at least) flattened, elliptic in cross-section or winged ...................................76


76(75) Pappus absent or reduced to a minute border or ring .................................64. Astranthium

76. Pappus of small bristles or scales plus 2 (in ray flowers, sometimes 3) longer awns, one over each shoulder of the achene .......................................................................65. Boltonia


77(72) Ray flowers either asexual or pistillate but sterile .................................................................78

77. Ray flowers, or many of them, pistillate and fertile ...............................................................89


78(77) Receptacle flat to weakly convex ...........................................................................................79

78. Receptacle convex to columnar (doubtful specimens should be keyed under both

alternatives) ............................................................................................................................82


79(78) Phyllaries graduated in size and all similar .....................................................66. Helianthus

79. Phyllaries of two very distinct sizes or shapes (dimorphic) ...................................................80


80(79) Inner phyllaries connate for at least 1/4 of their length; leaves finely dissected ......................

......................................................................................................................67. Thelesperma

80. Inner phyllaries all separate (or nearly so); leaves simple to pinnately compound but not finely dissected .......................................................................................................................81


81(80) Ray flowers present or absent, the ligules lanceolate to oblanceolate, entire or with small apical teeth; pappus awns ciliate, hispid, barbed, or sometimes reduced or absent ..............

. ...............................................................................................................................68. Bidens

81. Ray flowers present, the ligules roughly cuneate to oblanceolate, with 3 to 5 apical lobes (rarely entire); pappus of 2 smooth, barbed, or ciliate awns or teeth, sometimes reduced or absent ...........................................................................................................69. Coreopsis


82(78) Phyllaries scarious or scarious-margined ..........................................................70. Anthemis

82. Phyllaries neither scarious nor scarious margined ...............................................................83

83(82) Achenes with two distinct wings ........................................................................71. Verbesina

83. Achenes without wings, though sometimes with angles, ridges, or narrow margins ..........84


84(83) Ray flowers rose, pink, purplish, or white .........................................................72. Echinacea

84. Ray flowers yellow, often marked with red or brown ............................................................85


85(84) Receptacle weakly convex or nearly flat .........................................................66. Helianthus

85. Receptacle columnar or strongly conic, at least in well-developed heads ..........................86


86(85) Achenes terete; leaves sessile and clasping .....................................................73. Dracopis

86. Achenes angled or flattened; leaves petiolate to sessile but not clasping ..........................87


87(86) Plants sprawling; ligules elliptic, not toothed; involucre of a few spreading phyllaries so that all the florets are visible; pales fused to the achenes and so looking like their outer layers ............................................................................................................76. Sclerocarpus

87. Plants erect; ligules more or less 3-toothed apically; involucre more closely enclosing the florets; pales folded tightly around pales but not fused to them ...........................................88


88(87) Leaves entire to toothed; achenes 4-sided (sometimes obscurely so), marginless (Figure A at couplet 83) ................................................................................................74. Rudbeckia

88. Leaves pinnatifid to bipinnatifid; achenes flat, margined (Figure B at couplet 83) ..................

...............................................................................................................................75. Ratibida


89(77) Disc flowers, or many of them, fertile (look for developing achenes) ..................................90

89. Disc flowers mostly infertile (not producing mature achenes) .............................................94


90(89) Leaves alternate or mostly so ................................................................................................91

90. Leaves opposite ......................................................................................................................92


91(90) Pappus of 1 to 3 awns or teeth; leaves simple .................................................71. Verbesina

91. Pappus completely absent; leaves bi- or tripinnate .............................................77. Achillea


92(90) Rays whitish, minute (about 2 mm long) ...............................................................78. Eclipta

92. Rays yellow, larger (if whitish, then greater than 2 mm long) ..............................................93


93(92) Phyllaries of two sizes: 4 long, leaf-like outer bracts and several short inner bracts present; erect perennials ........................................................................56. Tetragonotheca

93. Phyllaries of one size only; sprawling annuals .........................................79. Calyptocarpus


94(89) Rays in 2 to 3 series .............................................................................................80. Silphium

94. Rays in 1 series ......................................................................................................................95


95(94) Leaves manifestly whitish or grayish beneath ...............................................81. Berlandiera

95. Leaves not whitish or grayish beneath .........................................................59. Lindheimera


96(1) Pappus made up at least in part of scales; bristles may or may not also be present OR pappus reduced to a small crown or absent ...........................................................82. Krigia

96. Pappus scales absent; pappus of delicate bristles, sometimes a ring of short basal hairs present as well .......................................................................................................................97


97(96) Achenes manifestly flattened; pappus bristles not plumose ................................................98

97. Achenes terete to angled, little if at all flattened; pappus bristles plumose or not ..............99


98(97) Achenes beaked (beak usually filiform, sometimes stout) .................................83. Lactuca

98. Achenes not beaked ...........................................................................................84. Sonchus


99(97) Lower stem and leaves with long, spreading, tawny hairs ..............................85. Hieracium

99. Lower stem and leaves variously pubescent but not as above ..........................................100



100(99) Ligules lavender or white; pappus bristles plumose ........................................................101

100. Ligules yellow or orange; pappus bristles not plumose ....................................................102



101(100) Mature achenes beaked (figure A at couplet 98) .....................................86. Hypochaeris

101. Mature achenes unbeaked (figure B at couplet 98) .................................87. Lygodesmia


102(100) Heads several to many per stem; achenes unbeaked (Figure A at couplet 98) ..................

....................................................................................................................... ......88. Youngia

102. Heads one to few per stem; achenes beaked (Figure B at couplet 98) .........................103


103(102) Heads on true scapes; achenes spiny near the apex .................................89. Taraxacum

103. Heads on scape-like stems bearing a few reduced leaves; achenes not spiny ....................

. ..................................................................................................................90. Pyrrhopappus




ASTERACEAE ANTENNARIA1. ANTENNARIA Gaertn. Everlasting, Ladies' Tobacco, Pussy's Toes


Perennial herbs from rhizomes or stolons, 3 to 40 cm tall. Stems unbranched, decumbent or ascending. Leaves alternate, more or less glabrous above and white-woolly beneath, lower leaves obovate to spatulate, the largest to 3 to 6 cm long, upper leaves reduced, narrower, and less crowded. Heads discoid, campanulate. Phyllaries in 1 to several series, linear, blunt, the distal portion thin, white, shiny, and translucent. Receptacle convex, naked or papillate. Plants dioecious. Flowers of staminate plants: Corolla 5-toothed; stamens 5, anthers caudate (with tail-like appendages); pappus of flattened, somewhat clavate, bristle-like members. Flowers of pistillate plants: Corollas tubular and filiform; style branches thread-like; pappus of slender, nearly capillary bristles. Achene about 1 mm long, narrowly elliptical, brown, minutely papillate.

A genus of about 100 species, with 2 in Texas and 1 locally. (See Bayer and Stebbins 1982).


1. Antennaria parlinii Fern. subsp fallax (Greene) Bayer & Stebbins Characters of the genus. Plants to 40 cm, colonial or mat-forming, with procumbent or decumbent stolons, silvery pubescent. Lower leaves ovate to widely elliptical, 3 to 6 cm long, to 4(5) cm broad, narrowed into a subpetiolar base, apices obtuse, persistently tomentose below, upper leaves reduced, becoming narrower and acute to acuminate. Heads in a tight or loose cyme; pistillate involucres 7 to 10(14) mm tall at maturity, staminate involucres smaller. Achenes 1.2 to 1.5 mm long. Sandy, forested areas. E. TX, W. to N. Cen. TX; common in E. U.S. Spring, but with occasional isolated plants flowering in winter. [A. fallax Greene; A. plantaginifolia (L.) Rich. var ambigens (Greene) Cronq.].




ASTERACEAE BACCHARIS2. BACCHARIS L. Groundsel Tree


Shrubs or subshrubs. Branchlets terete to striate or angled, smooth to glandular-scabrous. Herbage glabrous to hispid or sometimes glandular-punctate. Leaves alternate, crowded to widely spaced, linear to ovate, entire to serrate or dentate, sessile, with 1 to 3 primary nerves, reduced in the inflorescence and sometimes bract-like there. Flowers all disk type, unisexual, male and female flowers on separate plants (occasionally a plant with bisexual flowers). Inflorescence roughly racemose, paniculate, or narrowly cylindrical. Staminate heads with involucres hemispheric to cylindrical, outer phyllaries ovate to lanceolate and the inner phyllaries lanceolate to linear, obtuse to acute, commonly with scarious margins and the midvein dilated and smooth or else absent; receptacle smooth to pitted, naked to fringed, flat to slightly convex; corollas slender basally and enlarged and funnelform apically, white to yellow-brown, with 5 lanceolate lobes; style present, ovary abortive; pappus not exceeding style, usually 1 series of numerous bristles, somewhat plumose-tipped and usually barbed or crisped. Pistillate heads with involucres hemispheric to narrowly cylindrical; phyllaries in several series, the outer bracts ovate to lanceolate and the inner lanceolate to narrowly linear, obtuse to acuminate, usually with scarious margins, the midvein obvious to obscure; receptacle pitted to smooth, naked or fringed, flat; corollas filiform, with 5 apical teeth or lobes, yellowish-white to brown; pappus of many soft or rigid bristles in 1 to several series, equalling to much longer than the style. Achene yellow to reddish, 5- to 10-ribbed, glabrous to hispid, smooth or glandular.

This is a large (about 350 species) American genus complicated somewhat by the tendency of species to interbreed. Positive identification requires at least the pistillate flowers and a sample of leaves from both the upper and lower parts of the plant. Texas has 10 or 11 species, depending upon interpretation; 3 species are present here.


1. Achenes 3 to 5 mm long, with 5 or 6 ribs; plants rhizomatous subshrubs ...1.B.texana

1. Achenes 1 to 2 mm long, with 8 to 10 ribs; plants shrubs usually more than 1 m tall ..........2


2(1) Leaves linear to narrowly linear-lanceolate or very narrowly elliptic, about 7 to 35 times

longer than broad ...2.B.neglecta

2. Leaves obovate to oblanceolate, about 2 to 10 times longer than broad ...3.B.halimifolia

1.B. texanaASTERACEAE BACCHARIS texana (T. & G.) Gray Prairie Baccharis. Low shrubs or subshrubs, 25 to 60 cm tall, often rhizomatous, branched from the base; branches smooth, striate-angled. Leaves somewhat punctate, the lower sessile, linear to narrowly lanceolate, 2 to 4(5) cm long, 2 to 4 mm broad, 1-nerved, the margin very minutely undulate, upper leaves smaller, linear, acute the uppermost to ca. 10 mm long, sometimes crowded; bracteal leaves below the heads with margins slightly scarious and ciliate. Heads arranged more or less in a corymb. Pistillate flowers: involucre campanulate, with 20 to 30 flowers, 7 to 9(10) mm tall; phyllaries lanceolate, keeled, the midrib dilated, apex acute, margin narrowly scarious, usually ciliate; receptacle naked, flat, pitted; corolla 3.5 to 4 mm long, filiform, apically truncate, the 5 teeth obscure and erose, only ca. 0.2 mm long; pappus 11 to 13 mm long, bristles many, in several series (this hard to discern), dirty white to tinged with yellow or red-brown, microscopically antrorsely barbellate; achene 3 to 4.5 mm long, 5- or 6-ribbed, brown, sub-glandular. Staminate flowers: involucre 6 to 7 mm long, campanulate; phyllaries and receptacle similar to those of pistillate flowers; corolla tube ca. 5 mm long, filiform but enlarged at about the midpoint, lobes lanceolate, 1.6 to 2 mm long; pappus about as long as the corolla, usually plumose-tipped and the bristles crisped; ovary abortive. Open outcrop or mesa areas; in our area known from Grimes Co.; Ed. Plat., Rio Grande Plains, Plains Country, Trans Pecos, N. Cen., and SE. TX; OK to Mex. Aug.-Nov.


2.B. neglectaASTERACEAE BACCHARIS neglecta Britt. Roosevelt Weed, New Deal Weed, Jara Dulce. Shrub, 1 to 3 m. tall; branches striate and angled, glabrous. Leaves, sessile, punctate, somewhat glutinous, narrowly linear to narrowly elliptic, (7)10 to 20(35) times longer than broad, ca. 2 to 4(5) mm broad, lower leaves 3 to 5(8) cm long, acute, the upper leaves entire and the lower leaves entire to serrate, with 1 prominent nerve and 2 obscure nerves running parallel to the margins; leaves reduced and bract-like in the head-bearing region. Heads in clusters or sometimes solitary or few. Staminate heads with 10 to 15 flowers each, the involucres hemispheric to subhemispheric, 3.5 to 4 mm tall; phyllaries as described for the genus, the tips stramineous to green or red-brown; receptacle pitted; corollas 2.7 to 3.3 mm long, the lobes ca. 1 mm long; pappus shorter than the style. Pistillate heads with 15 to 30 flowers each; involucres campanulate, 4 to 4.7 mm tall; phyllaries green or with red-brown tips, erose; receptacle pitted; corollas 2.5 to 3.3 mm long, with 5 minute teeth to 0.3 mm long; pappus in 1 to 2 series, 6.5 to 9 mm long, exceeding the style; achenes 1.2 to 1.5 mm long, glabrous, 10-ribbed. Roadsides and other disturbed areas, preferring calcareous soils. Nearly throughout TX except the deep woods of E. TX and the higher elevations in the Plains Country; NC to AZ and S. to Mex. Late summer-fall (ours primarily Sept.-Oct.) [TX material often erroneously referred to B. angustifolia Michx.].

NOTE: This species and B. halimifolia, below, commonly interbreed in our area. Zanowiak (1991) found that most samples could not be positively identified without isozyme analysis, and even then most proved to be hybrids.

A yellow dye can be made from the leaves and flowers (Tull 1987).


3.B. halimifoliaASTERACEAE BACCHARIS halimifolia L. Sea Myrtle, Consumption Weed, Groundsel Tree, Silver-ling. Shrub 1 to 2(6) m. tall; branches striate-angled; herbage glabrous and glutinous. Leaves punctate, petiolate to sessile, obovate or oblanceolate to elliptic, elliptic-oblanceolate, or rhombic, 2 to 6 cm long and 3 to 20 mm wide, (2)2.5 to 5(10) times longer than broad, with a prominent midrib and 2 less-prominent, ascending lateral nerves radiating from the midrib somewhat above the base, entire or the proximal 1/2 of the blade entire and the distal 1/2 with 1 to 3 pairs of teeth, upper leaves reduced and more entire. Inflorescence roughly widely paniculate, each head with about 20 flowers. Staminate flowers with hemispheric involucres 4.5 mm long; phyllaries as described for the genus; receptacle pitted, fringed to naked; corollas 3 to 3.5 mm long, the lanceolate teeth about 1 mm long; pappus 3 to 4 mm long, flaccid. Pistillate flowers with campanulate involucres 4.5 to 5 mm long; phyllaries as described for the genus, greenish-brown tipped, the midrib sometimes indistinct; receptacle flat to somewhat convex, naked; corollas 2.5 to 2.8 mm long, the lobes about 0.1 mm long; pappus bristles in 2 series, soft, 10 to 12 mm long, much exceeding the style and the phyllaries; achenes 1.3 to 1.8 mm long, 8- to 10-ribbed, glabrous. Bisexual flowers sometimes present, corollas about 3 mm long. Often plentiful in sunny, sandy areas. E. and SE. TX, less abundant W. to N. Cen. TX; Coastal states, MS to TX and inland to OK. Late summer-fall.

See at B. neglecta, above.






ASTERACEAE IVA3. IVA L. Swamp-weed, Marsh-elder


Annual or perennial herbs, erect, branching. Leaves (in ours) opposite or opposite passing to alternate above, entire to serrate, glabrous to pubescent. Heads in spikes, racemes, or panicles, subtended by reduced leaves or bracts. Involucres hemispheric or turbinate. Phyllaries 3 to several, free or united. Receptacle flat and chaffy. Staminate and pistillate flowers in the same head, staminate in the center and pistillate on the periphery. Ray flowers absent. Staminate corollas funnelform. Pistillate corollas tubular, truncate (the lobes not well-developed). Pappus none. Achenes (in ours) 2 to 3 mm long, obovate to cuneate (Kindscher 1987).

A North American genus with 19 species; 8 in TX; 2 locally.

American Indians used the achenes for food and archaeological evidence suggests that they cultivated the plants.


1. Largest leaves usually greater than 1.5 cm wide, ovate; phyllaries distinct ...1.I.annua

1. Largest leaves usually less than 1.5 cm wide, lance-linear; phyllaries united .........................

...2.I.angustifolia


1.I. annuaASTERACEAE IVA annua L. Marsh-elder, Pelocote, Seacoast Sumpweed. Annual herb from a taproot; stems 4 to 20 dm tall, simple or branched. Leaves opposite, scabrous on both sides, 3-nerved from the base, lower leaves ovate, 3 to 12 cm long, remotely serrate, upper leaves becoming lanceolate upwards and reduced to bracts in the head-bearing region. Heads 4 to 5 mm broad, in terminal spikes, subtended by ovate-acuminate bracts with ciliate margins. Involucres turbinate; phyllaries 3 to 4, distinct, cuneate, with a truncate or rounded apex, hispid or ciliate. Staminate flowers 8 to 17 per head; pales linear and glandular; corolla about 2.5 mm long. Pistillate flowers 3 to 5 per head; pales filiform, clavate, or absent; corolla about 2.5 mm long; achene 2 to 4.5 mm long, brown, obovate, sparsely to densely resin-dotted. Common in the E. half of TX and most of the E. U.S. Late summer to fall. [Some sources recognize varieties, or Includes var. caudata (Small) R. Jackson; I. caudata Small; I. ciliata Willd.].


2.I. angustifoliaASTERACEAE IVA angustifolia Nutt. ex DC. Narrow-leaf Marsh-elder. Annual or sometimes short-lived perennial, 5 to 12 dm tall; stems much branched above; stems and leaves strigose. Leaves opposite below, passing to alternate upwards, lower leaves entire to remotely serrate, lance-linear, 2 to 4(5) cm long and 2 to 8(12) mm broad, the largest visibly 3-nerved, leaves becoming gradually narrower upwards, much reduced in the head-bearing region. Heads sessile, in spike or panicles; bracts in the inflorescence linear-lanceolate, ciliate. Involucres turbinate, 2 to 3 mm broad and about as high; phyllaries united, hispid to strigose on the outer surface and ciliate on the tips. Staminate flowers 1 to 3(5) per head; pales filiform-linear; corolla about 2 mm long. Pistillate flowers 1 or 2 per head; pales absent; corolla about 1.5 mm long; achenes black, rugose, sometimes pubescent, 2 to 2.8 mm long. Common nearly throughout TX, also LA, AR, and OK. Late summer to fall.

If varieties are recognized, our plants are var. angustifolia.



ASTERACEAE XANTHIUM4. XANTHIUM L. Cocklebur


Erect, taprooted annuals 2 to 20 dm tall. Leaves mostly alternate. Heads axillary, subsessile, unisexual, composed solely of disk flowers. Staminate heads: In axils of upper leaves; phyllaries in 1 to 3 series, free, herbaceous; receptacle conic, chaffy; florets several; corolla tubular; anthers free; vestigial gynoecium present. Pistillate heads: In axils of middle leaves; phyllaries united, enclosing the 2 pistillate flowers, maturing into a bur covered with uncinate spines; corolla and pappus absent; achenes 1 in each of the bur's 2 chambers.

A North American genus. The number of species varies from 2 to about 24, depending on the reference consulted. Two species occur in TX; 1 is found locally.

Cockleburs cause injury in livestock and the young foliage can cause poisoning in livestock (Tull 1987). The burs are useful for craft projects and children find them fun to throw.


1. X. strumarium L. Abrojo. Characters of the genus. Stem scabrous-strigose. Leaves long-petioled, ovate-deltoid with cuneate bases, scabrous-strigose on both surfaces. Burs to about 3 cm long. Common throughout TX and U.S. Summer to fall, with the burs remaining throughout the winter. [X. orientale L.; X. chinense Mill.; X. italicum Moretti; X. pensylvanicum Wallr.; X. saccharatum Wallr.; X. speciosum Kearn.; X. cenchroides Millsp. & Sherff.]

According to Cronquist (1980), ours are var. canadense (P. Mill.) T. & G.



ASTERACEAE AMBROSIA5. AMBROSIA L. Ragweed


Perennial or annual herbs, 2 to 30 dm tall. Herbage aromatic with glandular hairs, resinous. Leaves alternate to opposite, entire to pinnately lobed or dissected. Heads discoid, unisexual; plants monoecious. Staminate heads: In spikes or racemes, sometimes aggregated into paniculate inflorescences, usually nodding; involucres hemispheric; phyllaries in 1 series, united; receptacle flat, chaffy with slender or reduced pales; flowers several to many per head; anthers more or less free; rudimentary gynoecia present; pappus absent. Pistillate heads: In axils of leaves and bracts, inconspicuous, consisting of 1 to few flowers; phyllaries united, forming a hard, nut-like structure around the achenes; phyllary tips forming spines or tubercles on the mature involucre; corolla, androecium, and pappus absent.

About 43 species, mostly American. Texas has 8; we have 3. All are wind-pollinated and are the actual culprits causing hayfever when Goldenrod (which blooms around the same time) is blamed.

American Indians used the achenes for food and probably even cultivated the plants (Kindscher 1987).


1. Leaves simple or palmately 3- to 5-lobed; outer surface of staminate involucre with 3 darker green lines radiating outward from the peduncle ...1.A.trifida

1. Leaves pinnately lobed or pinnatifid; staminate involucre unlined .........................................2


2(1) Lower leaves petiolate; annuals from taproots ...2.A.artemisiifolia

2. Lower leaves subsessile; perennials from rhizomes ...3.A.psilostachya

1.A. trifidaASTERACEAE AMBROSIA trifida L. Giant Ragweed, Buffalo Weed. Erect, taprooted annual 1 to 3(5) m. tall; stem angled, bearing darker green lines (striations), scabrous or warty. Petioles sometimes slightly winged; leaves mostly opposite, scabrous on both surfaces, margins serrate, lower leaves orbicular-ovate in outline, 10 to 20 cm long, 3- to 5-cleft, the divisions sometimes having their own small lobes, upper leaves 3-cleft to simple, lanceolate-rhombic-ovate. Staminate heads in racemes, the inflorescence often branched and paniculate; involucre about 3 mm broad, with 6 to 8 lobes and 3 darker striations. Pistillate heads in axils of bracts or leaves below the staminate racemes; mature pistillate involucres obovoid, about 4 mm long, scabrous-reticulate, with 4 to 8 ridges and as many small spines, a terminal beak about 1 mm long also present. Usually in damp areas, often forming large stands. Common in N. and E. TX, the Midwest, and the central U.S.; adventive elsewhere. Summer to fall.

Almost all TX material is classifiable as var. texana Scheele [Includes var. aptera (DC.) Kuntze; A. aptera DC.].

The seeds of giant ragweed were used as food by Native Americans. The plant was even cultivated to assure a harvest (Kindscher 1987).


2.A. artemisiifoliaASTERACEAE AMBROSIA artemisiifolia L. Short Ragweed, Altamisa, Common Ragweed, Roman Wormwood. Erect taprooted annual 3 to 10 dm tall; stem well-branched, hispid-strigose. Leaves alternate below and opposite above, petioles 1 to 3 cm long, often winged, upper leaves short-petioled, becoming almost subsessile in the head region; blades to 10 cm long and 7 cm wide, pinnatifid to bipinnatifid, the forward-pointing divisions (especially the terminal one) also toothed and lobed, tips acute, sinuses rounded, hirsute-puberulent above and hispid-strigose below, especially along the veins. Staminate heads in racemes, inflorescence often paniculate; heads obconic, oblique, about 3 mm broad, hirsute to villous; pales filiform. Pistillate heads few to several in clusters below the staminate inflorescences; mature involucres obovoid, about 3 mm broad, hispidulous-strigose to glabrate, slightly angled and reticulate, with 5 to 7 small spines or tubercles, terminal beak about 1 mm long. Infrequent but found throughout most of TX except the far S. and W.; most of the U.S. except the far SW.; adventive elsewhere. Late summer to fall. [A. elatior L.]

Some sources, e.g., Kartesz (1998), recognize varieties.

This plant is responsible for much of the pollen that causes fall hayfever.


3.A. psilostachyaASTERACEAE AMBROSIA psilostachya DC. Western Ragweed, Perennial Ragweed. Perennials from rhizomes or runner-like roots; stems 3 to 6(10) dm tall, often branched above, striate and with short, ascending hairs. Leaves subsessile, lanceolate or lance-ovate in overall outline, pinnately lobed or pinnatifid with forward-pointing divisions, divisions acute to acuminate, some of the larger ones weakly toothed, leaves scabrous-hirsutulous with hairs that are inflated at the base, also with minute glands or granules. Inflorescence simple to much-branched. Staminate heads in racemose arrangements; involucre broadly obconic, about 2.5 mm long, oblique, crenate, with 2 larger teeth on one side, hispidulous with hairs with inflated bases. Pistillate heads few to several in clusters below the staminate inflorescence; mature involucres 2.5 mm long, obovate, rugose, minutely hirsute, 4 to 6 short teeth or tubercles present, terminal beak about 1 mm long. Abundant in the Coastal Plain and Cen. TX, forming extensive colonies; becoming less common westward; throughout the U.S. except in the Rockies and the E. Cen. states. Late summer to

fall. [Includes var. lindheimeriana (Scheele) Blank., var. californica (Rydb.) Blake, and var. coronopifolia (T. & G.) Farw., etc.; A.

californica (Rydb.); A. coronopifolia T. & G.; A. lindheimeriana Scheele; A. cumanensis of authors, but not A. cumanensis Kunth in H.B.K.].


ASTERACEAE ARTEMISIA6. ARTEMISIA L. Sagebrush, Sage, Wormwood, Mugwort


Perennial herbs, less commonly subshrubs or shrubs, from taproots, woody crowns, or rhizomes. Stems erect, somewhat branched, (2)5 to 10 dm tall. Herbage glabrous to tomentose, usually aromatic. Leaves alternate, entire to lobed or dissected. Heads small, in panicles, or spike-like or raceme-like inflorescences. Involucres urceolate at anthesis and narrowly campanulate in fruit, 3 to 6 mm tall. Phyllaries with scarious margins (this sometimes obscured by pubescence), in 3 series and of 2 sizes--the outer series 1/4 the length of the inner two, lanceolate and acuminate, forming a sort of calyculum, the inner 2 series equal in length, the middle series about as broad as long, blunt, the innermost series elliptic, blunt to acute. Receptacle convex to conical, naked (rarely pubescent). Heads inconspicuously radiate. Ray flowers 1 to few, yellowish-white, pistillate, fertile or infertile; corollas 1 mm long or less, cylindrical, ligule absent and zygomorphy represented by a v-shaped notch down the side of the tube next to the disk. Disk flowers 3 to 20 per head, about as long as the longer phyllaries, perfect, fertile or infertile; corollas yellowish-white. Pappus absent. Achenes about 1 mm long (or less), cylindrical or elliptic in cross-section, obovate to oblong in outline, weakly 1-ribbed, brown.

A few hundred species in cool, arid climates. There are 7 species in TX, mostly western, and 1 found occasionally in our area.

All species are wind-pollinated and have allergenic pollen (Correll and Johnston 1970). Many can cause contact dermatitis (Lampe 1985). One species, A. dracunculus L. var. sativa, is the Tarragon used in cooking (Mabberley 1987). Most TX species provide some forage for livestock (Correll and Johnston 1970).


1. A. ludoviciana Nutt. Western Mugwort, White Sage. Perennial herb from rhizomes; stems sometimes woody at the base. Stems several to many, (2)3 to 9(15) dm tall; herbage gray- or whitish-pubescent, but in some plants the upper surface of the leaves glabrescent. Leaves extremely variable in shape, often almost all entire, but often at least some dissected, the lobes and main axis usually 3 to 10 mm broad, commonly the entire ones elliptic to linear-lanceolate. Inflorescence a narrow panicle, somewhat spike-like, leaves included among the heads. Involucres usually densely gray or white woolly. "Ray" flowers present, as described for the genus, occasionally fertile but usually not. Locally common to infrequent throughout TX except the extreme S.; uncommon in our area; wide-ranging in temperate N. Amer. Summer-fall. [A. mexicana Willd.; A. gnaphalodes Nutt.; A. vulgaris var. americana Bess.; A. lindheimeriana Scheele; A. redolens Gray; A. rhizomata A. Nels; A. texana Rydb.; A. albula Woot. & Standl.] Further studies may prove this species to be conspecific with the Eurasian A. vulgaris. Several different subspecific divisions have been proposed.

Plains tribes used white sage both medicinally and ceremonially (Kindscher 1992).



ASTERACEAE PTEROCAULON7. PTEROCAULON Ell. Blackroot


4 species in the warmer parts of the Americas; 1 in TX.


1. Pterocaulon virgatum (L.) DC. Perennial herbs from woody crowns or rhizomes; stems 1 or 2 per plant, (5)8 to 15 dm tall. Leaves alternate, linear with revolute margins to 12 cm long, green above and white-tomentose beneath, resinous-glandular beneath the hairs, decurrent on the stem as wings. Heads of disk flowers only, campanulate, arranged in spike-like inflorescences. Involucres to 5.5 mm long. Phyllaries in several series, lanceolate, white-pubescent except at the tips, which are often reddish, outer bracts ca. 2.2 mm long, inner ca. 4.8 mm long; receptacle dark, flat to conical, naked, rough, or with a few hairs. Disk florets of 2 kinds: peripheral flowers with filiform corollas to 5.6 mm long, pistillate and fertile, and ca. 4 central flowers with tubular corollas, perfect but infertile; anthers with small tail-like appendages (caudate). Pappus of fragile buff to white bristles. Achenes ca. 1 mm long, brown, columnar to oblanceolate in outline, sometimes slightly curved, ribbed and pubescent. Uncommon in sandy soils. E., SE., and Coastal Plains in TX, but also found in our area. Also W.I. and E. Mex. Late summer to fall--Sept.-Oct. [Gnaphalium virgatum L.].



ASTERACEAE EVAX8. EVAX Gaertn. Rabbit-tobacco, Cotton-rose


Small, taprooted annuals. Stems branched or unbranched. All parts gray-white woolly or tomentose. Leaves alternate, usually spatulate and less than 2 cm long. Heads in dense and woolly glomerules, subtended and interrupted by leaves, individual florets within the heads mostly obscured by the pubescence. Involucres absent, the pales appearing like phyllaries. Receptacle convex, chaffy, the pales thin and woolly. Ray flowers absent. Disk flowers of 2 kinds: peripheral flowers with minute, tubular-filiform corollas, pistillate and fertile; and a few central flowers with flaring, 4-toothed corollas, perfect but usually infertile. Anthers with microscopic tails. Pappus absent. Achenes oblong to elliptic, flattened, less than 1 mm long, brown, often with minute papillae.

A genus of about 24 species, 3 in TX and 2 locally.


1. Heads mostly clustered at the ends of stems and branches; central florets of the heads usually infertile ...1.E.verna

1. Heads mostly clustered in the axils of the leaves; central florets of the heads usually fertile ...2.E.candida

1.E. vernaASTERACEAE EVAX verna Raf. Plants 3 to 12(15) cm tall, often branched. Glomerules of heads mostly all terminating stems and branches, subtended by leaves 3 to 10 mm long; leaves within the cluster of heads little if at all protruding above the surface of the cluster. Nearly throughout the state; also LA, OK, NM, and N. Mex. April-May. [E. multicaulis DC.; Filaginopsis drummondii T. & G.; Filago verna (Raf.) Shinners; F. nivea Small]. Some sources, e.g., Kartesz (1998), recognize varieties.


2.E. candidaASTERACEAE EVAX candida (T. & G.) Gray Plants 6 to 12(25) cm tall; stems 1 to several from the base. Leaves 15 to 22 mm long, 3 to 5 mm broad. Glomerules of heads in the axils of the middle and upper leaves. Achenes ca. 0.5 mm long. Sandy prairies in E. and SE. TX; rarer westward; endemic. April-May. [Calymmandra candida T. & G.; Filago candida (T. & G.) Shinners].



ASTERACEAE FACELIS9. FACELIS Cass.


4 South American species, one of which is adventive in TX.


1. F. retusa (Lam.) Schultz- Bip. Taprooted annual 3 to 30 cm tall; stems branched at the base, decumbent, densely gray-woolly. Leaves alternate, crowded, sessile, spatulate, 0.5 to 3 cm long and 2 to 4 mm broad, green above and white-floccose beneath, leaf apices retuse to truncate, mucronate. Heads in terminal glomerules, cylindric, 6 to 11 mm long. Phyllaries in several series, scarious, occasionally with reddish or purple tips. Receptacle flattish, naked. Ray flowers absent. Disk flowers of 2 kinds: peripheral flowers with truncate or scarcely-lobed corollas, pistillate; and central flowers with 5-lobed corollas, perfect and usually fertile. Anthers caudate (tailed). Pappus of whitish plumose bristles 8 to 10 mm long. Achenes obovate-oblanceolate in outline, flattened, whitish-pubescent, 1 to 1.5 mm long. Roadside weed of sandy soils in E. TX; widely adventitious in the SE. U.S. April-May. [F. apiculata Cass.].



ASTERACEAE GNAPHALIUM10. GNAPHALIUM L. (Including Gamochaeta and Pseudognaphalium) Cudweed,
Everlasting


Taprooted annuals or weak perennials. Herbage floccose-woolly nearly throughout. Leaves alternate, sessile, sometimes slightly decurrent on the stem. Heads cymose or in glomerules. Phyllaries in several series, scarious but with a wide, stiff, green midnerve. Receptacle flattish, naked. Heads composed of disk flowers--all fertile, but the center florets of each head pistillate. Corolla minutely 5-toothed; anthers minutely tailed. Pappus one series of short, scabrous capillary bristles. Achenes terete or flattened.

200 species worldwide, with 8 in TX and 4 locally. Cabrera (1961) retained G. obtusifolium in Gnaphalium, but removed G. falcatum, G. purpureum, and G. pensylvanicum to Gamochaeta on the basis of the united pappus bristles, a treatment recognized by Hatch, et al. (1990). Cronquist (1980), GPFA (1986), and Gandhi and Thomas (1989) followed the trend of recognizing only Gnaphalium. Recent work (as reflected in the treatment presented by Kartesz (1998) suggests that

the group may be paraphyletic, with Gamochaeta being valid and some species transferred to still another genus, Pseudognaphalium. Synonyms in the segregate genera are provided here.


1. Upper surfaces of mature leaves bright green, drying brown or dark olive green, scarcely pubescent; pappus bristles, if deciduous, not united, falling separately ...1.G.obtusifolium

1. Upper surfaces of mature leaves gray-green, drying the same color; pappus bristles united at the base and deciduous as a unit .............................................................................2


2(1) Leaves linear to narrowly oblanceolate, less than 6 mm wide, about equally pubescent on both sides ...2.G.falcatum

2. Leaves broadly oblanceolate to long-tapering obovate, more than 5 mm wide,

pubescence denser on the under surface than the upper ......................................................3


3(2) Pubescence of lower leaf surface of tightly-interwoven hairs (pannose), whitish; involucres mostly glabrous, woolly only at the base ...3.G.purpureum

3. Pubescence of lower leaf surface looser, villous, grayish; involucres woolly throughout ......

. ...4.G.pensylvanicum

1.G. obtusifoliumASTERACEAE GNAPHALIUM obtusifolium L.(= Pseudognaphalium obtusifolium (L.) Hilliard & Beard subsp. obtusifolium) Cat-foot, Fragrant Cudweed. Erect annual; stems 25 to 50(100) cm tall, slightly aromatic, at first woolly then becoming merely glandular-villous or puberulent with age. Leaves long-lanceolate to linear, 2.5 to 8 cm long and 2 to 8 mm broad, sessile, often with undulate margins, drying brown or dark olive green, upper surface of leaves viscid-puberulent or glandular with age, lower surface densely white-woolly. Heads in terminal glomerules; involucres 6 to 7 mm long, pubescent at the base; phyllaries scarious, oblong, obtuse to acute, of several lengths. Flowers in each head mostly pistillate. Pappus bristles not united basally, falling individually. Achenes 0.6 to 0.8 mm long, sometimes with 3 or 4 faint nerves, glabrous. Common in E., SE., and Cen. TX, less common westward to the Ed. Plat.; Gulf states N. to E. Can. Sept-Nov. [Gn. polycephalum Michx.].

This plant was used in a variety of medicinal preparations in both Native American and Anglo traditions (Kindscher 1992).


2.G. falcatumASTERACEAE GNAPHALIUM falcatum Lam. (= Gamochaeta falcata (Lam.) Cabrera) Erect annual 1 to 3 dm tall; stems densely to loosely villous. Leaves narrowly linear-oblanceolate or linear, to about 6 cm long and 6 mm wide, acute, upper and lower surfaces about equally woolly. Heads nearly sessile in the axils of the upper leaves. Involucres 4 to 6 mm high, woolly at the base. Pappus bristles united and deciduous as a ring. Achenes about 0.5 mm long. E. half of TX, introduced in other SE states. Mar.-Apr. [Gn. purpureum L. var falcatum (Lam.) T. & G.].


3.G. purpureumASTERACEAE GNAPHALIUM purpureum L. (= Gamochaeta purpurea (L.) Cabrera) Purple Cudweed. Annual 1 to 3 dm tall, usually with several stems from the base. Leaves narrowly oblanceolate, 2 to 7 cm long and 0.5 to 1.5 cm broad, sessile, entire or serrate to serrulate, especially at the tips, apices obtuse to rounded, lower leaf surfaces densely white-pannose, the hairs tightly intermeshed, upper leaf surfaces less densely pubescent, at most villous. Heads subsessile in the axils of the upper leaves, sometimes these leaves reduced and crowded so that a spike-like inflorescence results. Involucres 3 to 6 mm high, woolly at the base; phyllaries scarious, linear to lanceolate, acute, subequal, sometimes purple or pinkish. Pappus of bristles united at the base and deciduous as a unit. Achenes 0.5 to 0.7 mm long. E. half of TX, most common in sandy soil, less common W. to Ed. Plat.; wide-ranging N. to N. Eng., S. and W. to MO and KS; also OR. Mar.-May(July).


4.G. pensylvanicumASTERACEAE GNAPHALIUM pensylvanicum Willd. (= Gamochaeta pensylvanica (Willd.) Cabrera) Annual or biennial to 4 dm; stems usually several from the base. Leaves long-tapering obovate or broadly oblanceolate, 5 to 20 mm wide, obtuse to rounded at the tip, margins sometimes slightly crenate or serrate, especially at the tips, lower leaf surface grayish-green and loosely villous, upper surfaces of leaves less densely pubescent and less gray. Heads in axils of reduced upper leaves, resulting in a spike-like inflorescence. Involucres 3 to 4 mm high, woolly from base to tip, the phyllaries brownish or purple beneath the wool. Pappus bristles united at the base and deciduous as a unit. Widespread but not abundant in the E. half of TX; more common in the sands of the Rio Grande Plains; Gulf states N. to PA, also CA. Spring, but occasionally as late as July; one collection from Dec. [Gn. spathulatum of some authors, not Lam.; Gn. purpureum L. var. spathulatum (Lam.) Baker; Gn. peregrinum Fern.]




ASTERACEAE PARTHENIUM11. PARTHENIUM L.


Herbs or shrubs, bitter and aromatic. Leaves alternate, entire to much-divided. Heads small, hemispheric, radiate with inconspicuous rays, borne singly or in corymbiform or paniculate clusters. Phyllaries in 2 to 4 series, the inner ones slightly shorter and subtending the ray flowers. Receptacle flat to slightly convex, chaffy. Ray flowers 5, pistillate and fertile; ligules white, sometimes reduced or virtually absent; styles and corollas persistent on the achenes. Disk flowers perfect but functionally only staminate, corollas white. Pappus of 2 to 3 awns, 2 scales, or absent. Each disk floret (except the outer row of 10) falling as a unit together with its subtending pale. Achenes of ray flowers flattened perpendicular to the radius of the head, obovate to rotund in outline, the margins thickened into ribs. At maturity each ray achene, its subtending pale, and its 2 neighboring outer-row disk florets falling as a unit.

16 species in the Americas; 5 in TX (mostly Trans-Pecos); 1 in our area.


1. P. hysterophorus L. False Ragweed, Santa Maria Feverfew, Cicutilla, Ragweed Parthenium. Taprooted annual 3 to 10 dm tall; stems single at the base and branching above, hirsute, striate. Lower leaves with subpetiolar base, in a basal rosette, upper leaves alternate, nearly sessile or with short subpetiolar bases, blades pinnatifid to bipinnatifid, passing to pinnately lobed or nearly entire upwards, hirsute, the pubescence denser beneath. Heads in an open panicle-like arrangement, each head 3 to 4 mm broad. Outer phyllaries oblong and pubescent, inner ones ovate and scarious-margined. Pappus of 2 white, petaloid scales, ovate-oblong, entire to notched. Achenes 2 to 3 mm long, oblanceolate-obovate, black, glabrous below and papillate at the apex. Abundant in disturbed areas in the Rio Grande Plains and SE. TX, less common northward and elsewhere in the state; adventive as a weed in warmer parts of U.S. and the Americas; perhaps not native to TX. Typically summer to fall, but our collections range from March to December.



ASTERACEAE SOLIVA12. SOLIVA R. & P. Burweed


Low, taprooted annuals, mostly 2 to 10 cm tall, rosette-like, acaulescent or with many short, sprawling stems. Leaves alternate but often crowded below the heads, bipinnatifid to tripinnatifid or dissected, the ultimate divisions lance-linear or deltoid-linear. Heads in the axils of the terminal leaves, but stems often branching just below the heads, putting the heads in the middle of the plant. Involucres hemispherical, 3 to 5 mm high. Phyllaries numerous, lanceolate, in 2 series, margins scarious. Receptacle convex, hairy. Flowers several to many per head; peripheral flowers corresponding to ray flowers, but the corollas absent, pistillate and fertile. Central flowers perfect but infertile, corolla minutely 4-toothed. Pappus absent. Achenes flattened perpendicular to the radius of the head, with 2 lateral wings or appendages, these spine-tipped; style persistent on the achene as another spine.

9 species, chiefly in S.A., widely dispersed by man. 3 species in TX and 2 infrequently encountered in our area.

Cabrera (1949) separated these plants into Gymnostyles and Soliva--an approach followed by Kartesz (1998). Recent treatments such as Cronquist (1980) and Gandhi and Thomas (1989) maintain our two species in Soliva. Our two species are so similar that I fail to see the logic in separating them.

The mature heads of spiny achenes can cause injury if stepped on or handled.


1. Achene with 2 lateral wings that are indented, creating upper and lower lobes,

unornamented; leaves usually tripinnatifid ...1.S.sessilis

1. Achene with 2 lateral appendages that are not lobed but which have cross-ribs; leaves pinnatifid to bipinnatifid ...2.S.stolonifera


1.S. sessilisASTERACEAE SOLIVA sessilis Ruiz & Pavòn Low plants, rooting at the lower nodes, forming mats to 20 cm broad. Herbage loosely hairy. Leaves small, to 3 cm long and 1.2 cm wide, tripinnatifid; petioles expanded beneath the heads. Heads usually less than 1 cm broad; phyllaries ovate to elliptic, 3 to 4 mm long; receptacle conic-elevated. Achenes (2.5)3 to 4 mm wide, the body scabrous-hispidulous, with broad wings having upper and lower lobes, these spine-tipped and flanking the spiny style. Local in E. and SE. TX; collected infrequently in our area; native to S. Amer. and adventive elsewhere--NC to FL, LA, AR, and TX. April-May. [S. pterosperma (Juss.) Less.]


2.S. stoloniferaASTERACEAE SOLIVA stolonifera (Brot.) Lour. Carpet Burweed. Low, rooting under the head-bearing rosettes. Leaves to 2.5 cm long and 0.7 cm broad, usually once or twice pinnatifid. Phyllaries generally 2 to 3 mm long. Achenes with wings unlobed but cross-ribbed and ornamented, the terminal spiny processes less than 0.5 mm long. Rare and localized in E. and SE. TX, collected infrequently in Brazos and Leon counties; Urug. and Argen.; adventive elsewhere--SC to FL and TX. Mar.-Apr. [Gymnostyles stolonifera (Brot.) Tutin; G. nasturtiifolia Juss., though some sources (e.g., Kartesz (1998)) say G. nasturtiifolia of authors, but not of Juss.].



ASTERACEAE COTULA13. COTULA L.


7 species of the S. hemisphere, one of which has been introduced into this country from Australia. Collected a few times in the 1970's on the Texas A & M campus, perhaps not persisting.


1. C. australis (Sieb.) Hook. f. Annual herb to about 15 cm tall, branched near the base; stems sparingly pubescent. Leaves 1 to 3 cm long, pinnately divided, the ultimate divisions linear-elliptic. Heads 2 to 5 mm broad, hemispherical; phyllaries broadly elliptic-ovate, margins scarious; receptacle more or less flat. Disk flowers yellow, corollas 4-toothed. Marginal achenes flattened, about 1 mm long, white-ribbed, with 2 sharp, smooth edges and glandular faces. Achenes stipitate, borne above the receptacle on minute, peg-like stalks (stipes). Superficially similar to Soliva, but differing by the stipitate achenes and lacking the spiny persistent styles. This plant is a bothersome lawn weed in Australia and is adventive in S.A., CA, and apparently TX and OK as well. Mar.-Apr.



ASTERACEAE PLUCHEA14. PLUCHEA Cass. Marsh-fleabane, Stinkweed


Aromatic, rather unpleasantly-scented annuals or perennials. Leaves alternate, simple, sessile to petiolate, ovate to narrowly lanceolate, usually crenate to serrate, variously pubescent. Heads in terminal corymb-like arrangements. Involucres campanulate to hemispheric. Phyllaries much overlapping, green to papery or colored. Receptacle flat, naked. Ray flowers

absent. Disk flowers of 2 kinds: a few perfect, infertile ones in the center of each head, corollas 5-lobed; the rest pistillate and fertile, corollas 3-lobed. Corollas tubular, pink to rose or purple or white. Pappus 1 series of fine, minutely barbed bristles. Achenes cylindric, 4-angled or 4-ribbed, glabrous to setose or hirtellous, brown, with a minute white enlargement at the base.

9 species in the Americas, with 4 in TX and 3 for us.


1. Leaves not narrowed into the midrib at the base; leaves sessile with clasping to rounded or truncate bases; corollas creamy-white ...1.P.foetida

1. Leaves narrowed into the midrib at the base; leaves sessile or petiolate, if sessile then neither clasping nor truncate; corollas pink ............................................................................2


2(1) Lateral branches of each inflorescence ascending to the height of the terminal flower cluster, the inflorescence thus flat-topped or layered in overall appearance; under high magnification, outer and middle phyllaries short-pubescent dorsally, usually only sparsely glandular ...2.P.odorata

2. Lateral flower-bearing branches not growing to equal the terminal flower cluster, the inflorescence thus not flat-topped but paniculate in overall appearance; only the outer phyllaries short pubescent and ciliate dorsally, conspicuously glandular under high

magnification ...3.P.camphorata

1.P. foetidaASTERACEAE PLUCHEA foetida (L.) DC. Stinking-fleabane. Perennial herb 5 to 8 dm tall; stem one from the base, branched above, glabrate at base and becoming more pubescent upwards. Leaves sessile, 3 to 10(13) cm long and 1 to 3 cm wide, lower leaves lanceolate to oblanceolate with truncate or cuneate bases, upper leaves proportionately broader--usually oblong-elliptic, bases usually auriculate and clasping but sometimes truncate, margin unevenly apiculate-serrate, pubescent below, especially on the veins, glabrous and glandular above. Heads about 8 mm high, grouped at the ends of the branches, the overall inflorescence tending to be paniculate or flat-topped. Phyllaries pubescent. Corollas creamy-white. Achenes about 1 mm long, pinkish, pubescent on the angles. Uncommon in wet places and muddy soils in E., SE., and S. coastal TX; SE. U.S.; also W.I. Summer to fall, our collections July-Oct. Some sources recognize varieties.


2.P. odorataASTERACEAE PLUCHEA odorata (L.) Cass Canela, Purple Pluchea. Annual to 15 dm, glabrate below passing to densely short-pubescent above. Leaves sessile or petiolate, short-ovate to ovate-lanceolate or elliptic, 1 to 15 cm long and 1 to 4 cm broad, margins entire to serrate or dentate, blades glabrous to tomentose. Terminal cyme maturing first and the lateral branches of inflorescence usually growing long enough to equal it, the entire inflorescence thus flat-topped or layered. Involucres 4 to 7 mm high; outer and middle phyllaries short-puberulent and ciliate dorsally, the inner phyllaries somewhat less so, all often pink-tinged, only sparsely gland-dotted; corollas pink. Achenes 1 to 1.3 mm long, pubescent on the angles. Throughout TX in muddy areas; our most abundant species. S. half of U.S. to S.A. and W.I. Summer-fall (ours July to Nov.) [Long known as P. purpurascens (Sw.) DC., a name applied in error as a result of reference to the wrong type specimen. P. purpurascens is actually a shrubby plant (Gillis 1977).] Some sources recognize varieties.

The common name means "cinnamon" in Spanish. This plant is used in herbal teas in Mexico (Ajilvsgi 1984).


3.P. camphorataASTERACEAE PLUCHEA camphorata (L.) DC. Camphor-weed. Annuals or perennials to about 15 dm; stems glabrate below passing to pubescent above. Leaves elliptic to elliptic-oblong, 6 to 15 cm long and 3 to 7 cm broad, margins essentially entire to denticulate-serrate or repand, lower leaf surfaces and veins of upper leaf surfaces minutely pubescent, both surfaces with widely-scattered resin globules. Heads in cyme-like aggregations, the lateral "cyme"-bearing branches not growing to equal the terminal "cyme", the inflorescence thus paniculate in over-all appearance and not flat-topped. Involucres 4 to 7 mm high, broadly campanulate at maturity. Phyllaries pinkish, only the outer ones short-pubescent and ciliate, with abundant small resin globules. Achenes pinkish to tan, 0,6 to 1 mm long, pubescent. Common in moist places. E., SE., and N. Cen. TX; SE. U.S. Summer-fall.


ASTERACEAE ERECHTITES15. ERECHTITES Raf. Fireweed, Burnweed


15 species in the Americas, Australia, and New Zealand; 2 species found in CA and the Pacific NW.; 1 species in the SE. and S. part of E. TX; also E. portion of S. U.S.


1. E. hieraciifolia (L.) Raf.ex DC.var. hieraciifolia. Annual from a short taproot with many adventitious roots above, prop roots sometimes also present; stems usually 1, scarcely if at all branched, to 1 m. tall, striate-ribbed. Leaves alternate, narrowly ovate-lanceolate to lanceolate, irregularly doubly serrate, the upper leaves sometimes nearly lobed, lower leaves with a subpetiolar base, upper leaves sessile to clasping. Inflorescence branched, heads on slim peduncles 2 to 18 cm long. Involucres cylindrical to turbinate or campanulate; phyllaries of 2 kinds: a short (1 to 3 mm) outer series of filiform bracts forming a calyx-like calyculum and 2 inner series, longer, (9 to 13 mm), linear, not much overlapping, margins scarious; receptacle flat, naked. Flowers of 2 kinds. Peripheral flowers pistillate and fertile; corollas yellowish-white, filiform/tubular, with 5 minute teeth, the 3 teeth nearest the outside of the head longer than the other 2. Central flowers many, perfect and fertile (though sometimes no seed is set), corolla flaring and evenly 5-toothed. Pappus of white capillary bristles 10 to 13 mm long. Achenes of both types of flowers columnar, 2 to 3 mm long, ribbed, brown. Fields, waste places, and disturbed or burned-over ground, Aug.-Nov. [Includes var. intermedia Fern.].



ASTERACEAE CONYZA16. CONYZA Less. Horseweed


Weedy, taprooted annuals, branched at the base or above. Leaves alternate, oblanceolate to linear, entire to dentate to pinnately or bipinnately lobed. Inflorescence various. Involucres campanulate to hemispheric. Phyllaries in 2 or 3 series, all alike, little if at all overlapping, herbaceous to scarious, usually pubescent on the outer surface. Receptacle flat but with a depression in the center. Ray florets in 1 to 3 peripheral rows, pistillate, fertile, corollas cream-white, the ligule minute and often reduced to a small lobe on the opening of the corolla tube. Disk florets perfect, fertile, corollas cream-colored, the limb 5-toothed. Pappus of capillary to stiff white bristles, all alike or the outer bristles shorter than the others. Achenes flattened and 2-ribbed (outermost achenes in the heads sometimes nearly 3-ribbed or trigonous), narrowly obovate in outline, usually pubescent.

50 species in the tropics and subtropics, world-wide; TX has 6; we have 2.


1. Ray florets in 2 to 3 rows around the periphery of the head; corollas slightly shorter than the pappus ...1.C.bonariensis

1. Ray florets in 1 row around the periphery of the head; corollas longer than the pappus ........

...2.C.canadensis

1.C. bonariensisASTERACEAE CONYZA bonariensis (L.) Cronq. Annual to 1 m. tall; stems glabrate below, becoming villous-ciliate upwards. Leaves narrow, 3 to 12 cm long and 1 to 25 mm wide, ascending, slightly grayish, lower leaves prominently lobed or toothed, those in the head-bearing region mostly entire. Head-bearing region of the plant cylindrical, round-topped. Heads ca. 7 mm high. Pappus of white to tan bristles, minutely antrorsely barbed. Achenes yellow to tan, 1 to 1.3 mm long, tapered and slightly flattened. Rare in E. TX, very rare in Brazos Co. around College Station; widespread in warmer parts of the Americas, N. to FL and TX. July. [Leptilon bonariense (L.) Small; L. linifolium (Willd.) Small; Conyza floribunda H.B.K.; Erigeron bonariensis L].

If varieties are recognized, our plants are var. bonariensis. Our material of this species is doubtfully identified.


2.C. canadensisASTERACEAE CONYZA canadensis (L.) Cronq. Horse-weed. Erect, robust annual 1 to 2 m. tall, simple at the base and branched in the head-bearing region, glabrous to pubescent. Leaves linear-oblanceolate, those in the upper part of the plant 4 to 10 cm long and 3 to 6 mm wide, often toothed, pubescent. Inflorescence cylindric. Heads 3 to 5 mm high; corollas white to lavender. Pappus of tan to white antrorsely barbed capillary bristles. Achenes yellowish, shiny, slightly flattened, tapering from base to apex, 1 to 1.2 mm long. Almost throughout TX; widespread in temperate N.A. Aug.-Nov. [Erigeron canadensis L.; Leptilon canadense (L.) Britt.]

This plant was used in a number of different medicinal preparations by Plains Indian tribes and by European settlers (Kindscher 1992).

There are 3 varieties in TX. We have 2 as follows:


var. canadensis Stems with spreading pubescence. Common in N. Cen. TX and Plains country; common in N. and Cen. U.S.


var. glabrata (Gray) Cronq. Stems nearly glabrous; phyllaries stramineous (straw-colored). More common in TX and the SW. U.S. than the variety above.



ASTERACEAE VERNONIA17. VERNONIA Schreb. Ironweed


Perennial herbs. Stems usually 1 from the base, simple or branched above. Leaves alternate. Inflorescence a terminal corymb-like arrangement. Involucres cylindric to campanulate (especially when mature). Phyllaries in several series, imbricate. Receptacle flat to convex, virtually naked. Ray flowers none. Disk flowers many per head, perfect and fertile, corollas pink to purple or white. Pappus of coarse, gray-white to rusty bristles in 2 lengths: the inner bristles longer and thinner than the outer, which are shorter, broader, and almost scale-like. Achenes with 6 to 10 ribs, often with resin globules between the ribs.

About 1,000 species world-wide, but less common in Europe. There is a great deal of interspecific hybridization, making determinations difficult at times. The TX material belongs to 7 species, of which we have 2. (See Jones and Faust 1978; King and Jones 1975; and Faust 1972).


1. Leaves linear-lanceolate; stems (excluding head-bearing region) glabrous to only slightly pubescent ...1.V.texana

1. Leaves elliptic or elliptic-lanceolate; stems pubescent to tomentose ...2.V.baldwinii

1.V. texanaASTERACEAE VERNONIA texana (Gray) Small Texas Ironweed. Stems erect, 4 to 8 dm tall, branched only in the upper region, glabrous to minutely puberulent. Leaves ascending, linear or lance-linear, 6 to 12 cm long and 1 to 2 cm wide, upper leaves much narrower than the lower, margins minutely revolute, entire to toothed, attenuate at the base, sessile, upper leaf surface scabrous, pubescent along the veins; lower leaf surface essentially glabrous or with short, straight, conical projections and perhaps some longer hairs along the veins. Involucres broadly campanulate, 5 to 6 mm high; phyllaries imbricate, purplish, puberulent or glabrous, the outer ones ovate, the inner ones oblong, all acute, erect or spreading. Flowers as described for the genus, ca. 18 to 20 per head. Achenes about 2.5 mm long, pubescent on the ribs with resin globules present between the ribs. Uncommon in E. TX, rare in SE. and N. Cen. TX; also AR and LA. Summer, ours primarily in June and July.


2.V. baldwiniiASTERACEAE VERNONIA baldwinii Torr. Western Ironweed, Baldwin Ironweed. Stems erect, 6 to 15(20) dm tall, single from the base and branching chiefly in the head-bearing region, puberulent to tomentose. Leaves elliptic to lanceolate to ovate, 8 to 15(20) cm long and 2 to 6 cm broad, acuminate, serrate, sessile, narrowed and acute toward the base, but not as attenuated as those of V. texana, upper leaf surface glabrous to slightly scabrous or puberulent, lower surface glabrous to tomentose, small resin globules present. Involucres narrowly campanulate/cylindric; phyllaries in several imbricate series, purplish, tips sometimes spreading, dotted with shiny resin globules when fresh, margins pubescent. Flowers usually 18 to 34 per head. Achenes ca. 3 mm long, with resin globules between the ribs. Common in N. Cen. TX, the Ed. Plat., and Plains country; infrequent to rare in E. TX; IL to MN, S. to LA and TX. Summer, ours from June to Aug.

Most or all of our material is referable to subsp. baldwinii, with 23 to 34 flowers per head, involucres 4.7 to 6.7 mm tall, the bract tips recurved, generally acute to acuminate, recurved tips of the inner bracts pubescent on the inner surface. The other subspecies, subsp. interior