Trees, shrubs, vines, and herbs. Secondary woody growth is common. Leaves usually with pinnate or palmate venation. Floral parts usually in 4's or 5's, rarely in 3's or 6's. Cotyledons two (only very rarely 1, 3, or 4), first leaves of germinating plantlet opposite.
Dicots represent approximately four-fifths of the flowering plants. There are nearly 11,000 genera and 190,000 species in over 300 families (according to Cronquist's classification system.) Texas boasts about 180 families, 1,300 genera, and 5,000 species of dicots (Hatch, et al. 1990). This key treats 104 families, 495 genera, and 1,113 species of dicots in Brazos and surrounding counties.
Trees (rarely shrubs or vines, never so ours). Bark bitter and aromatic. Leaf buds covered by membranous or tough stipules. Leaves deciduous or evergreen, simple, alternate, pinnately-veined. Flowers perfect, solitary or clustered, usually large and showy, often fragrant, regular, floral axis typically elongate. Sepals 3 or more, petals 3 to many, perianth segments often similar, deciduous, imbricate in bud; stamens many, linear, anthers not well differentiated from filaments, deciduous, spirally arranged. Gynoecium superior, carpels free (at least at maturity), spirally arranged and covering the long receptacle; each carpel with 1 or 2 ovules, style and stigma often somewhat poorly developed. Fruiting structure a woody, conelike aggregate of follicles or samaras.
This primitive family has 7 genera and 200 species; 3 genera and 4 species in TX; 2 genera, each with one species, possible in our area.
Many species in this family are cultivated as ornamentals (especially the two described below) or for timber (Mabberley 1987).
1. Leaves 4-lobed, mitten-shaped; carpels maturing into indehiscent samaras .........................
2. Leaves entire; carpels maturing into dehiscent follicles ......................................2. Magnolia
Two species, one in the SE. U.S., the other in China. In our area the trees are not native, but ornamentals are sometimes found abandoned or persisting.
1. L. tulipifera L. Deciduous, pyramidal tree to 60 m or more. Trunk straight, to 3 m in diam., bark gray. Leaves stipulate, long-petioled, to 20 cm long and about as wide, tulip- or mitten-shaped: broadly retuse-truncate apically and with 2 acute lobes on each side (occasionally with additional smaller lobes or entire), base cordate to truncate, glabrous, somewhat aromatic. Flowers solitary at the ends of the branches. Sepals 3, pale green, 3.5 to 6 cm long, soon reflexed; petals 6, forming a cup, 4 to 5 cm long, green-yellow, with a large orange blotch at the base of each; stamens many, anthers extrorse; receptacle elongate, persistent; carpels many, coherent in bud, separating at maturity into woody, flattened, deciduous samaras 2.5 to 4.5 cm long; fruiting structure overall 3 to 5 cm long. Rich woods. FL to LA, N. to DE, MI, MO; becoming naturalized in E. TX. Spring. Fall color yellow.
This handsome tree is valued for its ornamental flowers and leaves. The wood is used for furniture, being similar to birch in grain but not as hard (Elias 1980).
Characters as described for the family. Leaves entire. Sepals 3. Petals 6 to 12. Fruit an aggregate of follicles, seeds arillate, often pendant on long funicular threads.
About 125 species, primarily in Asia; 4 species native to TX; 1 that may be found in our area.
1. M. grandiflora L. Southern Magnolia, Bull Bay. Pyramidal tree to 30 m tall. Trunk straight, bark smooth. Branchlets, buds, and petioles rusty-pubescent. Leaves evergreen but falling a few at a time throughout the year, elliptic, 10 to 30 cm long, 4 to 15 cm wide, rounded to acute apically, cuneate at the base, glabrous and glossy above, rusty-tomentose below, very leathery; petioles to ca. 4 cm long. Flowers large, to 20 cm broad, cup-shape, fragrant, bud coverings leathery and densely rusty-pubescent. Sepals 3, petaloid; petals 6 to 9 or 12, obovate to spatulate, 5 to 10 cm long and about as wide; filaments sometimes purplish. Fruiting structure ovoid in outline, 6 to 10 cm long and 5 to 6 cm broad, an aggregate of follicles, pubescent; seeds 1 or 2 per follicle, red obovoid, somewhat flattened or angled, ca. 1 to 2 cm long, hanging by a funicular attachment. Low, rich woods, especially near streams. E. and SE. TX, usually cultivated in our area but apparently persisting; VA to NC, SC, and FL, W. to LA and TX. July-Aug.
Cultivated for its beautiful flowers and foliage and its dense shade. The wood is used for furniture and railroad ties (Elias 1980).
Aromatic, evergreen or deciduous trees (occasionally shrubs or vines). Leaves alternate, simple, estipulate, entire to lobed. Flowers regular, perfect or unisexual, appearing before the leaves in deciduous species, usually in axillary inflorescences, 3-merous, perigynous, perianth undifferentiated or else interpreted as apetalous, the segments usually 6 in 2 whorls. Stamens generally 12 in 4 series of 3 each, often 1 or more of the series reduced to staminodia or absent, anther dehiscence by 2 or 4 valves or flaps. Ovary 1-celled, with 1 pendulous ovule, style simple. Fruit a 1 seeded drupe or berry.
About 45 genera and 2,200 species in warm and tropical regions; 5 genera and 5 species in TX; 2 genera, each with one species, here.
This family provides many spices and flavorings (e.g. cinnamon, bay leaf). Some species have wood suitable for timber (Mabberley 1987).
1. Leaves all unlobed, evergreen; fruiting pedicel slender.......................................... 1. Persea
1. Leaves lobed or unlobed, deciduous; fruiting pedicel thickened below calyx.. 2. Sassafras
3. PERSEA Mill.
We have the one species found in Texas.
1. P. borbonia (L.) Spreng. Red Bay, Swamp Bay. Evergreen shrub or tree to 20 m tall or more, often smaller. Bark brown, tinged with red, purple, or gray, deeply fissured. Young twigs dark green and pubescent, later glabrous and light brown; winter buds densely rusty-pubescent. Leaves coriaceous, lanceolate to elliptic or elliptic-oblanceolate, 5 to 20 cm long, 2 to 8 cm broad, tapered to a grooved petiole to ca. 2.5 cm long, apically rounded to short-acuminate or acuminate, midvein impressed above, upper surface bright green, lustrous, lower surface glaucous, thinly tomentose to pubescent; leaves with a spicy-aromatic odor when crushed. Flowers perfect, in small, few-flowered, axillary panicles, peduncles to 7 cm long, usually 3 cm long or less, pedicels and peduncle usually pubescent. Calyx greenish or pale yellow, with 6 lobes in 2 whorls, the outer series widely ovate and puberulent, the inner members about twice as long as the outer but their upper 2/3 early-deciduous; remainder of calyx persistent. Fertile stamens 9, in 3 series, anthers 4-celled and 4-valved, the inner 3 stamens biglandular at the base and with introrse anthers; 3 staminodia commonly present inside the 3 fertile series. Pistil single, simple, with a slender style and discoid stigma. Fruit a blue-black, lustrous drupe, subglobose or slightly pointed; flesh dryish. Moist woods, swamps, watercourses an shores. SE. TX, in our area only in the far E. portion; MD and VA, S. To FL, W. To TX. Flowering Apr.-June; fruit ripening late summer-fall. [Includes forma pubescens (Pursh) Fern; P. palustris (Raf.) Sarg.; P. pubescens (Pursh) Sarg.].
The leaves can be used as a bayleaf-like seasoning (Tull 1987; Elias, 1980).
We have the 1 species found in E. N. Amer. The other 3 species are Asian.
1. S. albidum (Nutt.) Nees Sassafras, Filé. Dioecious, deciduous tree to 35 m tall, often smaller and shrub-like. Bark aromatic-spicy, trunk to 6 dm in diameter. Young twigs glaucous or nearly so. Leaves ovate to elliptic in overall outline, 6 to 12(18) cm long, to 10 cm wide, tapered to a petiole to 4 cm long, apex obtuse to acute, entire or with 1 or 2 lateral lobes (entire, right-, left- and 2-thumbed "mittens" occurring on the same tree), lower surface at first soft-pubescent or tomentose, usually becoming glabrescent. Flowers yellowish-green, appearing before or with the leaves, few to many in axillary panicles, subtended by scaly bracts, pedicels silky-pubescent. Perianth segments 6, ligulate to ovate-lanceolate, 3 to 5 mm long, spreading. Staminate flowers with 9 fertile stamens, inserted at the base of the perianth in 3 rows, the inner row with pair of stalked glands at the base of each filament (representing staminodia?), anthers 4-celled, 4-valved. Female flowers with 6 rudimentary stamens, ovary ovoid. Fruit a blue-black drupe, subglobose to ovoid, ca. 1 cm long, pedicel reddish, expanded at the apex, somewhat fleshy. Sandy woods, old fields, fence rows, and road cuts. E. 1/3 TX; N. Eng. W. to MI, IA, MO, KS, and AR, S. to FL and TX. Mar.-Apr. Fall color red, yellow, or orange. [Incl. var. molle (Raf.) Fern; S. officinale Nees & Eberm.; S. variifolium (Salisb.) O. Ktze.].
The dried leaves are used as a flavoring in gumbos and other Cajun cooking. The bark of the roots is used for tea, but recent evidence suggests it is carcinogenic in large amounts. The plant also has some medicinal, pest-control,and timber uses (Tull 1987; Elias 1980).
Perennial herbs, often of wet places, more or less aromatic, usually with rhizomes or stolons. Stems erect to ascending, jointed. Leaves alternate, simple, entire, usually petiolate, stipules adnate to the petiole. Flowers perfect, hypogynous to epigynous, in dense or lax spikes or racemes, inflorescence subtended by an involucre and sometimes resembling a single flower if the bracts are petaloid. Perianth absent. Stamens 4 to 8, free or adnate to the ovary at the base. Carpels 3 to 5, fused completely or only at the base. Fruit a compound capsule or a fleshy, capsule-like berry.
Five genera and 7 species of E. Asia and N. Amer.; 2 genera and 2 species in TX; we have only 1.
One species in N. Amer. and another in E. Asia.
1. S. cernuus L. Lizard's-tail. Colonial perennial from a fleshy or slender creeping rhizome; stems to 1 m tall, simple to few-branched, sometimes zigzag, often naked below and leafy above, pubescent. Leaves cauline, stipules indistinct, blades cordate-ovate, 5 to 12 cm long, 2 to 9 cm wide, longer than the petioles, acuminate at apex, the major veins arising at the base and converging toward the apex, lighter green below. Spikes or racemes terminal or at the upper nodes opposite the leaves, peduncled, erect, nodding or scorpioid, pubescent, 7 to 15 cm long and to 15 mm broad, showy basal bracts lacking. Flowers white, crowded, to ca. 300 in number, each with a small bract adnate to the minute pedicel. Stamens 4 to 8, filaments slender; carpels 3 to 5, united briefly at the base, stigmas recurved; ovules 1 or 2 per carpel. Fruit a compound capsule, somewhat fleshy, wrinkled. Muddy soil or water of lakes, swamps, and streams, often in dense shade. E. and SE. TX; SW. Que. and S. Ont., S. to FL, W. to MN, IL, MO, KS, and TX. (E. 1/3 N. Amer.)
Sometimes cultivated for the ornamental flower spikes; also an ingredient in some herbal medicines (Mabberly 1987).
Perennial herbs or vines, leaves basal or cauline, alternate, sessile to petiolate, estipulate, mostly cordate at the base and entire. Flowers perfect, regular or irregular, 3-merous, apetalous, the sepals fused below into a tube, the tube straight to curved, with flaring lobes, usually brightly colored. Stamens 6 to 12, anthers adherent or united with the style. Ovary inferior to superior, usually (4-)6-celled, syncarpous. Fruit a septicidal capsule or berry with numerous seeds.
About 400 species in 7 genera, mostly tropical. There is 1 genus with seven species in TX; 4 species in our area.
Perennial herbs or vines, leaves alternate, sessile to petiolate, palmately veined. Flowers perfect, irregular, solitary or in fascicles. Calyx tubular, corolla-like, S- or U-shaped and more or less 3-lobed apically or else straight and with a single terminal lobe on one side. Stamens 6, anthers sessile and adnate to the short, stout, 3- to 6-lobed style. Ovary partially inferior to superior. Fruit a 6-lobed or 6-angled capsule opening from the base or apex; seeds many, horizontal.
About 300 species of the tropics; 7 from TX; 4 here. This treatment is based, in part on the work of Pfeifer (1966).
Many species are cultivated as ornamentals or for their medicinal properties--uses range from treating snakebite to easing childbirth. Many species are poisonous. Other species are the sole food source for the larvae of certain butterflies. The flowers trap flies for pollination (Mabberley 1987).
1. Plants erect herbs; flowers from the lowermost nodes; capsule dehiscing from the apex with thevalves spreading. .........................................................................................................2
1. Plants vines, vinelike, or spreading or prostrate; flowers from the upper nodes; capsule dehiscing from the base or irregularly ......................................................................................3
2(1) Leaves sessile, firm, reticulate below, oblong to oval or ovate ......................1. A. reticulata
2. Leaves petioled, thin, ovate to lanceolate; cordate to hastate basally ........2. A. serpentaria
3(1) Plants twining or climbing; leaves broadly cordate .......................................3. A. tomentosa
3. Plants sprawling or spreading, not twining; leaves linear to narrowly lanceolate ....................
................................................................................................................................4. A. erecta
1. A. reticulata Jacq. Texas Dutchman's-pipe. Perennial herb from a slender, fibrous-rooted rootstock. Stems to 40 cm tall, 1 to (usually) several, erect to decumbent, angled, simple or branched near the base, zigzag, hirsute with spreading hairs. Leaves very short-petiolate and nearly clasping, ovate to oblong, to 12 cm long and 6.5 cm wide, firm, obtuse to nearly acute apically, margins often undulate, lower surfaces conspicuously reticulate-veined, generally pubescent along the veins of both surfaces and along the margins. Flowers borne in several-flowered, bracted racemes arising from the lowermost nodes, often hidden by leaf litter, bracts ovate to suborbicular; pedicels slender, to ca. 1.5 cm long. Perianth 8 to 12 mm long, with a somewhat gibbous lower portion and an expanded, upturned limb to about 8 mm broad, dark brown, pubescent; anthers shorter than the 3-lobed style. Capsule reddish-brown,subglobose, 6-angled, to ca. 15 mm in diameter, dehiscent from the apex. Seeds ovate-cordate in outline, ca. 5 mm long, deeply concave with small, irregular tubercles on the rounded face. Humus soils (ours apparently from somewhat sandy soils) in pine-hardwood forests, pine savannahs, and rocky woodlands. E. TX, W. LA, and SW. AR. May-July. [author sometimes given as Nutt.].
2. A. serpentaria L. Virginia Dutchman's Pipe, VA. Snakeroot. Herbaceous perennial to 0.6 m tall, erect, from a knobby rhizome. Leaves petiolate, ovate to oblong or lanceolate, base cordate to sagittate or hastate, apically acuminate, to 15 cm long, lowermost leaves scale-like. Flowers solitary at the lowest nodes, on stalks to 8 cm long; perianth 1 to 2 cm long, purple to brown, curved, limb wide-spreading, 3-lobed. Capsule ellipsoid to subglobose, to 13 mm in diameter. Seeds obovoid, 4 to 5 mm long, minutely papillate. Upland woods and rocky, wooded slopes, and ledges. Ed. Plat. and E. TX; known from Madison Co. NY and CT, S. to FL, W to IL, OK, and TX. Apr.-June. [Incl. var. hastata (Nutt.) Duchatre; A. hastata Nutt.
3. A. tomentosa Sims Woolly Pipevine, Pipevine, Woolly Dutchman's-pipe. High-climbing, twining, woody vine to 25 m or more long, more or less densely white-pubescent throughout, especially on the younger portions. Leaves petiolate, the petioles of the largest leaves shorter than the blades, blades ovate to suborbicular or reniform, basally cordate (rarely truncate), apically obtuse to rounded, to 20 cm long and about as wide, commonly smaller, densely pubescent below, more sparsely so above, becoming less pubescent with age. Peduncles solitary or paired at the nodes, downy-pubescent, bractless. Perianth to about 4(7) cm long, densely beige-tawny tomentose, J-shaped, the orifice dark purple; limb 3-lobed, reflexed, rugose, greenish-yellow. Ovary inferior; capsules cylindrical, ca. 4 to 6(8) cm long and 3 cm broad, strongly ribbed, pubescent; seeds flat, triangular, ca. 1 cm long, grayish. In shade along streams and rivers, in bottomland woods and post oak woods. E. Cen. and N. Cen. TX; FL to TX, N. to OK, KS, MO, IL, and NC. Mar.-May. [Isotrema tomentosa (Sims) H. Huber].
4. A. erecta L. Swan-flower. Plants low, ascending to spreading or commonly prostrate, to ca 30 cm across, from a long, aromatic rootstock. Stems elongate, slender, with short internodes, pubescent. Leaves linear-elongate to setaceous, a few usually oblong-lanceolate, acute at the base, acute to acuminate apically, to 13 cm long and 1.5 cm broad, slightly pubescent above and nearly glabrous below. Peduncles short, tomentose. Perianth elongated, to 10 cm long and 1 cm wide; tube mauve, curved, to ca. 3 cm long, the throat grayish; limb purple-brown, to 7 cm long, limb one-sided, linear to narrowly lanceolate, the apex narrow and acute to obtuse. Capsule ca. 2 cm long, obovoid to nearly 4-angled, glabrous; seeds dark brown, triangular, ca. 5 mm long. Hard-packed sandy silt and sandy loam in open, grassy areas or in the shade. S. TX and adjacent Mex. Mar.-Nov. [A. longiflora Engelm. & Gray].
This is an inconspicuous and often-overlooked plant. It is a main food of the Pipevine Swallowtail butterfly larva. It is often easiest to find these plants by looking for the beautiful black and blue butterflies or the long-horned black and red caterpillars (these often eat most of the foliage).
Aquatic, rhizomatous herbs, vessels present only in the roots. Leaves peltate, without a basal sinus, usually held above the water on long petioles, apparently alternate, but each node also with a scale leaf on the lower side of the rhizome and another next to the foliage leaf, wrapped around the petiole base; branches axillary. Flowers solitary from the axils of the upper-side scale leaves, emergent. Perianth of ca. 20 to 30 parts, spirally arranged, the outer 2 members more or less sepal-like, the remainder more or less petaloid, usually in 2 series. Stamens many, spirally ararnged, filaments slender, anthers introrse, anther halves separated by a narrow, leaf-like connective. Ovary superior, with 12 to 40 distinct carpels sunken into the spongy receptacle, each carpel with 1 large ovule. Fruting structure consisting of the dry receptacle with separate, hard-coated, 1-seeded nuts loose in its cavities. Cotyledons 2; perisperm none.
One genus with 2 species.
1. NELUMBO Adans. Lotus, Sacred Bean
One species in each in the Old and New Worlds.
1. N. lutea Willd. Yellow Lotus, Water-chinquapin, Pond-nuts. Rhizomatous perennial rooting in mud; rhizomes slender or thicker and with banana-shaped lateral tubers. Leaves long-petioled, the blades floating or held well out of the water, blades orbicular, centrally-peltate, without a sinus, to 70 cm in diameter, the center depressed or cupped, margin sometimes undulate, the upper surface glaucous, persistently unwettable (water beads and runs off). Flowers solitary on stout peduncles, borne to about 1 m above the surface, showy, to ca. 25 cm across, pale yellow, hypogynous. Perianth segments many, intergrading, 20 or more total, ovate, obtuse to rounded, 2 to 12 cm long, the outermost green and sepaloid; stamens many, introrse, spirally inserted, anthers 1 to 2 cm long, with a terminal, hooked appendage, filaments elongate; receptacle obconic, to 10 cm across, pistils several to many, separate, sunken into pits in the receptacle, styles stout, stigmas capitate. Fruits ovoid, ca. 1 cm in diameter, each single-seeded, nutlike, hard-shelled and indehiscent, embedded but rattling loose in the receptacle. Still or sluggish water of ponds and streams. E. 1/3 TX; FL to TX, N. to MN, MA, Ont., W. to IA, NE. May-July.
The tubers and seeds are edible and were a major food of the plains Indians (Kindscher 1987). They may have extended the original range through cultivation. The dry receptacles and fruits are often used in floral arrangements.
Aquatic perennial herbs from horizontal rhizomes. Floating or emersed leaves cordate, with a basal notch, involute in bud. Flowers solitary, axillary, perfect. Calyx of 4 to 6(14) green to petaloid sepals, distinct to slightly united. Petals in ours 3 to many, usually showy, sometimes intergrading with the stamens. Stamens many, commonly introrse, sometimes grading from petaloid to well-developed. Gynoecium superior to inferior, pistils 5 to many, united at least basally, each with many ovules. Fruit a many-seeded berry, irregularly dehiscent; seeds arillate, cotyledon 1 or 2.
About 60 species in 6 genera; 2 genera and 4 species are listed for TX; there are 2 genera and 2 species here.
Almost all members of the family have some value as ornamentals.
1. Perianth of 4 sepals and numerous broad, showy petals; carpels sunken into the receptacle or ovary inferior ....................................................................................................1. Nymphaea
1. Perianth of 6 sepals and numerous narrow, stamen-like or scale-like petals; carpels not sunken into the receptacle .......................................................................................2. Nuphar
Rhizomatous herbs, rhizomes sometimes with lateral tubers. Leaves floating or just above or below the water surface, sub-peltate, more or less orbicular, with a sinus at the base extending to the petiole. Flowers solitary at or above the surface, usually opening in the morning and closing in the afternoon. Sepals 4, green, nearly completely distinct, spreading. Petals few to many, spreading, white, pink, blue, or yellow, the outer about equalling the sepals, the inner intergrading with the stamens. Stamens many, the outer petaloid; filaments broad, inserted on the sides of the ovary. Pistil compound, superior to nearly inferior, 12- to 35-celled, the top concave with a globose protrusion in the center from which the stigmas radiate, stigmas extended beyond the ovary and forming incurved sterile appendages. Fruit depressed-globose, berrylike, covered with the perianth bases, maturing underwater; ovules many, laminar, enveloped by sac-like arils. Usually growing in still, shallow water.
This largely-tropical genus has about 50 species; 3 species from TX; only 1 here. Many species are cultivated for the showy flowers; some have edible seeds (Mabberley 1987).
1. N. odorata Ait. subsp. odorata American Waterlily, White Waterlily, Fragrant Waterlily, Pondlily, Ninfa Acuática, Alligator-bonnet. Rhizomes horizontal, elongate, straight, generally 2.5 to 3 cm thick, without lateral tubers. Leaves arising from along the rhizome, blades suborbicular, to 25(30) cm broad, the sinus narrow, margins sometimes undulate, surface tinged reddish or purplish below. Flowers floating, very fragrant, to 12 cm broad. Sepals often purplish on back, elliptic to ovate or lanceolate, to 8 cm long and 2.5 cm broad; petals usually more than 25, white (rarely pink), thick, elliptic or somewhat broadened upward, nearly acute apically, 2 to 10 cm long; stamens numerous (generally more than 70), the petaloid outer ones 3 to 4 cm long; styles about 20. Berry generally 2.5 to 3 cm in diameter, the ellipsoid seeds ca. 2 mm long. Lakes, ponds, streams, and ditches in SE. TX; FL to TX, N. and E. to Newf., Man. and G.P. Mar.-Oct. [Includes var. villosa Casp. and var. gigantea Tricker; N. spiralis Raf.; N. lekophylla (Small) Cory; Castalia odorata (Ait.) Woodv. & Wood; C. lekophylla Small].
This is one of the species with edible seeds (Tull 1987).
20 species of temperate and cold regions. We have the 1 TX species.
1. N. lutea (L.) Sm. subsp. advena Kartesz & Gandhi Yellow Cowlily, Spatterdock, Yellow Pondlily. Perennial herb from procumbent, branched, cylindrical rhizomes. Leaves spirally arranged, petioles and peduncles with many small air cavities, petioles terete to flattened above, glabrous to pubescent. Leaf blades floating or emersed (in this subsp.), broadly ovate to suborbicular, to 30 cm long and 25 cm broad, with a deep basal sinus extending to the petiole in the center, basal lobes overlapping to divergent, glabrous to somewhat pubescent beneath; submersed leaves, if present, similar to emersed leaves but thin, translucent. Flowers solitary, at or just above the surface, sometimes fragrant, to 4.5 cm across and 2.5 cm high, hypogynous. Sepals usually 6, to ca. 2 cm long, rounded, concave, somewhat petaloid, the inner surface green to yellow or sometimes reddish; stamens many, the outer ones staminodial (representing petals but not petaloid), small, thickened, less than 1 cm long, yellow or reddish, oblong, emarginate; anthers 3 to 7 mm long, recurved, shorter than the stigmas, persistent; ovary compound, stigma with 5 to 25 rays, disklike. Fruit broadly ovoid, fleshy, slightly constricted below the depressed, sometimes reddish stigmatic disk, 3 to 5 cm broad, the stigma rays generally ending 1 to 2 mm from the margin of the disk, fruit irregularly circumscissile near the base; seeds many, broadly ovoid, 4 to 6 mm long, 3 to 5 mm wide, yellow to brown, smooth. Water or stranded on mud of ponds, lakes, streams, and springs. E. TX and on the Ed. Plat; the species throughout most of E. N. Amer.; also Mex., Cuba. Mar.-Oct. [Includes subsp. macrophyllum (Small) E. O. Beal and subsp. ozarkana (MIll. & Standl.) E. O. Beal; Nuphar advena (Ait.) Ait. f. and var. tomentosa Nutt.; Nuphar microcarpum (Mill. & Standl.) Standl.; Nymphaea microcarpa Mill. & Standl.; etc.].
Aquatic herbs, rhizomatous and with stems, vessels none. Leaves of two types, in Cabomba the submersed leaves deeply dissected and the floating leaves entire and linear; in Brasenia alternate, long-petioled, and ovate or elliptic, cordate to peltate; stipules none. Flowers solitary. Perianth of (2)3 to 4 segments in each of 2 whorls. Stamens 3 to 6 in Cabomba, 12 to 18 in Brasenia. Ovary superior, of (1)2 to 18 free carpels, stigmas terminal or decurrent, ovules (1) 2 or 3, attached near the dorsal suture. Fruit a collection of leathery, nutlike, indehiscent follicles. Embryo with 2 cotyledons.
There are 2 genera and 8 species in tropical and warm regions; 2 genera, each with one species in TX; 1 species here. However, Cabomba caroliniana Gray is found in E. TX and may eventually be found here--it has stems covered with gelatinous mucilage, submerged opposite or whorled leaves which are palmately dissected, few emergent leaves, and white flowers to about 12 mm long.
A monotypic genus.
1. B. schreberi J. F. Gmel. Purple Wen-dock, Water-shield. Stolons slender, rooting in mud. Stems slender, coated with a gelatinous mucilage. Leaves long-petioled, alternate, blades floating, elliptic to oval or suborbicular, rounded at both ends, centrally peltate, without a sinus, to 10(12) cm long and ca. 8 cm wide, the undersurface and petioles gelatinous. Peduncles axillary, flowers borne just above the water, small, to 2 cm broad, dull purple or reddish purple, hypogynous. Sepals and petals each (2)3 to 4, similar, linear to linear-oblong, 1 to 1.5 cm long, petals perhaps slightly longer and narrower than the sepals; stamens 12 to 36, filaments slender, anthers latrorse-extrorse; pistils 4 to 18, free, styles short, stigmas linear, ovules 1 to 4 per pistil, pendulous and parietal. Fruits indehiscent, clavate to ovoid, ellipsoid, or obovoid, beaked, (5)6 to 8(9) mm long including the style, 2.5 to 3 mm broad, each usually with 1 or 2 seeds. Ponds, lakes, and slow streams in E. TX; FL to TX, N. to Que., Ont., and MN, W. to B.C. and OR--most of N. Amer. except the G.P. and SW.; sporadic on all continents save Eur. Apr.-May in our area, collected with fruit as late as June.
A widespread family with 1 genus (2 if Podostemum is included rather than given its own family).
Submersed aquatic perennial herbs. Stems branched, the branches 1 per node, rootless, but lateral branches sometimes modified into anchoring rhizoids. Stems breaking easily and the pieces developing into new plants. Leaves whorled, sessile, generally 1 to 4 times dichotomously dissected into linear or filiform segments. Plants monoecious, flowers small, solitary and sessile in the axil of one leaf at a given node, each flower subtended by an 8- to 12-cleft involucre (perhaps representing the perianth?); perianth absent. Staminate flowers generally with ca. 4 to 16 stamens, filaments short and anthers with the connective ending in 2 or 3 sharp points or bristles. Pistillate flowers with 1 superior, unilocular, 1-seeded pistil. Fruit a warty or spiny to beaked or spineless achene, topped with the hardened style.
2 to 30 species, depending upon interpretation; 2 in TX; 1 here.
The plants provide some food and cover for fish, but also for mosquitoes and other pests (Mabberley 1987). Unequivocal identifications are often difficult without the achenes.
1. C. demersum L. Common Hornwort. Characters as described for the family. Stems to 3 m long, brittle to flexuous, often forming large masses. Lowermost leaves of the seedling simple, leaves of the main stem (3)9 to 12 per node, each usually forked 1 to 2(3) times, the segments capillary to linear, flattened, serrate along one margin, variable in length and width, but to about 1.5 to 2 cm long. Bracts of staminate flowers 1 mm long, stamens commonly more than 15, anthers 1 to 1.1 mm long, sometimes dotted with red glands. Bracts of pistillate flowers 1.2 to 2 mm long. Achene ellipsoid, compressed, 4 to 6 mm long, with 2 basal spines 2 to 5 mm long and a persistent style 4 to 6 mm long. Still water, lakes, ponds, and slow streams throughout TX, primarily in the E.; Que. to B.C., S. to Mex.; also Old World. Summer.
NOTE: C. muricatum Cham. (C. echinatum Gray) is found to our E. and may someday be found here. The leaves are usually forked 2 to 4 times, the divisions entire or serrulate along both margins; achenes have 3 to 5 lateral (not basal) spines.
Perennial (sometimes annual) herbs or occasionally herbaceous or woody vines or subshrubs. Leaves estipulate or with minute stipules, commonly basal, if cauline then alternate (in a few genera opposite or whorled), often with sheathing bases, entire to toothed, lobed, or compound. Flowers mostly regular, some irregular and hooded or spurred, solitary or in racemes or panicles, generally perfect, sometimes unisexual, most often with all parts free. Sepals usually imbricate, varying from small and caducous to petaloid. Petals present or absent, variable in number, often with nectaries. Stamens usually many, sometimes petaloid. Carpels superior, 1 to many, simple, free, style and stigma one, ovules 1 to many per pistil. Fruit a cluster of achenes, follicles, or berries (rarely pistils fused and fruit capsular--never so in ours); seeds with endosperm.
There are about 58 genera and 1,750 species in temperate and boreal regions; 11 genera and about 50 species in TX; 8 genera and 19 species here.
Many species have poisonous and/or useful alkaloids (Belladonna, Aconitum, etc.). Many, such as are grown for ornament--Buttercup (Ranunculus), Larkspur (Delphinium), Monkshood (Aconitum), Columbine (Aquilegia), Windflower (Anemone), and many more (Mabberley 1987).
1. Plants vines or rambling plants, herbaceous or woody ........................................1. Clematis
1. Plants herbs, not vining or rambling .........................................................................................2
2(1) Leaves linear and entire, all basal; cluster of achenes slender, 8 to 30 times longer than broad .....................................................................................................................2. Myosurus
2. Leaves not as above; cluster of achenes or follicles globular or cylindric, less than 8 times longer than broad .....................................................................................................................3
3(2) Flowers spurred, zygomorphic .................................................................................................4
3. Flowers not spurred, regular ....................................................................................................5
4(3) Carpel 1; petals united; leaf segments linear-filiform; plants annual ................3. Consolida
4. Carpels in 3's; petals free; leaf segments broader; plants perennial ..............4. Delphinium
5(3) Perianth whorls 2 at anthesis, clearly distinguishable as sepals and petals ..........................6
5. Perianth whorls only 1 (sepals) at anthesis OR two whorls present but not differentiated into petals and sepals ..............................................................................................................7
6(5) Corolla red to orange ................................................................................................5. Adonis
6. Corolla yellow to white ......................................................................................6. Ranunculus
7(6) Leaves alternate, mostly or largely cauline; sepals minute and falling early, not showy .........
7. Leaves mostly basal; sepals larger, persisting longer, petaloid andshowy ........8. Anemone
Herbaceous (sometimes slightly woody) vines climbing by twining petiolules, or herbs. Leaves petiolate to sessile, opposite, simple or pinnately compound. Flowers perfect or plants dioecious, flowers axillary or terminal, solitary or in panicles, inconspicuous to showy, often nodding. Sepals 4(5 to 6), separate to slightly joined at the base, thin to leathery, white or variously colored, petaloid, valvate in bud, spreading to cupped. Petals absent or minute and passing into the stamens. Stamens many, the outer staminodial or all modified into staminodia in pistillate flowers of dioecious species. Pistils many, maturing into achenes, styles persistent, often long and pubescent to plumose or sometimes nearly naked.
About 250 species in temperate and subtropical zones; 11 species in TX; 4 here. Synonyms for parts of the genus include Viorna and Atragene.
Many species are cultivated for ornament, and some have been bred with enormous flowers (Mabberley 1987). These do not do very well in our area--our summers are too hot and Clematis prefer a cool-root run.
1. Flowers in cymose panicles; sepals mostly less than 15 mm long, whitish .............................
..... .................................................................................................................1. C. drummondii
1. Flowers solitary or grouped on long peduncles; sepals usually longer than 15 mm, variously colored .......................................................................................................................2
2(1) Leaflets with 3 conspicuous veins from the base, but not conspicuously reticulate, thin; sepal margins widely expanded (more than 2.5 mm ) above the middle and crisped, ruffled, or undulate, sepals blue-purple ................................................................2. C. crispa
2. Leaflets fairly obviously reticulate-veined, somewhat thick; sepals not expanded above the middle or with a margin to 2.5 mm wide, rose-purple to violet or lavender ............................3
3(2) Achene beak strongly plumose with spreading hairs; leaf blades strongly reticulate, even
the quaternary veins rasied below, and ultimate closed areoles less than 2 mm in the
longest dimension (usually less than 1 mm) ..................................................3. C. reticulata
3. Achene beak silky to sparsely pubescent; leaf blades with primary and secondary veins
raised below but tertiary and quaternary veins not or only slighty raised, with ultimate
close areoles more than 2 mm long..................................................... ..............4. C. pitcheri
1. C. drummondii T. & G. Texas Virgin's Bower, Barbas de Chivato, Old Man's Beard. Foliage cinereous-pubescent. Leaves mostly with 5 to 7 leaflets, the uppermost ones simple but 3-cleft; leaflets 1.2 to 2.5 cm long, divergently 3-lobed or sometimes parted, primary lobes ovate-oblong to lanceolate or rhombic-ovate, acute to acuminate, entire or with 1 to 3 acute-acuminate teeth. Panicles few-flowered or sometimes flowers solitary with a pair of leaf-like bracts at the base, usually peduncles trichotomous and with small bracts on the lateral pedicels. Sepals greenish-white, ca. 12 mm long, oblong-lanceolate, sericeous on the outer surface. Fruiting flowers with many sterile filaments as long as the sepals. Achene tails plumose, very slender, eventually to 7.5 to 10 cm long; achene bodies to about 5 mm long, flattened, with 1 straight margin and 1 curved. Dryish soils, dry creek beds, and rocky canyons, often climbing over shrubs and trees. Cen., S., and W. TX; TX to AZ and Mex. Apr.-Sept. Known from "near Old Independence," this presumably near Independence in Washington Co.
2. C. crispa L. Blue Jasmine, Curly Clematis, Leather-flower. Freely-climbing or weakly-ascending herbaceous vine, often flowering early when only ca. 30 cm tall. Stems many-angled, glabrous or nearly so. Petioles 1 to 6 cm long. Leaves pinnately compound, with 2 to 5 pairs of leaflets, leaflets petiolulate, ovate to lanceolate or linear, 1.5 to 10 cm long and 0.3 to 5 cm broad, apically obtuse, acute, or apiculate or acuminate, bases cuneate to cordate, entire (to rarely cleft in 3), membranaceous, usually with 3 primary veins noticeable beneath and not strongly reticulate. Peduncles solitary and terminal on each branch, arranged above a pair of compound (or rarely simple) leaves. Calyx cylindric-campanulate or long-urceolate-campanulate; sepals bluish, varying from rosy to violet, 2.5 to 5 cm long, recurved or spreading above the middle, this spreading portion with a thin, broad, crisped or undulating margin several mm broad, apex acuminate. Achene bodies 6 to 9 mm long, suborbicular to rhomboid, slightly sericeous, style canescent to villous or pilose in flower, in fruit 1 to 3 cm long and either finely appressed-pubescent or glabrate. Wet soils, climbing over shrubs, along streams, and in low woodlands or on dry sandhills if water is available. E. TX W. to Calhoun and Williamson Cos.; FL to TX, N. to SE. VA, IL, and MO. Mar.-Oct. [C. cylindrica Sims; Viorna crispa (L.) Small; V. obliqua Small].
3. C. reticulata Walt. Netleaf Clematis. Herbaceous vine, sometimes woody at the base. Stems climbing, to 4 m long, 6-angled, glabrous to sparsely pilose, especially at the nodes. Cauline leaves with 7 to 9 leaflets, these long-petioluled, usually more or less elliptical, apically rounded, and mucronate but varying to ovate and acute, 1 to 9 cm long, 0.5 to 5(7.5) cm broad, entire or 1- to 3-lobed or the proximal ones sometimes 3-foliolate, decidedly leathery and strongly reticulate, even the tertiary and quaternary veins raised below and ultimate closed areoles less than 2 mm (usually less than 1 mm in the longest dimension, tip of rachis often tendril-like. Peduncles axillary, 1- to 3-flowered, bracts usually less than 4 cm long, located ca. 1/3 the distance from the base of the peduncle. Flowers nodding, urn-shaped, 15 to 25 mm long. Sepals equalling or slightly longer than the sepals, ovate-lanceolate, 1.2 to 3 cm long, leathery, recurved, purple-red to mauve or pinkish-lavender, the tips sometimes greenish, more or less densely yellow-candescent externally, margins densely tomentose, usually not expanded or with a border less than 2.5 mm broad. Head of achenes globose. Achene bodies suborbicular, ca. 4 mm broad, with a marked rim, appressed-pubescent; achene tails 4 to 6 cm long, plumose, pale yellow-brown, frequently broken off short. Sandy soils of woods in E. TX; we are on the W edge of the range; SC and FL, W. to TX, N. to AR. Apr.-July. [Incl. C. subreticulata Harbison ex Small; Viorna reticulata (Walt.) Small].
4. C. pitcheri T. & G. var. pitcheri Pitcher Clematis, Leather-flower, Bluebell, Pitcher-flower. Herbaceous vine, sometimes somewhat woody at base. Stems freely climbing, to several m long, simple or somewhat branched, 6-ribbed or -angled, reddish-brown, sparingly pubescent to glabrous. Cauline leaves with 3 to 5(11) leaflets, these variable in shape, simple to 2- to 5-lobed or 3-parted, to 3 to 9 cm long and 2.5 to 5 cm broad (broader if lobed), ultimate divisions ovate to elliptic to ovate-cordate, obtuse to acute, mucronate, stiffish or leathery, more or less reticulate-veined but the tertiary and quaternary veins scarcely or not at all raised beneath, the ultimate closed areoles more than 2 mm in the longest dimension, smaller leaflets and tip of rachis often tendril-like. Peduncles axillary, usually with a pair of simple bracts near the base, 1- to 7-flowered. Flowers nodding, somewhat urn-shaped to ovoid, generally less than 4 cm wide. Sepals from slightly longer than the stamens to twice as long, less than 3.5 cm long, slightly connivent at the base, leathery, rose-purplish to violet or brick red outside, colored or green inside, recurved above the middle, pubescent, margins not expanded or with a border less than 2.5 mm broad, not crisped or ruffled, white tomentose. Head of achenes globose, ca. 1.6 to 2.8 cm broad (minus the tails.) Achene bodies orbicular to suborbicular to somewhat quadrangular, rather inequilateral, 6 to 8 mm broad, with a thick margin and appressed pubescence; achene tails 1.5 to 3 cm long, silky near the base and appressed-pubescent above or sometimes glabrate or villous, never plumose, slender, tapered above and often contorted or curved, frequently broken off short. Climbing on other plants or sprawling along the ground. Thickets, open woods, along streams, and in lowlands. Primarily N. Cen. TX, Ed. Plat., and along the coast; IN to NE, S. to MO and TX. Apr.-Sept. [C. simsii and Viorna simsii of various authors; V. pitcheri (T. & G.) Small or (T. & G.) Britt.].
Small annual herbs from fibrous roots. Leaves in a basal tuft, linear-filiform or spatulate, entire. Flowers regular, solitary on scapes, yellowish or whitish. Calyx of 5 sepals spurred at the base. Corolla of 5 small, narrow petals, these clawed and with a small, nectariferous pit at the apex of the claw. Stamens 5 to 20. Pistils many, fruiting structure a slender spike (occasionally an oblong head in depauperate species) of many achenes, these somewhat 3- to 4-sided, crowded on the long, slender receptacle.
About 15 species in temperate regions; 1 known from TX; others perhaps to be found.
1. M. minimus L. Mousetail, Tiny Mousetail. Characters as described for the genus. Foliage 2 to 5 cm tall. Leaves narrowly linear to filiform, 1 to 10 cm long, 0.3 to 2.2 mm broad at the widest, equalling or shorter than the scapes, apically blunt, somewhat narrowed toward the base. Scapes 2 to 10 per plant, ca. 3 to 15 cm tall. Sepals oblong, (1.3)2 to 3.5(5.5) mm long, the acute, membranous spurs (0.7)1 to 3 mm long; petals linear to narrowly spatulate, 2 to 3.5 mm long, about equalling the sepal blades, often caducous and sometimes absent entirely; stamens (5)10(20). Fruiting receptacle (1.5)2 to 5 cm long and (1.5)2 to 3 mm thick, usually with many achenes. Achenes fitting closely together, somewhat quadrate-rhombic in face view, with a central stylar beak protruding from a short keel, rectangular or trapezoidal in side view, overall 0.9 to 2 mm long with beaks 0.5 mm long or shorter. Damp clay or calcareous soils, fallow fields; throughout TX but primarily the central region; FL to TX, N. to E. VA and S. Ont., W. to IL, MN, Sask., CA and B.C.; also Eurasia.
Naturalized annuals, stems simple to branched, leaves petiolate to sessile, pinnately or palmately divided, segments many, linear to filiform. Flowers in terminal, leafy-bracted or naked racemes, white to blue, violet, or pink (petals and sepals both colored), with a horizontal sepal spur, the other sepals 2 lateral and 2 lower; petals united. Gynoecium of a single carpel, maturing into a follicle.
About 40 species of the Mediterranean region and Europe; 2 species naturalized in TX; 1 here. Some or all of the species are sometimes referred to the genus Delphinium.
The flowers are said to have been used to garland mummies in ancient Egypt (Mabberley 1987).
1. C. ajacis (L.) Schur Rocket Larkspur, Espuella de Caballero. Erect, slender, branched annual to ca. 1 m tall. Stems crisp-puberulent, glandular at least in the upper portion. Leaves several to many, short-petiolate to sessile, pinnately or palmately dissected into numerous linear-filiform segments, usually pubescent. Terminal racemes spiciform, loosely 3- to 19-flowered, leafy-bracted or naked, lowermost bracts filiform-dissected, not simple. Flowers blue to violet, purple, pink, or white. Sepals 5, the uppermost extended backwards into a spur 1 to 2 cm long; lateral sepals rhombic to ovate to suborbicular, 6 to 11 mm wide; lower sepals 8 to 15 mm long; petals 2, united into a 4-lobed structure arching over the stamens and carpels and extending backwards within the sepal spur. Follicle ovoid, pubescent, 1 to 2 cm long; seeds covered with transverse broken ridges or ruffled scales. Roadsides, waste places, and old fields. Primarily Cen. and S. TX; native of Europe, escaped and spreading from cultivation in Can., U.S., and Mex. Apr.-Sept. [Delphinium ajacis L.; Consolida ambigua (L.) Ball & Heywood.].
Perennial herbs from fibrous or tuberous roots or rhizomes. Stems 1 to several, erect or ascending, simple to branched. Leaves alternate, at least the lower ones petiolate, blades often reniform in overall outline, palmately or rarely pinnately cleft or divided into 3 to 7 main lobes, each lobe usually further divided, ultimate divisions broadly linear to rounded or cuneate (not linear-filiform), commonly white-apiculate. Flowers few to many in terminal spike-like racemes or panicles, zygomorphic, blue to violet, purple, white or pink. Sepals colored, 5, the uppermost prolonged backward into a slender, horizontal spur. Petals 4, free, the upper pair extending backwards into spurs enclosed in the sepal spur, nectariferous, lower petals short-clawed, blades elliptical to orbicular, often bifid, bearded. Stamens usually many. Carpels 3, distinct, maturing into many-seeded follicles.
About 250 species of N. temperate regions; 3 or 4 species in TX (depending on interpretation). Dr. M. J. Warnock, the most recent monographer of our species (Warnock 1981), has identified all local specimens as subspecies of D. carolinianum.
Many species are cultivated for ornament. Most contain toxic alkaloids (Mabberley 1987).
1. D. carolinianum Walt. Blue Larkspur. Plants virgate from unbranched, cormlike, vertical rootstocks or a cluster of several woody-fibrous or tuberous roots. Stems simple, to 1.5 m tall, variously pubescent throughout or glandular above and glabrous below. Leaves mostly near the base or distributed rather evenly along the stem, blades palmately divided into several primary segments which are entire to repeatedly divided, variously pubescent. Racemes lax to dense, flowers showy, white to dark blue-purple, pedicels erect to appressed or spreading, bracts subulate and inconspicuous to obsolete. Sepals ovate to ovate-oblong, rounded to blunt or acute at apex, 11 to 15 mm long, the spur straight or curved; blades of lower petals short, included, upper petals various. Follicles erect, oblong, pubescent, seeds echinate. FL to TX, N. to CO and MN, S. and E. to IL, KY, TN, and GA.
Warnock recognizes four subspecies from TX. Two of these can be expected in our area.
subsp. vimineum (D. Don.) Warnock Plants usually from a single, vertical, swollen rhizome, 1.5 to 10.5 dm tall. Stem pubescence various, sometimes glandular. Petioles 3 to 15 cm long on both upper and lower leaves. Leaves basal or evenly distributed, 5 to 9 cm broad, 3-parted with few subdivisions, segments to 4 mm broad or broader, glabrous to sparingly pubescent. Inflorescences from axils of the upper leaves, often much-branched, pedicels more or less spreading. Sepals blue to white, spur 11 to 15 mm long, usually crossing the axis of the inflorescence at about 45o on the fully open flowers; upper 2 petals usually white (sometimes light blue), the lower pair blue or white. E., NE., Cen., and coastal TX; also LA (SW. AR?). Mar.-June. [D. vimineum D. Don; D. azureum Michx. var. vimineum (D. Don) A. Gray ex K. C. Davis; D. virescens Nutt. var. vimineum (D. Don) R. F. Martin].
This is by far our most common larkspur. M. J. Warnock has annotated all of our local specimens to this taxon.
subsp. virescens (Nutt.) Brooks White Plains Larkspur. Plants to 1.5 m, usually smaller, from a cluster of several deep-seated, woody-fibrous roots. Stems branched or unbranched, pubescent, often glandular. Petioles long on the lower leaves, reduced upwards. Leaves usually basal and cauline. Ultimate leaf divisions lanceolate or linear. Pedicels more or less appressed to axis of inflorescence. Sepals white or whitish to greenish or bluish, but not buff; spur 11 to 20 mm long, often curved upwards, crossing the stem nearly horizontally in fully open flowers; petals whitish. Sandy open slopes, woods, pastures, fields, and ditches. Ed. Plat. to N. Cen. TX and Panhandle, S. to Rio Grande Plains and coast; Alta. to MN and ND, S. to OK and TX. Apr.-July. [D. virescens Nutt. var. virescens and var. macroceratilis (Rydb.) Cory; D. macroceratilis Rydb.].
This plant usually occurs to our W. Included in this treatment on the basis of a hybrid between this subsp. and subsp. vimineum as annotated by M. J. Warnock--if a hybrid exists in an area, presumably both parents do also.
20 species of temperate Asia and Europe. Some species have digitalin-like alkaloids and some are cultivated for ornament (Mabberley 1987). We have the 1 ornamental established in TX.
1. A. annua L. Pheasant's Eye. Annual herb from fibrous roots. Stems (15)20 to 60 cm tall, much-branched. Leaves alternate, petioles with broad, sheathing bases, blades overall 2 to 5 cm long, much
dissected, the ultimate divisions linear-filiform. Flowers solitary, terminal, ca. 2 cm across, not opening wide but often pressed open in specimens, deep red with a dark center (color fading to yellow with purple-brown in some sheets). Sepals 5 to 8 in number, oblong, acute-cuspidate; petals 6 to 8, obovate, somewhat erose apically, concave, deciduous, longer than the calyx, but not much so; stamens many, filaments filiform. Gynoecium of many simple, 1-ovuled pistils. Head of achenes 1 to 2 cm long, cylindric. Achenes 3 to 5 mm long, glabrous. Native to Eur., cultivated for ornament and established in the E. 1/3 TX. Spring-summer.
Without flowers, this plant may easily be mistaken for an immature umbellifer such as Daucus.
Annual and perennial aquatic, terrestrial, or palustrine herbs, usually from fibrous roots. Stems erect to procumbent, branched or simple. Leaves basal and/or cauline and alternate, simple or more commonly ternate or variously lobed or divided, often variable on a single plant, petioles with expanded bases. Flowers terminal (sometimes appearing axillary), solitary or in a corymbiform inflorescence, regular, perfect, hypogynous, inconspicuous to showy. Sepals 3 to 5 (or more), green to yellowish. Petals commonly 5 (more in some species through conversion of stamens to petals), generally yellow or white (rarely reddish or green, not so in ours), each with a nectariferous scale and/ or pit near the inner base. Stamens (5)10 to many, anthers oblong or linear. Pistils (5)10 to many, usually many, uniovulate, gynoecium maturing into an ovoid-cylindric head of achenes, style short to long, straight to hooked, usually persisting as a beak on the achene, achenes variously glabrous to pubescent, muricate, and/or keeled.
About 250 species of temperate and boreal regions; 18 in TX; 7 here. Mature achenes are often necessary for positive identification.
All species are poisonous and are usually avoided by stock. Some species are weedy while others are cultivated for ornament (Mabberley 1987). The ornamentals are often from tuberous roots and frequently have double, brightly-colored flowers ranging from red and orange to yellow and white. According to childlore, if the glossy yellow of buttercup petals held under a person's chin is reflected on the chin, he or she "loves butter." Ranunculus is not to be confused with Oenothera (Onagraceae), often called Buttercup in our area.
1. Cauline and basal leaves entire to dentate, serrulate, or undulate ........................................2
1. Cauline or basal leaves lobed, parted, or divided ...................................................................3
2(1) Petals 5 to 9, about twice as long as the sepals; styles at anthesis 0.5 mm long; heads of achenes spherical to ovoid ..............................................................................1. R. laxicaulis
2. Petals 1 to 3, minute, shorter than to about equalling the sepals; styles at anthesis 0.1 to 0.2 mm long; heads of achenes hemispherical, ovoid, or cylindrical ...............2. R. pusillus
3(1) Mature achenes covered with spines or hooks ..............................................3. R. muricatus
3. Mature achenes smooth or with minute papillae, never with spines ......................................4
4(3) Petals 2 to 5 mm long; achenes essentially beakless, beak less than 0.1 mm long ..............
........................................................................................................................4. R. sceleratus
4. Petals more than 5 mm long; achenes at least shortly beaked, beak more than 0.1 mm long ...........................................................................................................................................5
5(4) Petals 8 to 18, 10 to 20 mm long; achenes (35)40 to 130 per head ........5. R. macranthus
5. Petals usually 5, 7 to 15 mm long; achenes 10 to 35 per head .............................................6
6(5) Achene beaks longer than 1.5 mm; style with stigma apical; pubescence of lower stem appressed; sepals ca. 6 to 8 mm long; roots tuberous and/or filiform ......6. R. fascicularis
6. Achene beaks shorter than 1.5 mm; stigma lateral; pubescence of lower stem spreading; sepals ca. 3 to 5 mm long; roots filiform ...........................................................7. R. sardous
1. R. laxicaulis (T. & G.) Darby Many-flowered Spearwort, Water Plantain, Spearwort. Terrestrial or shallow-water emergent. Plants annual, essentially glabrous, slender, from fibrous roots. Stems hollow, erect to reclining, sometimes rooting at the lowermost nodes, 1.5 to 5(8) dm tall, freely branched. Leaves all simple, petioles of basal and lower cauline leaves 1 to 7 cm long (longer if emergent), blades ovate to oblong, 1 to 5 cm long, 6 to 18 mm broad, shallowly dentate to serrulate or entire, rounded or truncate at the base, obtuse to truncate at the apex (rarely acute); stipular leaf bases ca. 1 cm long. Upper cauline leaves attenuate, sessile, linear-elliptic to lanceolate or oblanceolate, 1.5 to 4 cm long, 2 to 6 mm wide, acute, dentate to denticulate. Inflorescence often well-branched; pedicels to 2 cm long in flower and 6 cm in fruit. Sepals 5, greenish to yellow, ovate, spreading, 1.5 to 3 mm long, to 1.5 mm broad, glabrous to sparsely pubescent, promptly deciduous; petals 5 (rarely to 10), yellow, 3 to 9 mm long, 1.5 to 2.5 mm broad, exceeding the sepals; nectary scale adnate to the petal along the lateral edges, forming a pocket, glabrous, 0.3 to 0.5 mm long, truncate or with the margins prolonged; stamens 10 to 30. Receptacle pyriform or spherical, 1.5 to 2 mm long in flower, 1.5 to 3 mm in fruit, glabrous; heads 2 to 4 mm long, 2 to 2.5 mm in diameter, hemispheric to ovoid (rarely cylindrical), with 15 to 30(50) achenes. Achene body plump, obovate to subglobose, 0.6 to 0.7(1.1) mm long, about as thick as broad, smooth, glabrous, the margin inconspicuous, style 0.1 to 0.5 mm long. Boggy lakeshores, ditches, marshes, etc. SE. TX; CT to MO, IL, E. KS, S. to FL and TX. Mar.-May. [R. texensis Engelm.; R. pusillus of some authors, but not Poir.; R. obtusiusculus Raf.; R. oblongifolius sensu Small; R. mississippiensis Small].
Similar to R. pusillus, but with larger, showier flowers and rarely with cylindric fruiting heads.
2. R. pusillus Poir. Spearwort. Terrestrial or shallow-water emergent. Plants annual, glabrous or rarely pubescent, from fibrous roots. Stems hollow, weak, often reclining and commonly rooting at the lowermost nodes, simple or more often branched, 1 to 5 dm long. Basal and lower (and usually middle) cauline leaves with petioles 1 to 6 cm long, blades ovate to oblong or rarely cordate, entire to crenate or irregular, truncate to rounded basally, rounded to truncate or acute apically, to 5 cm long, 5 to 15 mm broad; stipular leaf bases to 1 cm long; upper cauline and bracteal leaves alternate, sessile, lanceolate to linear, oblanceolate, or narrowly elliptic, 1 to 5 cm long, 2 to 5 mm broad, entire to slightly dentate. Inflorescence often well-branched, more or less paniculate; pedicels to 15 mm long in flower and ca. 6 cm long in fruit, glabrous. Sepals 5, spreading, greenish-yellow, generally ovate, 1 to 2 mm long, 0.8 to 1 mm broad, glabrous to sparsely pubescent, promptly deciduous; petals 1 to 3(5), pale yellow, inconspicuous, obovate, 1.5 to 2.5 mm long and 1 mm wide; nectary scale glabrous, truncate, laterally attenuate and forming a pocket 0.2 mm deep; stamens 5 to 10; style 0.1 to 0.2 mm long. Receptacle pyriform or spherical, 1.5 to 2 mm long in flower, 1.5 to 3 mm long in fruit, glabrous. Fruiting heads hemispheric--to ca. 4 mm in diam., ovoid--2 to 4 mm long and 2 to 2.5 mm broad, or cylindric--5 to 8 mm long and 2 to 3 mm broad. Achenes as many as 125 per head, obovate or oblong-obovate, ca. 1 mm long, smooth or papillate (or finely reticulate), glabrous, the sides convex and the margins inconspicuous. Shallow water or mud in ditches, bogs, marshes, ponds, seeps, open woods, and thickets. E. 1/4 TX, W. to Burnet Co.; SE. NY to N. FL, W. to TX, N. again to MO, IN, OH; also CA. Mar.-May. [Includes var. angustifolius (Engelm.) L. Benson; R. tener Mohr.; R. lindheimeri Engelm.].
3. R. muricatus L. Annual, winter annual, (or perennial.) Plants terrestrial, glabrous or pubescent, somewhat cespitose. Stems not rooting, erect or reclining, not markedly hollow, freely branched, usually several from near the base, shorter than the basal leaves and plants 5 to 15 cm tall, or else more robust and with flowering stems 2 to 5 dm tall. Leaves basal and cauline, basal and lower stem leaves with petioles (2)4 to 15(18) cm long, variable, blades simple, orbicular to broadly cordate, subreniform or semicircular, 2 to 5 cm long, 2 to 6 cm broad, deeply 3-lobed, the lobes shallowly to strongly crenate or crenately lobed, basally cordate to truncate, apically rounded; stipular leaf bases 1 to 2 cm long. Cauline leaves alternate, similar to basal but smaller and petioles shorter; bracteal leaves sessile. Pedicels 0.5 to 2 cm long in flower, 2 to 6 cm long in fruit, glabrous. Sepals 5, greenish, ovate, spreading, 4 to 7 mm long, 2 to 3 mm broad, mucronate, with a few bristles, promptly deciduous; petals 5, yellow, spatulate or obovate, 5 to 8 mm long, 3 to 4 mm broad; nectary scale glabrous, truncate, attached laterally and forming a pocket narrower than adjacent part of petal. Stamens few. Receptacle subglobose, ca. 1 mm long in flower, to 2 mm long in fruit, hispid. Head of achenes globose, 1 to 1.3 cm in diameter. Achenes 10 to 20, asymmetrical, obovate, the body 4 to 5 mm long, 3 to 3.5 mm broad, faces covered with straight or curved spines, minutely punctate-alveolate between the spines, margin prominent, keeled; beaks straight or falcate, base stout, subulate tip 2 to 2.5 mm long. Grassy banks, roadsides, sandy marshes, along ponds and streams, or in shallow water. E. 1/4 TX; naturalized from Eur. from SC to N. FL, W. to AR, LA, TX. Apr.-June.
4. R. sceleratus L. Cursed Buttercup, Blister Buttercup, Celery-leaved Butter-cup. Annual or short-lived perennial. Plants of marshes or fully aquatic. Stems erect, hollow, stout and inflated at base, to 2 cm broad, to 1 m tall, rarely rooting from lower nodes, freely-branched, usually glabrous below and pubescent above on the branches. Leaves cauline and basal, the lowermost often absent at anthesis. Basal leaves with long petioles to a maximum of 25 cm long, blades simple, reniform in overall outline, to 6(10) cm long and 10(13) cm broad, deeply 3-lobed or parted, the divisions lobed, parted, or divided, apices of ultimate divisions rounded, sinuses rounded, leaf bases cordate to truncate or widely cuneate; stipular leaf bases 5 to 10 mm long; cauline leaves alternate, becoming more sessile and smaller upwards, pedately divided; bracteal leaves entire, lanceolate to oblanceolate or linear. Inflorescence often well-branched, pedicels to 1(2) cm long in flower and 3 cm in fruit. Sepals 5, green-yellow, spreading, ovate, 2 to 3 mm long, 1.5 to 2 mm broad, pilose externally or glabrous, quickly deciduous but remaining longer than the petals; petals 5, yellow, obovate to oblong, generally 2 to 5 mm long, 1 to 3 mm broad; nectary scale glabrous, with the margins prolonged along the blade of the petal, sometimes 1 or both margins with a free flap at the tip or the scale surrounding the nectary; stamens 10 to 25. Receptacle obovoid or cylindric, 1 to 2 mm long in flower, 2.5 to 9 mm long in fruit, pubescent or sometimes glabrous. Head of achenes cylindrical, 3 to 10(15) mm long, 2 to 6(10) mm broad. Achenes 40 to 300, somewhat asymmetrical or wedge-shaped to obovoid, 0.8 to 1 mm long, often with minute, irregular transverse ridges on the faces; margins and base corky-thickened at least somewhat, their surfaces with ridges or a circle of minute punctations at the inner margin of the thickened edge; achenes glabrous, keel obscure, style and achene beak practically lacking, not recurved. Margins of lakes, streams, and marshes. S. and SE. TX; Nfld. to Albta., S. to FL and TX; native to Europe and widely naturalized.
The sap of this species is said to cause blisters on human skin (Lampe 1985).
5. R. macranthus Scheele Large Buttercup. Perennial from stout or somewhat tuberous roots, terrestrial, glabrous or more commonly strongly hirsute. Stems hollow, densely hirsute, suberect to reclining, not rooting at the lower nodes, to 1 m tall but usually much shorter. Leaves basal and cauline, basal leaves petiolate, petioles hirsute, to 30 cm long; stipular bases ca. 1 to 3 cm long; blades pinnately compound or sometimes simple and lobed, overall oblong-ovate in outline, 4 to 23 cm long, 3 to 25 cm broad, leaflets usually 3 to 7, basally truncate to obtuse, acute to obtuse at the apex, variously crenate to serrate or lobed, appressed-hispidulous. Flowers commonly few per plant, pedicels to 11 cm long at anthesis and 30 cm long in fruit, appressed-pubescent. Sepals 5, yellow-green, ovate-attenuate, 6 to 10 mm long, 3 to 5 mm broad, reflexed, appressed-pilose externally, promptly deciduous; petals 8 to 18, yellow, oblanceolate (obovate), sometimes emarginate, 1 to 2 cm long, 2.5 to 10 mm broad, showy. Receptacle cylindric, 2 to 3 mm long in flower, 5 to 12 mm long in fruit, pubescent, often sparsely so. Head of achenes subglobose to cylindric, 7 to 14 mm long, 7 to 10 mm in diameter. Achenes 35 to 130, elliptic-oblong to obovate, 2.5 to 4 mm long, smooth, glabrous, the margin keeled; beak slender, straight, 3 to 5 mm long. Wet soils--swamps, drainages, wet woods, creeks, mud flats around pools, and seepage slopes. Cen., S., and W. TX; W. to AZ; also Mex. Mar.-June.
6. R. fascicularis Muhl. ex Bigel. Prairie Buttercup, Early Buttercup. Perennial from a fascicle of both short, tuberous roots to 5 mm in diameter and filiform, cordlike roots. Flowering stems several from the base, 1 to 3 dm tall, weak, erect to suberect, not rooting at the nodes, often scapose, not hollow, with silky-canescent appressed or ascending hairs; herbage appressed-pubescent. Basal leaves several to many, petioles to 10 cm long; blades ternate, compound, or at least 3-parted, ovate-oblong in overall outline, 2.5 to 5.5 cm long, 2 to 4 cm broad, definitely longer than broad, divisions 3 or 5, the lateral ones sessile to short-petiolulate, the terminal petiolule 1 to 2 cm long; divisions oblanceolate to narrowly elliptic, simple and rounded or few-toothed apically to 3-(to 7-) lobed and/or angularly toothed, leaves sometimes appearing bipinnate, ultimate divisions blunt or rounded; stipular leaf bases 15 to 35 mm long; cauline leaves usually 1 or 2, alternate, reduced, with 1 to 3 segments; bracteal leaves much-reduced, commonly undivided, linear to lanceolate. Pedicels 1.5 to 6 cm long in flower, elongating to 2.5 to 9 cm or more in fruit. Sepals 5, green-yellow, ovate-attenuate, spreading, 6 to 8 mm long, 2 to 3 mm broad, usually somewhat silvery-pubescent, promptly deciduous; petals 5 (always so in our region, elsewhere to 9), yellow, obovate-oblanceolate or obovate-orbicular, 6 to 15 mm long, 3 to 6 mm broad, rounded apically; nectary scale truncate, glabrous, free nearly its entire length. Stamens commonly 40 to 50. Receptacle fusiform to obovoid, 1.5 to 2.5 mm long in flower, 3 to 7 mm long in fruit, lightly hispidulous. Head of achenes subglobose, 4.5 to 8(11) mm long, 6 to 12 mm in diameter. Achenes 10 to 35 per head, obovate-orbicular, with a short, flat stalk, main body 1.5 to 3 mm long, glabrous, the margin keeled but not usually prominent, the faces somewhat convex, smooth, dull; style beak subulate, straight to curved, 2 to 2.3 mm long (if unbroken), filiform and stigmatic apically. Sandy soils in shallow water, low pinewoods, meadows, seepage slopes, and prairies. NE. TX S. and W. to SW. TX plains; MA to MN, S. to GA, AL, LA, and TX. Feb.-May. [Includes var. cuneiformis (Small) L. Benson and var. apricus (Greene) Fern.; (R. apricus Greene)].
7. R. sardous Crantz Terrestrial annual from filiform roots. Stems not rooting at the nodes, sub-erect, simple or branched from the base, not hollow, sparsely to densely spreading-hirsute, 1 to 5 dm tall. Basal leaves sometimes withered or absent at anthesis, petioles 3 to 16 cm long; blades ternate, broadly cordate to ovate in overall outline, 2 to 3(4) cm long and 2 to 4 cm broad, lateral leaflets sessile to very short-petiolate, terminal leaflet stalked, leaflets shallowly to deeply parted or lobed, to ca. 2 cm long, deltoid to ovate in outline, cordate to truncate basally, rounded apically, appressed-pubescent; stipular leaf bases 1 to 1.5 cm long; cauline leaves alternate, often with larger blades than the basal leaves, shorter-petioled to sessile, ternate to 3-lobed, reduced upwards to sessile bracteal leaves with 3 linear divisions. Pedicels 3 to 5 cm long in flower, 2 to 6(8) cm long in fruit, sparsely appressed-pubescent. Sepals 5, yellow-green, ovate-attenuate, 3 to 5 mm long, 1.5 to 2 mm broad, reflexed, often early-deciduous, pilose to sparsely hispid externally; petals 5, yellow, 8 to 9 mm long, 5 to 7 mm wide, obovate; nectary scale truncate to ob-ovate, lateral margins free ca. 3/4 their length; stamens 25 to 50. Receptacle pyriform, ca. 1 mm long in flower and 2 mm long in fruit, with long white hairs. Head of achenes subglobose to globose or ovoid, 4 to 8 mm long, 5 to 7 mm broad. Achenes 10 to 40 per head, nearly circular, 2 to 3 mm long, body flat, margin prominent, faces smooth to papillate, glabrous, style stigmatic along one side, beak deltoid, flattened, 0.2 to 0.5 mm long, blunt, sometimes curved at the tip. Moist grassy slopes, low wet areas, marshes. Sporadic in the E. 1/3 TX; native of Eur. and naturalized in the U.S., especially on the Atlantic Coast. Apr.-June.
Many of our plants were erroneously identified as R. fascicularis because R. sardous was omitted from keys to the local flora or described with only papillate achenes. R. sardous is much more common in our area than previously thought, and our plants represent a westward range limit not described in many books.
Herbaceous perennials, sometimes rhizomatous, stems hollow. Leaves alternate, the lowest long-petiolate, the petiole bases often expanded; blades ternate or 2 or 3 times compound, the leaflets usually cleft or shallowly lobed, commonly petiolulate, paler beneath; cauline leaves similar but progressively simpler, uppermost leaves often short-petiolate to sessile, sometimes all the leaves so. Plants dioecious or polygamo-dioecious, flowers usually many, pedicellate, in panicles (umbels), not individually very showy, usually unisexual, small. Sepals 4 or 5, greenish to purple or whitish, caducous; corolla absent. Stamens many, exserted, yellow to purplish, the filaments filiform to slenderly clavate, often tangled; anthers slender, apiculate. Carpels about 4 to 17 (at least in ours), free, styles short to lacking, stigmas unilateral, persistent on the 1-seeded achenes; achene bodies usually longitudinally ribbed.
About 85 species, chiefly of the N. temperate regions, tropical S. Amer., and S. Afr.; 4 in TX; 2 here.
Most species contain alkaloids; a few are used in herbal medicines. Some are grown as ornamentals (Mabberley 1987). One of ours is a rare endemic.
1. Leaflets somewhat thick, rigid, longer than 1 cm, generally longer than wide, usually with 3 acute, entire lobes .......................................................................................1. T. dasycarpum
1. Leaflets thin, less than 1 cm long, about as long as wide, usually with 3 obtuse to rounded, crenate, or notched lobes .................................................................................2. T. texanum
1. T. dasycarpum Fisch. & Avé Lall. Purple Meadow-rue. Plants polygamo-dioecious, from a stout, erect rootstock. Stems glabrous, 5 to 14(20) dm tall, often purple. Lowest leaves petiolate, middle and upper short-petiolate to subsessile; stipules suborbicular, brown' leaflets firm, obovate in overall outline, the longest 1.5 to 4(5.5) cm long and ca. 4 cm broad, entire to acutely 3-lobed, occasionally with a few extra lobes or teeth, veins prominent beneath; undersurfaces pale, glaucous, or with a few minute hairs, margins sometimes revolute. Inflorescence corymbose-paniculate, open. Sepals 4 to 5, lanceolate to narrowly ovate, acuminate, 2 to 5 mm long, purplish to whitish, commonly caducous; filaments 4 to 7 mm long, soon drooping and becoming tangled, anthers mostly 1.3 to 3.2 mm long, oblong to linear, the subulate tip 0.1 to 0.2 mm long; carpels 5 to 14, stigmas 2.5 to 5 mm long, about as long as the ovoid to lanceolate body. Fruits (2.5)3.8 to 5.5 mm long, dark, ribbed. Meadows, swamps, and moist woods, especially along slopes and streams. E. 1/3 TX and N. panhandle; Ont. W. to Albta., S. to OH, LA, TX, NM, and AZ. Mar.-July. [Incl. var. hypoglaucum (Rydb.) Boivin; T. hypoglaucum Rydb.; sometimes varieties of this species not recognized.].
2. T. texanum (Gray) Small Houston Meadow-rue. Plant from a somewhat swollen, irregular rootstock which often becomes black on drying. Stems stiff, erect, to 45 cm tall. Foliage delicate, lower leaves petiolate, upper leaves short-petiolate to sessile. Main leaves ternate to 3 times ternate; leaflets to about 8 mm long and 10 mm broad, obovate, entire to crenate, notched, or with 3 rounded lobes, veins visible below, lower leaf surface paler than the upper, somewhat glaucous; stipules less than 1 to 2 mm long. Inflorescence dense to more compact. Sepals of male flowers 1.7 to 3 mm long, in the female flowers 0.7 to 1.5 mm long; filaments ca. 1.5 mm long, becoming somewhat tangled, anthers 1.4 to 2 mm long; stigmas 0.5 to 1 mm long. Mature fruits ovoid or ovoid-fusiform, sometimes curved, apically acute, 2.7 to 3.7 mm long and ca. 1.4 to 2 mm broad, with a stipe 0.1 to 0.3 mm long, 6- to 8-ribbed. SE TX; endemic. Mar.-May. [T. debile Buckl. var. texanum A. Gray].
While this plant is not officially listed as endangered, it is under study to determine if it should be listed as threatened or endangered (U.S. Fish and Wildlife Category 2.) It is certainly rare, and difficult to find even if being deliberately sought.
Perennial herbs from tubers or slender to thickened rhizomes. Stems erect. Leaves divided or nearly compound, all basal except for several, sessile to petiolate, borne in a whorl on the stem as an involucre below the peduncle. Basal leaves sometimes absent; if present, long-petiolate. Flowers solitary (as in ours) or in cymes or umbels, peduncles of all but the central flower sometimes with secondary involucres. Sepals petaloid, 4 to 20, usually showy, white, red, yellow, blue, or purple. Corolla absent or petals present as gland-like staminodia. Stamens and pistils many in a cylindrical or spherical cluster. Fruiting structure an elongate or compact head of achenes, these flattened, pubescent or glabrous, with short or elongate style beaks.
About 120 species nearly worldwide, especially in the N. temperate zone; 4 species are found in TX and 2 here. Reference: Keener (1975). [Pulsatilla Adans. formerly included].
Many species, some with "double" flowers, are grown for ornament and have the sepals brightly colored or marked.
1. Involucre below the middle of the scape at anthesis, usually similar to some of the basal leaves; scapes densely pubescent above the involucre and nearly glabrous below; styles projecting beyond the pubescence of the head of achenes .......................1. A. caroliniana
1. Involucre above the middle of the scape at anthesis, usually dissimilar to the basal leaves; scapes similarly pubescent above and below; styles nearly hidden by pubescence of the achene cluster ................................................................................................2. A. berlandieri
1. A. caroliniana Walt. Carolina Anemone. Plants from a small tuber 6 to 15 mm in diameter with rhizomes and stolons. Stems/scapes 5 to 30 cm tall, glabrous below the involucre, villous to appressed-pubescent above. Leaves once or twice 3-parted or 3-cleft, at least some of the leaves with the ultimate divisions more or less linear and with acute tips, the largest blades overall 2 to 5 cm broad, longest petioles 2 to 6(10) cm long. Involucre below the middle of the scape at anthesis, bracteal leaves sessile, whorled, smaller than but similar to the basal leaves, 1 to 3 cm long, dissected, the ultimate divisions linear-acute. Flowers solitary, 1.5 to 4 cm across. Sepals 10 to 20(27), white to rosy or lavender to violet, often whitish above and colored below, linear-oblong, 1 to 2.7 cm long. Fruiting head ellipsoid (sometimes cylindrical), (8)13 to 19(22) mm long and 7 to 10 mm wide. Achenes nearly concealed by the pubescence of the head, ovate, plump, 1.5 to 2(2.5) mm long, styles 1.3 to 2 mm long, about equalling the achene body, with the slender tip often projecting beyond the woolly pubescence. Generally in sandy soils. E. 1/4 TX; FL to TX, N. to SD, MN, WI, IN. Feb.-Apr.
2. A. berlandieri Pritzel Tenpetal Anemone. Plants from a thick, clavate-to-oblong rootstock, not rhizomatous, 1 to 3(4.5) dm tall. Stems loosely villous below the involucre, becoming appressed-villous above. Basal leaves long-petioled, longest 5 to 16 cm long; blades variable in size and shape on a given individual, the largest 3 to 5.5 cm long, usually ternate, leaflets petiolulate to sessile, broad, ovate to oblong or obovate, the margins crenate-dentate or cleft, generally thinly appressed-pubescent or ciliate-hirsute. Leaves of involucre borne above the middle of the scape at anthesis, dissimilar to the basal leaves, generally 3, subsessile, 1.8 to 4.5(6) cm long, once (or rarely twice) palmately cleft, the ultimate segments linear, generally entire, acute. Flowers solitary, ca. 2 to 4.3 cm broad. Sepals 10 to 22(33), mostly white above and purplish to blue-violet below, 11 to 21 mm long. Head of achenes cylindric, (11)20 to 40 mm long, 5.5 to 11 mm broad, achenes nearly obscured by the pubescence of the head, orbicular, 2.7 to 2.5 mm long, flattened, style slender, ca. 1/3 the length of the achene body, eventually inflexed and not projecting beyond the woolly pubescence. Sandy clay or calcareous clay soils, throughout much of TX; NC to VA, SW. to KS, OK, TX, AR, AL, MS. Feb.-Apr. [A. decapetala Ard. var. heterophylla (Nutt.) Britt.; A. heterophylla Nutt. The latter, though widely used, has been determined to be an invalid name.].
Dioecious twining vines (elsewhere also scandent shrubs, rarely trees or herbs.) Leaves estipulate, simple, alternate, petiolate, palmately veined, entire to lobed. Flowers in racemes or panicles, unisexual, 3-merous, often small and inconspicuous, hypogynous. Calyx and corolla similar (at least in ours), free, in 2 to 4 whorls, imbricate in bud. Stamens 6 to 12(24), in some plants more or less connate (but not in ours.) Pistils free, 1 to 30, in our plants 2 to 6 (commonly 3), with only one maturing into a 1-seeded (by abortion) drupe (or nut); seeds often curved, coiled, or flattened.
About 78 genera and 520 species in tropical and subtropical areas, a few in temperate zones. Two genera and 3 species can be found in TX. We have 1 species.
Many species have alkaloid chemistries and are used in medicines, fish poisons, curare, sweeteners, etc. A few are grown for ornament (Mabberley 1987).
Largely as described for the family. Plants deciduous. Flowers in axillary or terminal panicles. Sepals, petals, and stamens each 6, in alternate whorls. Pistils 3 or 6, stigmas entire. Fruit a globose drupe; seed flattened, often rough.
11 species of the tropic, subtropic, and warmer temperate regions (excluding S. Amer. and Aust.); 2 in TX; 1 here.
Some species have edible fruit (not ours), some have medicinal uses; some are ornamental (Mabberley 1987).
1. C. carolinus (L.) DC. Red-berried Moonseed, Snailseed, Carolina Moonseed, Coralbead. Vine herbaceous to woody, stems usually climbing, to 3 or 4 m long, minutely pubescent. Leaves quite variable in shape, to 16 cm long and about as wide, ovate to cordate, entire or with a few shallow, sinuate or hastate lobes, apex mucronate, base truncate to cordate, lower surface sparsely pubescent or more commonly downy; petioles equal to or shorter than the blades. Panicles narrow,shorter than the leaves (except near the tip of the stem where the leaves are small), staminate panicles to 15 cm long, longer than the pistillate. Flowers greenish, 3 to 4 mm broad, petals in male flowers auriculate-inflexed basally around the filaments; pistils 3 (less commonly 6). Drupes in grape-like clusters, globose, red, yellow-pulped, 4 to 8 mm in diameter. Seed flat to bi-concave, the coiled embryo evident (hence "Snailseed"), often with radial ridges. Common in rich woods and thickets in the E. 1/2 of TX, rarer W.; KS, E. to WV., S. to NC, SC, FL, the Gulf States and TX. Flowering Jul.-Aug., often overlooked; fruiting in fall.
The curling vines and red fruit are pretty in arrangements. The fruits are variously listed as edible or poisonous and should probably be avoided (Tull 1987). Similar at first glance to Smilax (Catbriar, Greenbriar), but Smilax usually has glabrous leaves, black fruits typically in umbels, and prickles. Cocculus is never prickly.
In N. Amer. largely herbs (elsewhere sometimes shrubby), annual, biennial, or perennial, sometimes rhizomatous, glabrous to pubescent, with milky or colored latex in the roots, stems and leaves. Leaves in a rosette or alternate (very rarely opposite), simple, entire or pinnately or palmately lobed, estipu-late. Flowers solitary or occasionally clustered or paniculate, terminal or axillary, actinomorphic, perfect, hypogynous (a few species perigynous), sub-tended by bracts or leaves. Perianth 2- or 3-merous; sepals 2 or 3, sometimes slightly united below, enclosing the bud before anthesis and then commonly immediately deciduous. Petals twice as many as the sepals (sometimes 0, 6, 8, 12, or 16), free, often in 2 whorls, imbricate and crumpled in the bud. Stamens many, usually a multiple of sepal number. Ovary usually 1-celled with parietal placentation, carpels 2 to many; stigmas generally the same number as the carpels, fused into a discoid or lobed structure. Fruit mostly a valvate or poricidal capsule.
About 23 genera and 210 species, primarily in N. temp. regions; 4 genera and 12 species in TX; 2 species of 1 genus here.
The alkaloid chemistry of the plants gives some of them importance as drug or medicinal plants. From the latex of Papaver somniferum come opium and heroin, though the seeds are safe to eat and used in baking. Many species are grown as ornamentals. There are some oilseed crops (Mabberley 1987).
NOTE: Our poppies seem all to be white and fall into one genus. Should colored poppies be located in our area, they would likely be P. rhoeas L. if red, and Argemone sanguinea if lavender and moderately prickly (though our common A. polyanthemos may also rarely be lavender).
Annual, biennial, or rarely perennial herbs; latex yellow to orange or red; herbage with prickly bristles, often somewhat glaucous. Stems 1 to several, simple or cymosely-branched, generally 3 dm tall or more. Leaves cauline, alternate, sessile to auriculate or clasping, often mottled or with lighter veins, slightly to deeply pinnately lobed, the margins commonly spinose-toothed. Inflorescence cymose, the buds on short peduncles. Sepals 3 (2 to 6), imbricate, each with a prominent terminal cusp or horn, generally prickly, caducous at anthesis. Flowers white (as in ours), yellow, or lavender, showy, petals 4 to usually 6(12) in two whorls. Stamens usually many, the filaments filiform and anthers coiling after dehiscence. Carpels 3 to 6, the style practically lacking, stigma 3- to 6-rayed or -lobed. Capsules prickly, elliptic, somewhat fluted on the sutures, 3- to 6-valvate at the top 1/3, persistent, with the vascular elements visible within. Seeds subglobose, crested, blackish-brown.
28 species in the W. hemisphere; 8 in TX; our material referable to 2. Reference: Ownbey (1958).
Some species have ornamental value (though they are decidedly not pleasant to handle!) or are used in herbal medicines. A. albiflora can be made to yield a tan dye (Tull 1987).
1. Sepal horns 7 to 15 mm long; mature capsule surface clearly visible through prickles; stamens equalling the stigma at anthesis ..............................................1. A. polyanthemos
1. Sepal horns 3 to 6(10) mm long; mature capsule surface partly obscured by prickles; stamens equalling or shorter than the stigma at anthesis ...............................2. A. albiflora
1. A. polyanthemos (Fedde) G. B. Ownbey Prickly Poppy. Annual or biennial from a deep taproot; latex yellow. Stems 1 to 5, to 1.5 m tall, cymosely-branched, glaucous, sparingly prickly with stout, recurved or spreading prickles 2 to 7 mm long. Leaves succulent, glabrous above or with a few prickles on the veins, lower surface prickly on the main veins, margins spinose-toothed; lower leaves oblanceolate, 7 to 20(25) cm long and 3 to 10 cm broad, pinnately lobed to 2/3 the distance to the midrib, the lobes oblong, elliptic or obovate; margins undulate, base attenuate to a winged petiole; middle and upper cauline leaves elliptic to oblong or ovate, reduced upwards and becoming more shallowly lobed, bases usually sessile and clasping. Buds elliptic-oblong, to 22 mm long and 15 mm broad, sparsely to moderately covered with spreading prickles; pedicels 1 to 4 cm long. Flowers (3)5 to 10 cm across, subtended (generally closely) by 1 or 2 leafy bracts usually shorter than the sepals. Sepals widely elliptic, 10 to 25 mm long, 10 to 15 mm wide, prickly, the sepal horns terete, to 15 mm long, often completely prickleless, often recurved; petals white (very rarely lavender), sometimes drying yellow, the outer suborbicular, the inner whorl broadly obovate-cuneate, (1.5)2.5 to 5 cm long, 2.5 to 5.5 cm broad, glabrous, the apices minutely erose; stamens 150 or more, filaments lemon yellow, drying brownish, 6 to 12 mm long, anthers 1 to 1.5 mm long, bright yellow, about even with the stigma at anthesis; stigma 3- or 4-lobed, 3 to 4.5 mm broad, 2 to 3 mm tall, purple; carpels 3 to 4. Capsules broadly or narrowly elliptic, 2.5 to 4 cm long including stigma, 1 to 1.7 cm broad (excluding prickles); prickles simple, spreading or recurved, 3 to 10 mm long, yellow, the largest mostly on the ridges; capsular surface clearly visible. Seeds globose, 2 to 2.5 mm long, shiny dark brown, alveolate-reticulate, with a 2-horned crest on one side. Sand and gravel soils of prairies, hills, mesas, roadsides, waste areas, pastures, etc. N. 1/2 TX; W. ND and WY to MT, S. to TX and E. NM. Apr.-Jun. [A. intermedia of some authors, not Sweet].
2. A. albiflora Hornem. subsp. texana G. B. Ownbey White Prickly Poppy. Annual or biennial from a deep taproot; latex yellow (reported by some as white to clear--?). Stem usually 1, to 1.5 m tall or more, sparingly to moderately prickly, the prickles slender, spreading to recurved. Leaves glabrous or with a few weak prickles primarily on the midrib above, margins spinulose-dentate. Basal leaves lanceolate to obovate, 7 to 20 cm long, 3 to 9 cm wide, pinnately lobed ca. 4/5 the distance to the midrib; middle and upper leaves more entire. Bud broadly elliptic to subglobose, sparsely to densely covered with simple, slender, spreading prickles. Flowers usually subtended by 1 or 2 bracts near the flower or at a distance of up to 5 mm at fruiting time. Sepals to ca. 2 cm long, exclusive of horns, horns 4 to 6(10) mm long, smooth or sparsely prickly at base; corolla white, petals 3 to 6 cm wide, the inner ones obovate-cuneate; stamens many, equal to or shorter than the stigma at anthesis; stigma 2 to 3.5 mm broad, 1.5 to 2 mm tall, style sometimes visible, otherwise practically lacking. Capsules 3- to 5-carpellate, usually narrowly-elliptic, to 14 mm broad (excluding prickles), to 4 cm long including stigma, armed with usually slender, spreading, prickles that are herbaceous at the base, also with many unequally-sized smaller spines and prickles, the largest to 12 mm long; capsule surface at maturity partially obscured by the prickles. Seeds ca. 1.7 mm long, black-brown, crested. Sand or gravel soils of embankments, roadsides, fencerows, waste places, vacant lots, etc. Throughout most of TX: N. AR and S. MO to TX for this subsp. The other subsp. from FL to TX, N. to CT, IL, and MO, possibly native only in N. FL. Mar.-Jun. [A. intermedia of authors, not of Sweet].
Annual, biennial, or perennial herbs, erect or scandent, sap watery, foliage often glabrous and/or glaucous. Leaves basal or alternate, rarely sub-opposite, estipulate, commonly pinnately or ternately divided or dissected. Flowers 1 to several in bracted, terminal or axillary racemes or cymes, 2-merous, usually strongly zygomorphic. Sepals 2, small, bractlike, not enclosing the bud, often caducous, sometimes somewhat peltate. Petals 2 + 2, 1 or both of the outer petals saccate or spurred at the base, the inner pair connate apically over the stigma. Stamens generally 6, diadelphous, the two bundles of 3 each opposite the outer petals, 1 or both of the median stamens commonly spurred at the base and/or nectariferous; anthers of two types: the middle one of each group with four microsporangia and 2 pollen sacs, all others with 2 microsporangia and 1 pollen sac each. Carpels 2, superior, united, ovules 2 to many on 2 parietal placentae, style 1, stigmas 2 or 1 with several lobes. Fruit generally a capsule, sometimes with replum, dehiscing by 2 valves, rarely indehiscent or breaking into 1-seeded segments or, also rarely, fruit a nut.
450 species in 18 genera, primarily in the N. temp. region, some also in tropical and S. Afr.; 6 species in 2 genera in TX; we have 2 species of 1 genus here. The family is sometimes included in the Papaveraceae.
Some are cultivated, such as Corydalis (Scrambled Eggs) and Dicentra (Bleeding Heart.) Some species of Fumaria are weedy and may eventually be found here.
Annual, biennial, or perennial herbs from taproots, tubers, or rhizomes, stems 1 to several, erect, decumbent, or in some species scandent (not ours.) Foliage glabrous and/or glaucous. Leaves basal, cauline, or both, alternate, the lowermost (at least) usually petiolate, once or twice pinnately divided or dissected, the pinnae again divided and incised. Inflorescence a terminal panicle or raceme, crowded at first and becoming elongated, bracteate; flowers short-pedicellate, zygomorphic; cleistogamous flowers sometimes present. Sepals 2, minute, appressed, fugacious. Corolla yellow in ours, in others sometimes pink or purple; petals 4, outer pair free or slightly united basally, one of them with a blunt spur and the other at most sometimes gibbous at the base, both more or less with a hood or keel at the apex; inner petals alike, clawed, connate apically. Stamens as described for family. Style slender, distinct; stigma persistent, flattened, sometimes bilobed. Fruit capsular, many-seeded, 2-valved, with 2 persistent placentae, unilocular, often constricted between the seeds (torulose), topped with the persistent style. Seeds subreniform to circular, black, smooth to muricate, shiny, somewhat arillate or carunculate. Seedlings of some species have only 1 cotyledon.
About 300 to 320 species, primarily of the N. temp. region; 4 in TX, 2 here. This treatment is based, in part, on the work of Ownbey (1947).
Some are cultivated for ornament, many species have alkaloid chemistries, and some Asian species have edible tubers (Mabberley 1987).
1. Fruits densely covered with transparent, clavate pustules; spurred petal 16 to 22 mm long .................................................................................................................1. C. crystallina
1. Fruits glabrous; spurred petal 18 mm long or less .......................................2. C. micrantha
1. C. crystallina Engelm. Mealy Corydalis. Winter annual, foliage green or glaucous, glabrous. Stems 1 to several, simple or sparingly branched, 1 to 4 dm tall, erect to ascending. Petioles of the lower leaves to 6 cm long, upper leaves short-petiolate to sessile. Blades (1.5)2 to 8 cm long, pinnate, the (5)7 to 9 primary segments again pinnately divided and incised, ultimate divisions lanceolate to linear-lanceolate, subapiculate. Racemes usually surpassing the leaves, with up to 20 flowers, bracts ovate or narrowly ovate, 5 to 12 mm long, 3 to 6 mm wide, reduced upwards; pedicels ca. 1 to 2 mm long, erect, stout. Sepals to 2 mm long, broadly ovate to cordate, somewhat attenuate, sometimes incised; corolla bright yellow, spurred petal 16 to 22 mm long, with a high, undulate or toothed crest, wing margin broad, reflexed on the hood, spur 6 to 8 mm long, with a blunt, globose tip; spurless outer petal 12 to 14 mm long, the wide wing margin not reflexed, crest same as that of spurred petal; inner petals 9 to 11 mm long, claw 4 to 5 mm long, blade oblanceolate, apex twice as wide as base, basal lobes small; stamen spur 2.5 to 5.5 mm long, clavate, curved or bent apically. Capsules 10 to 20 mm long, 2 to 2.5 mm broad, erect, straight or incurved, densely covered with clavate, transparent pustules that often break at maturity. Seeds shiny, black, 2 to 2.3 mm in diameter, circular or nearly so, without a ring margin but finely muriculate in concentric rings. Prairies, fields, open woods, wastelands, roadsides, and disturbed areas. E. Cen. TX; W. MO and SE. KS S. to AR and E. TX. Mar.-May, ours sometimes as early as Feb. [Capnoides crystallinum (Engelm.) O. Ktze.].
2. C. micrantha (Engelm. ex Gray) Gray Scrambled Eggs, Slender Fumewort. Annual or winter annual, foliage glaucous or green, glabrous. Stems 1 to several, simple to sparsely branched, 1 to 6 dm tall, usually less than 3.5 dm, erect to ascending or becoming prostrate with age, sometimes striate when dry. Petioles of lower leaves to 6 cm long, upper leaves short-petiolate or sessile. Blades ovate to oblong, pinnate, the 5 to 7(9) primary segments again pinnatifid and incised, ultimate divisions oblong or elliptic to obovate, acute to subapiculate. Cleistogamous racemes, if present, inconspicuous, 1- to 5-flowered. Chasmogamous racemes exceeding the leaves at least somewhat, with 6 to 20 crowded to well-spaced flowers, not surpassed by secondary racemes; bracts ovate to elliptic, the largest 5 to 8 mm long and 2 to 4 mm broad, acute to acuminate, entire or slightly toothed, much reduced upwards; pedicels erect, 2 to 6 mm long on the lower flowers, reduced upwards. Sepals 1 to 1.5 mm long, ovate, often undulate or toothed, fugacious; corolla pale to medium yellow, spurred petal 11 to 15 mm long, sometimes arched, with a low, undulate or rarely obsolescent crest on the hood, wing margin well-developed, spur 4 to 7 mm long, the tip blunt to globose; spurless outer petal 9 to 11 mm long, curved, crest low; inner petals 7 to 10 mm long, claw 3 to 4 mm long, blade oblanceolate, tip twice as wide as base, basal lobes obscure; stamen spur 2 to 4 mm long, generally ca. 3/5 the length of the petal spur, straight to curved, sometimes clavate; stigma 2-lobed, rectangular. Capsules 10 to 30 mm long, erect, straight to slightly incurved, glabrous. Seeds circular to suborbicular, flattened but plump, 1.4 to 1.6 mm in diameter, shiny black, without a ring margin, edges obtuse, concentrically minutely and obscurely muricate. Species as a whole from the Great Plains to OK and TX, the Midwest, and the SE. U.S.
There are 3 varieties in TX. All of our material seems referable to the following. Another subspecies, subsp. micrantha, grows in NE. TX. It has shorter racemes, smaller flowers, a globose-tipped spur, and stout fruits 15 mm long or less. It may yet be discovered in the far NE. portion of our area.
subsp. australis (Chapm.) G. B. Ownbey Chasmogamous-flowered racemes elongate, usually greatly exceeding the leaves. Spur of upper petal short, saccate, never with a distinctly globose tip. Fruits slender, erect, 15 to 30 cm long, seeds nearly smooth. Disturbed soils, commonly sands, old fields, roadsides, open woods, and waste areas. E. and SE. TX; S. MO and SE. KS, S. to TX, E. to NC. Feb.-Apr. [C. micrantha Engelm. & Gray var. australis (Chapm.) Shinners; C. campestris (Britt.) Rydb.; C. halei (Small) Fern.].
Deciduous trees, bark commonly exfoliating, branches often widespread. Leaves alternate, simple, generally palmately lobed and veined; petiole inflated and hollow at the base to enclose the axillary bud; stipules encircling the twig, membranous, early-deciduous and leaving a circular scar. Plants monoecious; flowers small, actinomorphic, unisexual, in dense, long-peduncled globose heads. Sepals and petals each 3 or 4(7), usually reduced or sometimes absent, sepals distinct or basally connate. Staminate flowers with the petals, if any, alternate with the sepals; anthers many, opposite the sepals, sessile to subsessile, subtended by minute scales, linear, 2-celled, the connective with a peltate appendage at the apex; sometimes vestigial pistils present. Pistillate flowers without a corolla, often with 3 or 4 staminodia; carpels free, (3)5 to 8(9) in 2 or 3 whorls, subsessile, each pistil 1-celled, the apex incompletely sealed, stigma unilateral along the inner face of the linear-subulate style, ovules 1(2). Fruits achenes or nutlets, indehiscent, single-seeded, interspersed in the head with linear bracts and tufts of hairs, the cluster globose, dangling. Fruits wind-dispersed; seeds within linear-fusiform.
One genus with 6 or 7 species in the N. hemisphere; we have the one species found in TX.
The wood of some species is used for timber; some are planted for shade, especially in urban areas where they are tolerant of pollution (Mabberley 1987).
This species common in much of TX.
1. P. occidentalis L. Pyramidal trees to 50 m tall, trunk to 30 cm in diameter. Bark of young trees mottled green-gray and white because of large exfoliating patches. Older trees with shallowly-furrowed, gray-flecked bark; twigs tan or brown, commonly tomentose when young, soon becoming glabrate. Leaves broadly ovate to reniform in overall outline, generally 10 to 20 cm broad, palmately 3- or 5-lobed and -veined, the lobes broadly triangular, acuminate, entire or with a few remote teeth, the sinuses round and shallow, surfaces more or less glabrous or else floccose or stellate pubescent on the main veins and/or below; petioles 1/2 the length of the blade or less. Perianth reduced or absent. Staminate heads 8 to 10 mm broad, yellow-green; staminodia often present in male flowers. Pistillate heads reddish, 10 to 12 mm across, female flowers without corolla. Fruiting heads long-peduncled, dense, 2 cm or more in diameter, often remaining on the tree through at least part of the winter, light yellow to tawny, at last falling apart readily to release the brownish, clavate achenes. Achenes 7 to 8 mm long; seeds inside ca. 6 mm long. Generally in bottomlands, common along streams. Throughout most of TX E. of Val Verde Co.; ME W. to Ont., NE, S. to N. FL, LA, and TX. Mar.-May, collected with fruit throughout the year. Fall color dull tan. [Includes var. glabrata (Fern.) Sarg. and forma attenuata Sarg.].
Sometimes planted for its shade and peeling bark, but in our area this tree generally prefers a moister environment and often succumbs to an anthracnose infection before reaching maturity. The wood has been used for furniture, boxes, and interior wood work, but it is not very strong. The fruits, though plentiful, have little value as a wildlife food (Elias 1980).
Shrubs or trees. Leaves alternate, simple, often palmately lobed, commonly with stellate pubescence; usually stipulate. Plants usually monoecious, some-times polygamous; flowers in heads, clusters, or spikes, perfect or unisexual, usually regular and wholly or partly epigynous, wind- or insect-pollinated. Calyx of 4 or 5(-10) sepals, small, free or connate, adherent to the base of the ovary, sometimes reduced or absent. Petals 4 or 5, small and narrow, inserted on the calyx, sometimes absent. Stamens 4 or 5(10), alternate with the petals, sometimes alternating with an equal number of staminodia or else stamens up to about 32; sometimes staminodia or petal bases nectariferous; anthers dehiscent by long valves or slits, the connective extended. Carpels 2(3), united at least below, styles separate, placentation generally axile, each locule with 1(2 to several) ovules. Fruit woody, capsular, septicidal at the apex (sometimes also loculicidal), 2-beaked, 2-celled, usually with 1 or 2 seeds per cell. Seeds with a hard testa.
About 28 genera and 100 species in the subtropical and warm temperate regions of both hemispheres, largely in E. Asia; 2 genera and 3 species in TX: 1 species in our area.
Some species are valuable for timber; others are aromatic and have extracts used in perfumes or medicines (e.g. Witch-hazel, Hamamelis, from which an astringent is obtained.) Some are cultivated as ornamentals for the flowers (e.g. Hamamelis, which blooms in winter) or for shade and fall color (e.g. Liquidambar) (Mabberley 1987).
Roughly 1 American and 2 Asian species.
1. L. styraciflua L. Sweet-gum, Gumball Tree. Deciduous, pyramidal tree to 40 m tall. Bark gray-brown, deeply fissured; branches sometimes with corky ridges; twigs greenish and minutely pubescent when young, becoming light brown and glabrous, often angled; buds sharply ovate, scales reddish-brown. Leaves on slender petioles to 12 cm long; blades somewhat leathery, fragrant when crushed, generally orbicular in overall outline, to ca. 18 cm long and 15 cm wide, palmately (3-)5- to 7-lobed, star-shaped, basal lobes the smallest, lobes acute (rarely rounded), margins usually glandular-serrate, base cordate to truncate, upper surface smooth and shining, lower surface paler, sometimes with small tufts of hairs in the axils of the major veins. Plants monoecious, flowers unisexual and apetalous. Staminate flowers entirely without perianth, filaments ca. 1 mm long, grouped with small scales into globose heads arranged in more or less upright racemes to about 6 cm long. Pistillate flowers crowded and somewhat coherent in globose, capitate clusters 2 to 3(4.5) cm in diameter, pendant on peduncles to 6 cm long or more, each with a short, lobeless calyx tube about as long as the ovary, some staminodia present; carpels 2, styles 2, stigmatic along the inner surface. Fruiting structure woody, brown, ca. 3 cm in diameter, each capsule splitting open between the 2 persistent, hard styles to give the cluster an overall mace-head effect; each capsule with 1 or 2 brown, winged seeds 6 to 9 mm long and numerous abortive flake- or sawdust-like seeds. Wet places, low or wet woods, often with pine, oak, or hickory. E., S., and Cen. TX, W. to Lee Co.; FL to TX and Mex., S. to Cen. Amer.; N. to CT, NY, WV, OH; W. to IL and MO. Blooming Mar.-May, fruits maturing in the fall, some usually persisting on the tree through the winter. Fall color yellow through gold, orange, red, and maroon to purple--often all on the same tree.
Trees valued as a source of timber, the somewhat weak wood used mainly for veneer and plywood and as a source of resin. Commonly planted for fall foliage; cultivars are available with selected colors. The sap or gum is supposed to be pleasant to chew. The hard, spiky fallen fruits can become a nuisance on lawns, but can be used in dry floral arrangements. The seeds are not an important wildlife food, but are eaten by some birds and squirrels (Elias 1980).
Trees and shrubs with watery sap. Bud scales imbricate. Leaves alternate (very rarely opposite and never so in ours), commonly 2-ranked, simple, the base often oblique, entire to commonly serrate or doubly serrate; stipules paired, deciduous. Plants monoecious and/or with perfect flowers, flowers usually in fascicles or racemes or solitary in the axils of leaves or bracts, on previous or current season's growth, regular to slightly irregular, hypogynous to perigynous, small, greenish, individually not showy. Sepals (2)4 to 5(9), free or commonly fused at least partially, persistent. Corolla absent. Stamens usually as many as the sepals and opposite them (sometimes twice as many or up to 15), hypogynous or inserted on the calyx tube, filaments curved or sigmoid. Ovary superior, 2(3)-carpellate, 1-celled and 1-seeded (rarely bilocular and 2-seeded), style 2-parted and stigmas 2. Fruit a samara, drupe, or nutlet with a leathery coat.
16 genera and about 140 species from the tropics to temperate regions, especially in the N. hemisphere; 3 genera and about 12 species native to TX; 3 genera and 7 species in our region.
Some, especially elms (Ulmus), are important timber trees; others are cultivated for shade and ornament or have edible fruit (Celtis). A few species provide food for wildlife (Mabberley 1987).
1. Leaves with 3 major veins arising from the base; fruit a fleshy drupe ......................1. Celtis
1. Leaves pinnately veined; fruit a samara or nutlet with a spiky coat ........................................2
2(1) Fruit a samara; flowers perfect and appearing before the leaves or in the fall; leaf margins usually doubly serrate ................................................................................................2. Ulmus
2. Fruit a nutlet with spiky projections; flowers (at least some) unisexual and appearing with the leaves; leaf margins simply serrate ...................................................................3. Planera
Trees or less commonly shrubs; bark usually gray (sometimes pinkish in our C. laevigata), smooth or furrowed, often noticeably warty, branches sometimes spiny, winter buds small. Trees deciduous in our region. Leaves petiolate, with 3 main veins arising from the base, ovate to lanceolate, entire to serrate, sometimes coriaceous; stipules lateral, free, scarious. Plants monoecious and/or with perfect flowers; flowers appearing with leaves, produced on current season's growth; staminate flowers 1 to few in cymes or fascicles below, female or perfect flowers solitary or in few-flowered fascicles in the axils of the upper leaves. Calyx deeply to slightly (4-)5-(6-)lobed, the lobes imbricate. Stamens equalling the calyx lobes, inserted on the receptacle, incurved in bud, later exserted, anthers ovate, extrorse. Pistillate flowers usually with non-functional stamens; gynoecium of staminate flowers rudimentary, in female flowers with a 2-lobed style, the lobes entire to bifid, stigmatic on the inner surface, ovary unilocular, sessile. Fruit a glaucous, subglobose to ovoid drupe, thin-fleshed, somewhat dry, with a firm exocarp and a sweet to insipid pulp; seed bony, the surface smooth to rugose; maturing in fall and often persisting through the winter. About 60 mostly tropical species; 6 in TX; 2 here. No two sources agree on the classification of the species in the SE. U.S.
The fruits of many species are edible by humans and wildlife (Elias 1980) and were a common food of native Americans (Kindscher 1987). The wood is used for fuel and fence-posts. Some species will yield a yellow dye from the bark; ours yield tan from twigs and leaves. The flexible branches can be used for weaving baskets (Tull 1987). Some people are allergic to the pollen.
1. Leaves ovate-lanceolate to elliptic to lanceolate, the apex generally sharply acute to acuminate; upper surface usually smooth, sometimes scabrous ..................1. C. laevigata
1. Leaves broadly to narrowly ovate, the apex obtuse to abruptly long-acuminate; upper surface commonly scabrous ...........................................................................2. C. reticulata
1. C. laevigata Willd. Sugarberry, Hackberry, Palo Blanco. Broad-crowned tree to 30 m tall, branches spreading to pendulous. Bark sometimes smooth, in our area usually covered with corky warts or ridges (especially older trees), light gray or our trees commonly pinkish from a light film of lichen; twigs pubescent, becoming glabrous with age. Petioles 6 to 10 mm long; blades lanceolate to oblong-lanceolate or ovate-lanceolate, entire to somewhat serrate, the base rounded (to broadly cuneate), often oblique, apex typically long-acuminate or acute, often curved, texture from thin to coriaceous, both surfaces the same color, upper surface smooth, scabrous, or lightly hirtellous, leaves of fruiting branches usually less than 1/2 as broad as long, 4 to 10 cm long, 1.5 to 4.5 cm broad. Drupes subglobose, 5 to 8 mm in diameter, without a style beak, orange to brown or reddish at maturity, on pedicels 6 to 15 mm long, the stone 4.5 to 7 mm long, 5 to 6 mm broad. Various soils, along streams, in woodlands, thickets, etc. E. 2/3 TX; VA to FL and TX, N. to OK, KS, MO, IN, MN; also NE. Mex. Flowering early spring, fruit ripening in the fall. [C. mississippiensis Bosc.; C. berlandieri Kl.]
Some authors recognize varieties of this species, though the usefulness of this is somewhat doubtful. If varieties are recognized, our plants are probably mostly var. laevigata--leaf blades smooth, membranous, entire and petioles glabrous. Var. texana Sarg. has thicker, entire, scabrous leaves and pubescent petioles. Var. smallii (Beadle) Sarg. [= C. smallii Beadle] has thin, essentially glabrous leaves and petioles and toothed margins.
Our plants are often characterized by the presence of nipple gall--the leaves with small, raised circular or ring-shape swellings to ca. 5 mm across. The twigs also often bear semispherical, brown insect galls to ca. 1.5 cm broad, commonly mistaken for fruit. The real fruits are edible, but vary from tree to tree in palatability and are generally insipid.
2. C. reticulata Torr. Netleaf Hackberry, Palo Blanco. Small tree or large shrub 7(to 16) m tall, trunk to a rare maximum of 6 dm in diameter. Bark gray with corky ridges, branches often crooked or flattened, young branches villous. Petioles 3 to 8 mm long, pubescent, sometimes grooved above; blades thickish, ovate, 3 to 7 cm long, 1.5 to 4 cm wide, the base cordate to sometimes oblique, the tip obtuse to acute or subacuminate, margin entire or with some teeth in the distal half, upper surface scabrous, lower surface pubescent, paler and somewhat yellowish, strongly reticulate-veined. Drupes on pedicels 10 to 14 mm long, spherical, reddish or reddish-black at maturity, 8 to 10 mm in diameter, the flesh sweet. Our plants represent the larger-leaved phase with leaves generally longer than 4.5 cm and occurring in E. TX. The smaller-leaved phase is found from the Panhandle S. to the Trans Pecos, Ed. Plat., and Brown, Sutton, and Tom Green Cos. The two may someday be formally separated. The species as a whole on limestone hills, oak dunes, canyon slopes, mesas, and in arroyos, mesquite groves, and near water in dry regions. TX and OK, KS, and CO, W. to ID, WA, and CA; also N. Mex. Flowering in spring, the fruit maturing in the fall. [C. laevigata Willd. var. reticulata (Torr.) Benson; C. rugulosa Rydb.; C. rugosa Rydb. (not of Willd.)].
Kartesz (1998) submerges this species in C. laevigata, but in our area the two are distinct morphologically and show different habitat preferences.
Trees or rarely shrubs, deciduous or rarely semi-evergreen. Bark usually deeply furrowed, branches terete, slender, often with corky wings. Buds conspicuous, with many rounded-ovate, brownish, imbricate scales. Leaves petiolate, 2-ranked, usually oblique at the base, markedly pinnately-veined, mostly doubly, sometimes singly serrate. Stipules lateral, linear-lanceolate to obovate, entire, scarious, free or fused at the base, enclosing the leaf in the bud, later deciduous. Flowers perfect, in axillary cymes, fascicles, or racemesappearing before the leaves in spring or else in the fall. Calyx campanulate. Corolla absent. Stamens as many as the calyx lobes, filaments filiform to flattened, exserted after anthesis, anthers oblong, extrorse. Ovary sessile to stipitate, style 2-lobed. Fruit a 1-celled, 1-seeded, flattened samara, glabrous to pubescent and/or ciliate, ripening a few weeks after flowering.
18 species, primarily of N. temperate regions, to as far S. as N. Mex.; 5 species listed for TX; we have 4 plus one cultivated species apparently escaping. (See note below key.)
Some elm species have durable wood while others have wood with little practical use. Many have graceful forms and are planted as street and lawn trees. The best known elm of the U.S. is U. americana, once widely planted and with a stately habit, now almost entirely wiped out in many areas by Dutch elm disease (Mabberley 1987). Most elm fruits are only a minor source of food for birds; deer and rabbits may browse the twigs (Elias 1980). The pollen can be allergenic.
1. Leaves mostly less than 5 cm long; branches often with corky wings ...................................2
1. Leaves mostly longer than 5 cm; branches without corky wings ............................................3
2(1) Flowers appearing in the spring; leaf apices acute to acuminate, upper surface of leaf smooth ......................................................................................................................1. U. alata
2. Flowers appearing in the fall; leaf apices obtuse to rounded, upper surface of leaf
scabrous .........................................................................................................2. U. crassifolia
3(2) Flowers on slender, drooping pedicels; fruits glabrous on the faces, margins ciliate; leaf buds glabrous to slightly white-pubescent; leaves generally smooth above, sometimes scabrous ..........................................................................................................3. U. americana
3. Flowers short-pedicellate in dense fascicles, not pendulous; fruits pubescent but without marginal cilia; leaf buds with long rusty hairs; leaves usually very scabrous above ................
.................................................................................................................................4. U. rubra
Note: U. parvifolia Jacq., the Chinese Lacebark Elm, is cultivated in our area and apparently is capable of reproducing here. Native to Japan and China, it is easily recognized by patchy, exfoliating gray and orange bark on older trees. The leaves are semievergreen in mild winters, coriaceous, elliptic to elliptic-ovate, bases nearly equal, glabrous above, singly serrate, apices acute. Samaras glabrous, ovate-orbicular, with an apical notch. Flowering and fruiting in late summer to fall. U. pumila L. (Siberian Elm, Asiatic Elm, Chinese Elm) is commonly cultivated and sometimes escapes in the U.S. (including TX) but neither commonly planted nor escaping here.
1. U. alata Michx. Winged Elm, Wahoo, Cork Elm. Small to medium (rarely very large) tree, to ca. 20 m, branches spreading and crown round-topped, trunk to a maximum of 1 m in diameter. Bark light brown to reddish brown or gray, shallowly and irregularly furrowed. Twigs and branchlets pubescent when young, becoming glabrescent, often with thin, broad corky wings beginning the first or second year of growth, but some branches and some whole trees entirely without. Buds acute, glabrous to finely pubescent. Leaves short-petiolate, petioles 1 to 3 mm long; blades ovate, ovate-oblong, or oblong-lanceolate to elliptic, 3 to 6 cm long, 1 to 3.5 cm broad, sometimes larger, sometimes slightly curved, acute to acuminate, base oblique, rounded to somewhat cordate, doubly serrate, upper surface usually dark green, smooth, lower surface paler and minutely pubescent. Flowers in short, few-flowered racemes. Calyx 5- to 9-lobed; stamens usually 5, trees in full flower or fruit often with a "lacy" appearance. Samara ovate-elliptic to oblong, flat, 6 to 9 mm long, persistent style beaks long, slender, incurved, the whole villous, margins conspicuously ciliate with hairs longer than 1 mm. Woodlands, often with oak, along streams, fencerows; generally in upland areas. E. 1/3 TX; VA W. to IN, IL, and MO, S. and W. to KS, OK, and TX, E. to Cen. FL. Feb.-Mar., fruit maturing a few weeks later and soon falling. Fall color dull yellow to tan or occasionally yellow-orange.
Sometimes planted as an ornamental in our area, or left growing on lots cleared for construction and thus part of the final landscape. Not to be confused with species of Euonymus, also called Wahoo.
2. U. crassifolia Nutt. Cedar Elm, Olmo. Small to medium-sized tree to 20(25) m tall, crown spreading and round-topped, trunk 0.3 to 0.6 m in diameter. Bark light-brown to reddish-brown, with deep furrows and broad, scaly ridges. Branchlets pubescent when young, becoming glabrate, reddish-brown, often with two broad corky wings (this character cannot be used to separate this plant from U. alata). Buds small, acute, more or less glabrous. Petioles 6 to 9 mm long. Blades ovate to ovate-oblong or elliptic, 2.5 to 6 cm long, 1.9 to 3 cm broad, generally obtuse or even rounded apically, only sometimes acute, doubly serrate or sometimes nearly singly serrate, stiff, shiny green and usually decidedly scabrous above, soft-pubescent beneath. Flowers in short-stalked clusters of 3 to 5. Calyx deeply 4- to 8-lobed. Samara ovate, oblong, or sometimes nearly orbicular, 6 to 12 mm long, deeply notched at the tip, short-villous, margins ciliate with hairs usually less than 1 mm long. Woods, in our area especially bottomland woods with oak, ash, and hackberry; ravines, open slopes, and along streams, commonly on limestone soils but adapting to various types. July-Oct. with fruit remaining sometimes until Nov. Fall color generally tan, sometimes yellow. Cen. and S. TX; MS to LA, AR, and TX.
The fall flowering distinguishes this from all our other native elms. Often planted as a street tree.
3. U. americana L. American Elm, White Elm. At maturity a tall tree to 30(40) m, trunk often dichotomously branched, canopy "vase-shaped," wide-spreading, branches pendulous; trunk to a maximum of 3 m in diameter. Bark gray to gray-brown, scaly, with deep, irregular, intersecting fissures separating broad, flat ridges. Young branchlets pubescent or sometimes glabrescent, without wings. Winter buds acute, glabrous or only slightly white-pubescent. Petioles (2)5 to 8(10) mm long; blades ovate to elliptic or ovate-oblong, (4)7 to 15 cm long, (2)5 to 7.5 cm broad, base oblique, apex acuminate (sometimes abruptly so), doubly serrate, glabrous and smooth to more or less scabrous above, paler below and pubescent to glabrous, commonly with minute tufts of hair in the axils of the main veins. Pedicels drooping, 1 to 2 cm long, unequal in each cluster. Calyx 5-to 9-lobed; stamens 5 to 9, exserted, stigmas white. Samara flat, elliptic to orbicular, 0.8 to 1.5 cm long, deeply notched at the apex, notch reaching the seed, glabrous, but with marginal cilia more than 1 mm long. Lowland areas with rich soil along streams and rivers. E. 1/3 TX; FL N. to Newf., W. to Man., S. to TX. Feb.-Apr., the fruits maturing in late spring. Fall color yellow.
Once a common street and lawn tree, especially in the E. U.S., now almost completely destroyed by Dutch elm disease. A few resistant trees remain, and the tree is still found over much of its former wild range but individuals rarely surviving to reach their full potential and not nearly so numerous as formerly. Breeding and selection programs are underway to develop resistant strains. The wood, if properly seasoned, is durable and used for furniture, boxes, and shipbuilding. Baskets can be made out of elm splints. The seeds are an important food for game birds and rodents. Deer and rabbits browse the twigs (Elias 1980).
4. U. rubra Muhl. Slippery Elm, Red Elm. Medium-sized tree, 15 to 25(40) m tall, canopy spreading, broad and open. Bark gray-brown to dark brown or reddish-brown, with shallow, irregular fissures separating flat ridges; inner bark mucilaginous. Branchlets finely pubescent and scabrous when young, becoming glabrate, red-brown to orange or gray-brown. Leaf buds large, narrowly ovoid, obtuse. Flower buds with long rusty pubescence, 4 to 5 mm long. Petioles 4 to 9 mm long; blades obovate to ovate or elliptic, 8 to 20 cm long (sometimes longer), 5 to 7.5 cm broad, very asymmetrical at the base, the tip long-acuminate, sometimes abruptly so, margin doubly serrate, upper surface very scabrous, paler and minutely and somewhat densely soft-pubescent below. Flowers subsessile to short-pedicellate in few-flowered fascicles. Calyx with 5 to 9 rounded lobes, green, 0.5 to 0.7 mm long, fringed with red-brown hairs; stamens 5 to 8(9), filaments 4 to 5 mm long, stigmas pinkish. Samaras suborbicular to elliptic or broadly obovate,flat, 1 to 1.2(2) cm long, rusty-pubescent on the faces, without marginal cilia, only slightly notched at the tip. Woods, thickets, and along rivers and streams. E. 1/3 TX except the far SE.; FL and TX N. to ND, MN, Ont., and Que. Feb.-Apr., the fruit maturing in late spring. Fall color dull yellow. [U. fulva Michx.].
The mucilaginous inner bark was used in medicines to treat fevers and inflammations. The Indians used the bark as a substitute for birch bark in making canoes. The seeds are of minor value as a wildlife food. The wood can be used for some furniture, paneling, and boxes, but is inferior to American Elm (Elias 1980).
A monotypic genus of the S.E. U.S.
1. P. aquatica (Walt.) J. F. Gmel. Water-elm, Planer-tree. Small tree or large shrub to 15 m, branches spreading, crown low and broad; trunk short and with flaky, gray or light-brown bark; branchlets pubescent when young, becoming glabrous with age, reddish-brown to gray; winter buds 6 to 12 mm long, subglobose. Leaves simple, alternate, deciduous; petioles 3 to 6 mm long; blades ovate to ovate-elliptic, -rhombic, or -oblong, 2.5 to 8 cm long, 1.3 to 4 cm broad, pinnately nerved (though not so strongly as in Ulmus), base cuneate to rounded and oblique, apex acute, margin singly and irregularly serrate, smooth to scabrous above, paler below, glabrous at maturity. Plants polygamo-monoecious (primarily monoecious, but with some perfect flowers). Calyx deeply 4- or 5-lobed. Staminate flowers in clusters at the base of the young branchlets; stamens 4 to 5. Pistillate flowers 1 to 3 in the axils of the first leaves, sometimes with 1 stamen and then perfect. Fruit stipitate, ellipsoid or ovoid, ca. 8 mm long, the outer surface coriaceous and with irregular fleshy ribs and projections; seed inside flattened, broadest near the base. Swamps, floodplains, and along sluggish streams and lakeshores; withstanding periodic flooding. Uncommon, but often plentiful where found. FL to TX and AR, up the Mississippi floodplain to IL and KY, along the E. seaboard to VA. Blooming Feb.-Mar.(?), with the fruit maturing ca. 6 weeks later; our collections primarily April. Waterfowl eat the fruits. The wood is weak and subject to decay and so is not of much use (Elias 1980). Not much collected.
Erect or twining herbs with watery sap. Leaves simple and unlobed or palmately 3- to 7-lobed or else palmately compound, opposite or opposite passing to alternate above; stipules free, persistent; herbage with glandular hairs. Plants dioecious, rarely monoecious, flowers small, anemophilous, inflorescences in the axils of the upper (often reduced) leaves. Staminate flowers few to many in open cymes or panicles; sepals 5, corolla absent, stamens 5, opposite the sepals, floral parts spirally arranged. Female flowers few in more compact cymose, spiciform, or amentiform arrangements; calyx tube short, membranous, enclosing the ovary or else reduced to a minute ring; gynoecium superior, of 2 united carpels, unilocular, style 1, with 2 elongate, dry stigmas. Fruit a small nut or achene with or without the persistent calyx.
2 genera and 3 species native to the N. temperate region. No species are native to TX, but one species, Cannabis sativa, is occasionally found as an escape from cultivation. The other genus, Humulus, includes the hops used to flavor beer.
This family is very closely allied to the Moraceae and is sometimes included therein.
One species. Wild material cannot be pinpointed with certainty, but the plant has existed for centuries in cultivation (Mabberley 1987).
1. C. sativa L. Hemp, Marijuana, etc. Erect, annual, usually dioecious herb 0.5 to 4 m tall. Stem single, well-branched, antrorsely appressed-pubescent, pungent. Lower leaves opposite, upper chiefly alternate, petioles 2 to 11 cm long; blades palmately compound with 5 to 9(11) leaflets; leaflets linear to linear-lanceolate, 4 to 15 cm long, 3 to 20 mm broad, the middle leaflet the largest and the leaflets progressively shorter outwards; upper surface strigose, lower surface sparsely to densely pubescent, apex acute, margins serrate. Stipules 5 to 20 mm long, lanceolate to linear-lanceolate. Flowers in small cymose clusters in the axils of the upper leaves. Staminate flowers with pedicels 0.5 to 2 mm long, sepals 2.5 to 4 mm long, ovate to lanceolate, puberulent and usually with hyaline margins, stamens slightly shorter than the sepals; filaments 0.5 to 1 mm long. Female flowers sessile, partly surrounded to almost enclosed by the abruptly acuminate, stipitate-glandular bract; calyx usually very short, if longer, then hyaline and eventually adnate to the fruit. Achenes varying from greenish to buff or mottled with purple, 2.7 to 4.2 mm long, 2 to 2.8 mm broad, ellipsoid to ovoid, smooth. Blooming summer to fall. Naturalized or escaped from Asia. Que to B.C., S. throughout the U.S.
Cannabis exists as 2 cultivated subspecies. Subsp. sativa is a more northern element, grown for hemp fibers in various places throughout the world and sometimes escaping. For example, it is fairly common in the plains states (GPFA 1986). Subsp. indica (Lam.) E. Small. & Cronq. is the southern element and represents a lineage selected for high levels of psychoactive compounds. The plant and its compounds are used in various raw or refined forms--"pot", "ganja", hashish, etc. Oil from the seeds is used in paint, varnish, and foods, and the seeds are used in birdseed (Mabberley 1987). The plant is known to escape its illegal cultivation in TX. However, most plants encountered in our area will be under cultivation, often in small, hidden, wild-appearing plots in the middle of wooded country.
Trees, shrubs, vines, or rarely herbs, sap almost always milky (not so in Fatoua). Leaves alternate or opposite, stipulate and petiolate, simple, entire to lobed, our species deciduous. Plants monoecious or dioecious, flowers small, in axillary catkins, in oblong to globose clusters, inserted on a torus, or inside a hollow receptacle. Corolla absent. Staminate flowers usually with 4 or 5 free to united sepals; stamens as many as the sepals and opposite them. Female flowers often more or less united by the receptacles; sepals usually 4, more or less fused; gynoecium superior to inferior, bi- (or tri-) carpellate, usually 1-celled and uniovulate, style simple or 2-branched, stigmas 1 or 2. Fruit a small drupe or achene-like, usually enclosed by the fleshy calyx, often assembled into a multiple fruit which often includes the fleshy common receptacle as well .
About 1,200 species in 48 genera, generally in the tropics and warm temperate regions; 5 genera and 7 species in TX; 4 genera and 5 species here.
This family includes many fruits--e.g. figs (Ficus), breadfruit (Atrocarpus), and mulberry (Morus). Some species are useful for timber or paper pulp. Many are cultivated as ornamentals, including many species of Ficus (Mabberley 1987).
1. Plants herbaceous ....................................................................................................1. Fatoua
1. Plants trees or shrubs ...............................................................................................................2
2(1) Leaves entire; plants with some axillary thorns; fruits spherical, to 15 cm in diameter, wrinkled, milky-juiced ...............................................................................................2. Maclura
2. Leaves toothed and/or lobed; thorns absent; fruit not as above, less than 3 cm in diameter .
3(2) Mature twigs glabrous; pistillate flower cluster and fruit ovoid to cylindric; buds with 3 to 6 scales; lower surface of leaves green, glabrous to soft pubescent; style 2-branched ............
3. Mature twigs pubescent; pistillate flower cluster and fruit globose; buds with 2 to 3 scales; lower surface of leaves gray-green, velvety-tomentose; style entire ..........4. Broussonetia
One species native from Japan to Formosa; weedy elsewhere. First found in N. Amer. in 1950 (Darwin, et al. 1981), it quickly spread (Massey 1975) and reached TX in 1982 (Lipscomb 1984). This description owes much to that of Liu and Liao (1976).
1. F. villosa (Thunb.) Nakai Herbs, ours apparently annual, from a short taproot and fibrous roots. Stems to ca. 1 m (?) tall, erect, commonly solitary, with very short, curled, often capitate hairs and some longer pilose hairs; pith white-fibrous, especially in younger stems. Leaves alternate, stipulate, petiolate, petioles to ca. 1/2 as long as blades or more; blades ovate-cordate, to ca. 12 cm long, smaller on the younger, upper portions, margins crenate-dentate, apex acumiante, base broadly cuneate to obtuse at the point of attachment to the petiole but with a rounded lobe on either side, the basal sinus thus "M" shaped; surfaces with punctate cystoliths (mineral concretions); uppper surfacae scabrous-puberulent, lower surface somewhat paler, less pubescent, the hairs mostly on the veins. Plants monoecious, flowers in pubescent axillary cymes, these somewhat elongated or one-sided with the male flowers on one edge. Male flowers with 4 more or less ovate sepals; stamens 4, filaments and anthers commonly white. Female flowers with 4 more or less ovate sepals; style arms 2, but one of them very minute, the other pink-purple, plumose; ovary oblique. Fruiting calyx membranaceous, speals often purple-tipped. Achene golden brown, ca. 1 mm long, essentially trigonous and blunt, but with a smaller, flat fourth face with the attachment scar of the style; surface minutely tuberculate. Weedy, usually around greenhouses, nurseries, cultivated areas, etc. Native to Japan and Formosa, now apparently established in LA, AL, FL, GA, MS, NC, and TX; spreading rapidly. Supposedly introduced into our area in a pot plant from S. TX. Our specimens collected with flower and fruit in June. First collections here in 1991 (TAMU 22437).
This plant is a new member of the local flora. We will doubtless see more of it in the future. It is very reminiscent of the Urticaceae, and was in fact first described as an Urtica. Nakai (1927) placed it in the Moraceae. It is also superficially similar to some species of Acalypha.
A monotypic genus.
1. M. pomifera (Raf.) Schneid. Osage Orange, Bois d'Arc, Bow-wood, Naranjo Chino. Small to medium-sized tree to 12(20) m tall; crown open, irregular, round-topped; trunk to a maximum of 1 m in diameter; bark orange-brown to yellow-brown, with deep, irregular fissures; sap milky. Branchlets light green turning light brown, pubescent at first, soon glabrous; branches with thorns to 2.5 cm long, plants sometimes forming thickets. Leaves deciduous, alternate or clustered on lateral spur shoots, petioles 3 to 5 cm long, usually pubescent; blades 4 to 12.5 cm long, 2 to 6(7.5) cm wide, ovate to elliptic to lanceolate, entire, base broadly cuneate to rounded or subcordate, apex acuminate, bright green and lustrous, usually pubescent on midrib and veins above, somewhat paler and more pubescent beneath; stipules minute, caducous. Flowers 4-merous. Staminate flowers pedicellate, in axillary, pedunculate, globose clusters 1.5 to 3.5 cm long in groups of 1 to several on short spur shoots. Pistillate flowers sessile, coherent by the receptacles in pedunculate, globose heads 1.5 to 2.5 cm in diameter, clusters borne singly in the axils; styles conspicuous, elongate and filiform. Fruit a syncarp (multiple), globose, 6 to 15 cm in diameter, green, milky-juiced, with a wrinkled "rind" composed of the fleshy calyces. Achenes 8 to 12 mm long, buried deep within the ball. Fence rows, fields, roadsides, waste places. Common in the E. 1/2 TX; also AR, LA, OK. Apr.-May, fruits falling in autumn. Fall color yellow. [Toxylon pomiferum Raf.].
Widely cultivated as a fence and hedge plant outside its normal range in the SE U.S. (GPFA 1986). The wood is orange, very hard and strong; used for posts, railroad ties, and by the Osage Indians for bows (Elias 1980). A yellow dye can be made from the roots (Tull 1987). The leaves have been used to feed silkworms (Mabberley 1987). The sap causes contact dermatitis in some people (Lampe 1985). The large, "brain-like" fruits are distinctive and have great recreational value, though they are of virtually no importance as a wildlife food (squirrels will sometimes tear them apart to eat the achenes) (Elias 1980). The TAMU herbarium receives more identification requests for this plant than for any other.
3. MORUS L. Mulberry, Moral
Trees or large shrubs with milky sap. Branches often spreading, bark usually scaly or furrowed; twigs glabrous or only sparsely pubescent, winter buds with 3 to 6 imbricate scales. Leaves deciduous, simple, entire or palmately lobed (especially on young trees and vigorous shoots), with 3 to 5 major veins from the base, serrate to dentate; stipules lanceolate to linear-lanceolate, deciduous. Plants monoecious or dioecious (as in ours), flowers of both sexes in axillary, peduncled aments or the female in short to long cylindrical clusters (rarely, and not in ours, male and female flowers in the same cluster.) Calyx 4-parted. Stamens 4. Pistil 1, superior, style 2-branched, persistent, stigmas 2. Fruit an achene or nutlet, ovoid, flattened, enclosed by the succulent calyx to form a small drupelike fruit topped with the style; fruits combined in a sweet, juicy multiple fruit resembling a blackberry and white, pink, or purple to black in color.
7 to 12 species (depending on interpretation) of the subtropical and temperate N. hemisphere; 3 in TX; 2 here, 1 of which is naturalized from China.
Mulberries are grown in Europe and Asia for the edible fruits (Mabberley 1987), but in N. America, the mulberry is more a wildlife food--a favorite of songbirds and small mammals. The wood tends to be brittle, but some species are used for timber and small ariticles (Elias 1980). The leaves, especially of M. alba, are the principal food of silk-worms. The Tonkawa Indians of TX used the fibrous inner bark to make cloth, and the branches can be used in basketweaving. Tan or yellow-green dyes can be obtained from leaves and twigs, while lavenders to purples come from the berries (Tull 1987). Some species are grown as ornamentals or for shade, though many people are allergic to the pollen, and the fruits (which will stain anything) can be messy. The sap can cause contact dermatitis (Lampe 1985).
1. Lower leaf surface glabrous or pubescent on the major veins and/or in the axils of the veins only; upper surface usually smooth ................................................................1. M. alba
1. Lower leaf surface usually soft-pubescent throughout; upper surface often scabrous, sometimes smooth ..................................................................................................2. M. rubra
1. M. alba L. White Mulberry, Russian Mulberry, Silkworm Mulberry, Moral Blanco. Small tree to ca. 15 m or large shrub, branches spreading, crown round-topped. Young branchlets slightly pubescent or glabrous at first, becoming glabrous. Petioles 1 to 2.5(5) cm long; blades ovate to elliptic-ovate or (if lobed) rotund-ovate in overall outline, to 12(20) cm long, larger on more vigorous shoots, base somewhat oblique, rounded, or cordate, apex sub-obtuse to acute or acuminate; unlobed or with 2 to 5 palmate lobes, often mitten-shaped, margins serrate, glabrous and smooth or rarely very slightly rough above, lower surface lightly spreading-pubescent on the larger veins and/or with small tufts of hair in the axils of the major veins. Staminate aments pendulous, 1 to 4 cm long. Pistillate flower clusters 5 to 20 mm long, flowers small, greenish. Multiple fruits ovoid to short-cylindric, 5 to 25 mm long, white, pink, reddish, or blackish-purple when ripe, sweet but rather insipid. Woods, rocky areas, roadsides, fencerows, etc. Mostly in Cen. and S. TX; native to China, naturalized throughout much of Europe and N. America. Mar.-Apr. Fall color yellow. [M. nigra L.].
This species was introduced into the U.S. in a failed attempt to start a silk industry, the leaves being the chief food of the silkworm (Elias 1980). Often planted in yards, especially in far W. TX where it is the major shade tree in some towns. There it is able to withstand the hot, dry summers. A male, fruitless, clone is grown almost exclusively as it is much less messy than fruiting females.
2. M. rubra L. Red Mulberry, Moral. Smallish tree to ca. 20 m, trunk short, branches spreading, crown round-topped, bark smooth to scaly. Branchlets pubescent when young, becoming glabrous. Petioles (1.4)2 to 3 cm long, pubescent; blades ovate to ovate-oblong in overall shape, entire to lobed as described above for M. alba though perhaps less often lobed, to 20 cm long, usually smaller, truncate to subcordate basally, apex abruptly long-acuminate or cuspidate, margin serrate, smooth or commonly somewhat scabrous above (especially when young), soft-pubescent beneath, especially on all the veins. Staminate aments pendulous, 2 to 5 cm long. Pistillate flower clusters ovoid-cylindric, 8 to 25 mm long. Multiple fruit (1)2 to 3 cm long, dark purple to nearly black at maturity, juicy. Upland woods, floodplains, stream banks. Mostly E. and Cen. TX; VT to Ont., MN, and NE, S. to TX and FL. Mar.-May, the fruits maturing about 2 months later.
Sometimes cultivated. The wood is suitable for fence-posts and was once used for barrels (Elias 1980).
4. BROUSSONETIA L'Her. ex Vent.
7 or 8 species of E. Asia and Polynesia. We have the 1 species naturalized in TX.
1. B. papyrifera (L.) L'Her. ex Vent. Paper Mulberry. Dioecious shrub or small tree to 15 m, suckering freely from the base and colonial in some areas, branches spreading, crown round, bark tan-gray or mottled, smooth. Branchlets and twigs persistently pubescent. Leaves alternate or occasionally some opposite or very rarely whorled; petioles to 10 cm long; blades ovate or broadly ovate in overall outline, 6 to 20(25) cm long, unlobed or palmately 1- to 3-lobed, sometimes mitten-shaped, base rounded to cordate, often asym-metrical, apex acute to acuminate, margin serrate or dentate, scabrous and somewhat pubescent above, densely and softly short-pubescent and grayish below; stipules to 1.5 cm long, ovate-lanceolate, deciduous. Flowers unisexual, greenish, 4-merous. Staminate aments cylindrical, 4 to 8 cm long, pendulous. Female flowers in globose, tomentose heads 1 to 2 cm in diameter, styles entire. Multiple fruit a dense globose head 2 to 3 cm in diameter, the orange to red achenes protruding from the fleshy calyces. Roadsides, clearings, waste places, etc. Native of China, Japan, and Polynesia; cultivated for ornament and as a street tree and occasionally naturalized from NY to the SE. Great Plains, FL and TX; also Mex. Flowering spring and the fruits maturing in the summer. [Papyrius papyrifera (L.) Kuntze].
The fibrous inner bark of this tree is pounded to make tapa cloth in Polynesia and can also be used in paper-making (Mabberley 1987).
Annual or perennial herbs (some, but not ours, woody), sometimes succulent, sap watery. Foliage often with stinging hairs. Leaves alternate or opposite, simple, petiolate, usually stipulate. Flowers minute, individually inconspicuous, regular or rarely irregular, unisexual (some perfect in Parietaria), greenish, anemophilous, plants monoecious or dioecious. Inflorescences axillary, basically cymose but sometimes otherwise or reduced to a single flower. Sepals (3)4 to 5(6), united or free, calyx usually more deeply parted in the staminate flowers and sometimes absent in pistillate flowers; corolla absent. Stamens equalling the sepals and opposite them (rarely only 1,) inflexed and elastically reflexing to shed pollen at maturity, scale-like staminodia sometimes present opposite the sepals in female flowers. Vestigial pistil sometimes present in male flowers, in female flowers ovary superior, 1-celled, uniovulate, style 1, stigma usually 1, capitate or filiform. Fruit usually an achene, commonly enclosed in the persistent, accrescent calyx and/or tipped with the persistent style.
Ca. 1,050 species in 52 genera, primarily tropical, but a few temperate. There are 4 genera and 11 species in TX; 3 genera and 4 species here.
Some are important fiber plants, such as Boehmeria (ramie) and several species of nettles (Urtica). The young shoots of some, including nettles, are edible cooked like spinach. Many have stinging hairs which can cause a very painful reaction, even in dried specimens. A few, such as Artillery Plant (a Pilea), are grown as ornamental or greenhouse plants (Mabberley 1987).
1. Leaves alternate, entire ........................................................................................1. Parietaria
1. Leaves opposite (in ours), toothed ...........................................................................................2
2(1) Plants with stinging hairs; sepals of pistillate flower free or nearly so ......................2. Urtica
2. Plants without stinging hairs; sepals of pistillate flower fused, calyxcup-shaped ....................
NOTE: Pilea pumila (L.) Gray var. deamii (Lunell) Fern. is found in E. TX and may be found here eventually. Leaves opposite, glabrous, without stinging hairs and with small whitish lines; teeth 11 to 17 per margin. The flowers are in axillary cymes to 3 cm long. The calyx of pistillate flowers is deeply 3-parted, not enclosing the fruit. See also note about shade forms at description of Boehmeria.
Annual or (short-lived) perennial herbs; stems simple or well-branched and tufted, stinging hairs none. Leaves alternate, petiolate (sometimes shortly so), usually punctate, with 3 prominent nerves, stipules absent. Inflorescence a short cyme with 2 to 6 free to somewhat united bracts. Flowers mostly bisexual, though the first flowers female and the last male; bisexual flowers protogynous so style absent by anther dehiscence and flowers often appearing male. Calyx of 4 more or less united sepals. Stamens 4. Ovary included within calyx, style tufted. Achene nearly ovoid, enclosed in the persistent calyx; pericarp crustaceous and shiny.
Ca. 20 to 30 species, nearly worldwide; 3 in TX; 1 here.
Some species were formerly used in medicines (Mabberley 1987).
1. P. pensylvanica Muhl. ex Willd. Hammerwort, Pennsylvania Pellitory. Slender taprooted annual, stem 0.5 to 4.5 dm tall, sprawling to erect, simple or sparingly branched at the base, puberulent to pubescent with mostly hooked or curled hairs. Petioles slender, 5 to 25 mm long; blades thin and membranous, rhombic-, elliptic-, or linear-lanceolate, the larger ones 1.5 to 7 cm long and 0.8 to 2 cm wide (lowermost leaves sometimes much smaller, less than 1 cm long and nearly ovate), entire, base cuneate, apex tapered to an obtuse tip, punctate, glabrate to minutely scabrous, margins ciliate; stipiules none. Flowers in few-flowered clusters in all but the lowest leaf axils, bracts pubescent, linear to linear-lanceolate, 4 to 5 mm long, exceeding the flowers; staminate and pistillate flowers present, but most bisexual. Calyx deeply 4-lobed in staminate flowers, less deeply lobed in pistillate, brownish, 1 to 2 mm long, puberulent, calyx lobes acute. Stigma tufted, subsessile. Achene 0.9 to 1.2 mm long, minutely apiculate, ovoid, flattened, shining, light green to brown. Shady banks and in rocky areas, not common. E. and Cen. TX, N. to High Plains; FL to TX. Mex., and CA, N. to Ont., ME, NY, and B.C. Mar.-May.
Herbs (as ours), shrubs, or small trees, monoecious or dioecious. Leaves opposite or alternate, without stinging hairs, variously pubescent, stipules somewhat persistent. Flowers in axillary clusters, male flowers with a deeply 4-lobed calyx and 4 stamens. Female flowers with a 2- to 4-parted calyx enclosing the ovary; style 1, filiform-subulate, persistent, stigmatic and papillose on 1 side. Fruit an achene ca. 1 mm long, closely enveloped by the dry calyx.
Ca. 50 species of the tropical and N. subtropical regions; 2 (one naturalized) in TX; 1 here. Ramie, Boehmeria nivea (L.) Gaudin, has the longest, strongest vegetable fibers known and is used in rope and cloth (Mabberley 1987). Native to tropical Asia, it is naturalized in the SE. U.S.: FL to TX, N. to S.C. Not reported from our area, but possibly occurring here in the future. It has alternate, toothed leaves which are densely white-pubescent below. It is dioecious, with the flowers in axillary clusters.
1. B. cylindrica (L.) Sw. Bog-hemp, False Nettle, Button-hemp. Perennial herb; stems 4 to 10(12) dm tall, simple to rarely branched, glabrous to pubescent or scabrous. Leaves without stinging hairs, opposite or rarely alternate, petioles long, often to 1/2 the length of the blade; blades ovate to ovate-lanceolate or oblong-lanceolate, 4 to 15 cm long, 2 to 8 cm broad, with 3 prominent nerves from the base, base broadly cuneate to rounded or cordate, acuminate at apex, coarsely serrate, smooth to scabrous, sometimes minutely punctate above and/or soft-pubescent beneath, sometimes recurved; stipules distinct, brownish, lanceolate, deciduous. Plants usually dioecious, flowers ca. 2 mm broad, hispidulous, usually male and female on separate plants, but sometimes mixed in the inflorescence. Flowers in glomerules arranged in continuous or interrupted spike-like clusters (often the staminate interrupted and pistillate continuous), clusters solitary in the leaf axils, sometimes leafy at apex and flowers thus appearing to be borne on upper branchlets. Staminate flowers with a 4-parted calyx and 4 stamens. Pistillate flowers with a tubular, flattened to ovoid, 2- to 4-toothed calyx surrounding the ovary; style exserted, laterally papillose. Fruit ovate to rotund, compressed, very minutely winged, apiculate or beaked, pubescent, ca. 1 to 1.5 mm broad. Wet soils of bogs, marshes, swamps, and seeps, along rivers and streams, sun or shade. Primarily E. TX but W. to Val Verde and N. to Hemphill Cos. FL to TX, N. to Ont. and Que., W. to MN, NE, and IL. June-Oct. [Includes var. cylindrica and var. drummondiana Wedd.--see note below.].
NOTE: Plants in the shade tend to have broader, thinner, less pubescent leaves and were formerly treated as var. cylindrica. Plants of more open sites have narrower, more-lanceolate, thicker leaves with petioles shorter (2 cm or less), somewhat scabrous above and densely pubescent below, often recurved; flower spikes typically thicker and fruits larger than in shade forms. These plants were formerly treated as var. drummondiana Wedd. [B. drummondiana Wedd.; B. scabra Small].
3. URTICA L. Nettle
Annual or perennial herbs; stems simple or branched, erect, fibrous. Leaves opposite, stipulate, petiolate, dentate to serrate, glabrous to pubescent, with stinging hairs on the stems, petioles, and main leaf veins. Flowers unisexual, in panicles or branched spikes (rarely catkin-like arrangements or pseudo-heads) in the axils of the middle and upper leaves, plants monoecious or dioecious. Male flowers with a 4-parted calyx, the segments unequal, stamens 4, bent down in bud, springing up and the anthers dehiscing violently at maturity, rudimentary pistil often present. Female flowers with sepals unequal, the outer 2 small and inconspicuous, the inner 2 larger, accrescent; stigma sessile, tufted. Achene lenticular or flattened, ovate, enclosed by the 2 inner sepals.
30 to 50 species worldwide, especially in the N. temperate region; 4 listed for TX; 2 here.
Some species are a source of fiber for cloth, and the young shoots of some can be eaten like spinach (Mabberley 1987). The tips of the stinging hairs break off and the sharp remnants actually inject irritants. The redness and itching usually subside within 24 hours (Mabberley 1987). The hairs are capable of stinging even on dry specimens.
1. Cystoliths (mineral concretions) on upper leaf surfaces dark, round, and pimple-like; upper leaves similar to and only slightly smaller than the lower; marginal teeth triangular, straight-sided but blunt ...........................................................................................1. U. urens
1. Cystoliths on upper leaf surfaces scattered, irregular, usually linear and whitish;
uppermost leaves usually much smaller and proportionately narrower than the lower; teeth usually ovate-elliptic with convex sides, blunt ...........................2. U. chamaedryoides
NOTE: Some collections from N. Cen. TX show influence of both species with regard to vegetative characters (Correll and Johnston 1970).
1. U. urens L. Stinging Nettle, Burning Nettle, Dog Nettle, Ortiga. Annual, stems erect to ascending, to 6 dm tall, branched from the base or sometimes simple; herbage with stinging hairs, otherwise glabrous. Petioles about equalling the blades or shorter (only sometimes longer); blades suborbicular-ovate to ovate, ovate-oblong, or elliptic, to 4 cm long and 3 cm broad (to 16 cm long and 6 cm broad in extreme shade forms), rounded to acute, base subcuneate to cuneate, upper leaves similar in shape to lower leaves and proportionately as wide, only slightly smaller, marginal teeth triangular, usually straight-sided (sometimes convex), blunt, margin sometimes doubly-serrate; cystoliths small, round, pimple-like, usually dark. Stipules oblong, 1 to 4 mm long. Plants monoecious, inflorescences axillary, generally shorter than the subtending petiole, usually unbranched, of mixed male and female flowers, flowers white to pale yellow. Fruiting calyx with margins hispid-ciliate. Achene ca. 2 mm long. Clay or sand soils of open woods and waste areas. S. 1/3 TX; native of Europe, widely naturalized in N. Amer. Feb.-Aug.
2. U. chamaedryoides Pursh Heartleaf Nettle, Ortiguilla, Weak Nettle. Slender annual 0.5 to 10 dm tall, stems erect or sometimes supported by surrounding vegetation, branched from the base and sometimes with axillary branches, herbage with sparse stinging hairs, otherwise glabrous. Petioles shorter than to equalling the blade; blades 1 to 6 cm long, 1 to 4 cm broad, broadly ovate or subrotund to elliptic-lanceolate, much and usually abruptly reduced and narrower upwards (upper leaves lanceolate), obtuse to acute, base cordate to truncate or cuneate, membranous, sometimes purplish below, margins serrate, with usually rounded, convex-sided teeth, cystoliths visible when dried; stipules linear-lanceolate, 1 to 4 mm long, spreading or deflexed. Plants monoecious, flowers in globose to short-spicate clusters 3 to 6 mm long; peduncles shorter than the subtending petioles, slender, (1)2(3) in each axil. Inner 2 sepals 1 to 1.5 mm long, outer sepals less than half that length. Achene ovate to elliptic, 1 to 1.5 mm long, ca. 1 mm broad, tan to brown, enclosed by the 2 inner sepals. Among rocks, on wooded slopes in bottomlands, and on shell mounds. Primarily Cen. TX; FL to TX and Mex., N. to WV, KY, MO, and OK. Feb.-July. [Includes var. chamaedryoides and var. runyonii Correll].
Trees or rarely shrubs, aromatic and glandular, anemophilous. Leaves deciduous, generally alternate, pinnately compound (some, but not ours, trifoliolate), petiolate, estipulate. Plants monoecious, flowers individually inconspicuous, solitary in bracts of the drooping (or erect) catkins; catkins solitary or grouped into terminal panicles or fascicles; in some genera the catkins with male and female flowers, in others (such as ours) the catkins unisexual. Corolla absent. Male flowers with or without a (1-)4-(5-)parted calyx adnate to the bract and 2 bracteoles (bracteoles sometimes absent); stamens (3)5 to 40(100+), filaments short, anther dehiscence longitudinal; vestigial pistil sometimes present. Female flowers often solitary or in a small cluster or spike, each with 1 bract and 2 or 3 bracteoles, and with or without a 0- to 4-lobed calyx; gynoecium inferior, incompletely 2- to 3-celled below (sometimes the appearance of more cells caused by false partitions), unilocular above, 1-ovuled at the top of the partition, styles 2, distinct or united at the base or sometimes stigmas sessile; calyx, bract, and bracteoles often connate to form an involucre or husk on the fruit; fruit a nut or drupe-like (sometimes samara-like), enclosed by the dehiscent or indehiscent husk. Seed solitary, with no endosperm and oily, lobed and convoluted cotyledons.
7 genera and 50 to 60 species of the temperate and warm N. hemisphere to S. Amer. and Mal.; 2 genera and 12 species listed for TX; 2 genera and 7 species in our region.
The family is important for timber trees--Hickory (Carya) and Walnut (Juglans). These genera also produce nuts that people eat (walnuts, pecans, hickory nuts, and butternuts) and nuts that are important foods for wildlife, including deer, bear, peccaries, and many birds and rodents. Animals also distribute the fruits. Typically each tree will have a heavy crop after 1 or 2 years of lighter crops; trees in a forest are not synchronized (Elias 1980). Dyes, primarily browns, are obtainable from the nut husks of many species (Tull 1987).
1. Pith of older twigs chambered; husk of fruit indehiscent; staminate catkins borne on last year's wood ..............................................................................................................1. Juglans
1. Pith of older twigs solid; husk of fruit at least partially dehiscent at maturity; staminate catkins borne on new growth ......................................................................................2. Carya
Ours trees with furrowed, scaly bark, wood dark-colored; pith chambered in twigs 1 year old or more. Leaves usually once odd-pinnate, leaflets 7 to 23, primarily opposite but sometimes subopposite or alternate on the rachis, oblong-lanceolate or oblong-ovate, acuminate, falcate, serrate, some of the lateral leaflets the largest. Flowers produced in the spring; staminate catkins sessile, pendulous, solitary but superposed, at the end of the previous season's growth; sepals of male flowers (3)4(6), sometimes reduced or minute, adherent to the bracteole(s), floral receptacle adnate to the bract, stamens 8 to 40, filaments free, short, anthers green. Female flowers 1 to several in clusters or short spikes at the end of the new growth, each with a bract and 2(3) bracteoles; sepals 4, ovary inferior, pubescent, styles 2(3), elongate, the inner surface stigmatic and fringed. Fruits large, drupe-like, solitary or in groups of 2 to 3, ellipsoid or globose, pungent, with an indehiscent husk and a hard, irregularly-furrowed nut shell within.
About 20 species of the Medit., Eur., E. Asia, N. Amer. and the Andes; 3 species in TX; 1 here.
J. regia is the common walnut used in cooking. J. nigra is also valuable for nuts and timber. J. cinerea produces edible butternuts (Mabberley 1987). Many species are valuable for timber. Some produce dyes or stains. Some produce herbicide-like compounds and will kill other plants growing near them.
1. J. nigra L. Black Walnut. Large tree to 30 to 40(50) m tall, trunk to 1.2(2) m in diameter, straight often for half the height of the tree, crown open, round-topped; bark brown to blackish, with deep fissures and narrow, forking ridges; young branchlets brown to orange-brown, glandular-puberulent, becoming glabrous; leaf scars with 3 vascular traces; pith light brown. Petioles 6.5 to 14 cm long, the rachis and petioles glandular-pubescent; blades to 60 cm long, leaflets (9)11 to 23 (terminal leaflet sometimes absent), lateral leaflets the largest, sessile, ovate-lanceolate, to 15 cm long and 2.5(5.5) cm broad, long-acuminate, somewhat obliquely cordate at the base, glabrous above, paler and minutely pubescent below, serrate. Staminate catkins to 12 cm long, flowers pedicellate, each with 20 to 30 stamens. Female flowers 1 to 4 in short spikes 1 to 1.5 cm long, stigmas yellow-green. Fruits globose, to 4(6) cm in diameter, usually single or paired, yellow-green at first and gland-dotted, glabrous to slightly pubescent, turning brown, indehiscent. Kernel of nut sweet, oily. Rich woods, especially bottomlands, and fields. E. TX; sometimes planted; N. Eng. S. to FL, W. to MN and TX. Flowering in spring and the nuts maturing in the fall. Fall color yellow. [Wallia nigra (L.) Alef.].
The wood is used extensively for furniture, gunstocks, carving, etc., though it is becoming increasingly rare and valuable (Elias 1980). Owners of large trees have been known to return from vacation to find them stolen. The nuts are used in confections and ice cream. Brown to tan dyes can be made from the green husks and nutshells. No mordants are needed. Hair dyes are also possible. The leaf rachises can be used in baskets (Tull 1987). Some people experience contact dermatitis after handling the plants (Lampe 1985).
Trees, usually from taproots, bark sometimes shaggy; wood hard, dark-colored; pith solid; young twigs commonly pubescent. Leaves petiolate, usually odd-pinnately compound; leaflets 3 to 25, the 3 terminal ones usually the largest or at least as long as the laterals (cf. Juglans with lateral leaflets largest), often gland-dotted, sessile to short-petiolulate, opposite on the rachis, lanceolate to elliptic, oblanceolate, or obovate, acute to acuminate, glabrous above and pubescent or glabrous below with tufts of hairs in the axils of the veins. Flowers borne in spring from scaly buds, the staminate flowers below the new leaves and the pistillate above. Staminate catkins in fascicles of (usually) 3 in the axils of the bud scales; male flowers with (3)4(10) stamens adnate to the bract and the 2- or 3-lobed calyx/bracteole (interpretation of the structure varies), filaments very short or absent. Pistillate flowers 2 to 10 in clusters or short spikes on peduncles terminating the new growth; true calyx absent, bract and usually 3 bracteoles sepal-like in flower, surmounting the ovary; stigmas 2, sessile, sometimes divided, papillose. Fruits single or a few in a cluster, each with a 4-angled husk formed from the bract and bracteoles, husk splitting at maturity from 1/2 to all the way; nut inside with a bony shell, smooth to rough, incompletely 2-celled or 4-celled at the base.
About 17 species from E. N. Amer. to Cen. Amer, also E. Asia.; 9 listed for TX; 6 to be expected here. Positive identification is often impossible without mature fruit and is easiest if the tree can be observed in all seasons. Some species will hybridize, making the offspring even harder to recognize.
Hickories are valued for timber and nuts. The most important nut is the pecan, C. illinoinensis, the oil of which is also used in cosmetics and other products (Mabberley 1987). The wood of many hickories is strong and durable, though that of pecan is poorer than most. All species have value as a food source for deer, bears, small mammals, game birds, and songbirds. Large and smaller crops often alternate years (Elias 1980).
1. Sutures of husk winged or keeled at least half their length; bud scales valvate; bud scale scars separate, not forming a distinct ring ...............................................................................2
1. Sutures of husk not markedly winged or keeled; bud scales imbricate; bud scale scars forming a distinct ring ...............................................................................................................4
2(1) Nuts cylindrical, kernel sweet; cotyledons not deeply 2-cleft; leaflets 9 to 17 per leaf ............
......................................................................................................................1. C. illinoinensis
2. Nuts flattened to globose; kernel bitter; cotyledons deeply 2-cleft; leaflets 7 to 13 per leaf ..
3(2) Winter bud brownish; husk winged to the base, splitting to middle or base at maturity; terminal leaflet stalked; nutshell reddish- brown, wrinkled or furrowed ........2. C. aquatica
3. Winter bud yellowish; husk winged apically and splitting to just below middle at maturity; terminal leaflet basically sessile; nutshell smooth, gray .............................3. C. cordiformis
4(1) Margins of young leaflets densely ciliate; teeth of older leaves with persistent small tufts of hair; bark shaggy .....................................................................................................4. C. ovata
4. Margins of young leaflets not densely ciliate; teeth of older leaves without tufts of hair; bark generally not shaggy .................................................................................................................5
5(4) Leaves rusty-pubescent beneath (at least when young), the hairs not in groups; rachis glabrous to puberulent; husk shallowly grooved from apex to base, splitting to base at maturity ..................................................................................................................5. C. texana
5. Leaves pale-pubescent beneath, the hairs grouped; rachis tomentose; husk grooved at one end, splitting to middle or base ..........................................................................6. C. alba
1. C. illinoinensis (Wang.) K. Koch Pecan, Nogal Morado, Nuez Encarcelada. The tallest of our hickories, mature trees to 50(60) m, canopy spreading, open; trunk straight, to 2.5 m in diameter; taproot often very deep. Bark light brown, becoming darker with age, tinged with red, with irregular, shallow furrows. Twigs pubescent when young, becoming glabrous by the second year; lenticels elongated, orange; terminal buds pointed, flattened, to 10 mm long, yellow-brown, hairy, scales valvate and not leaving a distinct circular scar. Leaves aromatic, 30 to 50 cm long, leaflets 9 to 11(17), terminal leaflet stalked, nearly all the same size or terminal leaflet broader, 3 to 20 cm long and 1 to 7.5 cm wide, lanceolate to ovate-lanceolate or oblong-lanceolate, usually falcate, acuminate, bases cuneate to rounded, asymmetrical, margins coarsely serrate, sometimes twice so, glabrous; upper surface glabrous or pilose, lower surface paler and glabrous or pubescent. Staminate catkins in more or less sessile groups of 3 to 11. Fruits in clusters of 3 to 11, ovoid to ellipsoid, 2.5 to 6.5 cm long, 1.2 to 2.5 cm broad, rounded at base, pointed at apex, the 4 sutures winged and keeled, dark brown, at least partially covered with yellow scales. Husk brittle, thin (ca. 1 mm), splitting to base at maturity, often persisting on the tree. Nut cylindric to oblong, pointed at both ends, brown to reddish, often marked with black; shell thin, kernel sweet, cotyledons not deeply 2-lobed. Bottomland woods, along streams and rivers, and in moist woods. Primarily Cen. and NW. TX; OH to WI and IL, SW. to KS, OK, and TX; E. to AL.; also Mex. April-May. [also seen spelled illinoensis or illinoisensis; C. pecan (Marsh.) Engl. & Graeb. or C. pecan (Marsh.) Schneid.; Hicoria pecan (Marsh.) Britt.].
Widely grown for the nuts, of which there are hundreds of cultivated varieties. The trees bear from the age of 20, the best years being between 75 and 225 when annual crops of 500 lbs of nuts per tree are not unusual. The wood is used for tool handles, cabinetry, and veneer. The nuts are eaten by waterfowl, large game birds, deer, foxes, and squirrels (Elias 1980). This is the state tree of Texas. Sometimes one of the last trees to leaf out in spring. The pollen can be allergenic.
2. C. aquatica (Michx. f.) Nutt. Water Hickory, Bitter Hickory. Medium to tall tree to 35 m, crown narrowly rounded; trunk to 6(10) dm in diameter; bark light brown, sometimes tinged reddish, breaking up into long, loose, shaggy plate-like scales; winter buds pointed, slightly flattened, red-brown (sometimes slightly yellow), with caducous yellow glands, scales valvate, not leaving a distinct ring-scar. Branchlets reddish-brown or ash-gray, slightly glandular and tomentose when young, becoming gray and more or less glabrous. Leaves to 33 cm long, leaflets 7 to 13, sessile or petiolulate, ovate-lanceolate to lanceolate, 7.2 to 10(16) cm long, 1.2 to 4.0 cm wide, the laterals slightly to strongly falcate, acuminate, the base rounded to cuneate, more or less asymmetrical, margins coarsely to finely serrate or subentire; surface glabrous to glabrate above, puberulent to pubescent beneath at least along the veins. Staminate catkins in pedunculate cluster. Fruit in groups of 3 or 4, ovoid or ellipsoid to subglobose, much flattened, usually broadest above the middle, pointed at the tip and narrowed or rounded at the base, 4-winged along the sutures, 2.5 to 3.5(4) cm long, 2 to 3.2 cm broad, dark brown to blackish, with yellow scales. Husk brittle, thin, splitting late and usually only to just below the middle, rarely the entire length. Nut flattened, ovoid to obovoid, shell dark brown or reddish, somewhat wrinkled or uneven, apex abruptly sharp-pointed, kernel bitter, cotyledons deeply 2-lobed. Riverbottoms and swamps, especially where seasonally flooded; capable of growing on better-drained alluvial soils. E. TX; VA to FL, W. to E. TX, N. to S. IL and SE. MO. [Hicoria aquatica (Michx. f.) Britt.].
The wood is heavy and fine-grained, but brittle, which limits its use to fencing and firewood--the yield of heat per cord is high. The nuts are an important wildlife food (Elias 1980).
3. C. cordiformis (Wang.) K. Koch Bitternut Hickory, Pignut Hickory. Tall tree to 35 m, narrow but the top spreading with age, crown open and rounded; trunk straight, to a maximum of 9(10) dm in diameter; bark thin, somewhat smooth for a hickory, shallowly furrowed and scaly, light brown and reddish-tinged. Branchlets greenish and rusty-pubescent when young, becoming light gray to red or yellow-brown in the second or third year; lenticels pale; winter buds yellow-orange, ovoid, covered with minute scales, blunt, bud scales valvate, not leaving a perfect scar ring. Leaves 15 to 30 cm long, leaflets (5)7 to 9, ovate-lanceolate to lanceolate, obovate, or oblanceolate, to 15 cm long and 3(8.5) cm broad, the terminal 3 the largest, terminal leaflet sessile, apices acuminate, base of lateral leaflets rounded to subcordate, base of the terminal leaflet subcuneate, margins coarsely serrate except at the bases, without marginal hairs, glabrous above, paler and pubescent below. Staminate catkins in clusters, stalked. Female flowers usually 1 or 2 in terminal spikes. Fruits subglobose to obovate, slightly flattened 4-winged from the apex to near the middle, beaked, 2.5 to 3.5 cm in diameter. Husk brittle, thin, puberulent and with yellow scales, splitting to only about the middle. Nut globose to ovate, 1.5 to 3 cm long, somewhat flattened and angled, apex depressed, shell thin, smooth or uneven, gray; kernel bitter and bright reddish-brown. Bottomlands and low woods near streams and swamps, also on higher upland hills. E. TX; FL to TX, N. to NE, MN, WI, Ont. and N. Engl. Flowering about April and the nuts maturing in the fall. [Hicoria cordiformis (Wang.) Britt.; Hicoria minima (Marsh.) Britt.].
Collections from our area not seen, but our region well within the range of the species and the trees probably present in uncollected bottomlands.
The shock-resistant wood is used for tool handles and formerly for wooden wheels. It makes a good fuel. Oil from the nuts has been used in lamps and medicines. Wildlife seem to prefer the sweeter kernels of other hickories to these (Elias 1980). In some states, said to hybridize with C. illinoinensis and C. ovata (GPFA 1986).
4. C. ovata (P. Mill.) K. Koch Shagbark Hickory, Shellbark Hickory. Medium to tall tree to 20(45) m tall, crown narrowly cylindrical to somewhat spreading, canopy usually broadest above the middle; trunk to 9(15) dm in diameter; bark shaggy, exfoliating in curling strips or plates, light to dark gray. Branchlets reddish-brown and pubescent when young, becoming glabrous and gray with age; winter buds brown-gray, oviod, scales imbricate and leaving a distinct ring-shaped scar. Leaves 20 to 36 cm long, with (3)5(7) leaflets; leaflets stalked, ovate or sometimes ovate-lanceolate or obovate; to 20 cm long and 8 cm broad, generally broadest above the middle, terminal leaflet the largest, all rounded to acute or acuminate at the apex, bases usually cuneate, sometimes asymmetrical, margins ciliate, serrate above the base, with small tufts of hairs on either side of most of the teeth, at maturity the upper leaf surface glabrous, paler and generally glabrous or merely puberulent below. Clusters of staminate catkins peduncled, floral bracts elongated. Fruits solitary or in pairs (rarely in 3's), subglobose to depressed-ovoid, indented a the apex, to 6 cm long, unwinged but with 4 shallow grooves, dark red-brown to blackish, glabrous to pilose. Husk thick (to 15 mm), splitting at maturity to the base. Nut flattened, 4-angled or -ridged, subglobose, 2 to 3.5 cm in diameter, apex pointed, shell ridged, pale to whitish, thin; kernel light brown, aromatic, sweet. Rich woods, bottomlands and on slopes, common near swamps and streams but usually in well-drained soil. E. TX; GA and NC, W. to TX N. to NE, MN, WI, Ont., ME and S. N. Eng. Apr.-May, the nuts maturing in the fall. [Hicoria ovata (Mill.) Britt.].
Reported to hybridize with C. cordiformis, and often varying in characters of leaf pubescence and nut shape 9GPFA 1986). The wood is used for handles, baskets, wagons, and so on, and also for smoking meats. It has one of the highest heat values of any American wood. The nuts are eagerly eaten by wildlife and were a staple food of many Indian tribes (Elias 1980).
5. C. texana Buckl. Black Hickory. Small to medium tree to a maximum of 30 m, trunk to 60 or 70(90) cm in diameter; crown narrow and round-topped; bark of mature trees dark gray to blackish, deeply to shallowly furrowed, rough and with plates or scales. Twigs reddish-gray and pubescent when young, becoming dark-gray to reddish-brown and glabrous; lenticels pale and scattered; winter buds ovoid, pointed, rusty-pubescent, the scales imbricate and the scars forming a ring. Leaves to 30 cm long, leaflets 5 to 7(13), sub-sessile to short-petiolulate, to 12.5 cm long and 4 cm broad, the terminal 3 the largest, all lanceolate, or somewhat oblanceolate or rhombic-lanceolate when young, apex acuminate, often curved, cuneate at the base and usually asymmetrical, margins serrate, rusty- or bronze-pubescent beneath, especially on the veins and when young, the hairs not in groups, upper surface glabrate. Clusters (usually 3's) of staminate catkins pedunculate. Female flowers solitary or paired at the ends of the branches. Fruits subglobose to obovoid to somewhat oblong, slightly depressed or apiculate at the apex, to 5 cm long, with 4 shallow vertical grooves and sometimes slightly 4-winged at the base, dark brown and with numerous yellowish scales. Husk ca. 3 mm thick, splitting to the base at maturity (sometimes only to the middle), the valves of two different sizes. Nut globose to obovoid, somewhat 4-angled and flattened, pointed at both ends, shell rough and pitted, moderately thick, hard and brittle; kernel reddish, sweet. Dry, sandy upland woods and rocky slopes. E., Cen., and NE. TX; N. to OK, SE. KS, S. IL, and IN, E. to LA and AR. Spring, ours flowering about April, the nuts maturing in the fall. [C. buckleyi Durand; C. arkansana Sarg.; C. buckleyi Durand var. arkansana (Sarg.) Sarg.].
Because the trees are small, the branches crooked, and the wood brittle, the wood is used mostly for fuel. Hogs, small mammals, and larger birds such as jays eat the nuts, but the hard shell makes them inaccessible to smaller birds and rodents (Elias 1980).
6. C. alba (L.) Nutt. ex Ell. Mockernut Hickory, White Hickory, White-hearted Hickory. Small to large trees to 35 m tall, the crown narrow or broad, depending upon the competition; bark with shallow criss-cross furrows, dark gray and scaly. Branchlets and petioles tomentose with pale, gray-brown, or reddish fascicled hairs, becoming glabrous at maturity, lenticels conspicuous, pale; winter buds ovoid, pointed to rounded at the tip, rusty-tomentose, the 3 to 4 scales imbricate and leaving a ring-shaped scar. Leaves 14 to 35 cm long, glandular-resinous and fragrant, the rachis tomentose or pubescent with the hairs in groups; leaflets 5 to 9, sessile or short-petiolate, to 20 cm long and 12. 5 cm broad, the terminal 3 the largest, lanceolate to obovate- or oblong-lanceolate, apex gradually or abruptly acuminate, base cuneate to rounded, often oblique, margin coarsely or minutely serrate, lower surface pale-pubescent, glandular, the hairs on the margins in groups, upper surface lustrous, glandular-resinous, the midvein often pubescent. Male flowers in pedunculate clusters of 3. Fruits single or in small clusters, globose to ovoid, to ca. 5 cm long, slightly flattened, gradually narrowed to the ends, dark (reddish-) brown, pilose to glabrous, very slightly 4-ridged at one end and 4-grooved on the other. Husk thick, splitting at maturity (or tardily, later) to about the middle or nearly to the base. Nut ellipsoid to subglobose, 4-angled, 2.5 to 3.5 cm long, shell very thick, hard, commonly cracking transversely on drying, light red-brown, becoming darker; kernel small, dark brown, shiny, sweet. Moist or dry woods. E. TX: FL to TX, N. to IA, Ont., and N. Eng. [Authority sometimes given as (P. Mill.) K. Koch; C. tomentosa Nutt. or (Poir.) Nutt.].
The common name refers to the deceptively large nuts with thick shells and disappointingly small kernels. Nonetheless, a variety of wildlife consumes the nuts. The wood is strong, durable, and withstands compression well. It is used for tool handles, furniture, and wood splints (Elias 1980).
Aromatic, deciduous or evergreen shrubs or small trees. Leaves alternate, simple, usually estipulate, with long, colorless, unicellular hairs and peltate, usually yellow, multicelled glands. Flowers unisexual (very rarely perfect), plants monoecious or dioecious, both sexes in catkins or the female flowers in small clusters, typically each flower subtended by a bract; males sometimes also with 2 bracteoles; perianth absent. Stamens (2)4(5 or 6), fewest in the most acropetal flowers. Pistillate flowers with 2 bracteoles, gynoecium bicarpellate, unilocular, styles distinct or united only at the base, ovule 1. Fruit a drupe or nutlet, sometimes with accrescent bracteoles.
3 genera and about 50 species nearly worldwide; 1 genus with 2 or 3 species in TX; 1 species here.
Some species, chiefly Myrica, are sources of waxes or timber; a few have edible fruit (Mabberley 1987).
Shrubs, the roots usually with nitrogen-fixing bacteria. In ours, leaves coriaceous and evergreen or tardily deciduous, resin-dotted, entire to toothed or lobed, estipulate, usually petiolate. Plants dioecious, flowers in the axils of scale-like bracts; bracteoles not overlapping the fruit; perianth none. Staminate catkins ellipsoid to stoutly cylindric, from axillary scaly buds; male flowers with 2 to many stamens, filaments somewhat connate at the base. Pistillate catkins ovoid to cylindric, gynoecium as described for family, styles distinct, stigmas linear-lanceolate. Fruit a nutlet or (as in ours) drupelike, globose to ovoid, warty and commonly covered with wax or resin.
About 35 to 50 species (depending on interpretation) nearly worldwide; 2 or 3 species in TX (again, depending on interpretation); 1 here.
The genus includes plants used in the manufacture of bayberry candles and soap. The wax is removed from the fruit by boiling. Some species have edible fruit and some are grown for ornament or used in medicines, flavorings, or insecticides (Mabberley 1987).
1. M. cerifera (L.) Small Wax-myrtle, Candleberry. Shrub or small tree to 12 m tall, sometimes colonial (then plants usually to only 2 m tall), trunk to a rare maximum of 2 dm in diameter, usually less; young twigs waxy, glabrous or sometimes pilose. Leaves persisting for about 2 years, often rusty- or yellow-green-resin dotted on both surfaces, narrowly cuneate-oblanceolate to elliptic, obovate, or linear-spatulate, narrowed to a short petiole, generally less than 7 cm long, to 2.5 cm broad (leaves of colonial plants smaller), sometimes pubescent below, acute to obtuse, entire to serrate or toothed. Staminate catkins ovoid-cylindric, to 2 cm long. Fruit blackish, with a whitish waxy covering, 2 to 4 mm in diameter. Often on sandy soils, near streams, lakes, bogs, and wet or dry woods, often in mixed pine-hardwood forests. E. TX; FL W. to TX, N. to AR, DE, and NJ. Mar.-Apr. [Morella cerifera L.; Cerothamnus ceriferus (L.) Small].
Described above to include the formerly-separate M. pusilla Raf. An alternative, if varieties are recognized, is to treat those plants as M. cerifera L. var. pumila Michx. (C. pumila (Michx.) Small.) These are the smaller, colonial plants with leaves usually less than 5 cm long and fruit 3 to 4 mm broad--fruit is 2 to 3 mm in diameter for the typical form. The plants can be treated as 1 species, with or without varieties, or else as 2 separate species.
Wax and soap may be made from the waxy covering of the fruit. The leaves are said to have insect-repellent qualities and may be used to deter moths in stored woollens. Brown, tan, and gray dyes can be made from the leaves and berries. The leaves can be used as a substitute for bay leaves in soups, stews, and roasts, and also in teas (Tull 1987). Often planted in landscapes. Some birds eat the fruits (Elias 1980).
Trees and shrubs, often with a high tannin content. Leaves deciduous or evergreen, usually alternate (rarely opposite or whorled), simple, entire to lobed, straight-veined; stipules deciduous. Plants usually monoecious, anemo-philous (rarely dioecious or insect-pollinated). Male flowers usually in axillary catkins or head-like clusters, calyx with (4)6(7) small scale-like sepals, these sometimes connate or reduced or nearly absent; corolla absent; stamens (4)6 to 12(40), filaments slender, anthers dehiscing by longitudinal slits, vestigial pistil sometimes present. Female flowers solitary or clustered at the base of the male flowers or in the axils of other leaves, individually or collectively subtended by an involucre which later develops into a cupule; staminodia present or absent; ovary inferior, with 3 to 7 apical sepals (or sepals absent), carpels (2)3(7 to 12), united; styles as many as the carpels, distinct; locules as many as the carpels but the septa not complete at the apex of the ovary cavity; ovules 2 per locule, all but one aborting. Fruit a 1-seeded nut with a hard or leathery pericarp, subtended by the accrescent involucre/cupule (rarely fruit samara-like, but not in ours), seed with a starchy or oily embryo and no endosperm.
6 to 8 genera and roughly 800-1,050 species in the temperate regions of the world except southern and tropical Africa; 3 genera and 41 species listed for TX (Hatch, et al. 1990); 2 genera and 11 species here.
Many forests of the world are composed largely of members of this family, and it is one of the most important timber families. Beech (Fagus), Oak, (Quercus), and Chestnut (Castanea) provide hard, durable wood for construction and beautiful materials for furniture and cabinetry. Some species of chestnuts, beechnuts, and acorns are edible, and many species provide food and shelter for wildlife and domestic animals (Elias 1980). Many are cultivated as ornamentals.
1. Leaves with evenly dentate to serrate margins and manifest pinnate venation; nuts in groups of 1 to 3, each group completely enclosed within a spiny involucre ......1. Castanea
2. Leaves with entire, lobed, or remotely toothed margins and without manifest pinnate venation; nuts solitary, each usually only partly covered by an involucre; involucres spineless .................................................................................................................2. Quercus
Trees and shrubs, bark often fissured and brown. Leaves alternate, deciduous, petiolate, conspicuously straight-veined, dentate or serrate, the teeth usually tipped with bristles. Flowers appearing after the leaves, in axillary catkins at the ends of the branches. Staminate flowers in slender, flexuous, somewhat interrupted catkins, the upper catkins sometimes with a few pistillate flowers at the base; staminate flowers whitish to cream, often heavily scented; calyx 6-lobed, stamens up to 20. Pistillate flowers (those not on primarily staminate catkins) in clusters of (2)3(9) within a scaly, ovoid involucre, ovary 3- to 7-celled, topped with the 6-lobed calyx. Nuts 1 to 3 within the spiny, globose to ovoid, 2- to 4-parted involucre, shiny brown and leathery, borne on the current season's wood and maturing in the same year.
About 12 species of the temperate region; the two formerly-separate species in TX now treated as one, which we have (Johnson 1988).
C. sativa, the Common, Spanish, or Sweet Chestnut, is the one cultivated for nuts (eaten roasted, candied = marrons glacés, or ground into flour for soups) and for ornament. It is also cut close and the branches intertwined as hedge-fencing in Europe. Other species are also planted for timber or nuts (Mabberley 1987). C. dentata, American Chestnut, was once widespread and valued for wood and nuts, but has been almost exterminated by chestnut blight. Efforts are underway to develop disease-resistant cultivars.
1. C. pumila (L.) Mill. var. pumila Downy Chinquapin, Allegheny Chinkapin. Small tree or shrub, sometimes stoloniferous, to a maximum of ca. 15 m tall. Bark gray to gray-brown, slightly fissured, scaly. Branchlets brown, tan, or yellow-green, puberulent to tomentose with a mixture of simple, acicular, solitary or fascicled, stellate, and a few bulbous hairs. Petioles 2 to 22 mm long; blade size and shape variable, generally elliptic to oblong-ovate, 4.1 to 21.7 cm long, 1.5 to 8.3 cm long, apex acute to obtuse, base cuneate to cordate or oblique, margins serrate with 1 to 24 mucronate-aristate teeth per side, sinuses 1 to 5 mm deep, upper leaf surface green and glabrous to puberulent, lower surface whitish glabrate to tomentose; sun leaves smaller, more shallowly-toothed, more densely pubescent than shade leaves or leaves of vigorous shoots. Catkins 1 per axil at the ends of the branches, erect, horizontal, or drooping, yellow-white, slender, with a long main axis, the side axes small cymules, distal catkins androgynous, shorter, with 1 to 8 pistillate clusters at the base, short, wholly pistillate catkins sometimes present. Sepals in both sexes ovate to triangular, 6, imbricate, pubescent, ca. 1 mm long. Staminate flowers usually in groups of 7, each group subtended by a bract; male flowers with (6)12(18) stamens, each 2 to 3 mm long, reduced pistil sometimes present. Pistillate clusters usually consisting of 3 bracts and a single flower; flowers with (4)6(9) styles, these basally pubescent, 1 to 3 mm long; bracts of the pistillate involucre forming a densely or remotely spiny bur 1.5 to 4.6 cm long and about as broad, with 2 triangular valves fused till maturity and then splitting along 2 sutures to reveal the single nut. Nut 7 to 21 mm long, indehiscent, ovoid-conical, circular in cross-section or angled in the rare cases where there are 2 or more nuts in the involucre, shiny chestnut-brown except for the tan attachment scar, sweet, edible. Sandy hills and open woods and thickets in E. TX; E. MA S. to FL, W. TN, AR, and TX. Mar.-Jun.
Until recently it was thought that the chestnuts of the SE. U.S. were of several species and subspecies. It is now known that most of the variability is accounted for by differences in environment--the plants' response to fire, logging, and chestnut blight (Johnson 1988). Undisturbed stands are usually tree-like. These may be connected by stolons to different-seeming shrubby specimens. [Synonyms for var. pumila include C. pumila (L.) Mill. var. Ashei Sudw., C. pumila (L.) Mill. var. margaretta Ashe, C. alnifolia Nutt. and var. floridana Sarg., and C. paucispinna Ashe]. A second variety, var. ozarkensis (Ashe) Tucker exists, but is not found in TX.
The wood is used mostly for fencing as the plants are often small. The nuts are eaten by humans and wildlife.
Deciduous or evergreen trees or shrubs; wood hard, pith continuous, star-shaped in cross-section; branchlets often fluted; buds crowded at the ends of the branches, scales imbricate. Stipules usually deciduous, subulate to ligulate, associated with the buds rather than the leaves. Leaves petiolate, alternate, simple, pinnately veined, entire to toothed or lobed, the lobes or teeth with or without bristles. Staminate catkins pendulous; male flowers apetalous, with (2)5(8) sepals fused into a cup shape, enclosing (3)5 to 10(12) stamens, filaments slender, free, anthers short. Female flowers 1 to several in reduced catkins in the axils of the leaves of the new season's growth, each flower surrounded by an involucre of many scales (representing a complete, reduced inflorescence); calyx of 6 fused sepals adherent to the style bases, ovary inferior, 3-carpellate and 3-loculed, styles 3, free, stigmatic ventrally near the expanded apex; ovules 2 per locule. Fruit a 1-celled and 1-seeded nut (acorn) with a hard pericarp, at least partially enclosed by the hard cupule/ involucre of scales, before maturity nearly completely enclosed. Flowers appearing in the spring with the leaves and fruit developing in 1 or 2 years so that mature acorns appear on either the previous or current season's growth.
About 500 species in the N. hemisphere, in the W. hemisphere, from Canada to Colombia; ca. 38 species in TX (depending upon interpretation); about 10 here. (The number varying according to which varieties one choses to raise to specific rank; this is a conservative treatment.) Oaks are divided into two groups (White vs. Red or Black) on the basis of morphological characters. Some of the author's tentative work with enzyme chemistry and chloroplast DNA analysis suggests that there are also chemical and genetic bases for the division.
Oaks are the dominant component of many N. temperate forests. The wood is hard and durable and has been used throughout history for construction, furniture, veneer, shipbuilding, flooring, barrels, fuel, and so on. Cork is the bark of Q. suber. Many oaks are cultivated for ornament (Mabberley 1987). Acorns (and to a lesser extent, foliage) are an important wildlife food, eaten by deer, bear, hogs, raccoons and other small mammals as well as gamebirds, waterfowl, woodpeckers, and jays (Elias 1980). Some domestic stock feed on oak; foliage and acorns together should not form more than 50% of the forage as the high tannin content can be injurious. Some species produce acorns edible by humans and these have been a staple of many Native American and European peoples. The tannins must be leached out before the acorns can be roasted or made into flour (Tull 1987). The oil from the kernels can be used in cooking. Oak bark and the insect-formed galls on oak foliage are good sources of dyes ranging from green to orange, brown, and black. Tannic acid is a natural mordant and the colors often darken with exposure to light. Some species are used in the leather-tanning process (Tull 1987).
NOTE: Several species not collected in our region occur just outside our area and may someday be found here. Because they are quite similar to some of our species, they are included in the following key, marked with an asterisk, but not described in the text. The key is based on mature leaves and acorns. Oak leaves are highly variable and leaves from seedlings or fast-growing sprouts may not key here. Examine leaves from several branches.
Most oaks will hybridize with most others within their subdivision. Hybrids are often intermediate between the parents in appearance.
1. Leaves or lobes or teeth bristle-tipped, if round-lobed then aristate from the veins (bristles sometimes broken off); acorns maturing on the previous year's growth; cup scales usually tightly appressed apically; acorn shell tomentose within. (Red or Black Oaks) ...................2
1. Leaves or lobes or teeth not bristle-tipped, instead rounded to mucronate; acorns
maturing on current year's growth; cup scales loosely appressed apically; acorn shell glabrous within. (White Oaks) ...............................................................................................14
2(1) Leaves entire or with a few irregular teeth or undulations ......................................................3
2. Leaves distinctly lobed, parted, or cleft, or with 3 shallow terminal lobes ..............................7
3(2) Leaves much broader apically than basally .............................................................................4
3. Leaves not much broader apically than basally ......................................................................5
4(3) Petioles and twigs glabrous or glabrate; leaves generally less than 10 cm long .1. Q. nigra
4. Petioles and twigs pubescent; mature leaves more than 10 cm long .......2. Q. marilandica
5(3) Leaves persistently gray-pubescent beneath ......................................................3. Q. incana
5. Leaves somewhat pubescent to essentially glabrous beneath, at least not persistently gray-pubescent .........................................................................................................................6
6(5) Leaves thick, semi-evergreen, ca. 3 times longer than broad or shorter, flat when
emerging from the bud ........................................................................................* Q. laurifolia
6. Leaves thin, deciduous, at least 4.5 times longer than broad, revolute when emerging from the bud .........................................................................................................4. Q. phellos
7(2) Leaves apically 3-lobed or much broader apically than basally and clavate to spatulate in overall outline ............................................................................................................................8
7. Leaves not apically 3-lobed or much broader apically than basally, usually with a few to several lateral lobes ...............................................................................................................10
8(7) Petioles and new twigs glabrous; leaves glabrous below except for hairs in the axils of the major veins; cup covering base of acorn ................................................................1. Q. nigra
8. Petioles and new twigs tawny-pubescent; leaves variously pubescent below; cup covering about 1/3 of the acorn ..............................................................................................................9
9(8) Leaves densely tawny-tomentulose below ..........................................................5. Q. falcata
9. Leaves variously pubescent below, never tomentulose .............................2. Q. marilandica
10(7) Lateral lobes simple, perhaps with several bristles from the veins, but not with aristate secondary lobes or teeth ........................................................................................................11
10. Lateral lobes usually with a few to several teeth or secondary lobes, each of these aristate ....................................................................................................................................12
11(10) Terminal lobe elongate, usually with 2 lateral, aristate teeth; leaves often densely
pubescent below ...................................................................................................5. Q. falcata
11. Terminal lobe rounded to acute, without lateral teeth (but sometimes extra veins bristle- tipped); leaves glabrous below ...............................................................................1. Q. nigra
12(10) Leaves densely tawny-pubescent beneath OR leaf bases U-shaped ..............5. Q. falcata
12. Leaves variously pubescent to glabrous below; bases rounded to cordate or obtuse, not U-shaped .................................................................................................................................13
13(12) Acorn cup covering less than 1/3 of the acorn; buds 5 mm long or less, glabrous to sparsely pubescent (use a lens) ....................................................................6. Q. shumardii
13. Acorn cup covering 1/3 of the acorn or more; buds 5 to 10 mm long, densely tawny- or gray- tomentose or strigose ..................................................................................* Q. velutina
14(1) Leaves entire or only slightly toothed or undulate .................................................................15
14 Leaves distinctly lobed ............................................................................................................16
15(14) Leaves evergreen or nearly so; margins revolute; acorn fusiform to narrowly ovoid ..............
.........................................................................................................................7. Q. virginiana
15. Leaves deciduous; margins not revolute or only slightly so; acorn ovoid to elliptic to subrotund ...............................................................................................................* Q. durandii
16(14) Lower leaf surface glabrous, glaucous, or essentially so .....................................................17
16. Lower leaf surface pubescent (sometimes minutely so; use a lens) ...................................18
17(16) Leaves regularly and usually deeply lobed, the lobes rounded; cup covering less than 1/2 of the acorn ..................................................................................................................* Q. alba
17. Leaves usually irregularly lobed, lobes acute to rounded; cup covering 1/2 to all of the acorn ........................................................................................................................8. Q. lyrata
18(16) Mature acorn 1/4 to 1/2 enclosed by cup; leaves usually with 3 to 5 (total) deep, usually paired lobes, these often constricted at the bases or truncate or square, leaves often cruciform ...............................................................................................................9. Q. stellata
18. Mature acorn 1/2 to totally enclosed by cup, cup sometimes fringed along the rim; leaves variously shaped, often broadest above the middle, lobes in regular pairs or not, usually more than 5, leaves almost never cruciform .........................................................................19
19(18) Acorns 2 to 5 cm long, cup covering ca. 1/2 to 3/4 the acorn; leaf lobes generally rounded, usually regular and paired, terminal lobe often much larger than the others; leaves overall 7.5 to 15 cm broad ....................................................................................10. Q. macrocarpa
19. Acorns 1 to 2.5 cm long, cup covering most of the acorn, the orifice smaller than the diameter of the nut; leaf lobes acute to rounded, sometimes irregular or unpaired, terminal lobe not much larger than the others; leaves overall 2.5 to 10 cm broad ...........8. Q. lyrata
1. Q. nigra L. Water Oak. Medium to large tree to 15(25) m tall, canopy without competition broad and rounded; trunk to 0.5(1) 1 in diameter; bark dark gray to black, hard, smooth or with wide, scaly, rough ridges, in our area, at least, often with horizontal bands of white lichen giving the tree a striped appearance; branchlets dark red-brown the first year, 1 to 2 mm in diameter and fluted; by the second year becoming gray and glabrate; winter buds 3 to 7 mm long, ovoid, strongly angled and subacute, dark red-brown, the scales densely tawny-strigose above the middle; stipules 6 to 10 mm long. Leaves deciduous but coriaceous, usually 5 to 10 cm long and 2 to 5 cm broad, quite variable in shape, those of saplings, suckers, and fast-growing shoots often quite different from leaves of mature branches, typical leaves generally cuneate to clavate or spatulate in overall outline, with 3 obscure to shallow, rounded terminal lobes, these often without obviously aristate tips by the end of summer, sometimes leaves with 5 or more teeth or lobes, the lobes usually acute, sometimes rounded, sometimes leaves nearly entire, base cuneate or narrowly rounded, margins flat to minutely revolute, upper surface glabrate to minutely puberulent, usually glossy and glabrous, lower surface glabrate except for tufts of hairs in the axils of the major veins; petioles 3 to 12 mm long, sometimes somewhat winged with the decurrent blade, sparsely stellate-tomentose, becoming glabrous or retaining pubescence. Staminate catkins 4 to 7.5 cm long, loosely-flowered, densely to sparsely arachnoid or villous. Female flowers 1 to 3 in catkins 3 to 5 mm long. Fruits maturing on previous season's growth, single or paired on peduncles 0 to 5 mm long. Cup saucer-shaped, shallow, 9 to 15 mm broad and 2.5 to 5.5 mm high; base flat to slightly rounded, margins not incurved, scales narrowly ovate, appressed, densely tawny silky-pubescent. Acorns 8 to 18 mm long, 9 to 15 mm broad, hemispherical to sub-globose, the base very flat, apex rounded, minutely pubescent, dull brown beneath the hairs, sometimes faintly striped, covered by the cup only at the base. Woods, most often lower and moister woods and bottomlands, largest trees found in floodplains. E. TX; N. to MO and DE, E. to FL, essentially on the Gulf and lower Atlantic coastal plains. Mar.-Apr., acorns maturing in the fall. Fall color tan; new leaves in spring very bright acid green. [Includes var. tridentifera Sarg.; Q. aquatica Walt.].
The nuts are a good wildlife food, but the wood is inferior to that of other species (Elias 1980). Sometimes planted as a street or lawn tree and beautiful if given enough water.
2. Q. marilandica Muenchh. Blackjack Oak. Small tree to 10(12) m; canopy usually asymmetrical and the branches irregular; trunk seldom very straight, to 4 dm in diameter; bark dark, roughly furrowed, pinkish to ivory-colored within; branchlets of the current season to 4 mm thick, fluted or round, tomentose, by the second year glabrate and gray; winter buds 4 to 8(10) mm long, ovoid to lanceolate, acute, reddish-brown and tawny-villous or -strigose; stipules 6 to 8 mm long. Leaves deciduous, thick and stiff, 10 to 18(25) cm long and nearly as broad, commonly smaller, cuneate to clavate in outline, typically with 3 shallow apical lobes, these acute to rounded or rounded with minute acute tips, bristles from vein tips usually broken off by late summer, rarely leaves subentire or with paired basal lobes, base cuneate to cordate, margin revolute and thickened, both surfaces densely pubescent in spring, upper leaf surface becoming glabrate or slightly pubescent on the midrib at the base, lower surface with persistent pubescence or glabrate with tufts of stellate hairs in the axils of the major veins; petioles 1 to 1.5(2) cm long, pubescent at least at first and remaining so or becoming glabrate; new leaves in spring often tinged with red. Staminate catkins 5 to 12 cm long, usually loosely 30- to 50-flowered, stellate tomentose to villous. Female flowers 1 to 2 at the ends of the new growth. Fruits single or paired, maturing the second year. Cupule to 20 mm broad and ca. 12 cm deep, hemispherical to goblet-shaped, base rounded to constricted, margin not or only slightly incurved, scales relatively few, somewhat truncate at the apex, reddish-brown, all but the margins buff-colored silky or tomentose, loosely to tightly appressed. Acorns 1.8 to 2.4 cm long, 0.9 to 1.5 cm broad, ellipsoid, rounded at the base and apex, minutely villous at first, becoming light brown and glabrous, ca. 1/3 covered by the cup. Upland forests, usually on poor sandy or clay soils or on gravel hills, often along upland drainages, a common pioneer species. E., Cen., and N. Cen. TX; E. to FL, N. to IA, SE. MI, and S. NY. Mar.(Apr.) [Q. nigra of Coult., not of L.; Q. marilandica var. Ashei or var. ashii (Sudw.) Cory & Parks--this recognized by some authors].
The plants provide cover and food for animals. The wood is good only for fuel, railroad ties, and charcoal (Elias 1980).
3. Q. incana Bartr. Bluejack Oak, Sandjack Oak, Upland Willow Oak. Small tree or large shrub to 8(12) m tall; crown open and irregular; trunk short, to 15 cm in diameter; bark gray-brown to nearly black, breaking into thick, nearly square plates; branchlets 1 to 2 mm thick, somewhat to strongly fluted, yellowish- or reddish-brown and pubescent when young, becoming gray and eventually glabrous; winter buds 5 to 8 mm long, conic or narrowly lanceolate, reddish-brown, appressed pubescent or glabrate; stipules ca. 7 mm long. Leaves deciduous, thin but somewhat leathery, elliptic to oblong, lanceolate, long-ovate or long-obovate, 5 to 8(12) cm long, 2 to 3 cm broad, usually in the small end of the range, entire or with a few small lobes or teeth, especially near the usually acute, aristate tip (apices sometimes rounded, but always aristate), base rounded to cuneate, margin moderately revolute, upper surface dull pale green and lustrous but minutely stellate-pubescent, lower surface pale green-gray, with a felt-like tomentum of stellate hairs, rarely only sparsely stellate-pubescent; petioles 4 to 9 mm long, somewhat winged, entirely pubescent or the top and bottom glabrous and sides with pubescence like the twigs. Staminate catkins 4 to 7.5 cm long, moderately densely flowered, peduncles stellate-tomentose and perianths ciliate. Pistillate flowers 1 or 2 in a subsessile catkin. Fruits solitary or paired, short-pedunculate or subsessile, maturing on previous season's growth. Cup to 18 mm broad and 7 mm tall, goblet- or saucer-shaped, margins not incurved, scales ovate with narrow apices, closely appressed, gray or tawny tomentose except for the shiny brown margins. Acorns 1 to 1.4 cm long, 1.2 to 1.5 cm in diameter, subrotund, base flat, dull brown beneath the gray puberulence, ca. 1/4 to 1/3 covered by the cup. Typically on sandy uplands and sand hills, also on richer soils. Cen. and E. TX; VA to FL, W. to TX. Mar.-Apr. [Q. cinerea Michx.].
Bluejack provides food and cover for wildlife, but the trees are too small for much more than fuel and fence posts (Elias 1980).
4. Q. phellos L. Willow Oak. Medium to tall tree, to 20(30) m tall; canopy broad and rounded or else pyramidal; trunk short, 7.5 to 12 dm in diameter; bark reddish-brown to gray and smooth when young, becoming darker and rough when older, in our area commonly with horizontal stripes of whitish lichen; twigs 1 to 2 mm thick, fluted, reddish-brown and glabrous to stellate-tomentose the first year, becoming glabrous or sometimes persistently pubescent; winter buds 3 to 4 mm long, lanceolate or narrowly ovoid, acute, the scales dark chestnut, glabrous with paler ciliate margins; stipules 6 to 8 mm long. Leaves deciduous but coriaceous, 6 to 12(16) cm long, 1 to 2.5(4) cm broad, usually linear-lanceolate or lanceolate but occasionally narrowly oblanceolate, ovate, or obovate, apices rounded to acute, mucronate, cuneate to rounded basally, margins entire, flat to undulate or barely revolute, leaves when emerging from bud strongly revolute and elongate before unrolling, upper surface green, dull or lustrous, glabrous or with some pubescence on the midrib near the base, lower surface paler, villous to tomentose at first, becoming glabrous or retaining dense tufts in the axils of the veins or densely tomentose along the midrib; petioles very short, 1 to 4 mm long, stout, densely stellate pubescent at first, sooner or later becoming glabrate; spring color bright acid green. Staminate catkins 2.5 to 3.5(7.5) cm long, villous, moderately densely flowered. Female flowers 1 to 3 in a cluster 1 to 3 mm long. Fruits solitary or paired on peduncles to 5 mm long, maturing on the previous season's growth. Cup 1 to 1.5 cm broad, 4 to 8 mm high, saucer-shaped to goblet-shaped, base flat to constricted, margin not inrolled (or only very slightly so), scales narrowly ovate, apices attenuate, closely appressed, red-brown, all but the tips and margins minutely tawny or grayish pubescent. Acorns subglobose to ellipsoid, 1 to 1.5 cm long, flat-bottomed, the tip rounded, dull medium-brown beneath the puberulence or else glabrate, ca. 1/4 covered by the cup. Moist woods, the largest trees where water continuously available, often near swamps or streams but sometimes also on shaded upland sites. E. 1/4 TX; E. to FL, N. to OK, MO, IL, and S. NY. Mar.-Apr.
This oak has rapid growth, unlike most of its cousins. It is an important food source for deer, squirrels, and turkeys, and is of lesser importance to smaller birds. When the acorns fall into water, waterfowl will eat them. The wood is marketed as "red oak" and is used for furniture, interior cabinetry, and stairs (Elias 1980). Often planted as a street or lawn tree.
5. Q. falcata Michx. Spanish Oak, Southern Red Oak, Cherrybark Oak, Swamp Red Oak. Large tree to 20(4) m tall, canopy narrow and rounded in woods, broader and more open in cleared areas; trunk to 5 dm in diameter; bark roughly furrowed, black; twigs 1 to 3 mm thick, fluted, reddish-brown, sparsely to densely tawny-stellate pubescent, becoming glabrate by the second year; buds 3 to 8 mm long, narrowly ovoid, acute, round or somewhat quadrangular, dark red-brown, pubescent all over or mostly at the apex; stipules ca. 1 cm long. Leaves deciduous, thin, obovate to ovate in overall outline, to 23(30) cm long, and 15 cm broad, polymorphic; some leaves (usually on young wood) bell-shaped with 3 aristate apical lobes, other leaves (usually on older wood) regularly or irregularly pinnately 3- to 7-lobed, the lobes pointed and aristate, terminal lobe typically elongate and with two aristate teeth near the apex, sinuses deep and rounded, margins slightly to strongly revolute, upper surface densely stellate pubescent, becoming glabrate or glabrous and dark shiny green, sometimes somewhat persistently pubescent near the base, lower surface with persistent tawny or rusty-gray pubescence, some leaves loosely stellate-villous (especially shade leaves); petioles 1 to 3.5 cm long (sometimes shorter), glabrous above and pubescent below like the blade. Staminate catkins 7.5 to 12.5 cm long, densely pubescent. Female flowers 1 to 3 in catkins 5 to 10 mm long. Fruits solitary or in pairs on peduncles to 1 cm long, maturing on previous season's wood. Cup saucer-shaped, shallow, 1 to 2 cm broad, 6 to 8 mm tall, sometimes goblet-shaped or turbinate, base flattish to slightly rounded or con-stricted, margin not inrolled, scales ovate, broad-based, tips appressed, chestnut brown, pubescent except for the margins. Acorn ovoid or with a flat bottom, apex rounded, 1.2 to 1.5 cm long, 8 to 15 mm in diameter, dull brown, sparsely pubescent, about 1/3 covered by the cup. Moist or wet woods, river bottoms, or uplands. E. TX; E. to FL, N. to OK, MO, IL, OH, and NJ. Spring, the nuts maturing in fall. [Q. digitata (Marsh.) Sudw.; Q. rubra of Sarg., not of L., and vars. leucophylla Ashe, digitata (Marsh.) Cory & Parks, and triloba (Michx.) Sarg.].
TX material is not generally divided into varieties, but some sources, e.g, Elias (1980), list two which they claim are fairly distinct. Var. falcata is said to have broadly rounded or U-shaped leaf bases (sometimes inequi-lateral), leaf lobes less often falcate and more or less spreading at right angles, pubescence rusty, bark broadly ridged and fissured, lumber prone to crack in the sun and rot in contact with the soil. Var. pagodifolia Ell. has cuneate leaf bases and lobes more regular than in var. falcata and more falcate, pubescence light brown to whitish, bark with shorter ridges than var. falcata and somewhat reddish, trunk taller and branch-free for a greater length, lumber not cracking on exposure to the sun. [Also spelled pagodaefolia; Q. pagoda Raf.; Q. pagodifolia (Ell.) Ashe]. While specimens seen from our area would seem to be all var. falcata, it is possible that both types would be present if our material were analyzed at the varietal level. FNA (1997) and Kartesz (1998) recognize the second type as Q. pagoda Raf.
Like most red oaks, Southern Red Oak provides food and shelter for wildlife and wood for construction, furniture, and fuel (Elias).
6. Q. shumardii Buckl. Shumard Red Oak, Southern Red Oak, Swamp Red Oak. Medium to tall tree to 20(30) m; canopy broad, round and open; trunk to 1(1.5) m in diameter; bark of older trees with light-colored low ridges or plates and darker furrows, inner bark reddish-gray; twigs 1.5 to 3 mm thick, fluted, at first lightly stellate pubescent, soon glabrate and tan to brown or gray; winter buds 3 to 7 mm long, very narrowly ovoid, acute (rarely broadly rounded), glabrous or the apex pubescent, scales light brown to golden, erose and becoming ragged; stipules 6 to 10 mm long. Leaves deciduous, thin, 8 to 20 cm long, 6 to 15 cm broad, broadest above the middle, obovate to ovate in overall outline, regularly pinnately lobed, lobes usually 7(9 to 11), commonly toothed or slightly lobed, each tooth acute and aristate, sinuses narrow and rounded, often reaching half the distance to the midrib or more, leaf bases truncate to inequilaterally rounded, sometimes decurrent on the petiole, margins flat to slightly undulate, very minutely thickened-revolute, upper leaf surface sparsely stellate pubescent at first, soon glabrate and shiny, lower surface paler, glabrate except for slight pubescence on the major veins and tufts of hair in the axils of the major veins; petioles (0.5)2.5 to 5(6) cm long, becoming glabrate with the blades, sometimes rose-tinged at the base. Staminate catkins 8 to 10(18) cm long, only sparsely pubescent, sparsely flowered. Female flowers 1 or usually 2 in catkins 4 to 8 mm long. Fruits solitary or in pairs on peduncles 1 to 5 mm long, maturing on previous season's growth. Cup generally shallow and saucer-shaped, sometimes deeper and turbinate, 2 to 3.5 cm broad and 7 to 10 mm tall, the sides 5 mm high, base usually flat, scales somewhat thickened, closely appressed, ovate, light brown, grayish to yellowish or brownish pubescent except for the rounded tips. Acorns (1.5)2 to 2.5 cm long, 18 to 22 mm broad, ovoid, rounded at the tip and very flat-bottomed, shiny brown under the light-colored pubescence, only the basal 1/4 to 1/3 enclosed in the cup. Moist woods of the E. TX timber region, W. along river drainages to the edge of the Edwards Plateau. TX to FL, N. to KS, OK, SW. MI, and scattered to PA and NJ. Mar.-Apr. Fall color often red or scarlet.[Q. coccinea of Coult., not of Muenchh.; Q. rubra of Coult., not of L.; Q. palustris Coult., not of Muenchh.; Q. texana of Sarg., not of Buckl.; Q. schneckii Britt.; according to TX authors including Q. shumardii Buckl. var. macrocarpa (Torr.) Shinners and var. schneckii (Britt.) Sarg.--the latter variety is recognized by some authors.].
The wood is heavy, fine-grained, and used for furniture, flooring, veneer, and many other items. The trees provide food for wildlife (Elias 1980). Its beautiful shape makes it a good lawn or street tree; it has good fall color in our region in some years.
7. Q. virginiana Mill. Live Oak, Encino. Small to large tree to 20 m tall; crown broad and spreading to 40 m or more, branches sometimes bending to the ground; trunk to 1 to 2 m in diameter; bark dark brown to blackish, rough, shallowly to deeply furrowed and scaly; twigs 1.5 to 2.5 mm thick, rounded or fluted, with waxy stellate pubescence or tawny hairs, by the second year glabrate or sometimes persistently pubescent, gray to brown with age; buds to 2.5 (4) mm long, globose, red-brown, puberulent; stipules ca. 2 to 4 mm long. Leaves evergreen (in our area the old leaves falling in spring), quite leathery, 3 to 9(12)cm long, 1 to 2(6) cm broad, variable in shape, from oblong to elliptic to obovate or oblanceolate, apices rounded to acute or mucronate, bases acute to rounded or cordate, margins strongly revolute or somewhat flatter, entire or with a few teeth or small lobes, upper surface deep green, glabrous or sparsely stellate pubescent near the base, lower surface densely stellate pubescent, the hairs waxy and tightly appressed, veins anastomosing before reaching the margins; petioles 2 to 8(10) mm long, somewhat flattened and winged, glabrate to densely tomentulose. Staminate catkins 2 to 4.5(7.5) cm long, pubescent, peduncled. Female flowers in groups of 1 to several, catkins to 5.5 mm long, peduncled. Fruits solitary or in groups of a few on peduncles to 5(8) cm long, maturing in 1 year on current season's growth. Cup 7 to 20 mm broad, 7 to 12 mm high, goblet-shaped, constricted basally (rarely rounded), scales moderately thickened at the base of the cup, and somewhat loosely appressed, triangular, broadest at the base of the cup and narrower toward the rim, red-brown, densely tawny-gray pubescent or the tips reddish and puberulent. Acorns ovoid to fusiform, 1.5 to 3 cm long, 1 to 1.8 cm broad, dull to shiny brown, glabrous except sometimes with a ring of stellate hairs around the tip of the persistent style; acorn 1/4 to 1/2 included in the cup. March. Coastal TX inland to Cen. TX; E. to FL, along the Atlantic coast to VA; also OK and NE. Mex.
Live oaks have traditionally been treated as two species or as two varieties of Q. virginiana. Research at the University of Texas at Austin (Nixon 1984) suggests that all the plants belong to one species. There is still some debate about whether two species should be recognized; apparently the two are more distinct in other portions of their ranges (FNA 1997). Both types may be present in our area. The coastal variety can be recognized as Q. virginiana Mill. or Q. virginiana Mill. var. virginiana. This plant is usually a tree, with acorn cups rounded basally, fruit subsessile, ovoid or narrowly ovoid, not fusiform, leaves broadest near the apex, margins strongly revolute.
Our other plants may be treated as Q. fusiformis Small or Q. virginiana Mill. var. fusiformis (Small) Sarg. This is a plant of sandy soils of woods and stream banks in interior and central TX. Plants often somewhat shrubby, acorn cups narrowed basally, fruits fusiform and pedunculate; leaves broadest near the base, margins flat or only slightly revolute. [syn. = Q. virginiana Mill. var. macrophylla Sarg.].
Many birds and mammals--including jays, turkeys, and deer--eat the fruits. The wood is the heaviest native hardwood and is used for beams, shipbuilding, and posts and has a high fuel value (Elias 1980). Live oaks are often planted in landscapes but are best in large areas where the shape can develop fully; often seen covered with Spanish moss in the South. Once established, the plants are very intolerant of root disturbances such as heavy food traffic or construction.
8. Q. lyrata Walt. Overcup Oak. Medium-sized tree, 20 to 30 m tall; crown broad and rounded; trunk varying from short and crooked to tall and straight, to 1 m in diameter; bark red-brown to gray-brown, with flat, squarish plates and older trees with deep fissures, lower branches sometimes somewhat pendulous; twigs to 4 mm thick, fluted, at first minutely villous, becoming glabrate and grayish-, reddish-, or yellowish-brown; winter buds 2 to 4 mm long, subglobose to ovoid, obtuse, scales light brown, gray-puberulent; stipules ca. 5 mm long. Leaves deciduous, thin, 7 to 25 cm long, 2.5 to 12 cm broad, usually in the small end of the range, narrowly obovate or oblong-obovate in overall outline, shallowly to deeply pinnately lobed, the lobes irregular, usually acute to acuminate, smaller toward the base and the uppermost somewhat clavate, sinuses wide, rounded or squarish, leaf bases cuneate or attenuate-cuneate, apices acute or sometimes rounded, margins minutely revolute, upper leaf surface glabrous and shiny, lower leaf surface paler, dull green and minutely villous or glaucous appressed-tomentose; petioles 2 to 20 mm long, usually short, minutely pubescent to glabrate. Staminate catkins 4 to 10(15) cm long, densely to laxly flowered, sparingly stellate-pubescent. Female flowers 2 or 3 in short catkins. Fruits solitary or paired on glabrous to tomentose peduncles 0 to 4 cm long, maturing in one year on current season's growth. Cups to 1.5 to 3 cm long, to 2 cm deep, globose to hemispheric, usually flattened at the base, scales generally ovoid, brown, densely covered with tawny pubescence except for the sometimes fringed apices, loosely appressed, those at the base of the cup thickened, those near the greatly-constricted orifice small, thinner, more closely appressed; . Acorns hemi-spheric to ovoid, slightly beaked, light brown, usually 2/3 to nearly entirely enclosed by the scaly cup. Moist woods, along streams, and in bottomlands. E. TX; TX to FL, N. to MO, IL, and IN in the Mississippi valley, E. to TN, VA, MD, and NJ, primarily in the coastal plain. Mar.-Apr.
The wood is hard, heavy, durable, and similar enough to white oak to be sold as such. Large acorn crops, important to wildlife, occur every 3 or 4 years (Elias 1980).
9. Q. stellata Wang. Post Oak. Shrubby to medium-sized tree 10 to 20 m tall; crown broad and rounded; trunk straight or commonly crooked, branches usually irregular and canopy with dead wood and asymmetrical; bark red-brown to dark gray, rough and blocky, with deep furrows and broad ridges; twigs orange to reddish-brown and yellowish-pubescent or mealy when young or sometimes sparsely stellate pubescent, becoming brown or gray and glabrate with age, or sometimes the pubescence persisting and becoming gray; winter buds 3 to 6 mm long, subglobose to ovate, tips rounded, scales dark brown or reddish-brown, sparsely pubescent, becoming glabrate; stipules ca. 3 to 5 mm long. Leaves deciduous, thin to thick and leathery, ca. 10 to 15 cm long, 7.5 to 10 cm wide, generally broadest above the middle and with 3 to 5 short, broad lobes, sometimes cruciform, apex rounded and base cuneate, sparsely pubescent becoming glabrate above, lower leaf surface sparsely to densely tawny-tomentose, becoming less so with age; petioles commonly pubescent, ca. 1.2 to 2.4 cm long. Staminate catkins 3 to 8 cm long, pubescent. Female flowers 1 to 4 in short clusters in the axils of the new leaves. Fruits solitary to 4 per cluster, maturing in 1 season. Cup to 1.5 cm broad, cup shaped to hemispheric or pyriform, base thickened, margins thin and nearly incurved, scales oblong to triangular-ovate, finely pubescent. Acorn ovoid to subglobose to ellipsoid or fusiform, 1 to 3 cm long, rounded at the apex, light brown, glabrous to finely pubescent, 1/2 to 2/3 included in the cup or covered only at the base.
Post oaks are a dominant feature of many TX landscapes. They are drought-tolerant and grow on various soils. The acorns provide food for wildlife. The wood is sold as white oak--it makes a good fuel and is used for fencing, furniture, and interiors (Elias 1980). The trees are planted or left standing in landscapes, but do not have the regular silhouette of other oaks and are extremely intolerant of root disturbance.
Three varieties are recognized in TX, each sometimes treated as a separate species. We can expect all three. The differences are very slight. Determinations are difficult and often require seeing the whole plant in its habitat.
var. stellata Moderate-sized trees, twigs to 4 mm thick; leaves rather thick, generally (4)8(15) cm long and (2)6(10) cm broad, in overall outline obovate, elliptic, or obtriangular, apically rounded and basally cuneate, shallowly or deeply 2- to 4-lobed on each side, the main, upper pair of lobes oblong to clavate, at right angles to the midrib so that the leaf is cruciform. Acorn cups 12 to 25 mm broad, to 18 mm high, cup-or goblet-shaped. Acorns to 3 cm long and 18 mm broad, variously shaped, 2/3 included or covered at the base only. Dry upland woods, often on sandy soils, common and dominant in the Cross Timbers and Post Oak Savannah. Cen. TX and the timber region of E. TX; E. to the Atlantic, N. to MA and KS.
var. paludosa Sarg. = Q. similis Ashe Bottomland Post Oak. Medium to large trees; twigs 2 to 3 mm thick, persistently pubescent. Leaves thin, membranous, (5)12(16) cm long and 8 cm broad, obovate, usually with 2 pair of lateral lobes, the apical pair sometimes clavate but the blade not cruciform, lobes generally narrow. Acorns similar to those of var. stellata. Bottomland woods and along streams. E. TX; S. AR and LA, E. to SC. [Q. ashei Sterrett].
var. margaretta (Ashe) Sarg. = Q. margaretta (Ashe ) Ashe Sand Post Oak, Runner Oak. Low to medium shrubs or small to medium trees branched from the base, branches sometimes trailing; twigs 1.5 to 4 mm thick, pubescence absent to present and persistent. Leaves deciduous (sometimes semievergreen), thin and hard to soft, 5 to 12 cm long and 2 to 10 cm broad, obovate to elliptic, oblanceolate, or cuneate in overall outline, apices broadly rounded, 3-lobed at the apex or 2- or 3-lobed on each side, sinuses deep, rounded, broad to narrow or even closed, lobes rounded to clavate or truncate, sometimes toothed or undulate, not cruciform. Acorn cups 10 to 18 mm broad, 5 to 10 mm tall, deeply cup-shaped or shallower; acorn 10 to 17(20) mm long, 7 to 13 mm broad, ovoid to ellipsoid, ends rounded, ca. 1/3 to 1/2 included within the cup. Deep sands in low woods and in loblolly pine forests in E. and Cen. TX; TX, E. to the Atlantic, N. to VA and OK. [Q. drummondii Liebm.].
10. Q. macrocarpa Michx. Bur Oak. In our area medium to tall trees, 20 to 40 m tall; crown rounded and spreading; trunk straight, to 2 m in diameter, the lower 2/3 often without branches; bark dark or yellow-brown to red-brown, with irregular scaly ridges and deep furrows; twigs usually coarse, 3 to 5 mm thick, fluted, villous or pubescent and orange to reddish-green, becoming dark brown or grayish and glabrate; winter buds 3 to 7 mm long, ovoid or narrowly ovoid, acute to obtuse, reddish-brown or gray-brown, sparingly pubescent or tomentose; stipules persistent or deciduous, ca. 1 cm long. Leaves deciduous, variable, generally 15 to 30 cm long and 7.5 to 15 cm broad, obovate in overall outline, shallowly to deeply 5- to 9-lobed, usually symmetrically so, mid-lateral sinuses the deepest, often to near the midrib, lobes clavate, rounded apically, margins often undulate, terminal lobe largest and basal ones shortest, sometimes leaves somewhat cruciform or on stump sprouts or shade trees subentire, leaf bases rounded to cuneate, margins minutely revolute, upper surface dark green and glabrous or glabrate, dull to lustrous, lower surface gray- or silvery-puberulent, villous with mixed long and appressed-stellate hairs, sometimes the stellate hairs absent and the surface greenish, leaves sometimes more glabrate in autumn; petioles to 2.5 cm long, densely or sparingly pubescent. Staminate catkins 3 to 4(9) cm long, laxly-flowered, the rachis yellow-pubescent. Female flowers in clusters of (1)4(5). Fruits single or paired, sessile or with a peduncle to 2 cm long, maturing in one season. Cup deep, cup-shaped to hemispheric, 3 to 6 cm broad and 2 to 5 cm tall, scales broad-based and narrowed to the apex, basal scales short, apices of rim scales elongate and forming a definite fringe; dorsal surfaces of scales keeled, sometimes apparently fused to neighboring scales, cup as a whole hard, sturdy, grayish-pubescent. Acorn 3 to 5 cm long, 2 to 4 cm broad, ovoid, flat at the base and rounded apically, brown beneath the tawny-gray pubescence, usually 1/2 to 3/4 included in the cup (sometimes as little as 1/4 or entirely included.) Moist woods, along streams, rivers, and creeks. E. and Cen. TX: in a broad band down the N. Amer. continent: N.B. to Sask., S. to SE. MT, SE. NE, OK, and TX, then NE to AR, TN, and OH, sporadic in N. Eng. Mar.-Apr. Some floras list several varieties.
Bur oak wood is hard and heavy. It is cut and sold as white oak for cabinetry, furniture, flooring, boatbuilding, etc. Large seed crops occur every 2 or 3 years and are an important food source for wildlife (Elias 1980). The trees are tolerant of drought and various soils and climates. Sometimes planted on lawns and in landscapes for its beautiful shape and dense shade.
Deciduous trees or shrubs. Leaves simple, alternate, usually pinnately straight-veined, commonly serrate. Stipules deciduous. Flowers unisexual, plants monoecious. Calyx of 1 to 6 minute, scale-like sepals or absent. Male flowers small, in spreading or drooping catkins with imbricate bracts or scales subtending 1 to 3 flowers each; stamens equalling and opposite the sepals or else up to about 18, filaments short, free or connate at the base, bifid, pollen sacs of each anther separate (except in Alnus); rudimentary pistil present or absent. Pistillate flowers grouped in 2's or 3's in short, pendulous or erect, often woody catkins or the catkins reduced to clusters of 1 to few flowers; gynoecium typically inferior (sometimes nude), of 2(3) united carpels, styles distinct or nearly so, ovary 2-(3-)locular below, unilocular above, the ovules pendant from near the top of the incomplete partition, 1 to 2 per locule; staminodia absent. Fruit a 1-celled, 1-seeded nutlet, nut, or 2-winged samara, indehiscent, sometimes subtended or nearly enclosed by 2 or 3 foliaceous bracts.
6 genera and 150 species in the N. temperate regions and tropical mountains; 4 genera and 6 species in TX; 4 genera and 4 species found here.
The family is important for timber and furniture woods--especially Birch (Betula) and Alder (Alnus). Some, primarily Corylus (Hazelnut or Filbert), have edible nuts. The fruits, buds, or twigs of some species are moderately important wildlife foods (Elias 1980; Mabberley 1987).
1. Fruits nutlets borne in papery cones to 5 cm long; staminate flowers without a calyx; margins doubly serrate; leaves rounded to slightly cordate at the base ................................2
1. Fruits small samaras in catkins or woody conelike catkins; staminate flowers with a calyx; leaves serrulate to doubly serrate, narrowly to broadly cuneate to subtruncate basally .......3
2(1) Bracts of pistillate inflorescence becoming inflated and bladder-like, enclosing the fruits;
bark exfoliating in vertical strips; staminate catkins for next season exposed over the winter
.. ................................................... ................................................................................1. Ostrya
2. Bracts of pistillate inflorescene flat, 3-lobed; bark smooth and trunks sinewy; staminate
catkins enclosed in buds over the winter..... .........................................................2. Carpinus
3(1) Trees with exfoliating bark; leaves doubly serrate; bracts of solitary pistillate aments 3- lobed, papery; stamens 2, bifid ..................................................................................3. Betula
3. Shrubs with smooth or scaly bark; leaves serrulate; bracts of racemose pistillate aments unlobed, aments eventually woody and conelike; stamens 4, not bifid ....................4. Alnus
Small trees. Wood very hard, bark furrowed, rough, scaly or flaky. Winter buds acute. Leaves with petioles 5 mm long or less, open and concave in bud, usually broadest near or below the middle, doubly serrate. Flowers appearing with or before the leaves. Staminate catkins in groups of 1 to 3 from buds at the tip of the previous year's growth, exposed over the winter, flowers consisting of several bifid stamens in the axil of each bract, perianth absent. Pistillate catkins solitary, drooping on the short shoots of the current season, bracts deciduous, each subtending 2 flowers; ovary of female flowers enclosed by the fusion of perianth and bracts, topped with a border of adherent calyx, 2-celled, 2-ovuled, stigmas 2, long and linear. In fruit bracts and perianth forming a bladdery or papery involucre enclosing and much exceeding the small smooth or ribbed nutlet; involucres loosely overlapping to form a structure resembling the fruiting structure of hops (Humulus).
Ca. 10 species from N. temperate regions to Cen. Amer.; 3 species in TX; we have the 1 species found in E. TX.
1. O. virginiana (P. Mill.) K. Koch American or Eastern Hop-hornbeam, Leverwood. Small upright tree with straight trunk and wide-spreading crown, to 20 m tall, older bark rough and scaly or shaggy, branches slender, sometimes somewhat pendulous, branchlets 2-ranked, young branchlets pubescent, sometimes also glandular pubescent, becoming glabrate. Leaves ovate-oblong to elliptic-lanceolate, (2)7 to 12(15) cm long, (1.2)3.8 to 5(7.6) cm wide, broadest above the rounded to slightly cordate (sometimes tapering) base, apex acuminate, margins usually doubly serrate, teeth sharp, abruptly acuminate, sometimes with hooked tips, downy-pubescent beneath, especially on the veins, and sometimes with tufts of hair in the axils of the veins; petioles 0.2 to 4 mm long, pubescent, sometimes also glandular-pubescent. Staminate catkins clustered, 3 to 5 cm long and pendulous at anthesis, scales broadly triangular-ovate to ovate-subquadrate, to ca. 2.5 mm long, the apex rounded and abruptly pointed or awned, margins ciliate, perianth of male flowers absent, stamens several, bifid, immature male catkins visible in winter. Pistillate catkins on peduncles to 2.5 cm long, smaller than the male catkins, slender, ca. 1 cm long at anthesis, short-cylindrical, loose; flowers subtended by a tubular bracteole-perianth, grouped by 2's, each pair subtended by a deciduous, ovate scale. In fruit, the scales ovoid or ellipsoid, 0.8 to 2 cm long, inflated and papery; fruiting structure to 5 cm long, conelike. Nutlet flattened, ellipsoid or ovoid, 4 to 8 mm long, 2.6 to 4 mm broad, glabrous or puberulent, ribbed. Rich moist or dry woods. E. TX; FL to TX, N. to SE. MA, TN, IL, IA, and SD, N. in Can. to Man. and N.S. Mar.-Apr., specimens with fruit collected mostly Jun.-Aug.
If varieties are recognized or O chisoensis Correll included, ours plantsbecome var virginiana [=var. lasia Fern.], with densely, nearly-permanently villous branchlets.
2. CARPINUS L. Hornbeam
Aboaut 25 species of the N. temperate zone and higher elevations farther S. We have a subspecies of the 1 species found in N. America.
1. C. caroliniana Walt. subsp. caroliniana American Hornbeam, Blue Beech, Water Beech, Lechillo. Small tree with a short, often crooked, fluted, or sinewy trunk to 8 m tall; bark gray or brownish, smooth or slightly rough; branches spreading; wood very hard and heavy; branchlets 2-ranked, sometimes pendulous distally, pubescent when young. Leaves narrowly ovate to oblong-ovate, 3 to 8.5(12) cm long, 3 to 6 cm broad, broadest below the middle, acute to obtuse, margins doubly serrate with the secondary teeth small and blunt, very young leaves sometimes pubescent above, otherwise glabrous, lower surface slightly to moderately pubescent, especially in the axils of the veins; petioles ca. 6 to 9 mm long. Staminate catkins on previous season's growth, enclosed over the winter in quadrangular buds 3 to 4 mm long, solitary or in racemose clusters, 2.5 to 4 cm long, pendulous at anthesis; scales ovate, each subtending (1) 3 flowers on a common receptacle; perianth of male flowers none; stamens 3 (to 6), anthers usually divided into 2 half-anthers and apically pilose. Pistillate catkins produced on short, leafy new growth beyond the staminate catkins, ovoid to short-cylindrical, to 5 cm long; bracts foliaceous, more or less ovate, but with 1 or 2 elongate lateral lobes and so halberd-shaped, entire or with a few teeth on the midlobe, 2 to 3 cm long, 1.4 to 2.3 mm wide; pistillate flowers 2 per bract. Infructescence 2.5 to 7 cm long, rather loose, at maturity each bract subtending and deciduous with 1 small deltoid, ribbed nutlet which is often topped by the persistent sepals and style. Usually in the understory of moist places such as stream banks and bottomland woods. E. TX; uncommon in our area; this subspecies NJ to FL, W. to MO, OK, and TX; the other subspecies N. to Ont., Que., and MN. Mar.-May.
The wood is dense, heavy, and not prone to splitting. It has been used for tool handles and bowls, but the trees are too small to produce lumber of any size (Elias 1980).
Ca. 60 species of the N. Hemisphere; we have the 1 species found in TX.
In many regions more common and more important than here. Many species are valuable for timber or plywood. Twigs are used for brooms and the sap in sweetenings and botanical hair products. Birch beer, medicines, bark for canoes, and dyes come from other species. Birch bark is useful as tinder as it burns even when wet. Many have importance as wildlife food (Elias 1980; Tull 1987).
1. B. nigra L. River Birch. Small to medium tree to 30 m tall, crown pyramidal when young, broad and spreading at maturity; trunk to 8(10) dm in diameter. Bark of older trees and main branches shaggy and exfoliating, reddish, tan, gray, and salmon, oldest trees with gray, fissured bark; twigs reddish-dotted, pubescent or becoming glabrate. Petioles to 1.5(2) cm long, tomentose, slightly flattened and slender, blades ovate to deltoid or rhombic, broadly cuneate to truncate at the base, apex acute, 4 to 8(10) cm long and 8 cm wide, broadest below the middle, margins usually double serrate, at least above the middle, shiny green above, paler and pubescent beneath, sometimes grayish white or sometimes pubescent only on the veins or glabrate. Catkins forming in the fall and developing in spring. Staminate catkins in clusters of 2(3), sessile, 5 to 7.5 cm long, bracts ovate to suborbicular, each subtending 3 flowers, flower with 4 bifid stamens adnate to a 4-lobed calyx with 2 bractlets. Pistillate catkins solitary, upright, pedunculate, 2.5 to 4 cm long, pubescent, scales pedately 3-parted, 6 to 8 mm long, deciduous at maturity, pistillate flowers without a perianth or bractlets. Nutlets/samaras laterally winged, the body as wide as or wider than the ciliate wings, transversely oblong or suborbicular, flattened, 3.5 to 5 mm long, 2.5 to 5.5 mm broad, pubescent at the apex. Along streams and in bottomlands in E. TX; NH to MN and KS, S. to OK and E. TX, E. to FL. Mar.-Apr., fruits falling later and not present in winter as in some other species. Fall color yellow.
Deer browse the buds and twigs; game birds and rodents eat the seeds. The wood is durable but too knotty to use for large objects (Elias 1980).
35 species of the N. temp. region, SE. Asia, and the Andes; we have the one species found in TX.
Many species are useful for timber and carving, the wood of some being a good substitute for costly mahogany. The bark of some is used in tanning leather, some have medicinal properties, others are ornamental (Elias 1980; Mabberley 1987). The roots harbor nitrogen-fixing bacteria. The cones resemble miniature pine cones, are often gold-plated and sold as "pine cones", and can be used in craft projects.
1. A. serrulata Ait.) Willd. Smooth Alder, Hazel Alder, Common Alder. Shrubs or small trees to 5 to 10 m tall; trunk crooked, often spindly, to 10 to 15 cm in diameter, often multi-trunked, crown narrow, rounded, spreading or somewhat ascending; bark of trunk smooth, grayish-or reddish- brown, with darker lenticels; twigs rusty-pubescent; winter buds stalked. Petioles to 15 mm long, pubescent or smooth; blades obovate to obovate-elliptic or elliptic, to 10(12) cm long and 6(8) cm wide, base narrowly to broadly cuneate, apex acute to obtuse or rounded, margins serrulate or sometimes somewhat undulate, glabrous and green above, paler and somewhat pubescent below (especially on the veins), new leaves somewhat sticky and aromatic. Catkins formed in fall and developing in the spring. Staminate catkins in clusters of 3 to 5, pendulous, each scale subtending 3 flowers, flowers with a small 4-parted calyx and 4 not-bifid stamens. Female catkins in groups of 2 or 3, ovoid to ellipsoid, 0.5 to 2.5 cm long, pedunculate, bracts glabrous, 3 to 4 mm long, each subtending 2 flowers and subtending and adherent to 4 minute scales. Fruiting structure semi-persistent, a woody, cone-like structure, its scales cuneate to cuneate-obovate, truncate to lobed, each with 2 to 4 narrowly laterally-winged nutlets or samaras, these obovoid, flattened, 1.5 to 3 mm long, 1.5 to 2.5 mm broad, lustrous. Along streams and in bogs and swamps. Primarily E. TX; ME SE. to KS and E. OK, S. to E. TX and FL. Known at least from bogs in Leon Co. in our region. Mar.-Apr.
Herbs (as ours) or sometimes trees, shrubs, or vines, usually glabrous and sometimes somewhat succulent. Leaves alternate, entire to undulate, stipules absent or very minute; betalain pigments present and anthocyanins absent. Flowers perfect or unisexual, usually regular, solitary in the axils or in axillary or leaf-opposite spikes, racemes, panicles, or cymes. Calyx of 4 to 5(10) sepals, free or somewhat connate below, imbricate in bud. Corolla almost always absent. Stamens 4 to many, often twice as many as the sepals and in 2 alternate whorls, sometimes basally connate. Gynoecium usually superior, of (1)2 to many carpels, free or fused, usually with as many styles and locules as carpels, each carpel with 1 ovule; nectary often present as a ring on the receptacle within or outside the stamens. Fruit dry to fleshy, carpels often separating at maturity. Seeds with a curved, peripheral embryo and perisperm, endosperm absent.
There are 18 genera and variously 65 to 125 species, some of which are only loosely allied to the family. TX has 4 genera and 4 species; 2 genera and 2 species are found here.
The family is of not great economic value. A few are used as dye-plants, medicinals, or pot-herbs. Some are poisonous (Mabberley 1987).
1. Fruits multicarpellate; stamens 10; plants usually much more than 1 m tall ..........................
1. Fruits unicarpellate; stamens 4; plants usually less than 1 m. tall ...........................1. Rivina
NOTE: Agdestis clematidea DC. is known from E.-Cen. TX. No specimens have been seen from our area, but it may be found here in the future. It is a scandent herb with greenish-white to white flowers to 12 mm across and an inferior, dry, winged, 1-seeded fruit.
About 35 species of warm temperate and tropical regions; we have the one species found in TX. Some are edible or provide dyes, others are cultivated ornamentals or are poisonous (Mabberley 1987).
1. P. americana L. Pokeweed, Pokeberry, Inkberry, Pigeonberry. Herbaceous perennial from a large rootstock in 15 cm in diameter. Stems to 3 m (or more) tall, solitary to several from the base, branched above, often purplish (especially late in the season); foliage glabrous, odiferous. Leaves simple, entire; petioles 1 to 5 cm long; blades 10 to 30 cm long, 4 to 10 cm broad, elliptic to lanceolate or lance-oblong to ovate-lanceolate, base cuneate to broadly rounded or attenuate, apex acute to acuminate. Flowers in leaf-opposite racemes 10 to 20 cm long at full anthesis, pedicels 5 to 10 mm long, with 1 or more bracteoles near the middle or base. Flowers regular, bisexual or unisexual, hypogynous. Sepals 5, somewhat petaloid, pinkish to whitish or greenish, darker in fruit, ovate to suborbicular, 2 t 3 mm long; corolla absent; stamens 10; styles and stigmas usually 10; carpels 10, united. Berries in lax racemes, dark purple, 6 to 10 mm in diameter, subglobose (sometimes drying ribbed by the seeds within); seeds usually 10, oval, slightly flattened, 2.5 to 3.5 mm long. Fields, roadsides, waste places, woods, ditches, recent clearings, etc. Throughout most of TX; S. Ont. and S. Que. S. to FL and TX. July-Oct. [P. decandra L.; P. rigida Small].
Very young shoots (only!) may be safely eaten if boiled in several changes of water, otherwise, the foliage is poisonous (Lampe 1985). The roots are also poisonous and the berries should be considered potentially toxic, although pokeweed is an ingredient in some herbal remedies. The berry juice can be used in reddish to black dyes and inks, though the dyes tend to fade. The seeds are high in oils and may someday be economically important (Tull 1987).
About 5 species of tropical and warm temperate regions. We have the one species found in TX.
1. R. humilis L. Pigeonberry, Rouge-plant, Coralito. Herbaceous perennial from a thick rootstock. Stems erect or vinelike and scandent, branches spreading, to 15 dm tall, usually shorter; foliage glabrate or rarely pubescent. Leaves simple, alternate; petioles ca. 1 to 4 cm long; blades ovate to lance-ovate, to 15 cm long and 9 cm wide, base rounded to truncate, apex acute to acuminate, margin entire or frequently undulate. Flowers in terminal or axillary racemes, often the two leaves just below a raceme appearing opposite; pedicels 2.5 to 4 mm long, with a tiny bracteole subtending the calyx. Flowers white to green, rose, or red-purple; calyx of 4 sepals 2 to 2.5 mm long, linear to oblong, enclosing the fruit or becoming reflexed; corolla none; stamens 4. Fruit red (often on red-purple flowered plants) to orange-red (often on white-flowered plants), simple, single-seeded, 2 to 3.5 mm in diameter. Moist or alluvial soils, on chaparral hills, along fencerows, etc. S., Cen., and W. TX; rarer eastward; FL to AR and TX: throughout tropical America. Mar.-Oct. [R. laevis L.; R. portulaccoides Nutt.].
Dyes and inks can be made from the darker fruits (Tull 1987), but like its larger cousin, pokeweed, this plant should be considered poisonous if eaten (Lampe 1985). The two color forms (white flowers/orange-red fruits and red-purple flowers/red fruits) often occur together.
Annual, biennial, or perennial herbs, usually from a woody or fleshy taproot, some (but not ours) shrubs or trees. Stems variously procumbent to scandent or erect, commonly dichotomously or trichotomously branched, glabrous to pubescent, often with swollen nodes; plants producing betalain pigments and not anthocyanins. Leaves opposite, simple, petiolate to sessile, estipulate. Flowers generally perfect or sometimes functionally unisexual and the plants dioecious; flowers regular sometimes dimorphic or heterostylous, usually arranged in corymbose or panicoid cymes, sometimes solitary, racemose, spicate, capitate, or umbellate; inflorescence often subtended by an involucre of free or united bracts, these herbaceous or petaloid, often persistent and accrescent in fruit; sometimes the inflorescence reduced to a single flower and giving the appearance of one large flower with calyx-like bracts and a petaloid calyx. Calyx petaloid, of (3)5(8) fused sepals, tubular to funnelform or campanulate, plicate to valvate in bud, deciduous or persistent, the base often becoming hardened and enclosing the fruit (in some genera, e.g. Mirabilis, with 1-flowered inflorescences, the result is a flower with calyx-like bracts below the fruit and a corolla-like calyx above). True corolla none. Stamens (1) as many as the calyx lobes or sometimes as many as 30, hypogynous, filiform, unequal, often connate basally. Gynoecium included in the corolla tube but superior (appearing inferior in species where the calyx base hardens around the ovary), unicarpellate and unilocular; style 1, elongate, slender, stigma commonly capitate to peltate, ovule 1, nectary disk sometimes present around the ovary. Fruit an achene or nut, often enclosed by the hard or leathery calyx base and termed an anthocarp, fruits often clustered in the involucre; seed often with starchy perisperm.
About 350 species in 45 genera in warm and tropical regions, especially in the Americas, a few in temperate regions. There are 13 genera and 69 species in TX; 3 genera and 8 species in our area, including 1 endangered species.
Most members of the Nyctaginaceae are not economically important. A few have medicinal or food value. Some, notably Mirabilis (Four-o'clock) and Bougainvillea, are cultivated for ornament (Mabberley 1987).
1. Stigma linear or fusiform; flowers in heads ............................................................1. Abronia
1. Stigma spherical, hemispheric, or peltate; flowers in cymes, panicles, or axillary or terminal involucrate clusters ....................................................................................................2
2(1) Flowers less than 4 mm long; involucre absent or reduced, if present, then the bracts not united ....................................................................................................................2. Boerhavia
2. Flowers more than 4 mm long; involucre always present, of united bracts .........3. Mirabilis
Annual or perennial herbs; stems prostrate to erect, branched, sometimes clumped and seemingly acaulescent, pubescent, often viscid. Leaves petiolate, opposite, the two of a pair often of different sizes, blades succulent, entire to sinuate. Flowering stems sometimes scapose; flowers perfect, few to many in long-pedunculate heads subtended by 5 conspicuous, scarious, usually distinct bracts. Perianth funnelform to salverform, tube elongate, slender, and commonly constricted above the ovary, limb 5-lobed, often withering and persisting. Stamens usually 5, included. Ovary ovoid, style filiform and stigma linear or fusiform, included. Fruit turbinate to biturbinate, deeply 2- to 5-lobed or -winged, leathery, hardened, or papery. Embryo with one cotyledon aborted.
About 36 species of N. Amer; 5 species in TX (2 listed by Correll and Johnston (1970) since transferred to Tripterocalyx (Galloway 1975)); 2 species here, one endangered.
Some species, especially A. fragrans, are cultivated as ornamentals (Mabberley 1987).
1. Anthocarp 8 to 15 mm long, 5 to 12 mm broad, papery; leaf margins entire ..........................
......................................................................................................................1. A. macrocarpa
1. Anthocarp 7 to 9 mm long, 3 to 4.5 mm broad, leathery or hard, not papery; leaf margins sinuate ..................................................................................................................2. A. ameliae
1. A. macrocarpa L. A. Galloway Large-fruited Sand Verbena. Perennial herb from a long, fleshy to semi-woody taproot. Stems to 50 cm tall, ascending to semi-erect, glandular-viscid pubescent. Petioles 0.5 to 4 cm long; blades ovate to elliptic or rotund, 2 to 5 cm long, 1.5 to 3.5 cm broad, entire, viscid-pubescent. Floral bracts ovate to elliptic, 7 to 13 mm long, 4 to 6 mm broad; flowers 20 to 45(75) per head, heads to ca. 10 cm broad; flowers magenta, 18 to 30 mm long, tubular, limb 5-lobed, 4 to 10 mm broad, fragrant. Anthocarps turbinate, papery, scarious, 5-winged, somewhat notched at the apex, 8 to 15 mm long, 5 to 12 mm broad. Seeds fusiform, 2 to 4 mm long, lustrous. Bare, wind-blown sand dunes surrounded by post oak woodlands. Known only from small areas of Leon and Robertson Cos. First collected in 1971, described in 1972 (Galloway 1972). On Federal and State endangered species lists. Late Mar.-early June.
2. A. ameliae Lundell Amelia's Sand Verbena. Coarse perennial herb; stems to 60 cm high or long, spreading, thick viscid-glandular pubescent and villous with hairs of variable length, somewhat grooved when dry. Petioles 1 to 8.5 cm long, those of the lower leaves longer than the blades; blades ovate to elliptic or orbicular, 3 to 8 cm long, 2 to 6 cm broad, apices rounded, bases subcordate to subtruncate, decurrent at the base, margins noticeably sinuate, fleshy, leaves drying stiff and brittle, sparsely short viscid-pubescent above and below. Peduncles 2.5 to 13 cm long, slender, viscid-puberulent; floral bracts 5 to 7, thin, oblong to elliptic or obovate, 10 to 16 mm long, 5 to 10 mm broad, apex broadly rounded and acute, greenish to pale magenta, externally viscid-villous. Heads many-flowered, to 5 cm broad; flowers with perianths orchid/pale magenta, 18 to 25 mm long, villous-viscid, tube slender, limb to 1 cm broad, 5-lobed, the lobes emarginate; stamens 5, included. Anthocarp narrowly turbinate, apex rounded to truncate, not narrowed, attenuate at the base, 7 to 9 mm long, 3 to 4.5 mm broad, 5-winged, reticulate-veined, glabrous to sparsely pubescent at the apex; seeds oblong-oblanceolate, ca. 2.5 mm long. Sandy areas along roadsides in post oak belts. Rio Grande Plains to the Panhandle, reported as far E. as Leon Co.; endemic. Mar.-Jun.
Annual or perennial herbs; stems sprawling or trailing, branched, sometimes diffuse, radiating from a central crown, variously pubescent or glandular, internodes often villous. Leaves opposite, the two of each pair unequal, usually paler beneath, entire to sinuate, often acute. Flowers to 1.5 cm broad, perfect, not in involucres but each flower bracted, in cymose panicles or racemes, pedicels jointed just below the ovary. Calyx corolla-like, campanulate to nearly rotate, constricted above the ovary, limb deciduous, shallowly 5-lobed. Stamens 1 to 5, exserted or inclined, filaments of different lengths, connate below. Ovary stipitate, style filiform and stigma peltate. Anthocarp ovoid or obpyramidal, with 3 to 5 angles or, less commonly, 5 to 10 ribs, rarely with 3 to 5 wings, glabrous to pubescent; seed with a curved embryo. Boerhavia is Linnaeus' original spelling, but the name Boerhaavia appears in many works.
About 40 species, primarily tropical and subtropical; 11 in TX with the inclusion of formerly-separate Commicarpus scandens (L.) Standl. as B. scandens L. Two species are found in our area.
A few are used in medicines or as food, but not ours. Some are weedy (Mabberley 1987).
1. Anthocarp glandular, narrowly obovoid, rounded at the apex ............................1. B. diffusa
1. Anthocarp eglandular, narrowly obpyramidal, truncate at the apex ....................2. B. erecta
1. B. diffusa L. Scarlet Spiderling. Perennial from a fleshy or woody, sometimes stout root; stems few to many, decumbent, procumbent, or ascending, sparingly branched below, 2 to 15 dm long, brown or herbaceous, lower portions minutely puberulent to viscid-pubescent, especially near the nodes, upper portions densely glandular-puberulent to somewhat viscid or even glabrate. Petioles 3 to 40 mm long, commonly villous; blades thin or rarely leathery, suborbicular or widely ovate to ovate-oblong, 1.5 to 5.5 cm long, 8 to 50 mm broad, bases subcordate to rounded or truncate, apices rounded to acute, margins entire to sinuate, paler beneath, brown-punctate or not, glabrous to minutely puberulent, margins villous-ciliate, lower surface with veins sparsely villous, sometimes the entire leaf densely hirsute or hirtellous. Flowers in lax, much-branched terminal and axillary cymes, branches divergent or ascending and glabrous to puberulent or glandular, Involucres absent. Flowers sessile to short-pedicelled, in clusters of 2 to 4 at the ends of slender peduncles 3 to 10 mm long; floral bracts minute, less than 2 mm long, deciduous or persistent, ovate or lanceolate, viscid-puberulent. Perianths purple-red to red-green, limb 2 mm broad, all minutely glandular-puberulent; stamens 1 to 3, briefly exserted. Anthocarp narrowly obovoid, rounded at the apex, 2.5 to 4 mm long, densely glandular-puberulent (or glandular-pilose), 5-ribbed or -angled, angles and grooves smooth. Waste places, fencerows, riverbanks, fields, etc. S. and W. TX; W. FL to TX, and SE. CA; Mex. through Cen. Amer. to N. and W. S. Amer.; also W.I. and Caribbean; adventive elsewhere, including once in NC. July-Nov. [B. glabrata Blume; B. caribaea Jacq.].
2. B. erecta L. Erect Spiderling. Taprooted annual or perennial; stems erect to decumbent, usually branched at the base, 2 to 12 dm long or tall, branches spreading or decumbent, sometimes red-tinged, lower portions finely pubescent, middle internodes often with viscid brown bands, upper portions glabrous or minutely puberulent. Petioles 4 to 40 mm long, ca. 1/2 the length of the blades; blades ovate-rhombic or ovate-deltoid, or the upper ovate-lanceolate to linear, 2 to 8 cm long, 1.4 to 5 cm broad, apices acute or obtuse to rounded, apiculate, bases truncate to rounded or somewhat cuneate, margins entire to undulate or repand, both surfaces usually brown-punctate, glabrescent, lower surface paler than upper (or glaucous.) Inflorescences axillary and terminal, cymose, umbellate, or somewhat racemose, usually well-branched, branches filiform, erect to ascending, usually glabrous but sometimes viscid; involucre absent. Flowers in clusters of 2 to 6 on slender peduncles, pedicels 1 to 5 mm long; floral bracts linear to lanceolate, reddish, 0.6 to 1.2 mm long, persistent. Perianths white or tinged with pink to purple, 1 to 1.5 mm long, tube glabrous or glandular-punctate, limb campanulate, sparsely pubescent; stamens 2 or 3, exserted. Anthocarp green or yellow-green, primarily glabrous, narrowly obpyramidal, 3 to 4 mm long, 1 to 1.5 mm broad, apex truncate, 5-angled, the angles obtuse to acute, smooth, the grooves between coarsely cross-rugose. Dry fields, banks, waste places, and in cultivated ground. SC and FL W. to TX, AR, and S. AZ, S. through Mex. and Cen. Amer. to N. S. Amer.; also W.I. and Caribbean. Spring to fall.
Perennial herbs, usually from large tuberous roots; stems erect to ascending or sprawling, simple or branched from the base, branches forking, nodes often swollen; foliage glabrous to pubescent, glandular, or hispid. Leaves opposite, often fleshy, sessile to petiolate, variously-shaped, glabrous to pubescent or glaucous. Involucres terminal or axillary or both, in loose or congested cymose arrangements or involucres solitary in the axils. Involucres each with 1 to 10 flowers, 5-lobed, lobes equal or unequal, calyx-like, green or tinged with red or purple, more or less campanulate, in some species accrescent and papery in fruit, usually becoming rotate and noticeably veined. Flowers perfect, regular, calyx colored and corolla-like, tube elongate and constricted above the ovary, limb expanded, campanulate to salverform or funnelform, 5-lobed, deciduous in fruit but the perianth base persisting and becoming hardened around the ovary. True corolla absent. Stamens 3 to 6 (sometimes more), unequal, filiform, united at the base, anthers exserted. Ovary superior, unicarpellate, style filiform and stigma capitate. fruit a 1-seeded anthocarp (ovary enclosed by the leathery remnants of the calyx), 5-angled or 5-ribbed, surfaces smooth to tubercled, glabrous to pubescent. Seed filling the pericarp, pericarp fused to the testa; endosperm mealy.
About 45 to 60 species of warm parts of the Americas, especially SW. N. Amer.; 29 species listed for TX; 4 from our area. Positive identification often requires mature fruits, and those species from section Oxybaphus form a complex group without clear specific circumscriptions. The following is based on Lundell (1969). B. L. Turner (1993) proposed a different interpretation of TX species.
Some, notably M. jalapa, are grown for ornament; a few are used for dyes or in cosmetics or medicines. The tuberous roots of some are edible (Mabberley 1987).
1. Perianth more than twice as long as the involucre; involucre herbaceous and not enlarged in fruit; anthocarp scarcely angled, not ribbed ......................................................1. M. jalapa
1. Perianth less than twice as long as the involucre; involucre enlarged and papery or membranous in fruit; anthocarp with 5 prominent ribs ............................................................2
2(1) Leaves (except uppermost) obviously petiolate, petiole sharply differentiated from the blade ..............................................................................................................2. M. nyctaginea
2. Leaves sessile or nearly so (sometimes attenuate to a very short, stout petiole), petiole not sharply differentiated from the blade .......................................................................................3
3(2) Angles and sides of fruit tuberculate, tubercles with tufts of silvery hairs; leaves linear- lanceolate or broader .............................................................................................3. M. albida
3. Angles of fruit not tuberculate, sides transversely rugose, the whole fruit pubescent; leaves narrowly linear ......................................................................................................4. M. linearis
1. M. jalapa L. Common Four-o'clock, Marvel of Peru. Perennial from a large, tuberous or fleshy root. Stems erect or sometimes somewhat floppy, 4 to 10 dm tall, well-branched, branches erect to ascending, stout or slender, nodes commonly swollen. Foliage dark green, glabrous to slightly pubescent (rarely short-villous), often viscid, but not glandular. Petioles slender, ca. 1/2 as long as the blades, 0.3 to 5 cm long, reduced upwards and the uppermost leaves nearly sessile; blades ovate, ovate-deltoid, or rhombic to rhombic-elliptic, 4 to 14 cm long, 2 to 8.5 cm broad, bases truncate to subcordate, apices acuminate, glabrous or rarely pubescent. Peduncles 1 to 2 cm long, arranged cymosely at the ends of the branches, inflorescence with many reduced leaves. Involucres green, campanulate or turbinate, 7 to 15 mm long, lobes linear-lanceolate to ovate-lanceolate, acute, usually ciliolate and bristle-tipped, as long as the tube or up to twice as long, glabrous or short-pubescent to villous, enlarged only slightly in fruit and remaining herbaceous. Flower 1 per involucre, perianth salverform to trumpet-shaped, 2 to 6 cm long, deep red-purple to white (in cultivated forms also rose, yellow, and variegated), glabrous to lightly villous externally, tube ca. 2.5 mm thick and gradually expanded upward, limb 2 to 3.5 cm broad, spreading, shallowly 5-lobed, notched, or sometimes (especially in cultivated material) nearly entire; stamens 5, equalling the perianth or slightly exserted. Anthocarp ovoid or obovoid, dark brown to black, 7 to 10 mm long, 5-angled, wrinkled-warty, warty, or rugose, glabrous to puberulent (sometimes described as resembling a miniature hand-grenade.) Plains, prairies, and roadsides, often escaped from cultivation in W. TX, sometimes persistent where established. Native to tropical America, now found throughout Mex., Cen. Amer., and S. Amer. Spring or summer until frost.
Widely cultivated for the colorful flowers and often strongly night-scented. The common name refers to the plants' habit of blooming at dusk; flowers last one day. In cultivated strains the flowers are sometimes much larger and occasionally with extra perianth lobes or stamens (up to about 10); some of these plants are up to 2 m tall. Used by some cultures in medicines. The flowers are used as a food dye in China and the seed in cosmetics in Japan (Mabberley 1987).
2. M. nyctaginea (Michx.) MacM. Wild Four-o'clock. Perennial from a thick fleshy taproot, crown usually branched. Stems several to many, erect to ascending, 3 to 15 dm tall, branched below or above, branches divaricate and forking or sometimes simple, glabrous to pubescent or glaucous, sometimes ridged and/or purplish, nodes usually swollen. Uppermost leave sessile but lower leaves with petioles 1 to 10 cm long; blades ovate- or deltoid-lanceolate to ovate-oblong or cordate, (3)5 to 12(15) cm long, to ca. 5 cm broad, base truncate, rounded, or cordate, apex acute to obtuse or acuminate, glabrous or pubescent (ours apparently tending toward pubescent.) Peduncles 3 to 10 mm long, pubescent, arranged in umbels in forked terminal clusters (sometimes appearing paniculate). Involucres 5 to 6 mm long at anthesis, (8)10 to 15(17) mm long in fruit, membranous, persistent, widely campanulate to almost rotate, lobes 5, obtuse or obtuse-apiculate, united nearly 2/3 their length, margins pilose, sometimes pilose externally or near the base, commonly somewhat reddish and veiny in fruit. Flowers regular and perfect, 3 to 5 per involucre, perianth pink to purple or rarely white, campanulate, tube short (to 2 mm long at anthesis), limb (8)10 to 15(18) mm broad; stamens 3 to 5, exserted. Anthocarp grayish or brown to nearly black, 4 to 6 mm long, cylindric-ovoid or narrowly elliptical, pilose, warty, or rugose, with 5 ribs and 5 angles. Seeds pale brown, 3 to 3.5 mm long, obovoid. Roadsides, weedy areas, calcareous gravels of the Post Oak Belt and West Cross Timbers; Man. and WI, W. to MT, S. to TN, TX, and NM; also Mex.; adventive in the E. U.S. and CA. Spring-summer, ours mostly April and October. [Oxybaphus nyctagineus (Michx.) Sweet; Allionia nyctaginea Michx.; A. ovata Pursh].
3. M. albida (Walt.) Heimerl White Four-o'clock. Perennial from an often woody taproot and a short, branched rootstock. Stems 1 to few, erect to ascending, 2 to 12 dm tall, simple or branched, 4-angled below, nodes swollen, internodes below the inflorescence each with 2 long bands of short (less than 0.5 mm), incurved hairs or else bands of hairs absent and the stems nearly glabrous, often whitish. Leaves sessile or with pubescent petioles to 5(10) mm long; blades variable in shape, linear-lanceolate to oblong-or elliptic-lanceolate or ovate, 3 to 12 cm long, 3 to 30 mm broad, roughly 3 to 12 times longer than wide, thin or thick and succulent, base cuneate or attenuate, apex acute to short-acuminate, rounded or blunt, margins entire to subsinuate, with or without cilia, surfaces glabrous to pilose, upper surface bright green, lower surface often glaucous or whitened. Inflorescence in young plants of solitary pedunculate involucres, in older plants involucres in much-branched cymose-paniculate arrangements, branches of inflorescence and peduncles puberulent with short, incurved hairs or short-viscid pubescent or both, inflorescence with many reduced, bract-like leaves. Involucres with slender peduncles, 1- to 3-flowered, campanulate, 3.8 to 4 mm long at anthesis, persistent and enlarging to (8)10 to 12(15) mm long in fruit, 5-lobed, lobes acute to rounded, united about 2/3 their length, glabrous to sparsely or densely viscid-pilose. Perianth rose, pink, or whitish, (6)8 to 10 mm long, lobes shorter than the tube, glabrous to sparsely (short-) pilose; stamens 3 to 5, exserted, style exserted. Anthocarp dark olive-brown, 5 to 6 mm long, obovoid, broadly angled, angles with cylindrical or flattened tubercles, each tubercle topped with a tuft of silvery hairs, faces also tuberculate. Seed obovoid, yellow-brown, ca. 3.5 mm long. Dry soils of meadows, roadsides, fencerows, waste places, and so on. Most of TX except the Panhandle; SC and TN, N. to Man., W. to ND, KS, MO, LA, and TX. May-Nov. [Includes var. lata Shinners and var. uniflora Heimerl; Allionia bracteata Rydb.; A. albida Walt.; Oxybaphus albidus (Walt.) Sweet; some authors include also M. decumbens (Nutt.) Daniels (= A. decumbens Nutt.)].
See note at M. linearis, below.
4. M. linearis (Pursh) Heimerl Narrowleaf Four-o'clock. Perennial from a deep, elongated, woody taproot, rootstock branched. Stems erect to ascending or procumbent, simple or branched near the base and often branched above, 2 to 10 dm tall, usually glaucous, often whitish, lower portions glabrous to puberulent, upper portions viscid-puberulent or short-villous, especially in the inflorescence. Leaves sparse or numerous, sometimes crowded, linear to narrowly linear-lanceolate, sessile or attenuate to a short petiole, 3 to 10 cm long, 1 to 5 mm wide, generally about 15 to 30 times longer than wide, apex acute or obtuse, entire to (rarely) undulate or sparsely dentate, thickish, generally gray-green above, glaucous usually at least beneath, glabrous to viscid-puberulent. Inflorescences sometimes axillary, more commonly paniculate or cymose-paniculate. Involucres rotate-campanulate, 3.8 to 4.2 mm long at anthesis, persistent and enlarging to (6)8 to 10(12) mm long in fruit, lobes acute to rounded, glandular-pubescent or viscid-villous. Flowers usually 3 per involucre, perianth purplish-red to pink or nearly white, pilose, 8 to 12 mm long, limb deeply 5-lobed, lobes retuse or emarginate; stamens exserted. Anthocarp brownish-olive, 4 to 5 mm long, obovoid, 5-angled, the angles obtuse, densely pubescent to sparsely strigose, not tuberculate, sides transversely rugose. Seeds yellow-brown, 2.8 to 3.2 mm long, rounded-obovoid. Dray areas, especially on calcareous or igneous soils, also sandy plains. Common in the Panhandle, SE. to Hunt, Jack, Nueces, and Kleberg Cos. MN and SD to MT, S. to MO, TX, and AZ; adventive elsewhere. Spring-summer. [M. angustifolia (Nutt.) MacM.; Allionia linearis Pursh; Oxybaphus linearis (Pursh) Robins; some authors also include M. diffusa (Heller) Reed (= Allionia diffusa Heller)].
NOTE: The differences between M. albida and M. linearis, at least in our area, are subjective. Specimens can not be distinguished by leaf shape or pubescence, and even fruit morphology can be inconclusive. Most specimens from our area identified as M. linearis have tuberculate fruits and are probably M. albida. Others have fruits with elongate, transverse tubercles intermediate between the cylindrical tubercles of M. albida and the transverse wrinkles of M. linearis. It is possible that the true M. linearis is not present in our area. R. W. Spellenberg (1991), who has done work with the genus, describes the Oxybaphus section of Mirabilis (which includes these two species) as confused, consisting of entities which have diversified but not fully speciated.
Succulent annual or perennial herbs, less commonly shrubby, a few (but not ours) spiny. Leaves alternate or opposite, usually more or less succulent, often with a centric structure rather than with a top and a bottom, epidermis of stem and leaf often with bladder-like cells; stipules usually absent, if present (as in our Trianthema), usually scarious; some species (none of ours) leafless stem succulents. Flowers solitary or in small cymose clusters, perfect or sometimes unisexual and the plants monoecious. Calyx of (3)5(8) sepals, commonly succulent. Petals (actually staminodes) inserted on a hypanthium or on the apex of the ovary, free or sometimes fused at the base, usually many in 1 to 6 cycles, but sometimes absent. Stamens 4 to 5 or 8 to 10 or sometimes many, inserted on the hypanthium, the base of the corolla, or the apex of the ovary; all free or basally united into groups. Staminodes sometimes present. A ring of nectaries is commonly present at the inner base of the stamens. Ovary inferior or superior, of 2 to 5 (many) fused carpels, usually with as many free or distally-free styles and locules as carpels, rarely unilocular; ovules (1) to many in each locule, placentation axile, basal, apical, or parietal. Fruit usually a loculicidal capsule, often enclosed by the persistent hypanthium or calyx, occasionally fruit of another type. Seeds with the embryo coiled around the perisperm.
This primarily South African family has about 2,500 species in 12 genera, (more genera if the tropical Mesembryanthemum s.l. is broken up). There are 2 genera and 7 species in TX; 1 species in our area.
The Aizoaceae formerly included plants of the Molluginaceae. Cronquist (1981) and others separate the Molluginaceae because it has anthocyanins instead of betalains and because few of its plants are succulent.
Many species are cultivated for their bright, daisy-like flowers and their unusual, succulent leaves and stems (Mabberley 1987).
About 9 species in warmer parts of the world. We have the one species found in North America.
1. T. portulacastrum L. Horse Purslane, Verdolaga Blanca. Annual or perennial succulent herb, glabrous, stems usually decumbent, sometimes ascending, well-branched from the base, to 1 m long. Leaves opposite, the members of each pair unequal, broadly obovate to suborbicular-obovate, the smaller ones perhaps narrower, 1 to 4 cm long and 3 cm broad (the leaves of axillary branches smaller than those of the main stems), apices rounded, notched, or apiculate; petiole shorter than or about equalling the blade, somewhat scarious and dilated at the base; stipules scarious, entire. Flowers sessile (to short-pedicelled) and usually solitary in the axils, sometimes hidden by the petiole base. Calyx green externally and pink or purple within, the margins somewhat hyaline, sepals 5, ovate to lanceolate, concave, ca. 2.5 mm long, a small mucro or horn present on the dorsal apex; corolla none; stamens 5 to 10, in 2 perigynous whorls, if the same number as the calyx lobes, alternate with them, staminodes none; ovary superior, 1-celled or incompletely 2-celled, stigma 1. Capsule circumscissile near the middle, ca. 4 mm long, cylindrical, somewhat curved, with winged appendages at the apex; seeds 1 to several, black, rough, round, ca. 2 mm broad. Sandy soils of dunes, thickets, and stream margins. S. and W. TX; FL to CA, N. to OK, MO, and NJ; S. through Latin Amer.; also Africa and Australia. May.-Oct. (Nov.).
Ours all perennial stem-succulent herbs or shrubs (some others tree-like) with a thick cuticle. Roots usually shallow; stem variously unbranched, columnar, or sparingly branched or cushion-forming, sometimes segmented into "joints" which are cylindric, tubular, or flattened, variously smooth, ribbed, or tubercled; ribs, tubercles, or smooth stems with areoles representing modified axillary buds or short shoots, the leaves or bud scales replaced by spines, spines of a node often identifiable as either central or wider-spreading radial spines, sometimes the areoles also with minute barbed spines or hairs (glochids). True leaves none or scarcely developed, alternate and usually seasonal and fleeting on new growth if present. Flowers usually solitary at or near the areoles or at the base of the tubercles, rarely (and not in ours) at the branch tips or in a cymose arrangement, typically conspicuous and brightly colored, pollinated by birds, bats, bees and night moths, often large in relation to the size of the plant, usually perfect, regular except perhaps for the insertion of the stamens and pistil or a slight curvature of the perianth. Perianth parts many, spirally arranged, typically intergrading from green sepaloid to larger, petaloid members, all united below into a perianth tube or hypanthium, this sometimes with areoles and/or leaves. Stamens many, inserted on the hypanthium; nectary commonly present. Ovary with 3 to many united carpels, usually unilocular with as many parietal placentae as carpels, epigynous (except in Pereskia), in some, e.g. Opuntia, the ovary apparently recessed into the stem; style 1, stigmas usually as many as the carpels, radiating from the top of the style. Fruit usually a fleshy berry, commonly with areoles, sometimes (not in ours) dry and dehiscent; seeds many, with a curved embryo; perisperm present or absent. These plants have betalain rather than anthocyanin pigments.
Depending upon interpretation, 30 to 200+ genera. Many genera are ill-defined or segregated on arguably trivial characteristics. There are 1,000 to 2,000 species in the temperate and tropical New World, especially in arid regions. (Rhipsalis is present in parts of the Old World, whether it is native there is still under debate.) There are 13 genera and 78 species in TX; 2 genera and 5 species in our area. This treatment follows Benson (1982), which is complete for the family. Generic lines are constantly being shifted; newer names are indicated.
Many species are grown as ornamentals. Some, such as Opuntia, have edible fruit; others are somewhat weedy (Mabberley 1987). Many (but none of ours) grow in severely restricted or highly-specialized habitats and are rare or endangered.
1. Stems divided into cylindrical or flattened joints, not ribbed; glochids present in areoles; young joints often with reduced leaves ..................................................................1. Opuntia
1. Stems not divided into joints, ribbed; glochids absent; leaves absent 2. Coryphantha
Coarse perennial herbs and shrubs, often prostrate or clump-forming; stem divided into cylindrical segments (cholla cacti) or flat pads (prickly pears), ribs and tubercles absent or present. Leaves fleshy, cylindroid to subulate, soon deciduous. Areoles with 0 to 10 spines, or spines sometimes absent from an entire plant, barbed or smooth, circular in cross-section or flattened at the base, glochids (small, retrorsely-barbed hairs or spines) present, usually on all the areoles. Flowers at areoles on older stem segments, floral tube above the ovary very short, stamens borne close above the ovary. Fruit indehiscent, a fleshy berry (as in ours) or sometimes dry, spiny or smooth; seeds flat and nearly circular.
An indeterminate number of species, estimated at about 300 from B.C. and N. Eng. to the S. tip of S. Amer. Most recent treatments list about 25 species for TX; there are 4 in our area.
Many species have edible fruits or pads (nopales). The plants can also provide emergency forage for livestock if the spines are first burned off. Some are grown in arid climates as living fences; many are grown as ornamentals. Some have become serious weeds where introduced, as in Aust. and S. Afr. (Mabberley 1987) A dye, cochineal, is extracted from the bodies of mealybug-like insects which feed on Opuntia. It yields red, pink, magenta, violet, and rose colors (Tull 1987).
NOTE: Cacti are best keyed in the field. For flat-jointed species, the presence or absence of spines, thier color, and their position are critical. The one or two joints on herbarium sheets may not have typical spines, and in fact may have been chosen for their lack of spines.
1. Stem segments cylindrical ............................................................................1. O. leptocaulis
1. Stem segments flattened .........................................................................................................2
2(1) Spines absent from all areoles of the plant ....................................................2. O. humifusa
2. Spines present on at least some areoles ................................................................................3
3(2) Spines (at least some of the larger) flattened at the base, narrowly elliptic in cross-section (roll between thumb and finger); if any spines yellow, all yellow when fresh (darkening with age or when pressed), usually 1 to 6 per areole in all but the lowest areoles of a pad.................................................................................3. O. engelmannii
3. Spines not flattened at the base, needlelike, circular in cross-section (occasionally 1 or 2 somewhat flattened); at least some spines not yellow when fresh, 1 to 6(11) per areole ....4
4(3) Seed margin (over embryo) ca. 0.5 mm broad, smooth and regular; joints dark or light green; perianth parts yellow; spines few, near the upper edge of the joint, gray or
brownish, usually 1 per areole, 1.9 to 3 cm long, spreading; all roots fibrous .........................
..........................................................................................................................2. O. humifusa
4. Seed margin (over embryo) ca. 1 mm broad, irregular and corky; joints blue-green, glaucous; perianth parts yellow to creamy yellow, the bases often with some red; spines few to many, usually on the upper part of the joint, pale gray (rarely reddish or brownish), 1 to 6 per areole, 3.8 to 5.6 cm long, mostly deflexed; main root tuberous ............................
......................................................................................................................4. O. macrorhiza
1. O. leptocaulis DC. Pencil Cactus, Desert Christmas Cactus, Tasajillo, Tesajo, Pencil Cholla. Erect shrub to ca. 1.5 m tall, well-branched, joints cylindrical. Largest joints 30 to 40 cm long, the terminal joints much smaller, 2.5 to 7.5 cm long, all less than ca. 1 cm in diameter, smooth or with a few indistinct tubercles. Spines (occasionally lacking), 1(3) per areole, straight, not markedly barbed, commonly somewhat deflexed, the long ones 2.5 to 5 cm long, 0.5 mm in diameter at the base, gray, with loose, deciduous, tan sheaths; glochids few and small. Blossoms 1 to 1.5(2) cm broad; perianth green to yellow or bronze, the petaloid parts ca. 6 to 9, rather broad. Fruit persistent, bright red, fleshy, juicy, obovoid, ca. 1 to 1.5 cm long, smooth and spineless but with glochids; seeds tan, irregularly discoid. Heavy or bottomland soils. W. and S. TX; AZ to NM, OK, and TX; also N. Mex. Flowering late spring to early summer; collected with fruit throughout the year. Commonly disarticulating into component joints and losing color in herbarium sheets.
The fruits are edible, but must be peeled and de-seeded.
2. O. humifusa (Raf.) Raf. var. humifusa Eastern Prickly Pear. Low, creeping, clump-forming plant 7.5 to 10 cm tall, from fibrous roots. Joints usually dark green to lead-colored, sometimes reddish-purple in winter, flattened, mostly orbicular or broadly ovate, 5 to 7.5(12.5) cm long and 4 to 6.2(7.5) cm broad; leaves conical, to 4.5 to 6 mm long. Areoles few, ca. 3 mm in diameter, ca. 1 to 2 cm apart. Spines occasionally entirely absent, when present found on the upper areoles of a joint, 1 per areole, spreading at right angles to the joint, gray or brownish, the longest usually 2 to 3 cm long, 0.5 to 1 mm in diameter, circular or nearly so in cross-section, not markedly barbed; glochids yellow or brown, 3 mm long. Flowers 4 to 6 cm long and broad; sepaloid perianth parts green with yellow edges, ovate-acute, 5 to 25 mm long, 4.5 to 20 mm broad, short-acuminate or acute, undulate; petaloid parts yellow, cuneate-obovate, 25 to 40 mm long, 12 to 20 mm broad, apex rounded, entire; style 9 to 12 mm long, ovary smooth and with few areoles. Fruit fleshy and reddish or purplish at maturity, with glochids, obovoid to clavate, 2 to 4 cm long, 2 to 3 cm in diameter, the apex somewhat depressed, persisting on the plant until early spring; seeds pale tan or gray, flat, 4.5 mm in diameter, 1.5 mm thick, the margin over the embryo ca. 0.5 mm broad, hard, smooth, regular. This variety at 1.5 to 600 m on sandy or rocky soils of hillsides, valleys shores, and in prairies and deciduous forests. MT to the Great Lakes to MA, S. to NM, TX, and FL, along the Atlantic and Gulf coasts MA to FL. Flowering spring to summer. [O. compressa (Salisb.) Macbr.--a much-used but invalid name].
3. O. engelmannii Salm-Dyck var. lindheimeri (Engelm.) Parfitt & Pinkava Texas Prickly Pear, Nopal Prickly Pear. Sprawling to erect shrubs, solitary or forming thickets, 1 to 3(3.5) m tall; trunk, if any, very short; roots fibrous. Largest terminal joints green or in the western portion of the range blue-green, flattened, obovate or sometimes orbicular, 15 to 25(30) cm long, 12.5 to 20(25) cm broad, to 2 cm thick; leaves narrowly conical, 3 to 9 mm long. Areoles usually 2.5 to 4 cm apart, elliptic, ca. 4.5 mm long and 3 mm broad; spines usually in all but the lower areoles (rarely a plant spineless), 1 to 6 per areole, yellow or whitish-yellow, the very base sometimes black or brownish, not red, 1.2 to 4(5) cm long, basally 0.7 mm broad, narrowly elliptic in cross-section, not barbed, usually 1 per areole spreading and the others sharply reflexed; glochids yellow or becoming brown with age, 3 to 6 mm long. Flowers 5 to 7.5(10) cm in diameter, 5 to 8 cm long; sepaloid parts green-yellow or green-red, obovate, mucronate to acuminate, somewhat crisped or ruffled, 6 to 35 mm long, 6 to 15(25) mm broad; petaloid parts yellow (rarely red), cuneate-obovate, mucronate, undulate, 3 to 4(5) cm long, 1.2 to 2.5(4) cm broad; style green-yellow, 12 to 20 mm long, ovary spineless at anthesis. Fruit fleshy and purple at maturity, with small areoles and glochids, obovate to elongate, 3 to 7 cm long, 2.5 to 3(4) cm in diameter, shallowly depressed at the apex, not persistent; seeds tan, asymmetrically elliptic, 3 to 4 mm long, 2.5 to 3 mm broad, 1.5 mm thick. This variety at 1.5 to 300 m on sandy, gravelly, or occasionally rich soils of the Great Plains, Ed. Plat., Rio Grande Plains, and (rarely) the Chihuahuan desert. Trans Pecos and Big Bend region, along the Rio Grande to the coast, common in S. and Cen. TX; NM to Cen. OK, S. through Mex. Flowering spring to summer. [O. lindheimeri Englem. var. lindheimeri].
4. O. macrorhiza Engelm. var. macrorhiza Plains Prickly Pear, Grassland Prickly Pear. Plants low, clump-forming, commonly 7.5 to 12. 5 cm tall; primary root usually large and tuberous, the others usually fibrous. Larger terminal joints moderately glaucous, flattened, orbicular to obovate, (6)7.5 to 10 cm long, 5 to 6(7.5) cm broad, ca. 12 mm thick; leaves long-conical, 7.5 mm long. Areoles usually 1 to 2 cm apart, ca. 3 mm in diameter; spines mainly from the uppermost areoles, often only on the margins of the pads, white to gray (sometimes brown or reddish-brown), 1 to 6 per areole, mostly deflexed, straight or occasionally twisted or slightly curved, scarcely if at all flattened basally, not barbed, slender, ca. 0.5 mm in diameter at the base, the longer ones 3.8 to 5.6 cm long; lochids yellow or brown, 3 mm long. Flowers 5 to 6.2 cm long and broad; sepaloid parts with a green midrib and yellow or sometimes reddish margins, ovate, acute to short-acuminate, 5 to 25 mm long, 5 to 20 mm broad; petaloid parts yellow or with a tinge of red at the base, cuneate-obovate, apex rounded and entire, 2.5 to 4 cm long, 1.2 to 2.5 cm broad; style green-yellow, 9 to 12 mm long, 4.5 mm broad, the base swollen, ovary smooth at anthesis, with few areoles. Fruit fleshy, red-purple, obovoid, with some glochids, 2.5 to 4 cm long, 2.5 to 3 cm broad, apically somewhat depressed, persistent on the plant for several months; seeds tan to pale gray, irregularly discoid, ca. 4.5 mm in diameter, 1.5 to 2.5 mm thick, the margin over the embryo corky, ca. 1 mm broad, rough, irregular. This variety usually from 600 to 1,200(2,400) m, scattered over TX except for the deep SE, the far NE, and the SW. Ed. Plat.; CA to SD, ID and MI, S. to MO, AR, LA, TX and Mex. Flowering spring to summer. [Synonym for the species is O. tortispina Engelm.].
Stems 1 to several, sometimes branching and forming low mounds or colonies of up to 200 or more stems. Plants subglobose to cylindroid, 2.5 to 10(12.5) cm tall, 2.5 to 7.5 cm in diameter, covered with spirally arranged tubercles, these all separate and commonly with a long groove on the upper side of each. Leaves not evident. Areoles circular to elliptic, apical on the tubercles. Spines smooth, white to gray, pink, yellow, black, or brown, the central spines 0 to 10 or more, straight to curved, hooked, or slightly twisted, needle-like or subulate, grading into the ca. 5 to 40 radial spines. Flowers and fruits on current season's growth near the apex of the stem, at the base of the upper side of a tubercle and not on the spiniferous areole, the tubercle groove or isthmus connecting the flower with the areole. Perianth 1.2 to 6 cm in diameter, greenish to light yellow or pinkish, the floral tube above the ovary funnelform. Fruit fleshy, thin-walled, red or green at maturity, naked. Seeds tan to brown or black, shining and smooth or else punctate and reticulate, usually broader than long, 1 to 2 mm in the greatest dimension.
Variously 20 to 30 to 64 species from Alberta to Mexico. There are 14 species in TX; 1 here.
Some or all members of the genus have been variously treated as Mammillaria, Neomammillaria, or Neobesseya (Weniger 1988). Our species can now be placed in Escobaria, a genus with 16 species listed for North America (Kartesz, 1998)
Several species, including ours, are cultivated as ornamentals and are reportedly easy to grow (AHS 1982).
1. C. missouriensis (Sweet) Britt. & Rose var. caespitosa (Engelm.) L. Benson = Escobaria missousriensis (Sweet) D. R. Hunt var. missouriensis Stems solitary or branched to form colonies 5 to 10 cm high and 15 to 30 cm in diameter; larger stems dark green, hemispheroid to depressed-globose, 2.5 to 5 cm long, 3.8 to 5(10) cm in diameter. Tubercles elongate, 6 to 9 mm long and broad, protruding 12 to 15 mm, usually with a groove on the upper side; areoles 1.5 to 2 mm broad, usually ca. 6 mm apart, white-woolly when young. Spines rather dense, somewhat obscuring the stem, pubescent, yellow when young but becoming dark gray, central spines usually absent (sometimes 1, 2, or 4, but not well differentiated), radial spines usually 12 to 15 per areole, straight and spreading, slender, the longer ones 1 to 2 cm long, to 0.3 mm in diameter at the base, needle-like. Flowers ca. 5 to 6.2 long and broad; sepaloid parts with a green midrib and yellow to whitish margins, the larger ones linear-lanceolate, 12 to 40 mm long, 12 mm broad, sharply acute, fimbriate; petaloid parts yellow or sometimes red or pink or yellow tinged with red, the largest linear-lanceolate, 12 to 40 mm long, 1.5 to 4.5 mm broad, acuminate-attenuate, entire; style green, 1.2 to 1.5 cm long, 1 mm broad at the widest. Fruit red, fleshy, globular to ellipsoid, 1.5 to 20 mm long and wide; seeds black and punctate, 2 to 2.5 mm long, profile shaped somewhat like an old-fashioned diving-helmet, the hilum basal and slightly oblique. This variety at low elevations in the Great Plains and grasslands: NE. KS to Bexar Co., TX and NW. LA. Flowering spring and summer. [Neobesseya missouriensis (Sweet) Britt. & Rose; Neomammillaria similis (Engelm.) Britt. & Rose; Mammillaria similis Engelm.].
The red fruits against the dark green, circular clumps are reminiscent of a Christmas wreath. Sometimes cultivated as a desert landscape or pot plant (AHS 1982).
Herbs, shrubs, or rarely small trees, ours herbaceous though sometimes woody at the base. Stems commonly more or less succulent or jointed or both. Leaves simple, opposite below and alternate above or alternate throughout, estipulate and without scarious bracts, entire to lobed, sometimes scale-like and/or succulent, glabrous to pubescent or farinose. Flowers usually small and green or greenish, subtended by 1 bract and 2 bracteoles (or the bract, bracteoles, or both absent), borne singly to many and glomerate in the axils of leaf-like bracts or arranged in spikes, panicles, or cymes, rarely (and not in ours) flowers in terminal strobili or sunken into the stem, regular, perfect or imperfect and the plants monoecious or dioecious. Sepals (1)5(6), free or occasionally briefly united below, imbricate in the bud, herbaceous or membranous but not scarious, occasionally absent in pistillate flowers. Corolla absent. Stamens usually as many as the sepals and opposite them, sometimes fewer, filaments free or sometimes connate basally or inserted on an annular disk or the base of the calyx, unisexual flowers usually with nectaries reduced or absent. Gynoecium superior (half-inferior in Beta), of 2 to 3(5) fused carpels, unilocular, styles 2(3 to 5), free to more or less connate, ovule 1. Fruit a 1-seeded thin-walled utricle, achene, or nutlet, indehiscent, dehiscent irregularly, or circumscissile, often enclosed by the persistent calyx or bracteoles, occasionally (as in Beta) several fruits borne together in their calyxes to form a multiple fruit. Seed usually lens-shaped with a curved or spirally twisted embryo, nutritive tissue usually perisperm. Betalain pigments rather than anthocyanins present.
About 100 genera and 1,300 to 1,500 species worldwide; 17 genera and 67 species in TX; 5 genera and 10 species here.
Many species are weedy and/or have allergenic pollen, including Lambsquarter (Chenopodium), and Tumbleweed or Russian Thistle (Salsola). Spinach (Spinacia) and beet (Beta) are grown for food. Some species of Chenopodium have edible fruits, e.g. quinoa, a pseudo-cereal of the Incas. Some genera such as Atriplex (Saltbush) are important members of desert floras (Mabberley 1987; Tull 1987).
1. Leaves spine-tipped, main ones filiform; in addition to the bract, flowers subtended by 2 leaf-like bracteoles equal to or longer than the perianth; embryo spirally coiled ..1. Salsola
1. Leaves not spine tipped, main ones broader; flowers ebracteolate or with minute
bracteoles shorter than the perianth; embryo curved to annular ...........................................2
2(1) Stamen solitary; sepal 1; leaves triangular to rhombic with 1 lobe on each side ....................
2. Stamens 3 to 5; sepals 3 to 5(8); leaves variously shaped ....................................................3
3(2) Flowers solitary or in small clusters in the axils of the bracts or leaves; main stem leaves linear or lance-linear ..................................................................................................3. Kochia
3. Flowers in axillary or terminal spikes, clusters, or panicles, these sometimes with bracts; main stem leaves not linear or lance-linear .............................................................................4
4(3) Persistent calyx with a wing encircling the mature fruit .....................................4. Cycloloma
4. Persistent calyx wingless in fruit ...................................................................5. Chenopodium
About 150 species, worldwide in distribution. The species found in TX can be expected to occur here.
1. S. tragus L. Russian Thistle, Tumbleweed. Taprooted annual herb, tumbleweed in habit, much branched from the base, 3 to 10 dm tall, becoming nearly hemispherical, without a main axis but with many equal lateral branches, often tinged or streaked with red, stout, ascending to spreading, hirsute, scabrous, or short villous to glabrous. Leaves sometimes blue-green, alternate, clasping or sessile, filiform to linear, subterete and succulent, 1.2 to 4.5(8) cm long, 0.4 to 1 mm broad, reduced upwards, margins entire to denticulate, scabrous or glabrous, apex strongly spinose, base broadened, thickened, and hyaline-margined, especially on the upper leaves. Inflorescence of terminal and axillary spikes 1 to 6(10) cm long, 10 to 18 mm broad, with 2 or 3 flowers per node but only the lowermost developing fully, floral bracts narrowly lanceolate or narrowly deltoid, 3 to 15 mm long, 1.5 to 2.5 mm broad, spreading or often recurved, strongly spinose, margin entire to denticulate-crenulate; the two bracteoles beneath each flower similar but smaller. Flowers perfect, sessile. Calyx 5-lobed, lobes oval to oblong-lanceolate below the middle protuberance and narrowly deltoid above, 1.5 to 1.7 mm long, 1.1 to 1.8 mm broad, acute to acuminate, persistent, at maturity (3)6 to 10 mm broad, becoming hard below and remaining membranous above, strongly transversely winged, wings thin, membranous, crenate-dentate, whitish or tinged with red or pink, conspicuously veined, those of the lowest flowers often only carinate and not winged; stamens 5 (or fewer), included at first and later exserted, anthers 0.5 to 1.2 mm long; stigmas 2, 1.0 to 1.4 mm long, erect with curling tips, style shorter. Fruit obconic or obovoid, apex concave to convex, membranaceous, closely enclosing the seed and nearly the same size; seed obconic, 1.5 to 2 mm long, 2 to 3 mm broad, black, lustrous, horizontal (rarely inverted, oblique, or erect); embryo spiral or cochleate, endosperm none. To be expected in vacant lots and cultivated fields and along roadsides and railroads. Native to N. Eurasia--S. Europe and Russia, now established worldwide; in N. Amer. from Que. to B.C., S. to CA, TX, and MO. July-Oct. [S. pestifer A. Nels; S. iberica Senn. & Pau; S. kali L. var. tenuifolia Tausch and var. ruthenica (Iljin) Soó in Soó & Jvorka; S. ruthenica Iljin]. Our plants for years have been treated as S. kali, which is apparently a species primarily of beaches and sandy coastal soils. S. australis R. Br. has been applied to S. tragus, but is conspecific with S. kali (Clements 1992). S. tragus is the earliest available name for our taxon.
Salsola has allergenic pollen. The stems of S. kali, and presumably our species also, were once burned and the ash used in the manufacture of glass and soap (Clements 1992). Young shoots are edible by humans and animals, though when mature, the plants can accumulate nitrates and oxalic acid and can be toxic to livestock that eat them in large quantities (Tull 1987).
About 3 to 6 species in parts of the temperate N. Hemisphere. We have the one species found in TX.
1. M. nuttalliana (Schult.) Greene Povertyweed, Nuttall Monolepis. Annual herb, well-branched from the base; stems prostrate to ascending, 1 to 3 dm tall, succulent, somewhat mealy when young, becoming glabrate with age, pale green. Leaves fleshy, alternate, subsessile or short petiolate below and sessile above, blades 1 to 6.5 cm long, reduced upwards, triangular to lanceolate or ovate, with 1 divergent lobe on each side near the base and sometimes a few teeth above, otherwise entire, apically more or less acute, basally cuneate. Flowers perfect or a few pistillate, often reddish, arranged in dense, sessile, axillary clusters. Sepal 1, persistent, bract-like, herbaceous to somewhat fleshy or coriaceous, oblanceolate or spatulate, entire, acute to acutish, 1.5 to 2.5 mm long; corolla absent; stamen 1 or absent in pistillate flowers; styles 2, filiform and short; ovary and fruit compressed-ovoid, pericarp adherent to seed, finely pitted at maturity. Seed vertical, flattened, dark brown to black, 0.8 to 1.4 mm wide, margin entire; embryo annular, endosperm mealy, radicle inferior. Dry or moist, often alkaline or saline, soils of roadsides, waste places, fields, etc. Throughout the state with the possible exception of deep E. TX. Man. to B.C., S. to CA, TX, MO, Son., and on old ballast dumps in ME; also Siberia, Patagonia. Mar.-Sept.
Annual or perennial herbs woody at the base, or small shrubs; plants often rounded or pyramidal. Stems erect, well-branched, pubescent to rarely glabrous. Leaves alternate (to opposite), linear to linear-lanceolate, entire, often terete, often fascicled. Flowers sessile, solitary or in small clusters in the axils of leaf-like bracts (bracteoles present but very minute), perfect or sometimes some only pistillate. Calyx 5-parted, herbaceous, persistent, in fruit developing horizontal membranous or scarious wings. Stamens 3 to 5, filaments compressed. Utricle depressed-globose, membranous, pericarp free from the horizontal seed. Embryo nearly annular; endosperm absent.
Variously treated as ca. 25 to 100 species, nearly all from the Old World; 2 listed for TX; 1 here. In some recent treatments (Mabberly 1987; Clements 1992). Kochia is included in Bassia, which has fruit with spines or other protuberances, but no wings.
Some including ours, are cultivated for ornament. The pollen is allergenic (Clements 1992).
1. K. scoparia (L.) Roth. ex Schrad. Summer-cypress, Belvedere-cypress, Kochia, Fireweed, Mock-cypress, Mexican Firebush. Annual herb, stems erect, 3 to 20(40) dm tall, well-branched from the base, branches spreading to erect; stems and branches greenish-yellow, green, or streaked with red (sometimes plants entirely red or red-purple in fall), glabrous to glabrescent, villous, or pilose with rusty or silvery hairs. Leaves alternate, 2 to 7(10) cm long, 0.5 to 8(12) mm broad, with 1 to 3(5) prominent veins, lower leaves linear to lanceolate, oblanceolate, to narrowly obovate, apically acute to obtuse or rounded tapered to a distinct petiole to 3 mm long, upper leaves linear, elliptic, narrowly lanceolate, or oblanceolate, apically acute to acuminate, tapered slightly to a sessile base; all leaves flat, entire, ciliate, glabrate (especially above) to sometimes villous or pilose with hairs to 6 mm long. Inflorescences from remote-flowered and long-spiciform to short, compact-cylindric or oblong-claviform, some plants floriferous along most of the length of the stems; flowers sessile, (1)2(3 to 5) in the axils of leaflike, reduced bracts 3 to 18 mm long and with only exceedingly minute bracteoles, perfect or some functionally pistillate or staminate, or perfect and pistillate on the same plant, each subtended and enveloped by tufts of short to long hairs. Calyx campanulate to urceolate, 0.3 to 0.6 mm long at anthesis, glabrous except for the 5 ciliate lobes, at maturity 1.5 to 3 mm broad, with sepals incurved over the fruit and with 5 dorsal horizontal lobes or wings, wings ranging from short and tubercle-like to semi-membranous, flat, and oblong-rotund to triangular and obtuse, these larger wings variously lobed, 0.6(2) mm long, cellular-reticulate, often striate, but not nerved; stamens 5, included or exserted; styles 2(3), free or commonly united for ca. 0.3 mm. Fruit a depressed-globose utricle with pericarp free from the horizontal seed; seed obovate, (1.5)2 to 3 mm long, with faces concave, dark brown to black, dull and smooth to granular. Wasteland weed naturalized from Eurasia and also sometimes escaped from cultivation, cosmopolitan; infrequently collected here. June-Aug. [Bassia scoparia (L.) A. J. Scott; K. alata Bates].
The variety culta Farw. or forma trichophila (A. Voss) Stapf ex Schinz & Thell, Mexican Firebush, is commonly cultivated. It has a dense globose form and bright purple-red fall color. It is possible, though, that this form is the plant described by Linnaeus and that wild plants (or at least those of the Great Plains of the U.S.) are referable to K. silversiana (Pall.) C. A. Mey. (GPFA 1986)
Livestock have been known to graze this plant, which has a protein digestibility on the same order as alfalfa, but may cause photosensitization in cattle. (GPFA 1986).
A monotypic genus native to North America.
1. C. atriplicifolium (Spreng.) Coult. Tumble Ringwing, Winged Pigweed, Plains Tumbleweed. Taprooted annual herb, freely branched and bushy, 1 to 8 dm tall and about as broad; stems erect to spreading, divaricately branched, striate, branches slender, with obtuse angles, loosely and finely woolly, becoming glabrate with age except around the flowers. Leaves alternate, sessile to short-petiolate, petioles 0 to 15 mm long; main stems leaves 6 to 7 cm long, 1 to 1.5 cm broad, usually absent at fruiting time, the remaining leaves smaller, from 2 cm long and 6 mm broad, all lanceolate to ovate, oblong, or narrowly oblong, coarsely and irregularly sinuate-dentate, teeth acute and mucronate, basally cuneate, apically acute, young leaves whitish tomentose, becoming glabrate. Flowers perfect or a few pistillate, in interrupted spikes arranged more or less in panicles, bracts narrowly oblong to narrowly elliptic, 0.3 to 1.0 cm long, toothed; bracteoles absent. Calyx lobes 5, triangular-ovate, 0.4 to 0.6 mm long, 0.4 to 0.8 mm broad, inflexed, keeled, calyx in age developing below its lobes a continuous, horizontal, membranaceous or white hyaline, irregularly lobed and toothed wing, calyx then 4 to 5 mm broad in overall diameter, covering the utricle, becoming purplish or reddish in age and more or less villous; stamens 5; ovary densely tomentulose; styles 2 or 3, partly united, erect to spreading. Utricle depressed-globose, pubescent, enclosed by the calyx; seed 1.3 to 1.7 mm in diameter, 0.8 to 1.0 mm thick, horizontal, black, smooth and with a few scattered, white, silky hairs or glabrous. Embryo annular, surrounding the mealy perisperm, radicle centrifugal. Plant with tumbleweed habit. Weedy in sandy places. Throughout the state except the coastal and S. TX plains; Man. to IN, S. to AZ and TX, W. to WY and NM; adventive in the E. U.S. and Europe. Fruiting in the summer.
Annual or perennial (sometimes biennial) herbs, rarely suffrutescent; herbage usually farinose (mealy-coated), sometimes glandular or glabrate, sometimes strongly-scented. Stems erect to spreading, solitary or branched above or below. Leaves simple, alternate, petiolate to sessile, blades variously linear to hastate or deltoid, entire to toothed or lobed. Inflorescences axillary or terminal spikes or glomerules (rarely, and not in ours, large heads of dichotomous cymes.) Bracts present or absent, blade-like to needle-like, usually beneath the glomerules; bracteoles absent. Flowers small, inconspicuous, usually greenish or mealy, perfect or rarely unisexual. Sepals (3 to 4)5, usually united at least basally, flat or keeled, persistent in fruit. Stamens 5 or fewer, filaments sometimes basally connate. Ovary superior, unilocular; style 1 or lacking, stigmas 2 to 5, filiform. Fruit an utricle with a fleshy or membranous pericarp separable or attached to the single horizontal or vertical, lenticular seed, seed coat smooth to rough or alveolate; embryo curved to annular, surrounding the mealy perisperm; radicle centrifugal or inferior.
About 70 to 150 species of temperate regions, many in xeric habitats; 25 species in TX; 6 here.
Many species are weedy. Some are grown as pot herbs or eaten raw like spinach. The plants have been eaten this way by such diverse peoples as Medieval Europeans and the Plains Indians of the U.S. C. ambrosioides, Mexican Tea or Wormseed, is used in medicinal preparations, usually as a vermifuge, and as a seasoning. Some species have edible fruits and were grown by Native Americans of both hemispheres. C. quinoa is an important pseudo-cereal native to the Andes. (Kindsher 1987; Mabberly 1987; Clements 1992).
In many cases, a complete specimen with mature fruit and lower stem leaves is necessary for confident identification.
1. Plants variously pubescent and/or glandular, neither farinose nor completely glabrous; usually with a turpentine-like smell; pericarp smooth .............................1. C. ambrosioides
1. Plants farinose (especially inflorescence) or glabrous, not glandular or pubescent; odorless or fishy-smelling; pericarp smooth to striate, pitted, or alveolate ............................2
2(1) Flowers in each glomerule in different stages so mature fruits and young flowers found together; leaves usually lanceolate to narrowly ovate, entire or the larger ones with a few basal teeth; pericarp smooth ...................................................................2. C. standleyanum
2. Flowers in each glomerule at about the same stage of development; leaves ovate to deltoid, rhombic, or trullate, usually dentate to variously lobed; pericarp smooth or
3(2) Fruit turbinate, with a hump or conical point on each face; uppermost leaves not much reduced or less toothed; inflorescence not extending much beyond leaves; pericarp smooth ...................................................................................................................3. C. murale
3. Fruit not turbinate; uppermost leaves reduced; inflorescence usually exceeding leaves; pericarp alveolate or smooth ....................................................................................................4
4(3) Pericarp distinctly alveolate or honeycombed; sepals with a winged keel about half the width of the sepal; plants often with a fishy odor ...........................................4. C. berlandieri
4. Pericarp usually smooth; sepals if keeled with a keel less than 1/4 the width of the sepal; plants without a fishy smell .......................................................................................................5
5(4) Leaves broadly rhombic, the largest 6 to 14 cm long and as wide, apex rounded to obtuse, not apiculate; tip of plant often with some purplish pigment ........................5. C. giganteum
5. Leaves ovate-rhombic or narrower, ca. 1.5 times longer than broad, the largest 2.5 to 8 cm long, acute to obtuse, apiculate; tip of plant usually without purple pigmentation .............
................................................................................................................................6. C. album
1. C. ambrosioides L. Mexican Tea, Wormseed, Spanish Tea, Epazote. Taprooted annual or short-lived perennial herb; stems erect or ascending, 3 to 10 dm tall, usually with several ascending branches; herbage with an unpleasant turpentine- or kerosene-like odor, covered with glandular resin dots, glabrous to puberulent or pubescent but never farinose. Petioles of lower leaves to ca. 18 mm long, blades ovate to oblong-lanceolate, 2 to 8(14) cm long, (0.5)1 to 5.5(6) cm broad, sinuate-dentate or -pinnatifid to serrate, teeth obtuse to acute, occasionally nearly entire, basally cuneate, apically obtuse to attenuate, usually markedly glandular-punctate (rarely eglandular), upper leaves becoming shorter, narrower, less-toothed, and eventually sessile. Flowers usually glomerate (occasionally solitary) in slender, dense or interrupted, leafy-bracted spikes; bracts lanceolate, oblanceolate, spatulate or linear, apically obtuse to attenuate, 0.3 to 2.5 cm long, 0.5 to 0.7 mm broad, occasionally absent. Flowers perfect; perianth (4)5-lobed, 0.7 to 1.0 mm long, 0.4 to 0.5 mm wide, the lobes ovate-rounded and obtuse, usually glandular, completely enclosing the fruit at maturity; stamens 4 to 5. Fruit ovoid, pericarp thin, deciduous, rugose to smooth, free of the seed; seed horizontal or vertical, ovoid, 0.6 to 1.0 mm broad, 0.4 to 0.5 mm thick, reddish brown to nearly black, testa rugose to smooth. Cultivated fields and waste places, also salt marshes and shores. Throughout TX. E. of the Trans-Pecos and Rolling Plains; Ont. and ME S. to FL, TX, CA, Berm., Mex., W.I, and S. and Cen. Amer.; naturalized in Eur., Asia, and Afr. Summer-fall. [Includes var. anthelminicum (L.) A. Gray; C. anthelminicum L.].
The leaves are used in Mexico as a seasoning for beans. An oil from the seeds is used in folk medicine as an anthelmintic but contains a toxic alkaloid. Overdoses have caused the deaths of infants; the leaves in quantity can also be toxic (Tull 1987).
2. C. standleyanum Aellen Standley Goosefoot. Taprooted annual herb; stem to maximum of 1 m tall (to 2.5 m in wooded areas), erect, slender, often branched, striate-angular, glabrous. Petioles (2)5 to 25 mm long; leaves of main axis larger than those of branches, blades lanceolate to ovate-lanceolate, ovate, or sometimes rhombic, (1)2 to 4.5(10) cm long, (0.3)0.5 to 1.5(5) cm broad, basally cuneate, apically acute to acuminate, entire or with a few teeth near the base, thin, greenish to sparsely farinose above, paler and moderately to sparsely farinose below. Flowers perfect, single to few in glomerules 0.5 to 2 mm broad arranged in terminal and/or axillary interrupted spikes, spikes sometimes arranged in panicles, flowers within each glomerule at different stages of development. Perianth 5-lobed, lobes obovoid, 0.5 to 0.7 mm broad, obscurely if at all keeled, apically rounded, sparsely to moderately white-farinose, only partially covering the fruit at maturity; stamens 5; style absent. Fruit horizontal, depressed-ovoid, 1.3 to 1.6 mm in diameter, pericarp fragile, smooth, membranaceous, readily separable from seed; seed ovoid, lenticular, 0.9 to 1.5 mm in diameter, 0.6 to 0.8 mm thick, black, lustrous, smooth to reticulate-alveolate or finely striate. Dry open woods, stream banks, waste areas, especially if disturbed, also on cliffs. E. TX and the Ed. Plat.; S. Que., NY, CT, and IN, S. to SD, VA, NM, AR, and E. TX. Summer-fall. [C. boscianum Moq., in part; C. album L. var. boscianum (Moq.) A. Gray; C. polyspermum L. var. spicatum A. Gray].
3. C. murale L. Sowbane, Nettle-leaf Goosefoot. Taprooted annual herb; stems 1 to 6(10) dm tall, erect to decumbent, somewhat stout or succulent, simple or branched near the base, lower branches decumbent; herbage somewhat ill-smelling, glabrous to sparsely farinose. Petioles 10 to 25 mm long, equalling or shorter than the blades; blades ovate to rhombic-ovate, 2 to 8 cm long, 1 to 5 cm wide, apically acute to obtuse, basally cuneate to subcordate or subtruncate, margins coarsely sinuate-dentate with sharp teeth, thin, glabrate to rather densely farinose (at least below), upper surface often lustrous, uppermost leaves not much reduced or less toothy. Flowers perfect, sessile in subglobose glomerules 2 to 4 mm broad or occasionally solitary, glomerules arranged in dense or lax terminal and axillary panicles (or cymes) 6 to 7 cm long, not or only slightly exceeding foliage; bracts and bracteoles absent. Perianth deeply 5-lobed, lobes oblong or ovate, 0.5 to 0.8 mm long, 0.6 to 0.7 mm broad, acute to obtuse, green, herbaceous, abaxially keeled, more or less farinose, partially enclosing the fruit at maturity; stamens 5; styles absent or very short. Fruit depressed-ovoid, green, pericarp membranaceous, pustulate but becoming smooth at maturity, adherent to the seed coat; seed horizontal, sharp-angled, turbinate, with hump or conical point on each face, 1.1 to 1.5 mm in diameter, 0.6 to 0.8 mm thick, black, minutely rugose or puncticulate. Sandy waste places, draws, cultivated areas, and yards. Native to Eur., Asia, and Afr., now widely adventive and established in N. Amer. from Que. and B.C. to Mex., Guat., and W.I. Mar.-Oct.
The young stems and leaves can be eaten as a salad herb (Clements 1992).
4. C. berlandieri Moq. Pigweed, Pitseed Goosefoot, Lamb's-quarters. Taprooted annual herb 2 to 10(15) dm tall; stems erect to nearly so, simple to much-branched, obtusely angled and striate, glabrate or farinose; herbage often with pronounced "fishy" odor. Petioles 7 to 70 mm long; blades narrowly to broadly rhombic, rhombic-ovate, or oblong, 1.2 to 4(15) cm long, 0.6 to 1.3(5) cm broad, apically acute or obtuse and mucronulate, basally cuneate to truncate or rounded, margins irregularly sinuate-dentate with acute or obtuse teeth (mostly near the base) to serrate, dentate, or occasionally entire in shade-grown plants, usually thick or even somewhat fleshy, rarely thin, commonly densely farinose when young, becoming glabrate in age, leaves reduced upwards, the uppermost ovate to linear-lanceolate, acuminate, and commonly entire. Flowers in irregularly-rounded glomerules 4 to 7 mm in diameter, the glomerules arranged in stout or slender, dense or interrupted, erect to drooping, terminal panicled spikes 5 to 15 cm long, usually exceeding the foliage; bracts and bracteoles absent. Perianth deeply 5-lobed, 2.1 to 2.5 mm broad, lobes ovate to deltoid, acute to obtuse, 1.0 to 1.5 mm long, 0.9 to 1.3 mm broad, moderately to densely farinose, more or less prominently keeled, enclosing and obscuring the fruit at maturity; stamens 5; styles united below, 0.1 to 0.2 mm long. Fruit depressed-ovoid, horizontal, distinctly alveolate-rugose, adherent to the seed coat; seed horizontal, (1)1.5 to 2 mm broad, 0.7 to 0.9 mm thick, ovoid, lenticular, brown to black, rugose to puncticulate or smooth. Dry soils, open areas. Throughout TX and N. Amer. [C. texanum Murr; C. palmeri Standl.].
Four varieties have been listed for TX (Hatch, et al. 1990). These varieties have been described based on size of the sepal keels, size of style base, and shape of the inflorescence. A more useful, higher-level distinction, however, may be between cultivated and free-living subspecies (Wilson and Heiser, 1979). In this case, our plants would belong to subsp. berlandieri.
Raw or cooked leaves and dried or cooked fruits are highly nutritious and have been extensively cultivated and consumed by Native Americans throughout the plant's range (Kindscher 1987). The leaves may be used in salads or as a leaf vegetable (Tull 1987).
5. C. giganteum D. Don. Taprooted annual herb; stems 1 to 3 m tall, erect, well-branched, branches erect or ascending from a spreading base, strongly angled, green, often red-striped, finely and sparsely farinose. Petioles to about as long as the blades; blades broadly rhombic, the largest 6 to 14 cm long and the lowermost about as broad, basally cuneate to subtruncate, apically obtuse to rounded, not apiculate, margins often shallowly to obscurely 3-lobed, usually irregularly and rather bluntly sinuate-dentate, rather thick, upper surface bright, deep green, under surface finely farinose when young, becoming glabrate with age, upper leaves smaller, rhombic-ovate to oblong or elliptic, dentate or sinuate-dentate, rarely entire, upper portion of plant often with some purplish or bright red pigment, at least when young. Flowers in small glomerules arranged in stout, dense panicled spikes. Perianth 5-lobed, usually reddish, farinose, lobes slightly keeled. A European plant sometimes escaping cultivation. Adventive in S. GA, N. FL., TX, Cuba, and Arg. Not much collected from our area.Treated by Kartesz (1998) under C. album L. var. abum
6. C. album L. Lamb's-quarters, Pigweed, Goosefoot. Taprooted annual herb; stems erect to sprawling, simple to well-branched, (1)6 to 25(30) dm tall, green or sometimes tinged with red at maturity, angular or sulcate, densely to sparsely farinose, branch ends sometimes pinkish. Petioles 1.0 to 2.5 cm long; blades rather thick and fleshy, trullate or ovate-lanceolate to ovate-rhombic or oblong, rarely ovate or lanceolate, 1 to 5.5(12) cm long, 0.5 to 3.8(8) cm broad, usually at least 1.5 times longer than broad, basally cuneate, apically acute or obtuse to rounded, apiculate, often with 3 shallow lobes, irregularly sinuate to dentate, shallowly serrate, or entire, upper surface pale green and glabrate, lower surface finely and usually densely farinose, upper leaves smaller, ovate to lanceolate, entire and not 3-lobed, acute and mucronate. Flowers perfect, grouped into subglobose glomerules 3 to 4 mm in diameter, these arranged in dense axillary or terminal, erect or ascending panicled spikes 2 to 19 cm long, occasionally single-flowered peduncles present; bracts and bracteoles absent. Perianth deeply 5-lobed, lobes ovate, 0.7 to 1 mm long, 0.7 to 1.1 mm broad, obtuse, green, white-margined, sharply keeled on the back, finely and densely farinose, nearly completely covering the pericarp at maturity; stamens 5; style absent. Fruit depressed-ovoid, pericarp smooth to papillate, non-alveolate, adherent to or free from the seed; seed horizontal, black, lustrous, lenticular, orbicular in outline, 0.9 to 1.6 mm broad, 0.5 to 0.7 mm thick, smooth, papillose, or with faint rugose-reticulate ridges or minutely pitted, margin obtuse. Open areas, especially if disturbed, yards, shores, cultivated areas, etc. Native to Eurasia, widely naturalized from Newf. to FL, W. to Yuk. and B.C., S. to Mex. and S. Amer., also N. Afr.--nearly cosmopolitan except in extreme desert conditions. Perhaps one of the most widely distributed angiosperms. Apr.-Sept. [C. viride L.].
Several species have at one time or another been treated as varieties of C. album. Most recent treatments list no more than var. album and var. missouriense (Aellen) Bassett & Crompton (Clements 1992; Hatch, et al. 1990). Our plants are referable to var. album with fruits 1.1 to 1.5 mm in diameter and lower leaves more than 1.5 times longer than wide. [C. album L. var. lanceolatum (Muhl.) Coss. & Germ.; C. lanceolatum Muhl.].
This plant has been cultivated since ancient times. The leaves are edible raw or cooked as a vegetable, but may accumulate oxalic acid which can bind calcium in the body. They should therefore not be consumed in large quantities. There are some reports of livestock poisonings. The seeds are edible cooked or dried and ground into flour. A high ratio of plant to fiber produces a bright, lightfast yellow on wool (Tull 1987).
Ours annual or perennial herbs, occasionally woody at the base. Stems erect to decumbent, trailing, or scandent. Leaves alternate or opposite, petiolate or sessile, simple, usually entire, estipulate. Plants monoecious, dioecious, or with perfect flowers, flowers entomophilous or anemophilous, small, regular, solitary or usually in cymose, glomerate, racemose, spicate, or paniculate inflorescences, each flower subtended by a bract and 2 bracteoles, these scarious and often spinose; bract persistent; bracteoles deciduous with the fruit or remaining. Sepals (0-1)2 to 5, scarious, membranous, or indurate at maturity, distinct or sometimes connate. Corolla absent. Stamens as many as the sepals and opposite them (rarely fewer), filaments free or usually united basally into a tube, this often with lobes or "staminodia" alternate with the filaments; andthers 1- to 4-celled; nectary ring often present with the base of the stamen tube. Gynoecium superior, of 2 to 3(4) fused carpels, unilocular, style terminal, usually 1, stigmas elongate or capitate, ovule usually solitary and basal (in a few genera ovules several on a basal or short free-central placenta.) Fruit in ours a dry, 1-seeded, circumscissile capsule (pyxis), achene, nutlet, or utricle, often enclosed by the persistent calyx. Seed lenticular, oblong, or reniform-orbicular, usually smooth and lustrous; embryo annular, surrounding the starchy perisperm. Betalain pigments present; anthocyanins absent.
About 65 genera and 900 species distributed nearly worldwide, commonest in tropical and subtropical areas; 11 genera and 57 species reported for TX; 5 genera and 16 species in our area.
Many species are weedy. Some, including species of Celosia, Gomphrena, and Amaranthus, are cultivated as ornamentals. Some species of Amaranthus are important pot herbs or pseudo-cereals. Amaranth flour is a common ingredient in organic or natural foods; the genus is receiving attention as a grain-substitute crop in the U.S. and other countries (Clements 1992; Mabberley 1987).
1. Leaves alternate; flowers imperfect ................................................................1. Amaranthus
1. Leaves opposite; flowers perfect or imperfect ........................................................................2
2(1) Flowers unisexual, arranged in panicles; plants rhizomatous perennials ...............2. Iresine
2. Flowers perfect, solitary or in spikes or clusters; plants annuals or perennials .....................3
3(2) Inflorescences usually terminal; erect herbs ......................................................3. Froelichia
3. Inflorescences usually axillary; prostrate herbs ......................................................................4
4(3) Staminodia present; sepals glabrous or with long, glochidiate hairs ..........4. Alternanthera
4. Staminodia absent; sepals densely woolly ................................................5. Gossypianthus
NOTE: Globe Amaranth (Gomphrena globosa L.) and Cockscomb (Celosia argentea L. or C. cristata L.) occasionally escape from cultivation but do not persist in our area.
Taprooted annual or perennial herbs. Stems herbaceous to suffrutescent, prostrate to erect, simple or usually well branched. Leaves alternate, simple, petiolate, entire to sinuate or crisped, estipulate but one species with spines present at the leaf bases. Flowers unisexual or a few appearing perfect, plants monoecious or dioecious. Inflorescences axillary or terminal clusters, spikes, or compound spikes, each flower subtended by 1 usually lanceolate bract and 2 bracteoles, bract and bracteoles often colored and/or obscuring calyx. Calyx of (1)2 to 5 tepals, subequal or the outer longer than the inner, sometimes rudimentary or absent in pistillate flowers, scarious or membranaceous, free to the base, midribs inconspicuous to broad, often excurrent or spinose. Stamens (1 to 3)5, free, anthers 4-celled but appearing 2-celled after dehiscence. Ovary 1-celled; style 1 or absent; stigmas 2 to 3(4), subulate; ovule 1, erect, basal. Fruit a circumscissile, irregularly dehiscent, or indehiscent utricle; seed 1, vertical, lenticular to subglobose, smooth and usually lustrous, embryo annular.
About 90 species, cosmopolitan, but most abundant in the New World; 24 listed for TX; 9 here. Includes Acnida L. and Acanthochiton Torr.
Several neotropic species are important grain crops. The seeds are high in well-balanced, high-lysine protein. Amaranths are thought to be among the first domesticated crops in Europe and the New World, the seeds having been found in archeological sites as much as 8,000 years old. Native Americans at the seeds raw, cooked into mush, or ground into flour. Many species are eaten as pot herbs or leaf vegetables. Several species are noxious weeds. A few, such as A. tricolor, have highly colored bracts or leaves and are grown as ornamentals (Mabberley 1987). Some species can accumulate nitrates in the foliage under certain circumstances and can lead to livestock poisoning if eaten in large quantities (Clements 1992). Dyes can be made from a few species (Tull 1987).
1. Plants with spines present at leaf bases; terminal spikes with staminate flowers at the apex and pistillate flowers at the base ......................................................................1. A. spinosus
1. Plants without spines; terminal spikes (if present) unisexual or with staminate and pistillate flowers intermixed .....................................................................................................................2
2(1) Plants dioecious ........................................................................................................................3
2. Plants monoecious ...................................................................................................................4
3(2) Sepals of pistillate flowers 1 to 2(3), all but 1 rudimentary; bracts of staminate flowers 1 to 2.5 mm long, shorter than to equalling the sepals; terminal spikes usually lax ....2. A. rudis
3. Sepals of pistillate flowers 5, the outer longer than the inner; bracts of staminate flowers 4 to 6 mm long, ca. twice as long as the sepals; terminal spikes usually dense .3. A. palmeri
4(2) Flowers in axillary glomerules only ...........................................................................................5
4. Flowers in terminal and/or axillary spikes, often paniculate, sometimes axillary glomerules present as well ..........................................................................................................................7
5(4) Sepals of pistillate flowers spatulate, narrowed into a claw basally, 3-nerved ........................
...................................................................................................................4. A. polygonoides
5. Sepals of pistillate flowers oblong to obovate or lanceolate, without a claw; with 0 or 1 conspicuous nerves ..................................................................................................................6
6(5) Bracteoles of pistillate flowers 2 or more times as long as the sepals; sepals of pistillate flowers 3; seed 0.7 to 1 mm broad; plants usually erect, pale-stemmed, with a tumble- weed habit ...............................................................................................................5. A. albus
6. Bracteoles of pistillate flowers about as long as the sepals; sepals of pistillate flowers 4 or 5; seed 1.4 to 1.7 mm broad; plants usually prostrate, not noticeably pale-stemmed, not usually of a tumbleweed habit ..........................................................................6. A. blitoides
7(4) Sepals of pistillate flowers 3; utricle indehiscent, quite rugose when dry .............7. A. viridis
7. Sepals of pistillate flowers 5; utricle circumscissile, smooth or rugulose on the upper half .............................................................................................................................................8
8(7) Sepals truncate to rounded or obtuse, usually emarginate and often mucronate, longer than the utricle; spikes usually short, thick, and spreading; utricle smooth .............................
........................................................................................................................8. A. retroflexus
8. Sepals acute to acuminate, shorter than to slightly longer than the utricle; spikes long and/or slender, usually ascending; utricle somewhat rugulose on the upper half ..................
...........................................................................................................................9. A. hybridus
1. A. spinosus L. Spiny Amaranth, Thorny Amaranth, Quelite Espinoso. Weedy annual herb from a long, sometimes branched taproot; stems erect or ascending, stout or somewhat succulent, branched, 1.5 to 12 dm tall, sulcate, often reddish, glabrous below and sparsely pubescent or puberulent above, most nodes with a pair of divergent spines 2 to 11(25) mm long. Petioles slender, 0.5 to 5(9) cm long, equalling or longer than the blades; blades ovate to ovate-lanceolate or rhombic-ovate, (1.5)3 to 10(12) cm long, to ca. 3.5 cm broad, apically obtuse and mucronate, sometimes emarginate, basally broadly cuneate, glabrous to sparsely pubescent, essentially entire. Plants monoecious with occasional perfect flowers, inflorescences axillary and terminal leafless spikes, often drooping, 3 to 15 cm long, each spike 5 to 10 mm broad, glomerules of flowers unisexual, staminate glomerules 0.7 to 1.1 cm broad, making up nearly the entire terminal spike and the distal portion of the others; bracts ovate-lanceolate, 0.8 to 2.5 mm long, 0.5 to 0.8 mm broad, spinose-tipped; bracteoles ovate, 0.7 to 0.9 mm long, 0.4 to 0.5 mm broad; staminate flowers: sepals 5, lance-oblong, acute to acuminate, the outer subulate-tipped, 1 to 1.5(2.1) mm long, 0.6 to 0.7 mm broad; stamens 5, 1.5 to 2.1 mm long. Pistillate glomerules 0.7 to 10 mm broad, globose, making up the basal portion of each spike and also some in the axils; bracts ovate to lanceolate or subulate, 0.5 to 3 mm long, 0.2 to 1.1 mm broad, tapered to a subulate green tip; pistillate flowers: sepals 5, oblong to acutish, 1.0 to 1.7 mm long, 0.6 to 0.7 mm broad, with 1 excurrent nerve; stigmas 3. Utricle ovoid to ellipsoid, 1.5 to 2 mm long, 0.7 to 1.0 mm broad, smooth below, spongy and roughened above, indehiscent or bursting irregularly (described by Clements (1992) as circumscissile); seed suborbicular, lenticular, 0.7 to 1 mm broad, 0.5 to 0.6 mm thick, black to dark reddish brown, smooth and lustrous. Waste ground, overgrazed pastures, etc., especially on sandy loam soils. E. 1/3 of TX, W. to Dallas, Travis, and Cameron Cos.; probably of tropical origin, now widespread in warm regions of the world, in N. Amer. N. to MO, IN, PA, NY, ME, and Man., often adventive even further north. June to Sept.(Nov.)
The shoots are edible when young and the spines still soft. The plant is featured in numerous herbal remedies in Latin America (Clements 1992).
2. A. rudis Sauer Water Hemp. Taprooted annual herb; stem usually stout, erect or ascending, 5 to 20 dm tall, simple or usually with ascending branches, striate-angular to smooth, glabrous or nearly so, green or somewhat glaucescent. Petioles slender, 0.6 to 5 cm long; blades oblong to lance-oblong, rhombic-oblong, ovate-oblong, ovate-lanceolate or lanceolate, 1 to 10 cm long, 0.4 to 3.5 cm broad, apex rounded to obtuse, sometimes notched, base acute to attenuate, glabrous, upper leaves much reduced, narrowly oblong. Plants dioecious. Inflorescences terminal and axillary, spicate to paniculate-spicate, usually leafy, axillary clusters also present. Male inflorescences 5 to 20 cm long, 8 to 10 mm broad, glomerules sessile, 3 to 5 mm broad, bracts ca. 1 to 2 mm long, 0.2 to 0.6 mm broad, midrib moderately heavy, excurrent; male flowers: sepals 5, oblong ovate, outer sepals 2.6 to 3.1 mm long, 0.6 to 0.8 mm broad, acuminate with conspicuous excurrent midvein, inner sepals 2.2 to 2.5 mm long, 0.6 to 0.7 mm broad, obtuse or emarginate; stamens 5, filaments 1.2 to 2.5 mm long. Female inflorescences 3 to 40 cm long, 6 to 15 mm broad; bracts 1.8 to 2.2 mm long, 0.3 to 0.5 mm broad, linear to lanceolate, attenuate, with a stout, excurrent midrib; female flowers: sepals 1 or 2(3), the shorter one rudimentary, 0.3 to 1.0 mm long and 0.3 mm broad, longer sepals 1.3 to 2 mm long, 0.3 to 0.4 mm broad, narrowly lanceolate, acuminate, midrib sometimes branched, excurrent; stigmas 3 to 4. Utricle globose to ovoid, 1.2 to 1.6 mm long, 0.9 to 1.1 mm broad, thin, smooth to rugose, tuberculate, or with faint ridges of tubercles corresponding to the style branches, circumscissile at the middle or slightly above, often reddish; seed round, lenticular, 0.9 to 1.1 mm broad, 0.4 to 0.5 mm thick, reddish brown to black, smooth and lustrous. Sandy waste areas, fields, usually in moist soils of swamps, alluvial areas, etc. Throughout TX except for the far W.; WI to ND and CO, S. to NM, TX, and LA; occasionally introduced or adventive elsewhere, e.g. WV, CA, NY; also naturalized in Europe. Mar.-Oct. [A. tamariscinus Nutt.; Acnida tamariscina (Nutt.) Wood].
The pollen of this plant is allergenic.
NOTE: A. australis (Gray) Sauer [Acnida cuspidata Spreng.] is a similar species found from the coast inland to Cen. and NW. TX. Not known from our area, but it may eventually be found here. It can be distinguished from A. rudis by having pistillate tepals rudimentary (less than 1 mm long) or absent, and without prominent midveins; staminate tepals lack heavy midveins and are all nearly the same size.
3. A. palmeri S. Wats. Palmer's Amaranth. Weedy annual herb from a long, stout taproot to 15 mm in diameter; stems erect, 6 to 10(30) dm tall, branched at the base and with ascending branches above, glabrous or occasionally villous-pubescent, striate-angled. Petioles slender, equalling to longer than the blades; blades rhombic-ovate to rhombic-lanceolate, 1 to 6(10) cm long, basally cuneate to rounded, apically acute to abruptly acuminate, entire to slightly undulate. Plants dioecious; flowers in dense or interrupted, erect or drooping, simple or racemose spikes 1.5 to 7(12) dm long, each spike to 1.2 cm broad, terminal on leafy branches or, if on leafless branches, loosely arranged and each subtended by a leaf, lateral spikes shorter, some axillary clusters also usually present, large mature inflorescences stiff and somewhat spiny; bracts 4 to 6 mm long, about twice as long as the sepals, midrib excurrent into a spine, those of staminate flowers usually erect and the midrib moderately heavy, those of pistillate flowers somewhat recurved, the midrib very heavy. Staminate flowers: sepals 5, the outer 3.5 to 4 mm long, acuminate, the midvein long-excurrent, inner sepals 2.5 to 3 mm long, obtuse or emarginate; stamens 5. Pistillate flowers: sepals 5, recurved, each with a branched midvein, outer 3 to 4.5 mm long, acute, the midvein excurrent into a spine, inner sepals 2 to 3 mm long, spatulate, emarginate, and often denticulate; stigmas 2(3). Utricle subglobose, 1.5 to 2.2 mm long, thin, circumscissile near the middle, somewhat rugose; seed obovate in outline, lenticular, 1 to 1.3 mm in diameter, dark reddish brown to black, smooth and lustrous. Sandy, silty, or gravelly soils of banks, arroyos, drainages, ditches, dumps, gardens, roadsides, etc. Nearly throughout TX; NE, KS, and CO S. to TX, NM, and Mex., W. to CA; introduced or adventive to MO, AR, LA, PA, WV, SC, FL, NY, etc. Summer-fall, as late as Dec. in S. TX. [Treated under A. torreyi S. Wats. in some works].
This plant hybridizes with A. rudis and A. retroflexus, among others (GPFA 1986).
4. A. polygonoides L. Tropical Amaranth. Taprooted annual herb; stems erect to ascending, 1.5 to 3 dm tall, branched from the base, branches slender, sulcate-angular, pubescent to glabrate, foliage deep green. Leaves crowded at the ends of the branches, with relatively short petioles usually shorter than the blades; blades oblong to oblong-lanceolate or rhombic-lanceolate, 15 to 30 mm long, basally acute to acuminate, apically obtuse to notched, entire, glabrous. Plants monoecious, flowers in crowded axillary glomerules, small leaves in the inflorescence sometimes spinose-tipped; bracts about half as long as the sepals, lanceolate, acuminate, midvein excurrent. Sepals delicate, more or less papery, ca. 2 mm long, spatulate, narrowed into a claw at the base, with a prominent midnerve and 2 conspicuous lateral nerves, often apically recurved, rounded to obtuse or occasionally cuspidate but not spinose. In male flowers, stamens 2. Utricle turbinate, thin walled, included in the calyx, 1.3 to 2 mm long excluding the 3 styles, indehiscent; seed ca. 1 mm long, obovoid, black, lustrous. In the Rio Grande Valley and Plains and Ed. Plat.; also NE. Mex. Collected several times on the Texas A&M campus; perhaps not a persistent member of our flora. Our collections from Oct. and Nov. [Includes subsp. berlandieri (Moq.) Thellung; A. berlandieri (Moqu.) Uline & Bray].
5. A. albus L. Tumbleweed, Tumbleweed Amaranth. Taprooted annual herb with a tumbleweed habit; stems usually erect, 2 to 12 dm tall, striate, whitish or noticeably pale, bushy branched, branches divaricate to ascending, glabrous to sparsely puberulent or villous, especially distally. Petioles slender, 3 to 35(50) mm long; blades elliptic to oblanceolate, obovate, rhombic-ovate, spatulate, or even lanceolate, 0.5 to 7 cm long, 0.4 to 2.5(3) cm broad, basally cuneate, apically obtuse to rounded, often retuse, emarginate, mucronate, or cuspidate, glabrous, the veins conspicuous, whitish on the lower surface, larger leaves sometimes purplish below and often deciduous before flowering time. Plants monoecious with occasional perfect flowers, flowers in axillary glomerules usually shorter but sometimes longer than the subtending petioles, plant often floriferous nearly to the base, male flowers fewer than female; bracts lanceolate or oblong-lanceolate, 2 to 4 mm long, green, rigid, spreading, acuminate, sharp-pointed. Staminate flowers: sepals 3, oblong, 1.2 to 2.3 mm long, cuspidate, scarious; stamens 3, filaments 0.5 to 1 mm long. Pistillate flowers: sepals 3, oblong to linear, 0.8 to 1.2 mm long, 0.3 to 0.5 mm broad, acute to acutish, 1-nerved, nerve not excurrent, green along the midnerve and often tinged with red; stigmas 3. Utricle obovoid to subglobose, exceeding the perianth, 0.9 to 1.5 mm long, 0.9 to 1.1(1.3) mm broad, circumscissile, sometimes with transverse ridges near the septum, rugose, stramineous or dark grayish brown, sometimes tinged with red; seed circular in outline, lenticular, 0.6 to 1(1.2) mm in diameter, 0.5 to 0.6 mm thick, dark reddish brown or black, lustrous. Waste places, cultivated areas, railroad tracks, sometimes near water or in ditches and then quite tall. Throughout TX but not abundant; widespread in N. Amer., Can. to Mex.; adventive in Eur., Asia, Afr., and S. Amer. July-Dec.
This plant has been treated under A. graecizans L. in many American works, but A. graecizans is an exclusively European species (Clements 1992).
This tumbleweed is a nuisance as the dry, windblown plants can accumulate in large piles.
6. A. blitoides S. Wats. Prostrate Pigweed, Quelite Manchado. Weedy taprooted annual herb usually without a tumbleweed habit; stems stout, prostrate or rarely ascending, freely branched at ground level or somewhat above, 1.1 to 6 dm long, commonly forming mats, smooth to striate, whitish or pale green, rarely tinged with red or purple, glabrous to sparsely pubescent. Leaves many, usually crowded; petioles stout, 0.2 to 2 cm long, glabrous; blades obovate to spatulate or elliptic, 8 to 40 mm long, 0.2 to 1.4 cm broad, basally cuneate to attenuate, apically rounded to acutish or very minutely mucronate, glabrous, prominently veined, veins white below, smaller leaves narrowly white-margined. Plants monoecious with occasional perfect flowers, flowers in axillary glomerules usually shorter than the subtending petioles, branches often floriferous to near the base; bracts erect, oblong to lanceolate, acute to acuminate, midrib excurrent into a short spinose tip, 1 to 3 mm long, 0.4 to 0.9 mm broad. Staminate flowers: sepals (4)5, oblong, acute, scarious, 1.3 to 2(3) mm long, 0.4 to 0.6 mm broad; stamens 3, filaments 0.7 to 1.7 mm long. Pistillate flowers: sepals (4)5, ovate to oblong or narrowly oblong, acuminate, 1.2 to 2.7(3) mm long, 0.5 to 0.7 mm broad, 1-nerved, green along the midnerve and with white margins; stigmas 3. Utricle obovoid to subglobose, 1.5 to 2.1 mm long, 1.4 to 1.8(2) mm broad, usually smooth, stramineous or sometimes tinged with red, circumscissile; seed circular in outline, lenticular, 1.3 to 1.6(1.8) mm in diameter, 0.9 to 1.0 mm thick, smooth, black, dull to lustrous. Waste grounds, roadsides, etc., on various soils. Throughout much of TX, but more abundant in the W. 1/2; W 1/2 of N. Amer., WA to Mex., E. to KS and TX; now also adventive in E. N. Amer. from S. Can. S.; also W.I. and Eurasia. July-Oct.
This species has been treated in some American works under the name A. graecizans L. True A. graecizans does not occur in N. Amer. (Clements 1992). Hatch, et al. (1990) placed A. blitoides under A. albus, but the two species are certainly distinct, having if nothing else different habits and a different number of sepals.
7. A. viridis L. Green Amaranth. Taprooted annual herb; stems erect or sometimes prostrate, slender, to 1 m tall, branched, often near the base, sulcate, glabrous. Petioles 1 to 4 cm long; blades deep green, broadly ovate or rhombic-ovate, 3 to 7 cm long, to ca. 5 cm broad, base acute to obtuse or rounded, apex acute or shallowly to deeply emarginate. Plants monoecious with occasional perfect flowers, flowers in spikes to 8 mm broad, the lateral not much shorter than the terminal, forming overall a panicle 10 to 20 cm long; bracts ovate to lanceolate, acute, equalling or shorter than the sepals. Staminate flowers: sepals 5, oblong, acute; stamens 3. Pistillate flowers: sepals 3, oblanceolate, acute, shorter than the utricle; stigmas 3, short. Utricle obovoid, compressed, ca. 1.5 mm broad, conspicuously rugose when dry, indehiscent; seed orbicular, lenticular, sharp-edged, ca. 1 mm in diameter, dark brown or reddish brown, lustrous, minutely puncticulate. Weedy in waste places, beaches, etc. S. TX and the Rio Grande Plains, well-known from our area; native of tropical S. Amer. and W.I., occasionally adventive in the E. U.S. and locally in the Gulf States; widespread in tropical regions, also Eur., Asia, Afr., and Pacific Islands. Mar.-June, in fruit as late as Nov. in our area. [A. gracilis Desf.].
NOTE: One recent checklist (Hatch, et al. 1990) lists our plants under A. blitum L. A more recent work on the Amaranthaceae (Clements 1992) lists both A. viridis and A. blitum and provides a detailed description and illustration of the latter. It is clear from that study that our plants do not belong to A. blitum, which is similar to A. viridis but has much shorter spikes and utricles which are smooth.
8. A. retroflexus L. Redroot Pigweed, Rough Pigweed, Quelite. Weedy annual from an often reddish taproot; stems erect or ascending, stout, 3 to 30 dm tall, simple or usually well branched, green or whitish, obtusely angled, usually villous-puberulent, at least above. Petioles slender, 1.5 to 8 cm long, equalling or somewhat shorter than the blades, usually villous; blades lanceolate or lance-ovate to obovate-oblanceolate, 2 to 8(12) cm long, 1.5 to 7 cm broad, basally acute to obtuse, apically acute to obtuse or often emarginate, more or less glabrous above and villous to puberulent beneath, especially along the whitish veins, margin entire or sometimes slightly crisped. Plants monoecious with occasional perfect flowers, inflorescences dense, usually crowded, terminal compound spikes and axillary spikes 5 to 20 cm long, each spike 10 to 20 mm broad, often dense glomerules present in the axils of the upper leaves as well, glomerules all bisexual, congested. Staminate flowers: bracts narrowly lanceolate, longer than the sepals, 3.7 to 7 mm long, 0.7 to 1.0 mm broad, acuminate, 1-nerved, somewhat villous; sepals 5, ovate-oblong to lanceolate, 2 to 3 mm long, acute or acutish, scarious, the midnerve excurrent; stamens 5, filaments 0.9 to 1.5 mm long. Pistillate flowers: bracts lanceolate to ovate, longer than the sepals, 3.7 to 7.0 mm long, 0.7 to 1 mm broad, tapered to a subulate tip, 1-nerved, somewhat villous; sepals 5, linear-oblong, 2.2 to 3.2 mm long, 0.7 to 1.0 mm broad, apically rounded or truncate, usually emarginate and/or mucronate, scarious, base sometimes thickened in age, whitish except for the green midnerve; stigmas 3. Utricle subglobose, shorter than the sepals, 1.2 to 1.8 mm long, 1.0 to 1.3 mm broad, circumscissile, more or less rugose on the upper half, stramineous; seed orbicular to obovate or oval, lenticular, 0.9 to 1.2 mm in diameter, 0.5 to 0.6 mm thick, smooth, lustrous, black to dark reddish brown. Sandy clay soils of fields, fallow land, railroads, stream banks, etc. Plains Country to S. Cen. TX and the Trans Pecos; temperate cosmopolitan weed, a pioneer species of Cen. and E. U.S., SE. Can., and NE. Mex.; spread to the Near East, N. Afr., and Asia. Jun.-Oct. [Includes var. salicifolius I. M. Johnst.].
Hybridizes with at least A. rudis, A. palmeri, A. hybridus, and A. powellii (GPFA 1986).
This plant is a noxious weed in many states and may also be an alternate host for some plant diseases. It can significantly reduce crop yields through competition. This is one of the species which can accumulate nitrates in the foliage in levels toxic to livestock, though under certain circumstances and in some areas, the foliate can be palatable and nutritious (Clements 1992). Native Americans ate both the seeds and the young plants (Kindsher 1987).
9. A. hybridus L. Slender Pigweed, Green Amaranth, Slim Amaranth,. Weedy annual herb from a stout taproot; stems erect or ascending, rather stout, 3 to 15(26) dm tall, usually branched, pale green, often tinged with red, striate, glabrous or rough pubescent below, often somewhat villous above. Petioles shorter than to longer than the blades, slender, 1.5 to 9 cm long, pubescent; blades rhombic-ovate to ovate or lanceolate, to ca. 15 cm long and 7 cm broad, basally cuneate to rounded, apically acute to rounded, darker green above and paler below, prominently veined, glabrous or pubescent below. Plants monoecious with occasional perfect flowers, inflorescences terminal compound spikes and axillary spikes, each 10 to 25 cm long, ca. 1 cm broad, some axillary clusters also present, all glomerules bisexual, congested. Staminate flowers: bracts lanceolate to ovate, 2 to 2.7 mm long, 0.4 to 0.6 mm broad, spinulose-tipped; sepals usually 5, narrowly oblong to ovate, acute, 1.7 to 2 mm long, 0.2 to 0.3 mm broad, the midvein usually excurrent; stamens 5. Pistillate flowers: bracts lanceolate to ovate, 2.2 to 3.4(4.2) mm long, 0.6 to 0.7 mm broad, tapered to a short spinose tip; sepals 5, oblong or linear-oblong, lanceolate, or even ovate, acute, 1.2 to 2.1 mm long, 0.4 to 0.5 mm broad, scarious, midrib usually excurrent; stigmas 3. Utricle obovoid or subglobose, 1.2 to 2.1 mm long, longer than or equalling the sepals, 0.9 to 1.3 mm broad, circumscissile, usually smooth, stramineous; seed circular in outline, lenticular, 0.8 to 1.1(1.5) mm in diameter, black or dark reddish brown, lustrous. Waste areas, railroads, river flats, cultivated areas, usually a pioneer of riverbanks, etc. In TX primarily in the W. 1/2, a naturalized weed, said to be originally native to tropical America, now naturalized from the milder areas of Can. S. through Mex. and Cen. Amer. to N. S. America; also the Medit., E. Asia, Austr., S. Amer. May-Oct. [A. chlorostachys Willd.; A. incuvatus Ti. ex Gren. & Godr.; A. patulus Berol.].
This plant hybridizes with A. retroflexus, A. powellii, A. rudis, A. hypochondriacus, and A. cruentus. It has been suggested that much of the variation present in this species is due to introgression with A. retroflexus (Clements 1992). The illegitimate name A. tamariscinus formerly applied to A. rudis is based on a hybrid between the two species.
Annual or perennial herbs and semi-shrubs, stems erect to decumbent or scandent, glabrous or pubescent. Leaves opposite, petiolate, entire or serrulate. Plants perfect or usually unisexual and the plants monoecious or dioecious, blossoms subtended by 1 bract and 2 bractlets, white, scarious, small, arranged in clusters or spikes aggregated into branched panicles. Calyx deeply 5-parted, often with long wool. Stamens mostly 5, united below. Ovary compressed, style short, stigmas 2 or 3, commonly filiform or sometimes capitate; ovule 1, pendulous. Utricle, small, subglobose, compressed, indehiscent, membranaceous. Seed inverted, smooth; embryo annular, endosperm mealy, radicle superior.
About 80 species in tropical and temperate regions of the world, especially America and Australia; 5 in TX; 1 here.
Some species are cultivated for their ornamental foliage; others are used in herbal medicines in various parts of the world (Mabberly 1987).
1. I. rhizomatosa Standl. Bloodleaf. Perennial herb, stoloniferous and with slender rhizomes; stems erect, usually simple below the inflorescence, 5 to 15 dm tall, glabrous to sparsely pubescent, sometimes short pilose at the slightly swollen nodes. Leaves opposite, thin, petioles slender; blades ovate to ovate-lanceolate, acute to long-acuminate, narrowed at the base and decurrent into the petiole, the larger ones 6 to 15 cm long and 2 to 7 cm broad, glabrous or sparsely pubescent. Plants dioecious, panicles terminal and from the upper axils, lax or somewhat pyramidal, 0.7 to 3 dm long. Staminate panicles usually lax and somewhat plumy, flowers in short spikelets, bracts and bractlets ovate, silvery-white, shorter than the sepals; sepals ovate-lanceolate, 1.2 to 1.5 mm long, silvery-white, obscurely 1-nerved; stamens 5. Pistillate panicles pyramidal, usually with erect or ascending branches; spikelets dense, 5 to 20 mm long, bracts and bractlets ovate, silvery-white, shorter than the sepals; sepals lance-ovate, 1.2 to 1.5 mm long, silvery-white, 1-nerved, subtended by wavy hairs 3 to 5 mm long. Utricle round, compressed, 2 to 2.5 mm long, equalling or longer than the sepals; seed suborbicular, 0.5 mm in diameter, dark red to reddish-black, smooth, lustrous. Sandy alluvial soils of creeks, floodplains, ditches, etc. E. 1/2 of TX W. to Comal and Denton Cos.; MD to S. IL and KS, S. to E. VA, GA, FL, AL, LA, and TX. Aug.-Oct. [I. celosioides Michx., non L.].
Annual, biennial, or perennial herbs from sometimes semiwoody taproots. Stems erect to ascending or procumbent, simple or branched, woolly to sericeus or pubescent, sometimes viscid above. Leaves opposite, sessile to petiolate, entire, variously tomentose, sericeus, or pubescent, rarely glabrate. Flowers perfect, sessile, in woolly spikes on long peduncles, each subtended by 1 scarious bract and 2 scarious bractlets. Calyx synsepalous, tubular, very densely woolly, with 5 glabrate lobes, becoming flask-shaped or conic at maturity, somewhat flattened, each edge with a longitudinal entire to dentate crest or row of teeth or spines, the faces with tubercles or spines at maturity. Corolla none. Stamens 5, filaments fused into a tube equalling the calyx tube, with 5 unilocular anthers and 5 sterile ligulate appendages. Ovary ovate, style elongate, stigma capitate, ovule 1, pendulous. Fruit an ovoid, indehiscent, membranaceous utricle included within the stamen tube and persistent, indurate calyx. Seed inverted, lenticular or obovoid, smooth; embryo annular, endosperm mealy, radicle superior.
About 12 species of the warmer parts of the Americas; 6 listed for TX: 3 here.
Determination to species requires microscopic examination of several mature calyces after the removal of the surrounding wool. Even then, differences between species are often very minor or arbitrary and putative hybrids are common. Published illustrations of calyces for our species differ widely from source to source. The genus is in need of further study. It is quite possible that such a study would reduce the number of recognized species.
1. Calyx tube at maturity with lateral rows of stout, rather sharp spines, each free to the base; stems slender, usually less than 1 to 2 mm thick, commonly well branched at the base .......
..............................................................................................................................1. F. gracilis
1. Calyx tube at maturity with lateral crests, these erose to deeply dentate, if dentate at least some teeth coalescent; stems stouter, to 7 mm thick, usually sparing branched .................2
2(1) Calyx crests deeply dentate ..............................................................................2. F. floridana
2. Calyx crests erose ........................................................................................3. F. drummondii
1. F. gracilis (Hook.) Moq. Slender Snake-cotton. Annual with slender stems 1 to 2 mm thick, simple or usually well branched at the base, (1)2 to 6(7) dm tall, branches ascending or procumbent, densely or sparsely villous-tomentose, sometimes viscid above. Leaves sessile or short petioled, often clustered near the base of the plant, linear-lanceolate, elliptic-lanceolate, or narrowly oblanceolate, 3 to 7(12) cm long, 2 to 7(12) mm broad, entire, acute to acuminate, basally cuneate, silky or tomentose on both surfaces, in our plants, at least, more pubescent beneath than above. Spikes 0.7 to 3 cm long, (5)7 to 8(10) mm broad, stout or slender, lateral spikes sessile. Bracts acuminate, scarious; bractlets rotund or sometimes acuminate, stramineous, blackish, or gray-brown. Mature calyx tube densely woolly, conic to flask-shaped, with 2 lateral rows of distinct, rather stout, round, sharp spines, faces with 1 to 3 blunt tubercles near the base; calyx lobes oblong-linear, obtuse or acute, (0.5)1.2 to 2 mm long, glabrate. Seed lenticular,1.2 to 1.6 mm broad, yellowish- or reddish-brown, lustrous. Sand and gravel soils of roadsides, pastures, etc. Throughout TX except the extreme E.; S. ND, IN, IA, and CO, S. to AZ, AR, TX, and Mex.; adventive E. to NY, NJ, MD, VA, KY, OH, SC, GA, and AL. Mar.-Nov.
Reported to hybridize with F. floridana in parts of its range (GPFA 1986).
Very similar to F. braunii Standl. which differs primarily in having, in addition to the lateral spines, the faces of the calyx with one or more sharp spines rather than tubercles, and lower leaves with petioles often half as long as the blades. All specimens of F. braunii the author has seen from our area are referable to A. gracilis. F. braunii is probably not present here. It is possible that the two species are not truly distinct.
2. F. floridana (Nutt.) Moq. Field Snake-cotton. Annual herb; stems rather stout, erect, wand-like, (3)4 to 18 dm tall, to 7 mm thick, usually simple at the base and sparsely branched above, canescent or tomentose, branches sericeus-tomentose, puberulent, or tomentose with whitish or brownish hairs, often viscid above. Leaves short petiolate, oblanceolate to spatulate, oblong, or elliptic, (2)3 to 10(14) cm long, (0.5)1 to 2(3) cm broad, obtuse to acute, basally cuneate, canescent or silky to subscabrous above, sericeus-tomentose below with pale or tawny hairs. Spikes dense, white-woolly, 1 to 10 cm long, 1 to 1.5 cm thick, often some of the lateral spikes pedunculate. Bracts scarious, acuminate to cuspidate; bractlets rotund, whitish to stramineous, fuscous, or blackish, shorter than the calyx. Mature calyx densely woolly, flask-shaped, somewhat flattened, 5 to 6 mm long, with 2 lateral deeply dentate wings or crests, at least some of the teeth coalescent, 1 or both faces of the tube with 1 or 2 tuberculate or spiny ridges or with a single basal spine; calyx lobes oblong to narrowly rhombic, obtuse, 1 to 2 mm long, glabrate, greenish-white to pinkish. Seed 1.5 to 2 mm long, reddish-brown, lustrous. Sandy soils of fields, hillsides, etc. Throughout the state.
Two varieties are listed for TX, both of which are found here. Some specimens appear to be intermediate between the two.
var. floridana. Branches with short, often viscid whitish or brownish hairs; leaves usually subscabrous to canescent above; peduncles with short hairs less than 2 mm long; one or both faces of the mature calyx with 1 or 2 tuberculate or spinose ridges in addition to the lateral crests. By far the more common variety in our area. S. GA and FL to MS and TX, N. to DE, MD, and NJ. May-Nov.
var. campestris (Small) Fern. Branches sericeus-tomentose with longish, white to brownish hairs, slightly viscid above; leaves canescent to silky above, sericeus-tomentose below with long, usually tawny hairs; peduncles lanate with hairs often 2 mm long; one or both faces of the mature calyx tube with a single basal spine in addition to the lateral crests. Dry fields, hillsides, plains, etc. MN, IL, IN, and WI to SD, NE, OK, CO, TX, AR, and NM. May-Nov. [F. campestris Small].
3. F. drummondii Moq. Drummond Snake-cotton. Annual. Stems stout, erect or ascending, 3 to 12 dm tall, branched at the base, branches sericeus-lanate with brownish hairs, often viscid above. Petioles of lowest leaves half as long as the blades, upper leaves short petiolate. Blades oblong to oblong-lanceolate, 5 to 14 cm long, 1 to 4 cm broad, apically obtuse, acute to acuminate basally, canescent above, densely sericeus-tomentose below with brownish hairs. Bracts broadly ovate, acuminate, stramineous or fuscous; bractlets stramineous or fuscous. Calyx tube at maturity with 2 erose wings, one or both faces with 1 or 2 low dentate ridges; calyx lobes lance-oblong, acutish. Seed 1.5 mm long, fuscous. Dry sandy soils of S. OK and TX.
Reported at least from Brazos, Robertson, and Madison Cos. in our area, but all specimens examined by the author seem referable to F. floridana, having at least 1 definite tooth on one or both lateral crests and not merely erose. Our plants, also, tend to the subscabrous condition of the upper leaf surface, which is typical of F. floridana. F. drummondii is treated in many works as a race of F. floridana. I would tend to agree with this interpretation. If F. drummondii is a separate species, it is quite possible that it is absent from our area or only occasional in occurrence.
Annual or perennial herbs. Stems prostrate to decumbent, ascending, or erect. Leaves opposite, sessile or petiolate, entire. Flowers perfect, sessile, in silvery-white, pedunculate, head-like spikes. Sepals 5, equal or unequal, 2 more concave than the others, distinct, scarious. Stamens (3)5, filaments united below into a cuplike tube, anthers 1-celled; staminodia present, shorter than to longer than the filaments. Stigma capitate. Ovule 1. Fruit a flattened, indehiscent utricle; seed lenticular, smooth.
About 80 species of tropical and warm regions of the world; 5 in TX; 2 here.
Several species are used as bedding plants for their colorful foliage, especially A. tenella Colla (syn.= A. ficoidea), Jacob's Coat, which has leaves of yellow, red, bronze, maroon, etc. Some species, but not ours, are eaten like spinach (Mabberley 1987).
1. Heads sessile; stems prostrate; plants of waste areas, lawns; etc............ 1. A. caracasana
1. Heads pedunculate; stems prostrate to ascending; plants of wet areas.................................
........ ..........................................................................................................2. A. philoxeroides
1. A. caracasana H.B.K. Mat Chaff-flower, Verdolaga. Perennial from an elongate, tuberous root; stems prostrate, 1 to 5 dm long, branched at the base, forking, hirsute with spreading or ascending hairs or glabrate. Leaves opposite but appearing clustered; blades spatulate to orbicular, 8 to 20 mm long, lightly pilose to glabrate, obtuse or abruptly pointed, tapered into a short petiole 1 to 5 mm long. Flowers in axillary, ovoid to short-cylindric heads; bracts whitish, ovate. Sepals 5, unequal, 3 to 5 mm long, lanceolate to ovate, stiff and awn-tipped, 1-nerved, back and margins with glochidiate hairs; stamens (3)5; staminodia about equalling the filaments, entire-margined. Utricle ovoid, flattened, winged below the apex; seed ovate-orbicular, 1 to 1.6 mm long, reddish-brown, lustrous, minutely pitted. Waste places, lawns, cultivated areas, etc. N. TX S. to the Rio Grande Plains, W. through the Ed. Plat. to the Trans-Pecos; SC to FL, W. to OK, TX, and CA; also S. America. Apr.-Nov. [A. peploides (H. & B.) Urban; A. repens of some American authors].
2. A. philoxeroides (Mart.) Griseb. Alligator-weed. Perennial from fibrous roots; stems prostrate to decumbent or ascending, 0.3 to 1 m long, branched and stoloniferous or mat-forming, glabrous except for hairs within the leaf bases, sometimes pinkish when fresh, in age becoming hollow and somewhat flattened. Leaves opposite, more or less fleshy; blades oblanceolate to linear-lanceolate, -elliptic or ovate, to ca. 11 cm long and 1.5 cm broad, with scattered hairs when young, glabrous in age, acute to mucronate, entire or with tiny teeth, these sometimes with long hairs, tapered to a sessile base or with a very short dilated and clasping petiole-like base. Heads terminal and/or axillary, on peduncles to 7 cm long; bracts glabrous, broadly ovate, ca. 1/4 as long as the perianths. Sepals 4 or more, subequal, whitish, 5 to 6 mm long, lanceolate to oblong-ovate, faintly 4-nerved, obtuse to acute or serrulate apically; stamens 5, the filaments partially fused below into a tube, anthers versatile; staminodia 5, fringed or petaloid, longer than the anthers and ca. 1/2 as long s the sepals; style elongate, stigma entire, capitate and papillate. Mature fruit apparently uncommon. Wet or aquatic habitats. SE TX; known from Madison Co.; Native to S. Amer; naturalized in Cen. Amer. And on U.S. coastal plain from NC to FL, W. To TX. Mar.-Aug. [Achyranthes philoxeroides (Mart.) Standl.]
Introduced into the U.S. as an aquatic ornamental and crayfish food. It is now a troublesome aquatic weed, nearly as bad as water hyacinth.
Perennial herb from a simple or branched, taprooted caudex. Stem procumbent to decumbent, variously pubescent. Leaves tapered to winged, sheathing petioles, basal leaves linear-oblanceolate to elliptic or spatulate, cauline leaves opposite or subopposite, members of each pair unequal and separated by ca. 120o, reduced upwards, ovate to oblong-elliptic, midvein impressed above, raised below, glabrous to pilose-villous, the wings of opposite leaves connate around the nodes. Flowers perfect, congested in alternate axils, 1 to 20 or more, spikes with zig-zag rachises, each flower subtended by 1 bract and 2 bractlets, all falling with the fruit. Calyx of 5 unequal sepals, reflexed at anthesis and closing after, woolly-villous. Stamens 5, free of the sepals, filaments united basally into a short tube, anthers 2-locular. Ovary superior, oblong-obovoid; style short, bifid, stigmas 2; ovule 1, basal. Fruit an irregularly dehiscent utricle. Seed 1, embryo annular, surrounding the perisperm, radicle erect.
Two species of the S. U.S., Mex., and W.I.; we have the one found in TX.
Formerly included in Guilleminea.
1. G. lanuginosus (Poir.) Moq. in A. DC. Perennial herb; stems prostrate to decumbent, plants 1 to 7 dm in diameter, internodes longest at the base, slightly zig-zag, yellow, sparsely pilose to woolly-villous with straight to crinkled hairs. Basal leaves linear-oblanceolate to spatulate, 1.9 to 7 cm long, (1)2 to 10(17) mm broad, acute to obtuse and apiculate apically, basally cuneate to a winged, sheathing petiole 5 to 20 mm long, cauline leaves ovate or elliptic-ovate to oblong-ovate, (1.5)5 to 18(33) mm long, (1)2 to 7(13) mm broad, glabrous to pilose to strigose with straight hairs or glabrate above, densely pilose-sericeus below, petioles winged and sheathing-clasping. Spikes globose to cylindrical, 1 to 10(15) mm long, with 6 to 12 flowers, bracts ovate to narrowly ovate, 1 to 2 mm long, 1 to 1.5 mm broad, bractlets 1.5 to 2.6 mm long, 1 to 1.4 mm broad. Calyx 2.4 to 3.5(4) mm long, densely woolly-villous with hairs to 2 mm long, sepals lanceolate to linear-lanceolate, the outer larger sepals 2.5 to 4 mm long, 0.6 to 0.8 mm broad, green medially and with scarious margins, inner sepals 2.5 to 3.6 mm long, 0.4 to 0.5 mm broad; filaments fused basally into a tube 0.25 to 0.4 mm high, the free upper portion of each ovate, dilated; styles 0.1 to 0.33 mm long at maturity. Mature ovary 1.4 to 2 mm long, 0.8 to 1.0 mm broad; seed reddish-brown, lustrous, 0.8 to 1.2 mm long.
As described in the most recent treatment (Henrickson 1987), there are two varieties which sometimes intergrade. Either might be found here, but our plants seem largely to belong to the first.
var. lanuginosus Woolly Cottonflower. Radical leaves oblong-oblanceolate to spatulate, cauline leaves ovate to oblong-ovate, leaves pilose to sericeous beneath, pilose-setose to glabrate above; flowers 2.4 to 4 mm long; mature outer sepals usually with 3 prominent raised veins on the abaxial surface. OK through TX S. to NE. Mex.; also Haiti and Dom. Rep. Apr.-Oct. [Guilleminea lanuginosa (Poir.) Hook f.; includes Gossypianthus lanuginosus (Poir) Moq. in A. DC. var. sheldonii Uline & Bray; Gossypianthus sheldonii (Uline & Bray) Small; Guilleminea lanuginosa (Poir.) Hook. f. var. sheldonii (Uline & Bray) Mears; includes Gossypianthus rigidiflorus J. W. Hook.; Guilleminea lanuginosa (Poir.) Hook. f. var. rigidiflora (Hook.) Mears].
var. tenuiflorus (Hook.) Mears ex Henrick. Lanceleaf Cottonflower. Radical leaves linear to linear-oblanceolate, cauline leaves oblong-elliptical to oblong, leaves glabrous or with a few scattered marginal hairs near the base, flowers (2.5)3 to 4 mm long; outer mature sepals with the central vein more prominent than the lateral. Coastal and E. TX, S. OK, and local in AR; probably in NE. Mex. [Gossypianthus tenuiflorus Hook; Guilleminea lanuginosa (Poir.) Hook. f. var. tenuiflora (Hook.) Mears].
Herbs, scarcely if at all succulent, stems prostrate to ascending, herbage glabrous to pubescent. Leaves simple, opposite to whorled or alternate, stipules absent or minute and deciduous. Flowers generally small, not showy, usually perfect, regular, in loose cymose inflorescences or solitary in the axils. Sepals (4)5, persistent, usually distinct or sometimes partially united. Petals (staminodial in origin) minute or more often absent, rarely connate (not so in ours.) Stamens (2)5 to 10 (sometimes more), filaments distinct or basally connate, anthers opening longitudinally. Nectary ring present between stamens and pistil, at the base of the ovary or inside the filament tube. Ovary superior, usually of 2 to 5(rarely more) united carpels, multilocular below but the partitions not always reaching the apex; styles distinct or style solitary (as in Glinus); placentation usually axile, ovules 1 to usually many per locule. Fruit an achene or usually a 3- to 5-valved capsule dehiscing longitudinally or sometimes opening by transverse slits, often enclosed by the persistent calyx. Seeds with the embryo curved around perisperm, sometimes arillate.
About 10 genera and 100 species of the tropics and subtropics, primarily Africa. There are 2 genera and 4 species in TX; 2 genera and 2 species here. This family used to be treated as part of the Aizoaceae. It differs from Aizoaceae in the strict sense in having generally free stamens, herbage scarcely if at all succulent, and anthocyanin rather than betalain pigments.
1. Flowers sessile or subsessile; our plants finely pubescent; calyx split only to about the middle .........................................................................................................................1. Glinus
1. Flowers on slender pedicels; plants glabrous; sepals wholly distinct ...................2. Mollugo
Annual herbs, stems commonly prostrate or ascending, plants at least partially pubescent (sometimes glabrous, but not ours.) Leaves whorled, the members of each whorl unequal, entire. Flowers in dense, short-peduncled clusters in the axils of the upper leaves. Sepals 5, distinct. Corolla absent. Stamens 3 to 10(more). Fruit a 3-valved, loculicidal capsule with numerous smooth or minutely tuberculate seeds.
Roughly 6 to 12 species of warm temperate and tropic areas; 2 in TX; 1 here.
NOTE: G. lotoides L, with tuberculate seeds, is an Old World species introduced to TX. It may someday be found here.
1. G. radiatus (R. & P.) Rohrb. Annual, stems generally prostrate and radiating from the center, to 5 dm long; herbage more or less stellate-tomentose, especially in the inflorescence. Leaves elliptic to obovate or widely spatulate, apices acute, rounded, or apiculate, to 2.5 cm long and 1.5 cm broad, tapered to petioles to 6 mm long. Flowers in clusters of 10 or more. Calyx stellate-tomentose, lobes ca. 2.7 to 4 mm long, oblong-elliptic to lanceolate with acuminate tips; stamens 3 to 5, shorter than the calyx, filaments slender, ca. 1 mm long, anthers ca. 0.5 mm long. Capsule ellipsoid, 3 to 3.5 mm long. Seeds many, brown to reddish-brown, smooth and lustrous or sometimes pebbly, but not tuberculate, ca. 0.4 mm long. Muddy soils or sands, also a weed of wet lawns. S. and E. TX; TX through Mex. to S. Amer. and W. I. Apr-Oct, with a few collections as early as Feb.
Annual herbs, herbage not succulent, stems well-branched. Leaves in a rosette, opposite, or verticillate, those of each node unequal. Flowers small, perfect, with filiform pedicels, axillary or in cymes. Calyx of 5 persistent, scarious-margined sepals. Corolla absent. Stamens 3 to 5(more). Ovary superior, 3-celled. Fruit a thin-walled ovoid to ellipsoid capsule, dehiscing longitudinally and with many seeds.
About 15 to 20 species of the warm regions of the world; 2 in TX; 1 here.
Some, but not ours, are used as potherbs or in medicines (Mabberley 1987).
1. M. verticillata L. Carpetweed, Indian Chickweed. Decumbent, prostrate, or ascending annual; herbage totally glabrous. Stems slender, wiry, radiate from the base and a short taproot, dichotomously branched, to 2 dm long, nodes slightly swollen, not rooting. Leaves pinnately-veined, in whorls of 3 to 8, those of a group unequal, broadly to slenderly oblanceolate to spatulate or sometimes linear, entire, 1 to 3 cm long and a maximum of 1 cm broad, apices obtuse, bases attenuate to short-petiolate or else sessile, base of plant sometimes with a whorl of broader leaves. Flowers small, 2 to 5 in sessile, minutely-bracted cymes at the nodes, pedicels filiform, 5 to 15 mm long; sepals pale green to white, or green with white margins, oblong to elliptic, to 2.5 mm long and 1 mm broad; stamens commonly 3(4 or 5), alternate with the sepals and carpels, united at the base by a ring of filament tissue: stigmas 3. Capsule slightly longer than the sepals, ovoid to ellipsoid; seeds minute, numerous, smooth, shining reddish-brown. A weed of waste ground and cultivated areas, roadsides, open sandy woods, brushlands, and dunes. Throughout TX and temperate and tropical America; apparently native to tropical America. Blooms all year.
Annual or perennial herbs (as ours) or rarely shrubs or subshrubs. Stems commonly somewhat succulent, usually glabrous or sometimes pilose at the nodes. Leaves alternate, opposite, or in a basal rosette, entire, commonly fleshy. Stipules scarious, modified into hairs, or lacking. Flowers terminal or axillary, solitary, racemose, cymose, or paniculate, regular (or nearly so) and perfect. Sepals 2, persistent or deciduous, herbaceous or scarious, sometimes interpreted as bracts. Petals commonly 4 or 5 in non-cultivated material, often dehiscent or fugacious, sometimes interpreted as sepals. Stamens 4 to many, usually as many as the petals, inserted with and opposite the petals, free or sometimes united to the corolla or else grouped in bundles; anthers 2-celled, longitudinally dehiscent. Carpels 2 to 3(9), ovary 1-celled, superior to partly or fully inferior, styles 2 to 7, more or less united, ovules many on a free-central or basal placenta. Fruit a circumscissile or longitudinally dehiscent capsule with as many valves as styles. Seeds 3 to many or 1 or 2 by abortion, usually reniform-orbicular, compressed, often smooth or sometimes roughened, tuberculate, and/or strophiolate (with an appendage at the hilum); embryo peripheral, curved around the perisperm. In nearly all, betalain pigments present and anthocyanins absent.
About 21 genera and 400 species worldwide. Texas has 4 genera and 17 species. We have 3 genera and 5 species.
The family is of importance primarily for ornamentals in a few genera, including Portulaca. Some, including species of Portulaca, are edible. (Mabberly 1987).
1. Calyx adherent to the base of the ovary; fruit a circumscissile capsule ....................1. Portulaca
1. Calyx free of the ovary; fruit a capsule opening by valves .............................................................2
2(1) Sepals persistent; plants from corms; flowers in racemes .......................................2. Claytonia
2. Sepals early-deciduous; plants from fleshy roots; flowers in cymes ............................3. Talinum
Succulent, prostrate to ascending or erect, annual (as in ours) or perennial herbs. Leaves flat to terete, alternate or opposite, the uppermost often crowded or appearing whorled beneath the flowers. Stipules scarious, reduced to tufts of hair, or absent. Flowers perfect, solitary or clustered at the tips of the stem and branches. Calyx 2-cleft. Petals (4)5(6), variously colored. Stamens 7 to many, inserted at the base of the petals. Ovary partly to wholly inferior, adnate to the calyx tube below, styles 3 to 9, ovules many. Capsule unilocular, membranous, circumscissile. Seeds many, reniform, smooth to tuberculate or echinate.
About 40 species worldwide, especially in the tropics and subtropics; 6 species in TX; 3 here.
Some, e.g. P. oleracea, can be very weedy. Some, including subspecies of P. oleracea, can be eaten as potherbs or are used medicinally by some cultures. P. grandiflora, a native of Argentina, is commonly called Moss Rose and is widely cultivated for its pink, orange, yellow, red, or white flowers (Mabberley 1987). It occasionally escapes cultivation, but usually does not persist for long. It may distinguished from our native species by its larger (3 to 5 cm broad) flowers which have more than 5 petals.
1. Lower valve of capsule with an expanded circular wing just below its rim ...............................
......................................................................................................................1. P. umbraticola
1. Lower valve of capsule without a circular wing .......................................................................2
2(1) Inflorescence and leaf axils glabrous or inconspicuously pilose; leaf blades fleshy but flat; corolla yellow .....................................................................................................2. P. oleracea
2. Inflorescence and leaf axils conspicuously white-villous; leaf blades fleshy and terete to subterete; corolla red-purple ..................................................................................3. P. pilosa
1. P. umbraticola H.B.K. Plants annual, succulent, glabrous, prostrate to partially ascending; stems angled, to ca. 20 cm long. Leaves relatively few, sessile, blades 1 to 3 cm long, 2 to 11 mm broad, flat, lower leaves spatulate to obovate, apices obtuse to rounded, upper leaves oblanceolate to oblong, apices acute to rounded. Flowers grouped at the tips of stems and branches. Sepals ovate, minutely keeled; petals 5, yellow to orange and/or partially red, spatulate or obovate, apices acute or cuspidate; stamens 7 to many; styles 3 to 6. Capsule circumscissile at or above the middle, lower valve with a membranous circular wing just below the rim, upper valve flattish. Seeds gray, cochleate, tuberculate. Sandy soils of prairies, mesquite thickets, roadsides, and salt marshes. Most of TX; GA W. to OK, TX, AZ and Baja CA; also Jamaica, Cuba. Mar.-Nov. Some sources recognize subspecies. [P. coronata Small].
2. P. oleracea L. Common Purslane, Verdolaga. Plants annual, fleshy, glabrous; stems prostrate or sometimes ascending or mat-forming, usually radiating from the base, sometimes purplish or reddish, branches 6 to 30 cm long. Leaves alternate (occasionally opposite), blades succulent but flat, oblanceolate to spatulate or cuneate-obovate, 6 to 30 mm long, 0.2 to 11 mm broad (sometimes larger), apices rounded or nearly truncate or retuse, bases attenuate,; axils glabrous or inconspicuously pilose. Flower buds flattened, acute, flowers solitary or in terminal clusters, sessile, without conspicuous subtending hairs. Sepals broadly ovate to orbicular, 2.8 to 4.5 mm long, 2.8 to 3.8 mm wide, keeled, apices more or less acute; corolla yellow, 5 to 10 mm broad, petals 3 to 4.6 mm long, 1.8 to 3 mm wide; stamens 6 to 10; style lobes 4 to 6. Capsule globose, 5 to 9 mm long, circumscissile at or below the middle; seeds 0.7 to 0.8(1) mm broad, black, granulate. Common weed of sandy waste and cultivated areas, grassy slopes, salt marshes, and margins of seasonal pools. Throughout TX and much of the temperate to tropical regions of the world; perhaps native originally to W. Asia. May-Nov. [P. neglecta Mack. & Bush].
The stems and leaves of this purslane are edible raw, boiled, steamed, fried, or pickled. It is high in vitamin A and also contain vitamin C, iron, and calcium. The juice can be used to thicken soups. The seeds are also nutritious and can be ground and mixed half-and-half with wheat flour. Purslane contains oxalic acid which can bind calcium in the diet and lead to kidney stones, so the plant should not form a major part of the diet. There have been rare cases of sheep poisonings in Australia as a result of the animals' eating large amounts in a short time or moderate amounts for several weeks (Tull 1987).
3. P. pilosa L. Chisme. Annual from an often-thickened taproot; stems succulent, 3 to 6, prostrate to somewhat erect, 5 to 15 cm long, upward-pointing branches somewhat shorter and with shorter internodes. Leaves alternate, often crowded, fleshy, terete to subterete, linear to oblanceolate-linear, 5 to 15 mm long, 0.5 to 1.5 mm broad, those of the branches often smaller than those of the main stems, axils conspicuously villous or woolly with white or whitish kinky hairs to 5 or 7 mm long. Flowers terminal, subsessile in moderately to densely villous clusters of 2 to 8, subtended by 6 to 10 linear, succulent leaves 5 to 12 mm long. Calyx remaining on upper part of capsule, 4(6) mm long with triangular to triangular-oblanceolate lobes; corolla red-purple, petals obovate, 6(7.5) mm long, 3 to 4.5 mm broad, apices retuse; stamens 10 to 15(30); styles 3 to 5. Capsule ovoid-globose, circumscissile below the middle, the upper part of the capsule hemispheric and shiny, invested with the persistent remains of calyx, corolla, and stamens, lower half of capsule 2.5 to 3.5 mm in diameter, short-stipitate; seeds many, black, 0.3 to 0.5 mm in diameter, stellate-tubercular. Sandy or gravelly soils. Cen. TX and Trans Pecos, less common eastward; SE. U.S.: FL, BA, AL, and MS, W. to AR, OK, TX, and NM; also Mex. and the Caribbean. Spring-fall. [P. mundula I.M. Johnston and var. pilosa (I. M. Johnston) Legrand].
This species exhibits a great deal of variation in habit, leaf shape, and pubescence, and at one time was divided into P. pilosa and P. mundula. Drier conditions favor the development of smaller, more terete branch leaves; wetter conditions favor larger leaves which tend to be semicircular in cross-section. Plants of temperate regions are more erect then plants of the tropics. Diploid and tetraploid populations are known, but there is no correlation between chromosome number and morphological variation. Matthews, et al. (1992) determined that the two names apply to extremes in variation of a single species, P. pilosa.
About 20 species of the N. Hemisphere. We have the 1 species found in TX.
1. C. virginica L. Spring Beauty, Virginia Spring Beauty, Fairy Spuds. Perennial herb from a rather deeply buried globose corm 1 to 3 cm in diameter, plants somewhat succulent, glabrous, erect or ascending, 10 to 30 cm tall. Leaves somewhat grasslike, basal leaves petiolate, 6 to 20 cm long, linear-oblanceolate, blade to twice as long as the petiole, 5 to 15 mm broad, acute at both ends, inconspicuously 3-ribbed, stem leaves short-petiolate, opposite, 9 to 15 cm long. Raceme erect to somewhat nodding, (5-)6- to 15-(19-) flowered, with 1 small ovate bract beneath the lowest pedicel, pedicels 1.5 to 4 cm long, recurved in fruit. Flowers regular, perfect; sepals 2, ovate-rounded or oval, apices usually rounded or obtuse, 5 to 7 mm long, herbaceous, persistent; petals 5 (occasionally 6 in aberrant blossoms), oval, 9 to 14 mm long, rounded to obtuse or rarely retuse apically, white or rosy with darker pink or purple veins; stamens 5, opposite the petals and basally adnate to them, anthers commonly rose or reddish; ovary 1-celled, styles 3, united to near the apex, ovules 6. Fruit a rounded-ovoid, membranaceous, loculicidal capsule ca. 4 mm long, the edges of the valves inrolling at dehiscence; seeds 1 to 6, orbicular, shining blackish-brown, ca. 2 mm in diameter. Sandy soil of rich woods, clearings, thickets, lawns, vacant lots, etc. E. 1/3 of TX; N. S. and S. Que. to MN, S. to GA, MS, LA, TX, OK, and MO. Jan.-Apr.
These are among the first spring plants to bloom in the College Station area. The leaves are edible raw or cooked. The crisp tubers are starchy, edible raw or cooked, and were consumed by Native Americans (Kindscher 1987.) Many feel that the beauty of this plant is greater than its food value and that it ought to be left undisturbed.
Herbs (as in TX material) or shrubby plants, commonly with fleshy, tuberous roots. Stems elongate or sometimes very short. Leaves fleshy, alternate to subopposite, entire, flat to terete. Flowers commonly showy, produced in pedunculate cymes or sometimes 1 to several in the axils. Sepals 2, distinct, deciduous. Petals 5 (rarely more), soon withering. Stamens 5 to many. Ovary wholly superior; styles 3, united to near the apex. Capsule unicellular, 3-valved, many-seeded. Seeds round-reniform, flattened, smooth or minutely roughened, tuberculate, or papillose.
About 50 species in the warmer parts of the world, especially N. America. Texas is home to 9 species, mostly in the SW.; we have 1 species.
1. T. parviflorum Nutt. Prairie Flame-flower. Herb 5 to 19(20) cm tall, roots fleshy; stem short or plants acaulescent. Leaves terete or subterete, linear, 1.5 to 5 cm long, 0.8 to 2.5 mm wide, broadened at the base. Flowers in a terminal cyme with small bracts at the forks, peduncles slender, 3 to 15 cm long, usually much exceeding the leaves; pedicels slender. Sepals ovate or oval-ovate, 2.7 to 4 mm long, 1.5 to 2 mm broad, deciduous; petals pink to purplish, elliptic or obovate, (4)5 to 7 mm long, 2.2 to 2.6 mm broad; stamens 4 to 8, anthers 0.9 mm long, oblong; style exceeding stamens, stigmas capitate or subcapitate. Capsule ellipsoid, 3.5 to 5 mm long, 2.8 to 3 mm in diameter; seeds 0.8 to 0.9 mm broad, smooth. Sandy soil in woodlands and over rock outcrops. In most of TX; MN and ND S. to AZ, TX, and AR. May-July.
This plant is not much collected in our area, perhaps because it is easily overlooked and blooms during the summer lull in plant collecting and not because it is really rare.
Ours annual, biennial, or perennial herbs, occasionally somewhat woody at the base if persistent; stems often with swollen nodes. Leaves opposite or sometimes appearing whorled, very rarely (and not in ours) alternate, entire, leaves of a pair often united at the base or connected by a transverse line, generally sessile, estipulate or stipulate; stipules if present often scarious or hyaline. Flowers regular, usually perfect, ours all hypogynous, in dichasial cymes, axillary, or terminal and solitary, (4)5-merous except perhaps for the gynoecium. Sepals (4)5, persistent, free or united to some degree. Corolla (perhaps of staminodial origin) of as many petals as sepals, sometimes fewer or reduced, occasionally absent; petals if present free, often clawed, toothed, and/or bifid. Stamens (1 to 4)5 to 10 in 1 or 2 whorls, usually as many as or twice as many as the sepals or petals, if the same number as the petals then usually alternate with them, filaments free or basally united to the corolla in a short tube. Nectary disk sometimes present. Gynoecium of 2 to 5 united carpels, styles as many as the carpels, free or united to some degree, ovary 1-celled or partially 2- to 5-celled at the base, placentation free-central, the central column reaching the ceiling of the locule or not (axile in partitioned ovaries, not so in ours.) Fruit a many-seeded, thin-walled capsule dehiscing by as many or twice as many entire or bifid valved or teeth as there are styles, rarely, as in Paronychia, fruit a 1-seeded utricle. Seeds flat to plump, smooth or ornamented, occasionally with marginal wings. Betalain pigments are lacking, the red or blue colors instead produced by anthocyanins.
Between 70 and 90 genera and 2,000 to 2,100 species worldwide, especially in the temperate and warmer northern hemisphere. There are 17 genera and 55 species in TX; 10 genera and 16 species here.
Many species are ornamental, such as Dianthus--carnation, Gypsophila--Baby's breath, and Saponaria--soapwort. Some are common weeds, such as Stellaria--chickweed and Cerastium--mouse-ear chickweed. A few have medicinal uses or are edible (Mabberley 1987).
1. Stipules present ........................................................................................................................2
1. Stipules absent ..........................................................................................................................5
2(1) Sepals, at least some of them, with a bristly tooth on each margin; leaves sharp, stiff, and scale- or bristle-like ..............................................................................................1. Loeflingia
2. Sepals entire, without marginal teeth (sometimes with apical teeth); leaves not scale- or bristle-like, though sometimes narrowly linear ........................................................................3
3(2) Sepals awn-tipped, mucronate, or cuspidate; petals absent or reduced and bristle-like; fruit a 1-seeded utricle .......................................................................................2. Paronychia
3. Sepals not awn-tipped, mucronate, or cuspidate; petals present; fruit a several- to many- seeded capsule .........................................................................................................................4
4(3) Lower leaves usually appearing in whorls of 4 because of short internode length, petiolate; styles united at the base ....................................................................................3. Polycarpon
4. Lower leaves opposite, sessile or nearly so; styles entirely free .....................4. Spergularia
5(1) Flowers and flower clusters subtended by 2 or 3 pairs of ovate-oblong scarious bracts.......
.. .........................................................................................................................5. Petrorhagia
5. Flowers not subtended by scarious bracts..............................................................................6
6(5) Sepals united; foliage often sticky-viscid ....................................................................6. Silene
6. Sepals free; foliage not sticky-viscid ........................................................................................7
7(6) Petals present at anthesis ........................................................................................................8
7. Petals absent at anthesis .......................................................................................................11
8(7) Styles 3 (sometimes varying from 3 to 6); valves of capsule as many as or twice as many as styles, usually 3 or 6 ............................................................................................................9
8. Styles 4 or 5, as many as the sepals; valves of capsule twice as many as the styles, usually 8 or 10 .........................................................................................................................10
9(8) Petals deeply cleft or bifid, sometimes almost to base .........................................7. Stellaria
9. Petals entire or merely notched or lobed ..............................................................8. Arenaria
10(8) Petals rudimentary, entire; leaves linear-subulate ...................................................9. Sagina
10. Petals well-developed, notched or cleft; leaves not linear-subulate ..............10. Cerastium
11(10) Leaves linear ..............................................................................................................9. Sagina
11. Leaves wider: oblong, oblanceolate, obovate, ovate, etc .....................................................12
12(11) Styles 3(sometimes 3 to 6); valves of capsule twice as many as the styles, usually 6 ...........
12. Styles usually 5; valves of capsule twice as many as the styles, usually10 ...10. Cerastium
NOTE: A species not yet found in our area deserves mention. Vaccaria hispanica (P. Mill.) Rauschert, Cow-herb, Cow-cockle. A native of Eurasia, introduced into parts of TX and also occasionally cultivated. Sepals united into a strongly 5-angled tube, petals pale rose, clawed, stipules none. Reported for our area but not yet seen outside of cultivation.
Seven species of N. Amer. We have the one species found in TX.
1. L. squarrosa Nutt. Spreading Loeflingia. Taprooted annual herb to 15 cm tall and about as broad, well-developed plants nearly globose; herbage with minute, blunt, glandular hairs more or less throughout. Stems freely branched, usually from near the base, rather stiff, ascending, often recurved. Leaves mostly crowded on short branchlets or at the tips of the branches stiff, straight or recurved (squarrose), subulate-setaceous, 4 to 10 mm long, the tips subspinose; stipules ovate-lanceolate to linear and setaceous, longer than wide, hyaline, adnate to the leaf at the base. Flowers sessile, solitary or in few-flowered fascicles in the axils, inconspicuous among the leaves. Sepals 5, free, shorter than the leaves, narrowly lanceolate, keeled, awn-tipped, somewhat recurved, the outer 2 or 3 somewhat larger than the others and resembling the leaves but with a hyaline setaceous tooth on each margin; petals 3 to 5, commonly minute and sometimes absent; stamens 3 to 5; style short or absent, the 3 stigmas nearly sessile. Ovary unilocular. Capsules slenderly ovoid, 3-valved, shorter than or equalling the sepals; seeds several, obovate, flattened, brown, ca. 0.3 to 0.5 mm long. Dry soils, especially deepish sands, also grasslands or open woods. E., Cen., and W. TX; NE. WY to OR, S. AZ, CA, and TX. Feb.-May. [L. texana Hook.].
Annual or perennial herbs, the annuals from taproots and the perennials from taproots and rootstocks, often somewhat woody at the base, plants often tufted or mat-forming. Leaves simple, opposite, sessile, entire, relatively small, linear to elliptic or oblong, apices acute to obtuse. Stipules minute to conspicuous, scarious or hyaline, often silvery, variously shaped. Flowers small, inconspicuous, perfect, in loose or dense terminal clusters or cymes or in the forks of the stems (flowers solitary or paired in P. sessiliflora), subtended and often exceeded by bracts. Sepals 5, free or briefly united basally, 1- to 3-nerved, apically cucullate (hooded), often apically awned or mucronate as well, white to greenish, if greenish then often with whitish or scarious margins, persistent. Corolla absent or reduced to bristle-like staminodia. Stamens (2 to 4)5, opposite the sepals, attached to a small perigynous disk within the basal portion of the calyx, included within the calyx. Ovary superior, ovoid-subglobose, 1-celled; style 1 basally, 2-parted apically; stigmas 2, included within the calyx; ovule 1, basal. Fruit a membranaceous utricle enclosed by the calyx.
About 50 species of temperate and tropical regions of the world; 13 recorded from TX; 3 here. The genus includes the former Anychia. This treatment follows Core (1941), Chaudri (1968), and Turner (1983).
The plants were formerly used to treat whitlow, a disease of the fingers. A few species are cultivated as rock-garden plants (Mabberley 1987).
1. Plants perennial, from thickened, often branched rootstocks .........................1. P. virginica
1. Plants annuals from taproots ...................................................................................................2
2(1) Sepal hood conspicuous, whitish and hornlike (even in dry material); leaves elliptic to oblanceolate, at least some more than 2 mm ............................................2. P. drummondii
2. Sepal hood not conspicuous, whitish, or hornlike (especially when dry); leaves linear, seldom more than 1.5 mm broad ....................................................................................3. P. setacea
1. P. virginica Spreng. Parks' Nailwort, Broom Nailwort. Perennial herb from a much-branched woody base, plants becoming reddish- or yellowish-brown with age and upon drying. Stem densely cespitose or matted, procumbent, branching from the base and simple above, leafy, internodes short; flowering stems erect, 15 to 40 cm tall, glabrous to puberulent (or pubescent), branching pseudo-dichotomously above in the inflorescence. Leaves linear-subulate, ascending, 15 to 25(30) mm long and 0.5 to 1 mm broad, longer than the internodes, acute and minutely mucronate, glabrous to glabrate, margins minutely ciliate and serrulate, often with a conspicuous midrib or else grooved; stipules silvery, linear-lanceolate, acuminate-attenuate, entire, the pair at a node often more or less united, ca. 8 to 13 mm long, shorter than the leaves. Cymes terminal, many times dichotomous, many-flowered, diffuse and fastigiate; bracts similar to the leaves and somewhat shorter than the flowers. Flowers sessile, ca. 3 to 4 mm long excluding the calyx awns, oblong-cylindrical, light brown above, reddish brown below, glabrous to sparsely hirtellous; sepals linear or lanceolate-oblong, 3-ribbed, 2.25 to 2.75 mm long, with very narrow membranous margins, slightly hooded apically within, awn or cusp stout, 0.45 to 1.25 mm long, scabrous to nearly smooth, almost erect to somewhat spreading; petals 0.5 mm long, if present. Fruit ovoid, ca. 2 mm long. Limestone outcrops and gravelly, limey soils of the Ed. Plat. to N. Cen. TX; Cen. TX N. to E. OK and W. AR. July-Nov. [P. dichotoma (L.) Nutt.; P. virginica Spreng var. scoparia (Small) Cory and var. parksii (Cory) Chaudhri; P. parksii Cory].
2. P. drummondii T. & G. Drummond Nailwort. Annual (or biennial?) herb from a taproot; stems 10 to 25(35) cm tall, deflexed- or retrorsely-hirsute, branched just above the base, well-branched, branches erect to spreading, minutely pubescent; internodes relatively long, to 3 cm. Leaves stiffish, linear-oblong to spatulate-oblanceolate, tapered to the base, apically obtuse to subacute or slightly mucronate, the upper leaves 10 to 17(20) mm long and 2 to 35 mm broad, lower leaves larger, 2 to 3 cm long, 4 to 7 mm broad, pubescence variously strigose to appressed pubescent to scabridulous or sometimes 1 or both sides glabrous to glabrescent, at least some leaves with midrib prominent near base, margins somewhat papillate and yellow- or pale-green; stipules lanceolate to lance-ovate, long-acuminate, shorter than the leaves, 5 to 10 mm long and 2 to 5 mm broad, silvery. Cymes terminal or subterminal and much-branched, nearly leafless, 10 to 20 mm in diameter, the bracts scarious, ciliate, shorter than the flowers, ovate, ca. 2 mm long and 0.75 mm broad. Flowers subsessile, (1.5)2 to 2.25(2.5) mm long, calyx turbinate and with short, hooked hairs at the base, contracted slightly just above the base; sepals ascending, cuneate to obovate-spatulate (1)1.25 to 1.4 mm long, conspicuously hooded and with a spreading-ascending awn 0.3 to 0.45 mm long, sepal tip, hood, and awn ca. 1 mm long together, white, rigid, and remaining so in pressed specimens, body of the sepal reddish-brown, sepals as a whole short-pubescent to glabrous or glabrate, 3-nerved dorsally; petals to 0.5 mm long, if discernable. Fruit subglobose (0.6)0.8 mm long and (0.6)0.7 mm broad, papillose. Sandy soils of dry oak woods and in sand dunes. S. Cen. TX, NE. to NE. TX and SE. to about Aransas Co. Endemic to TX. Apr.-Oct.
If subspecies are recognized, our plants are subsp. drummondii.
3. P. setacea T. & G. Bristle Nailwort. Annual from a taproot; stem erect to erect-ascending, 5 to 20(30) cm tall, simple at the base and branched pseudodichotomously above, glabrous to puberulent (occasionally hispid), sometimes glaucous. Leaves linear to subulate-setaceous, erect or ascending, 5 to 20 mm long, 0.3 to 1 mm broad, lower ones narrower and shorter than the upper, strongly ribbed, minutely cuspidate, upper leaves longer and a little broader, obtuse, glabrous; stipules lanceolate, long-attenuate or acuminate, 3 to 8 mm long, shorter than the leaves, silvery. Cymes dichasial and diffuse, individual cymules compact, flowers subsessile, equalled or exceeded by bracts, brownish to yellowish-green, (16)1.75 to 2 mm long excluding the sepal awns, tubular-oblong to flask-shaped, receptacle not distinctly turbinate or enlarged, lower portion of the flower often densely pubescent or strigose. Sepals narrowly oblong or oblong-linear, 1.25 mm long, ribbed dorsally, more or less glabrous or slightly ciliate at the tips, scarious margins very narrow and membranous, especially in the upper portion, not whitish or rigid, hood distinct and with an obvious tip, neither whitish, awn whitish or yellowish, ca. 0.65 to 1.25 mm long, more than half as long as the body of the sepal or about as long, stiff, scabridulous to nearly smooth, spreading, usually conspicuous; petals 0.6 to 0.65 mm long, if discernable. Fruit ovoid-oblong, 1 mm long and 0.8 mm broad, smooth and not papillose. Gravelly, sandy, or silty soils and on limestone hills or flats. S. Cen. TX and mid-W. TX, S. to Rio Grande Plains (W to Trans-Pecos?). Endemic to TX. Apr.-Aug.
If varieties are recognized, ours are var. setacea.
For years our plants were misidentified as P. chorizanthoides, a species which B. L. Turner (1983) determined to be the same as P. lindheimeri. P. lindheimeri is very closely related and quite similar. It is found on the Edwards Plateau. It may someday be found on the limestone outcrops in our area. It may be recognized by sepal awns only 1/3 to 1/2 as long as the body of the sepal, lack of scarious sepal margins, and a receptacle somewhat enlarged at the base. Characters of stem pubescence do not seem to be useful in separating the two species--both exhibit variation from glabrous to scabridulous-puberulent.
Sixteen species worldwide. We have the 1 species naturalized in TX from Europe and the Mediterranean.
1. P. tetraphyllum (L.) L. Four-leaved Allseed. Taprooted annual; stems prostrate or ascending, diffusely much-branched (sometimes pseudodichotomous), to 15 cm long, glabrous. Leaves opposite or in fours, flat, ovate to obovate or elliptic, 2 to 15 mm long, 1 to 8 mm wide, entire, apices rounded to obtuse or mucronate, basally attenuate or abruptly narrowed to a short petiole to 1.2 mm long or else leaves subsessile; stipules and floral bracts scarious, ovate-lanceolate to lance-acuminate or aristate, shorter than the leaves. Flowers small, numerous, in dense cymes, pedicels 0.5 to 3 mm long. Sepals 5, 1.5 to 3 mm long, the outer 2 usually shorter than the others, all scarious-margined and green-keeled, keel obscurely serrulate, all acute to acuminate (aristate); petals 5, white, very thin, oblanceolate, smaller than the sepals, 0.5 to 0.8 mm long; stamens 3 to 5; styles united, short, stigmas 3, ovary 1-celled. Capsule broadly ovoid, shorter than the sepals, 3-valved; seeds angled-ovoid, yellowish or reddish, white-papillose, ca. 0.5 mm long. On sandy or silty soils of beaches, wood margins, paths, etc. Cen. and S. TX; adventive or naturalized from Europe in at least TX, SC, and CA. Mar.-July, ours primarily Apr.
Annual or perennial herbs, low and branched. Leaves opposite or sometimes fascicled, linear to filiform, setaceous or fleshy. Stipules scarious. Flowers terminal in racemose, bracteate or leafy cymes. Calyx of 5 sepals. Petals 5 or fewer or sometimes absent. Stamens 2 to 10. Ovary unilocular. Styles 3. Capsule 3-valved to base, the many seeds globose-reniform or flattened, smooth or rough, sometimes winged or echinate.
About 40 species worldwide, commonly halophytic; 3 species in TX; 1 some-times found here.
1. S. salina J. & K. Presl. Salt-marsh Sand-Spurrey. Low annual, 5 to 20(35) cm tall; stems tufted, simple and erect or branched at the base and spreading, prostrate, or decumbent, often diffuse, fleshy, glabrate or usually glandular-pubescent or -puberulent throughout. Leaves sessile, opposite, rarely fascicled, linear, 5 to 25(40) mm long and to 1.5 mm broad, fleshy, apices blunt, acute or mucronate; stipules scarious, broadly triangular, about as wide as long or slightly longer, 1.5 to 4 mm long, shortly acuminate, margins entire to lacerate, members of a pair at a node often connate. Flowers small, perfect, usually many in bracteate, lax, racemose or scorpioid cymes, inflorescence often making up the majority of the plant; pedicels 1 to 10 mm long at maturity, often reflexed, stipitate-glandular. Sepals ovate to ovate-lanceolate, scarious-margined, obtuse, 2.5 to 4(5) mm long; petals 5, white or pink, less than half as long as the sepals; stamens 2 to 5. Capsule ovoid, as long as or exceeding the calyx, 3 to 6.5 mm long, membranaceous, 3-valved from the apex; seeds yellowish, reddish, or pale brown, obliquely ovate, 0.5 to 0.9 mm long, opaque, smooth or minutely papillose, not sculpted, wingless or occasionally with a thin, erose wing to 0.4 mm wide. Saline or brackish soil of dunes and tidal flats. E. TX, S. along the coast to S. TX, also isolated at El Paso along the Rio Grande. Collected at least once from College Station. AK to N.S., S. to CA, TX, IL, and FL, reported from ND; also Eurasia. Mar.-June. [S. marina (L.) Griseb.; Tissa marina (L.) Britt.].
5. PETRORHAGIA (Springe) Link Childing Pink
About 20 species native to the Mediterranean region; we have the 1 species found in Texas.
1. P. dubia (Raf.) G. Lòpez & Romo Childing Pink, Hairy Pink. Taprooted annual herb; stems simple or sparingly branched, to 5 dm tall, glabrous or scabrous. Leaves rather widely spaced, estipulate; blades linear to linear-oblanceoalte, to 5 cm long and ca. 3.5 mm broad, acute to broadly obtuse, 3-veined, the base a sheath 1 to 2.5 mm long. Flowers and cymes cubtended by scarious tan bracts, the longest 6 to 15 mm long, to ca. 10 mm broad, obtuse or obtuse-apiculate. Calyx synsepalous, 10 to 13 mm long, the lobes 3-veined, obtuse; petals pink or purplish, 10 to 14 mm long, 2 to 3.5 mm broad. Seeds 1.3 to 1.9 mm long, 0.8 to 1.1 mm wide, surface reticulate. Roadsides and fields. E. and NE TX.; native to Eur. and N. Afr. and now aggressively spreading in our state. [P. velutina (Guss.) P. Ball & Heyw.; Dianthus velutinus Guss.; erroneously treated as P. prolifera (L.) P. Ball & Heyw. by Correll and Johnston (1970)].
Annual, biennial, or perennial herbs, stems solitary or several. Leaves opposite or in some species (not ours) whorled, sessile to short-petiolate, estipulate. Flowers hypogynous, perfect or imperfect (not ours), solitary or in leafy-bracted cymes, cymes sometimes appearing one-sided. Calyx cylindrical to ovoid or campanulate, 5-toothed, with about 10 to 20 nerves, often accrescent in fruit. Petals 5 (occasionally absent in S. antirrhina), white to poink or red, conspicuous or not so, usually clawed, the blades commonly cleft or toothed, most often with paired, scale-like appendages at the ventral junction of blade and claw. Stamens 10, briefly united to the petals below to form a short tube. Styles 3(4 to 5). Ovary elevated on a short carpophore, unilocular or sometimes incompletely 2- to 4-celled. Capsule usually about as long as calyx and enclosed within, dehiscent by twice as many valves as styles (less often valves as many as styles). Seeds many, reniform to globose, often tubercled or papillose in rows.
About 500 to 600 species worldwide, depending upon interpretation; 6 in TX; 2 here. Some species have synonyms in Wahlbergella, Lychnis, or Melandrium.
Some of the species with showier flowers are used as garden plants. Others are weedy (Mabberley 1987).
1. Herbage pubescent with glandular hairs but without mucilaginous bands on the stems; 1 flower and 1 branch per node in the inflorescence, the resulting inflorescence raceme- like ..........................................................................................................................1. S. gallica
1. Herbage puberulent to glabrous but stems commonly with mucilaginous bands on the stems; 1 flower and 2 or 3 branches per node in the inflorescence, inflorescence not racemelike ........................................................................................................2. S. antirrhina
1. S. gallica L. Forked Catch-fly. Taprooted annual or biennial; stems erect and simple to decumbent and much-branched, slender, 10 to 45 cm tall; herbage hispid to hirsute, glandular-pubescent in the upper portion. Leaves basal and cauline, basal leaves spatulate to oblanceolate, obtuse and usually mucronate, ca. 3.5 cm long, cauline leaves lanceolate to linear-oblong, 1.5 to 3(3.5) cm long, usually a little shorter than the basal leaves, to ca. 8 mm broad at the widest, bases clasping, ciliate, apices obtuse to acute or apiculate. Cymes reduced to racemelike inflorescences, flowers 1 per node, short-pedicelled in the axils of leafy bracts. Calyx tubular-fusiform at anthesis, becoming ovoid, to 1 cm long, 10-nerved, teeth lance-attenuate, ca. 1.5 to 3 mm long, overall hirsute or villous and with glandular hairs; petals white to pale pink, entire to crenate, 8 to 12 mm long, exceeding the calyx, often slightly twisted. Capsule ovoid, 6 to 8(10) mm long on a carpophore 1 to 2 mm long; seeds ca. 1 mm broad, blackish, the faces pitted or papillose. Roadsides, vacant lots, other open areas. E. 1/3 TX; N.S. and MI, SW. to MO and TX, S. to SC and FL. Native to Europe. Apr.-May. [S. dichotoma of TX authors, not Ehrh.; S. anglica L.].
2. S. antirrhina L. Sleepy Catch-fly. Taprooted annual; stems erect or ascending, simple or branched from the base, (0.5)1.5 to 8(9) dm tall, essentially glabrous or the lower portion retrorsely puberulent, internodes, at least the upper, with mucilaginous bands, the bands often dark, stems and leaves without glandular pubescence. Leaves basal and cauline, basal and lower cauline leaves spatulate to oblanceolate, generally 3 to 6 cm long, 2 to 15(20) mm broad, cauline leaves linear to lanceolate or narrowly oblanceolate, 2 to 5(7) cm long, 1 to 15 mm broad, acute, ciliate near the sessile base. Flowers few to many in an open to compact forked cyme, 1 flower and 2 or 3 branches at each node of the inflorescence; pedicels slender, 4 to 25 mm long, with small bracts. Calyx 4 to 10 mm long, tubular-fusiform at anthesis, becoming ovoid, often somewhat constricted at the mouth, glabrous, 10-nerved, teeth short-triangular, ca. 1.5 mm long or shorter, often purple-tipped; petals 5, fleeting or occasionally absent, white to pinkish or purplish, 5 to 10 mm long, shorter than to exceeding the calyx, somewhat emarginate, appendages very small or absent; styles 3, included. Capsule ovoid, about as long as calyx, 5 to 8(10) mm long on a carpophore ca. 1 mm long or less, 3-loculed except near the apex, 6-toothed; seeds 0.5 to 0.8 mm long, subreniform, brownish or gray-black, papillose. Open sandy or grassy areas, roadsides, open woods, and waste areas. Throughout TX; Que. W. to B.C., S. into Mex., except perhaps where very dry. Mar.-Sept., ours collected primarily in May. [Includes f. deaneana Fern., f. bicolor Farw., and f. apetala Farw.].
Low annual or perennial herbs. Stems often slender, sometimes 4-angled. Leaves opposite, estipulate. Flowers relatively small, in terminal or axillary bracteate cymes or solitary and axillary, terminal, or in the forks of the stem, hypogynous, perfect, pedicellate. Calyx of 5 free sepals. Petals 5, sometimes reduced or absent, white, bifid to deeply cleft or emarginate, sometimes so deeply cleft as to appear 10 in number. Stamens usually 10, often 5 to 10 or even fewer. Styles 3(4 to 5), separate. Fruit 1-celled, several- to many-seeded, usually dehiscent by twice as many valves as styles. Seeds smooth to tuberculate or rugose.
About 120 species worldwide; 3 in TX; the 1 here native to Eurasia. [Alsine (Tourn.) L.; Myosoton Moench].
Some species are cultivated for ornament; others are weedy. Some, including, ours, are edible (Mabberley 1987).
1. S. media (L.) Vill. Common Chickweed. Low, weak annual or winter annual, sometimes overwintering in sheltered places; stems 0.5 to 8 dm long, decumbent, trailing, or loosely matted, bent and rooting at the nodes, usually well-branched from the base, terete, pubescent in 1 or 2 longitudinal lines in the upper parts. Leaves sessile or the middle and upper leaves with ciliate petioles to about as long as the blade; blades ovate to obovate, elliptic, or subrotund, basally rounded to cordate, acute to short-acuminate, (0.4)1 to 3 cm long and 3 to 15(20) mm broad. Flowers numerous, axillary or 3 to 7 in terminal cymes; pedicels slender, 3 to 20 mm long, ascending, generally recurved in age. Calyx pilose, villous, or hirsute, often somewhat glandular, sepals ovate to oblong-lanceolate, 3.5 to 7 mm long, acute to obtuse, scarious-margined; petals shorter than the sepals, 2.5 to 3.5 mm long, cleft nearly to the base, occasionally absent; stamens 3 to 10; styles 3, placentation free-central. Capsule ovoid, generally pendulous, longer than the calyx, usually by 1 to 2 mm, valves 6; seeds roundish, ca. 1 to 1.2 mm long, tuberculate. Lawns, moist woods, waste places, etc. E. 1/3 of TX; native to Eurasia, widely naturalized in N. Amer., and widespread worldwide. Jan.-Mar. or Apr.; beginning in Dec. in mild years. Some sources recognize subspecies [Alsine media L.].
The leaves and stems are edible raw in salads or used as a potherb. The plants taste best before flowering.
Annual or short-lived perennial herbs, often low, if perennial often mat-forming or tufted. Stems slender or more or less wiry. Leaves opposite, estipulate, sessile or very shortly petiolate, the blades broad or linear to acicular. Flowers pedicellate, regular, hypogynous, few to many in loose or compact to capitate, terminal, cymose or cymose-racemose inflorescences, rarely solitary in the axils. Sepals 5, distinct or united very briefly below. Petals 5 (sometimes absent), generally white, apices entire to notched or bifid (but not as deeply cleft as those of Stellaria). Stamens 5 to usually 10, inserted with the petals on the rim of a poorly- to well-developed perigynous disk. Fruit an ovoid capsule dehiscent by 3 or 6 valves (as many as or twice as many as the styles), placentation free-central. Seeds globose to reniform or flattened, smooth to papillose or tuberculate, commonly reddish or yellowish.
About 150 species, primarily in the N. temperate zone; 10 species in TX; 3 here.
The genus is not of great economic importance. There are a few ornamentals and some are weedy (Mabberley 1987).
Many authors have moved to split the genus into several parts (e.g., McNeill, 1980) based on the number of valves, etc. This would put two of our species into Minuartia. However, as Wofford (1981) points out, this does not correlate with other diagnostic features such as seed morphology, nor has the complex yet been studied in its entirety on a world-wide basis. Until it is, it is perhaps best to retain the genus as a whole. Synonyms in Minuartia are provided for those who wish to recognize the split.
1. Capsules dehiscent by (4 to 5)6 valves or teeth; leaves ovate, 2 to 6 (8) mm long ................
......................................................................................................................1. A. serpyllifolia
1. Capsules dehiscent by 3 valves or teeth; leaves linear-filiform to lance-linear, to about 4 cm long .....................................................................................................................................2
2(1) Sepals ovate, apically more or less obtuse, not ribbed ..............................2. A. drummondii
2. Sepals lance-elliptic, apically acute, with 3 to 5 distinct ribs ................................3. A. patula
1. A. serpyllifolia L. Thyme-leaved Sandwort. Taprooted annual; stems 3 to 35 cm tall, tufted, erect to spreading or decumbent, simple to intricately forked, retrorsely puberulent, sometimes cinereous, often glandular in the upper portions. Leaves sessile or the lower very shortly petiolate, ovate, ascending, acute to acuminate, 2 to 6(8) mm long, 1 to 4 mm wide, often scabrous or minutely pustulate, margins ciliate, otherwise puberulent or glabrous. Flowers several to many in open, bracteate cymes; pedicels straight, 3 to 7(12) mm long. Sepals 2 to 4 mm long, lanceolate to lance-ovate, somewhat inconspicuously 3-to 5-ribbed, acute to acuminate, glabrous to minutely scabrous and with minutely scarious margins; petals ca. 1/2 to 3/4 as long as the sepals, 1.5 to 2.5 mm long, oblong or narrowly spatulate. Capsule dehiscing by 6 teeth or valves, ovoid-flask shaped or pyriform, equalling or longer than the sepals, 3 to 3.5 mm long, light greenish-yellow; seeds (0.6)0.7 to (0.8) mm long, globose-reniform, rugose or with concentric rows of low tubercles. Sandy or rocky soils of fields, roadsides, and open areas. E. and Cen. TX; naturalized from Eurasia in N. Amer. from FL to TX, N. to Que., MO, and TN. Mar.-July.
2. A. drummondii Shinners Drummond Sandwort. Taprooted annual; stems 5 to 20 cm tall, single or few from the base, erect to ascending, commonly forked above, mostly glandular pubescent. Leaves few to many per stem, linear to less commonly linear-oblong or linear-oblanceolate, mostly 10 to 35 mm long, 2 to 7 mm wide, obtuse to somewhat acute, the upper leaves smaller than the lower, all somewhat succulent, glabrous or nearly so. Inflorescences few- to many-flowered cymes, glandular pubescent; pedicels to ca. 25 mm long, spreading at anthesis and reflexed before anthesis and in fruit, flowers relatively large for the plant. Sepals ovate to elliptic, obtuse, 5 to 7 mm long, scarious-margined, glandular pubescent and with very faint nerves; petals white or sometimes greenish at the base, obcordate, the apices shallowly notched, 9 to 15 mm long, usually much longer than the calyx. Capsule ovoid, about as long as the calyx, 4 to 6 mm long, dehiscing by 3 valves, straw-colored or tan; seeds ca. 1 mm long, brownish, muriculate. Sandy soils of fields, roadsides, openings in woods, etc. E., S., and S. Cen. TX; also AR, OK, and LA. Feb.-June, ours mostly Mar.-Apr. [Minuartia drummondii (Shinners) McNeill; Stellaria nuttallii Pax; Alsinopsis nuttallii (T. & G.) Small].
This plant is a sand-indicator. It often grows in beautiful profusion with other sand-lovers such as Pointed Phlox (Phlox cuspidata).
3. A. patula Michx. Taprooted annual; stems 5 to 30 cm tall, few to many from the base, slender, usually diffusely branched (sometimes only sparingly), erect to decumbent, glabrous or sometimes with scattered glandular hairs. Leaves slightly fleshy, sessile, linear-filiform to linear-lanceolate, to 4 cm long and 3 mm broad, attenuate to acute, the shorter ones spreading-ascending, the long spreading or reflexed. Flowers commonly many in open, bracted cymes; pedicels divergent, 5 to 50 mm long, commonly glandular. Sepals (3)4 to 6(7) mm long, lanceolate-attenuate to elliptic, acute, strongly 3- to 5-ribbed, with very narrow scarious margins, sometimes somewhat glandular; petals white, spatulate, emarginate to obcordate, up to 3 times as long as the sepals, 5 to 8(9) mm long. Capsule greenish-yellow, oblong, shorter than to longer than the sepals, 3-valved, splitting to about the middle; seeds nearly round in outline, 0.5 to 0.9 mm long, dark brown, with low tubercles. Sand, clay, or gravel soils of fields, meadows, pastures, and on rock outcrops. Primarily in E. TX; TX to AL, N. to MN, IN, and VA. Mar.-May. [Minuartia patula (Michx.) Mattf.; Sabulina patula (Michx.) Small, including A. patula f. pitcheri (Nutt.) Steyerm. and f. meadia Steyerm.]
Two varieties occur in TX, both of which are likely here; ours seem to belong mostly to the latter.
var. patula. Plants generally less than 15 cm tall, leaves narrow, 1.5 mm broad or less; sepals prominently 5-ribbed; petals little if at all longer than the calyx; capsule generally shorter than the calyx; seeds 0.5 to 0.7 mm long. [Alsinopsis patula (Michx.) Small.].
var. robusta (Steyerm.) Maguire. Plants larger and more open than in var. patula, generally more than 15 cm tall; leaves broader, generally more than 1.5 mm broad; sepals 3-ribbed; corolla obviously longer than calyx; capsule usually longer than calyx; seeds 0.7 to 0.9 mm long. [Minuartia patula (Michx.) Mattf. var. robusta (Steyerm.) McNeill].
About 25 species native to the N. Hemis. We have the one species found in TX. As with many other Caryophyllaceous genera, some are weeds while some are cultivated for ornament (Mabberley 1987).
1. S. decumbens (Ell.) T. & G. Trailing Pearlwort. Low annual or winter annual; stems 2 to 15(17) cm tall, branched from the base, commonly tufted, erect, ascending, or decumbent and spreading, primarily glabrous or sometimes with a few glandular hairs on the upper portion, sometimes purple-tinged. Leaves estipulate, linear or linear-subulate, 3 to 15 mm long and to 1 mm wide, often spreading-recurved, aristate or mucronate, the bases of each pair connate to form a short tube around the stem. Flowers bracteate, usually many, axillary and in scorpioid cymes which appear racemose, perfect, hypogynous, 5- or 4-merous (sometimes both on the same plant); pedicels filiform, 3 to 25 mm long, glabrous or glandular-pubescent above, erect to ascending. Sepals oval to ovate or elliptic, obtuse, 1.4 to 2.5 mm long, margins minute and scarious or hyaline and sometimes with purple at the tip, nerves obscure; petals (0 or 1 to) as many as the sepals, white or scarious, about equalling the sepals or slightly longer, ovate to obovate or elliptic, entire , OR petals sometimes rudimentary or absent; stamens (3 to) 10, usually twice as many as the sepals; styles short, as many as the sepals and alternate with them. Capsule 2 to 3.5 mm long, ca. 1.5 times as long as the sepals, narrowly ovoid, valves 4 or 5(6), apically recurved after dehiscence, placentation free-central; seeds reddish- or yellowish-tan, obovoid-triangular or angled-ovoid, 0.25 to 0.3(0.4) mm long, with minute ridges or tubercles. Prefers sandy soil in our area--moist or dryish fields, along paths, in wood openings, disturbed areas, etc. FL to TX, N. to MA, VT, KY, ILL, MO, KS; with disjunct populations in AZ, Alta., and N. B. Feb.-June.
Some authors recognize varieties.
Low annual or perennial herbs, stems erect to spreading. Herbage usually pubescent, often glandular or viscid, occasionally nearly glabrous. Leaves opposite, sessile, subsessile, or clasping, estipulate, commonly pubescent on both surfaces. Flowers few to many per plant in terminal, bracteate, compact to open cymes, sometimes in glomerules or solitary in the axils, flowers perfect, regular, hypogynous, pedicellate. Calyx of (4)5 free sepals. Petals as many as the sepals, rarely absent, shorter than to longer than the sepals, white, apically 2-lobed or -cleft to some degree. Stamens 5(10). Styles as many as the sepals and opposite them, rarely fewer. Fruit a yellowish, 1-celled membranous capsule, more or less cylindrical, straight or curved, dehiscent by as many teeth as styles. Seeds many, obovoid-reniform or obovoid-angular, dorsally grooved, papillose or tuberculate, reddish, less than 1 mm long.
About 60 species, mostly in the temperate zones; 6 in TX; 2 found here.
The genus is not very important economically. Some species, including ours, can be weedy; a few are grown as ornamentals, especially in rock gardens (Mabberley 1987).
1. Sepals with long, eglandular hairs on the back which project beyond the tip of the sepal (use a lens); pedicels generally shorter than the capsules .......................1. C. glomeratum
1. Sepals with hairs becoming shorter towards the tip and not projecting beyond the tip; pedicels generally longer than the capsules ..........................................2. C. brachypodum
1. C. glomeratum Thuill. Annual or winter annual; stems simple or branched at the base, 5 to 30 cm long or tall, erect to ascending or decumbent, sometimes somewhat matted; herbage finely hirsute and glandular pubescent to viscid, especially in the upper parts. Leaves 5 to 30 mm long, 3 to 15 mm broad, lower leaves spatulate to obovate, becoming oval to ovate or ovate-elliptic upwards, midstem leaves commonly the largest, bases rounded to tapered and somewhat clasping, apices obtuse to rounded, both surfaces pubescent. Flowers in dense glomerules at the tips of dichotomous branches, the inflorescence becoming somewhat more open with age but the tips remaining dense; bracts of inflorescence herbaceous, hirsute and glandular pubescent; pedicels shorter than the calyx and usually shorter than the capsule, only rarely to 5 mm long. Sepals ovate-lanceolate, acute, 3 to 5 mm long, with minute scarious margins, hirsute and glandular pubescent, with non-glandular hairs at the tips which project beyond the tips, hairs sometimes slightly purplish and the tips of the sepals themselves occasionally purplish; petals shorter than to slightly longer than the sepals, ca. 3 to 5 mm long, oblong, deeply notched, the margins of each not overlapping the petal on either side, occasionally petals absent; stamens 10. Capsule yellow, slightly curved, 5 to 9 mm long; seeds light brown, muriculate. Fields, roadsides, disturbed and waste areas, and cleared woods. E. 1/3 of TX; naturalized from Europe from FL to TX and CA, N. to N.Eng., NY, IL, SD, and B.C. Feb.-May. [C. viscosum L.].
2. C. brachypodum (Engelm. ex Gray) Robins Shortstalk Chickweed. Taprooted annual 5 to 35 cm tall; stems single or sparingly branched from the base, erect to ascending, finely pubescent or puberulent, glandular pubescent, sometimes viscid, occasionally nearly glabrous. Leaves chiefly lanceolate to linear-oblong to oblanceolate, or the lower sometimes ovate to obovate or spatulate, 5 to 30 mm long, 2 to 8 mm wide, acute to obtuse. Flowers (1)few to many in usually open to somewhat compact dichotomously-branched cymes, when flowers many, usually clustered near the tips of the branches; bracts herbaceous, glandular-pubescent; pedicels 1/2 to 1/4 as long as the calyx at anthesis, to 3 times as long as the calyx in fruit, straight or slightly curved with age. Sepals 3 to 4.5 mm long, lanceolate or elliptic, acute, usually very narrowly scarious-margined, glandular puberulent (usually sparsely) to glabrate, dorsal surface with short hairs which do not project beyond the sepal tips; petals elliptic, to 6 mm long and 2 mm broad, slightly shorter than to twice as long as the sepals, apex with a notch ca. 1 mm deep, the two sub-apices acute, petals sometimes absent. Capsules about 2 to 3 times as long as the calyx, 6 to 12 mm long, cylindrical, straight or upcurved; seeds 0.4 to 0.7 mm long, golden-brown. Meadows, prairies, vacant areas, open woods, and on slopes. Primarily in Cen. TX but also S. and W. TX; Que. to N.T. S. to GA, TX, AZ, OR and Mex. and Cen. Amer. Feb.-Apr. [Includes var. compactum Robins; C. nutans Raf. var. brachypodum Engelm.].
Ours herbaceous, suffrutescent, or herbaceous vines; elsewhere also trees and shrubs. Stems erect to sprawling, nodes often swollen. Leaves alternate (rarely opposite or whorled), simple, entire, or occasionally toothed or lobed, usually petiolate. Stipules present and usually obvious, commonly forming a scarious sheath (ocrea) around the node, this sheath entire to fringed or lobed, sometimes apparently absent as in Brunnichia and Eriogonum. Flowers sessile or pedicellate, pedicels sometimes jointed and often emerging from sheathing bracts (ocreolae), apparently primitively 3-merous, usually appearing 2-, 3-, or 5-merous, usually relatively small, hypogynous, perfect or sometimes unisexual (as in Rumex), if unisexual, plants monoecious or dioecious. Perianth not differentiated into calyx and corolla, tepals 4 to 6, in 2 similar to somewhat-dissimilar series or in 1 whorl of 5, herbaceous to petaloid, nearly entirely free to distinctly connate below into a short floral tube, 1 or both series sometimes enlarging or becoming more conspicuous in fruit or
developing keels, wings, or tubercles. Stamens 2 to 9, usually 1, 2, or several in front of each tepal (or in front of the outer series of tepals only), rarely more; filaments distinct or connate at base, often of 2 lengths, sometimes dilated basally. Nectary disk sometimes present at base of ovary or several nectaries present between the stamen bases. Gynoecium (2-)3-(4-) carpellate, unilocular (occasionally with vestigial partitions), styles free or united at base; ovule 1. Fruit an achene or nutlet, usually trigonous or lenticular, enclosed or exserted from the persistent perianth (and sometimes persistent stamens.) Seed with peripheral embryo and well-developed endosperm. Plants producing anthocyanins rather than betalain pigments.
About 30 to 50 genera, depending upon interpretation, ca. 1,000 species, primarily of the N. temperate regions; 6 genera and 57 species in TX; 4 genera and 25 species in our area.
The family is economically important chiefly for buckwheat (Fagopyrum esculentum) and rhubarb (Rheum). Others, including members of Cocoloba, Rumex, and Polygonum, are edible. Many members of the family are weeds--especially Polygonum and Rumex. Some of the woody species are used for timber or charcoal. There are a few ornamentals, including Antigonon (Queens' wreath) and Polygonum (Fleece vine, Bistort, etc.) A few are dye plants or sources of tannins (Mabberley 1987).
1. Plants tendril-bearing vines ................................................................................1. Brunnichia
1. Plants herbs; if viny, then without tendrils ................................................................................2
2(1) Flowers or groups of flowers within involucres of bracts; ocreae absent; stamens 9 .............
2. Flowers not in involucres (bracts, if present, borne singly); ocreae present; stamens 4 to 8. .
3. Tepals similar, in 1 or 2 whorls; flowers not in verticils; stigmas not tufted .....4. Polygonum
NOTE: Antigonon leptopus H. & A., Queens' wreath, is an ornamental vine with bright pink flowers. It is commonly planted in E. TX and occasionally volunteers, but the author is unaware of any persisting populations in our area.
A monotypic genus native to N. America.
1. B. ovata (Walt.) Shinners Perennial herbaceous vine, stems usually at least partly woody, climbing by tendrils terminating short lateral branches. Leaves alternate, entire, ovate to ovate-lanceolate, basally cordate to truncate or rounded, apically acute, acuminate, or abruptly short-acuminate, glabrous on both surfaces, 3 to 15 cm long, petiolate. Ocreae absent. Flowers in terminal spiciform racemes grouped into paniculate inflorescences. Flowers perfect, greenish or whitish, perianth of 5 basally connate tepals; stamens 8; pistil tricarpellate. Achenes trigonous, to ca. 6 mm long, enclosed in the persistent calyx, the calyx somewhat leathery, enlarging to become pinkish and showy, subtended by a winged pedicel-like base about 1.5 cm long and 5 mm broad, calyx and pedicel together to ca. 3 cm long. Wood edges and openings, especially near ponds, lakes, or streams. E TX; Gulf states N. to SC, MO, and TN. Flowering spring-summer; collected with fruit summer to fall. [B. cirrhosa Banks ex Gaertn. This name was published at nearly the same time as B. ovata and it is not clear which has priority.].
Annual, biennial, or perennial herbs, or some (but not ours) subshrubs or shrubs. Stems erect, sometimes tufted; herbage glabrous to tomentose. Leaves cauline and/or basal, alternate, entire, without ocreae. Inflorescence often borne well above the leaves, simple or more commonly much-branched, racemose, cymose, umbellate, or capitate, flowers in groups of 1 to several subtended by an involucre of (4)5 to 10 scale-like, foliaceous, or membranous bracts, involucres toothed or calyx like, tubular to campanulate, solitary or variously grouped, sessile to pedunculate. Flowers perfect or unisexual, pedicellate, the joint in the pedicel just below the rather persistent perianth. Perianth white to pink, red, purple, or yellow (often changing color after anthesis), ovoid to obovoid or oblong, glabrous to pilose or tomentose, composed of 6 tepals in 2 series of 3, free or briefly united basally and with the base very slender and pedicel- or stipe-like, or else only narrowed. Stamens 9, filaments pilose at the base. Ovary 3-carpellate, styles 3. Achene trigonous (or lenticular), in some species winged, glabrous to tomentose.
About 250 species, mostly in W. N. America; 19 in TX; 3 in our area. This treatment follows Reveal (1968).
A few (but not ours) have edible leaves or roots or are useful as ornamentals. E. ovalifolium is said to have been useful as an indicator plant for silver-bearing earth (Mabberley 1987).
1. Plants perennial from a thick, woody caudex; at least some leaves more than 7 cm long; perianth pubescent externally; achenes .......................................................1. E. longifolium
1. Plans annual or biennial, taprooted; leaves less than 7 cm long; perianth glabrous externally; achenes glabrous ...................................................................................................2
2(1) Involucres 2 to 2.5 mm long, glabrous to sparsely tomentose; outer perianth segments oblong-cordate ..............................................................................................2. E. multiflorum
2. Involucres 2.5 to 4 mm long, densely tomentose; outer perianth segments obovate .............
.............................................................................................................................3. E. annuum
1. E. longifolium Nutt. Longleaf Wild Buckwheat. Erect perennial herb to 1 to 2 m tall, stems pubescent to glabrous. Basal leaves petiolate, with the petiole base expanded or somewhat winged at the base; blades oblong to lanceolate or oblanceolate, (5)10 to 20 cm long, the largest up to 3 cm broad, upper surface glabrous to sparsely pubescent above, thinly to densely whitish/grayish tomentose beneath, cauline leaves similar to basal, but sessile and reduced in size, those near the inflorescence less than 1 cm long, sometimes revolute. Inflorescence a paniculate, dichotomously-branching cyme, open and with few branches, overall about 1/3 to 1/2 the height of the plant; involucres sessile or with peduncles to 3 cm long, turbinate to campanulate, 4 to 6 mm long, 2.5 to 4 mm broad, silvery pubescent to tomentose externally; pedicels 5 to 10 mm long. Perianth narrow and stipe-like below, densely silvery-white tomentose outside, glabrous and yellow within at anthesis, tepals alike, lanceolate to oblong-lanceolate in fruit, becoming narrower and the perianth overall (including stipe) becoming 5 to 8 mm long. Achenes 4 to 6 mm long, densely white-tomentose. Sandy or gravelly soils, commonly at the edges of pine or oak woods. May-July (Oct.?).
In TX, varieties may be distinguished only with difficulty. If varieties are recognized, ours as described above are probably var. longifolium [Incl. var. plantagineum Engelm & Gray and E. vespinum Shinners] found in E. TX, MO, N. AR, E. OK, and W. LA. This variety is combined by some with var. lindheimeri Gand. of W. and Cen. TX, the plants differing only in characters of pubescence and branching, leaf size, and habitat. [Synonyms for var. lindheimeri: E. coriaceum Coult. & Fish; E. texanum Scheele; E. longifolium Nutt. subsp. diffusum S. Stokes]. Other varieties occurring outside TX are also sometimes distinguished.
2. E. multiflorum Benth. Heart-sepal Wild Buckwheat. Annual (biennial) herb from a (sometimes very long) taproot; stems 4 to 20 dm tall, erect, whitish- to brownish floccose or tomentose almost throughout or the older parts glabrate with age. Leaves largely on the lower 3/4 of the plant, becoming smaller upwards, sessile to very short-petiolate, lanceolate to oblanceolate or elliptic, to 4(5.5) cm long, 4 to 15 mm broad, densely tomentose beneath, loosely floccose above, margins entire to undulate and or revolute. Inflorescence generally less than 1/3 or 1/4 the height of the plant, relatively compact, cymose and sometimes rather flat-topped; involucres turbinate, 2 to 2.5 mm long and about as broad, generally 5-toothed, glabrous to sparsely pubescent or tomentose; peduncles 0 to ca. 4 mm long. Perianth segments white, maturing to tan or cream with a broad rose-brown to brownish midrib, 1.5 to 2 mm long, externally glabrous, sparsely whitish-tomentose within, segments dissimilar: outer series oblong-cordate, inner series oblong to lanceolate. Achenes 1.5 to 2 mm long, brown and glabrous. Sandy or gravelly soils of oak and pine woods, roadsides, occasionally open areas. Cen. and E. TX; also SE. OK, LA, NE. Mex. Sept.-Nov.
The inflorescences can be lace-like. They are pretty in pressed arrangements and might be nice in the garden as well.
3. E. annuum Nutt. Herbaceous annual or biennial from an often-stout taproot; stems erect, solitary to few, simple or sparingly branched, 5 to 20 dm tall, floccose to whitish- or silvery-tomentose almost throughout or the older portions glabrate. Basal leaves sparse, oblanceolate, 2 to 5 cm long but often withering and commonly absent at anthesis, remaining larger leaves generally on the lower 3/4 of the stem, upper leaves few and smaller or absent, all oblong to oblanceolate, the largest to 7 cm long and 3 to 15 mm wide, densely tomentose below and sparsely pubescent above, sometimes revolute, with petioles 5 mm long or less. Inflorescence relatively loose and open, terminal, or occasionally smaller inflorescences present on lateral branches, usually cymose, di- or tri-chotomously branched, the branches ascending and sometimes nearly racemose or helicoid, inflorescence occasionally compact-cymose or umbellate; bracts minute; involucres sessile or on peduncles 5 mm long or less, turbinate-campanulate, 2.5 to 4 mm long, 2 to 3 mm broad, apically shallowly toothed, densely tomentose. Perianth white to rose, becoming dark reddish-brown at maturity, 1 to 2 mm long, glabrous externally and densely long-pubescent internally, the segments connate 1/2 their length basally, members of the outer series obovate, broader than the inner, narrowly obovate to oblong ones. Achenes 1.5 to 2 mm long, brown, glabrous but sometimes tangled in the hairs of the perianth and appearing pubescent. Sandy or gravelly soils of open grasslands or disturbed fields, also along roadsides. Throughout TX except on the plains along the coast; ID to W. NE, S. to TX and NE. Mex. Apr.-Nov., our collections primarily made in late summer and fall. [E. lindheimeri Scheele; E. cymosum Benth.].
Annual or perennial herbs from fibrous roots, taproots, or rhizomes. Stems usually erect, leafy, generally glabrous, often reddish-tinged. Leaves basal and/or cauline, alternate, entire to shallowly toothed, flat to undulate and/or crisped. Ocreae brownish and conspicuous, thin, papery or brittle, often disintegrating into fibers. Plants monoecious, dioecious, or with perfect flowers. Inflorescence terminal, of verticils in racemes, arranged in a panicle, often dense in fruit, bracts small or absent. Individual flowers small, pink to green at anthesis, pink to tan or red-brown in fruit, pedicels jointed to the peduncles. Perianth parts 6 in 2 series of 3, connate below, the 2 whorls more or less similar at anthesis, the inner 3 usually enlarging (and then called valves) and developing entire to spiny-toothed wings, wings membranous and usually reticulate, in fruit much larger than at anthesis, commonly enclosing or coherent to the achene, one or all of them developing a grain-like median tubercle, 3 outer perianth parts smaller, usually lanceolate to subulate and somewhat spreading to arcuate. Stamens 6. Styles and stigmas 3, stigmas peltate and tufted or feathery. Achene trigonous.
About 200 spp. of the N. temperate zone; 15 in TX; 6 here. Keying usually requires mature calyx and as much of the stem as possible.
The sap of many species is acid. Some, especially R. hymenosepalus (Canaigre), have been used to tan leather; others are dye sources, producing mostly tans, browns, and golds. The leaves of many, including R. acetosa, R. alpinus, and our own R. hastatulus, have edible leaves or petioles. Many are weedy, the seeds of some being very long lived. A few have medicinal uses, for example, as treatments for nettle-stings or poison ivy. The achenes of some are eaten by birds (Mabberley 1987).
1. Plants dioecious or polygamous; leaves (at least some) hastately lobed ...1. R. hastatulus
1. Plants monoecious or with perfect flowers; leaves not hastately lobed .................................2
2(1) Valve margins denticulate or spinulose at maturity ............................................2. R. pulcher
2. Valve margins entire or only minutely erose at maturity .........................................................3
3(2) Leaves flat, not undulate, entire or nearly so; stems branched or with well-developed axillary shoots below the inflorescence ...................................................................................4
3. Leaves undulate and/or crisped, sometimes denticulate; stems with or without branches or axillary shoots ............................................................................................................................5
4(3) Pedicels, including the parts above and below the joint, much longer than the mature valves, usually sharply reflexed at the immediate base and nearly straight thereafter ...........
......................................................................................................................3. R. verticillatus
4. Pedicels, including the parts above and below the joint, shorter than the mature valves, gently reflexed throughout ..............................................................................4. R. altissimus
5(3) Stem simple below the inflorescence; pedicel usually slender, obviously longer than the mature valves; panicle commonly leafy-bracted to about the middle; verticils usually crowded in flower and fruit; usually some leaves longer than 15 cm ...............5. R. crispus
5. Stem often with axillary shoots; pedicel shorter than or equalling the mature valves; panicle usually without leafy bracts; verticils commonly well-spaced in flower and fruit; leaves generally 12 cm long or less ........................................................6. R. chrysocarpus
NOTE: R. acetosella L. is reported from our area, but no collections have been seen by the author. Reports are usually based on misidentified specimens of R. hastatulus. The two are similar, but R. acetosella has valves equalling or shorter than the achenes, while the valves of R. hastatulus are usually longer than the achenes.
1. R. hastatulus Baldw. Heart-sorrel, Heartwing Sorrel. Short-lived perennial herb from a taproot; stems erect, slender, to maximum of 1 m tall, but usually much less, branched or unbranched. Leaves crowded near the base of the plant, pale green, basal leaves long-petiolate, 2 to 10 cm long and 3 to 18 mm broad, blades linear to oblanceolate to hastate, if hastate the middle lobe linear to oblanceolate and larger than the basal, divergent lobes, cauline leaves similar to basal but with shorter petioles or else narrowly lanceolate and sessile. Inflorescence 10 to 30 cm long, slender, with a few small bracts, branches spreading to ascending; pedicels slender, about as long as the valves at maturity, joint located below the middle but obscure. Flowers unisexual, plants dioecious; perianth pink, white, purple, yellowish, or red, valves 2 to 3 mm long and about as broad, basally cordate, surpassing the achene, without tubercles, margins erose, accrescent but not adherent to the achene, outer tepals reflexed in fruit. Achene 0.9 to 1 mm long, 0.6 to 0.9 mm broad, brownish. Sandy soils in open areas--pastures, vacant lots, roadsides, etc. E., SE., and N. Cen. TX; MA S. to FL and TX, N. in Mississippi valley to OK, KS, IL. Spring, our collections primarily Apr. [R. hastatulus Ell. or Ell. ex Baldw.; R. engelmanii Meisn.].
The leaves and young stems are good in salads and soups. They are high in oxalic acid, so it is undesirable to consume too much or too often. They are safe and delicious if eaten occasionally (Tull 1987). The flavor is best before the plant flowers.
2. R. pulcher L. Fiddle Dock. Taprooted perennial herb; stem usually simple, sometimes branched near the base, erect, ribbed, 40 to 80(85) cm tall. Lower leaf blades indented above the base, fiddle-shaped to oblong, basally cordate to truncate, apically obtuse, margins usually more or less undulate, sometimes also crisped, to ca. 20 cm long and 6 cm wide, petioles shorter than the blades, ower stem, lower petioles, and under surfaces of leaves commonly pubescent, especially on the veins, upper leaves smaller, becoming lance-oblong but still retaining the fiddle-shaped base, often less pubescent than the lower leaves. Ocreae obvious, thin, crumbly-brittle with age. Flowers in remote verticils in the axils of upper leafy bracts or the inflorescence paniculate with widely spreading, slender, sometimes tangled branches and widely spaced verticils, inflorescence often to 3/4 the height of the plant; pedicels shorter than the mature calyx or as long, stout, jointed near the base or middle, gently recurved. Valves ovate-deltoid, 4 to 6 mm long, 2.5 to 4.5 mm broad, strongly reticulate-veined, margins spinulose-dentate with subulate teeth to ca. 3 mm long, all 3 valves usually with tubercles, these equal or unequal in size, oval at maturity, golden-yellow, puffy-looking, rough or warty, outer tepals 2 to 3 mm long, not recurved. Achenes (2)3 to 4 mm long, trigonous-ovate, apically acute, dark brown and lustrous. Moist and disturbed areas such as ditches, stream banks, etc. E. 1/2 TX; NY to FL, W. to TX and MO, also CA, OR, Mex., and weedy on other continents; native to the Mediterranean area and widely naturalized in warm temperate areas. Spring, ours flowering primarily in Apr.
3. R. verticillatus L. Swamp Dock, Water Dock. Perennial herb from a stout root and, if growing in water, with fibrous roots from the lower nodes; stems erect or basally decumbent, usually single, sometimes with axillary shoots or branches below the inflorescence, 4 to 10(15) dm tall, striate, glabrous, often purplish. Leaves narrowly to broadly lanceolate or strongly lance-elliptic, tapered to both ends, glabrous, lower leaves to 40 cm long and 5 cm broad, upper smaller and more linear-lanceolate, margins of all flat and entire (only rarely slightly undulate); petioles shorter than the blades. Ocreae conspicuous, to 8 cm long, thin, brittle. Inflorescence an open panicle of ascending racemes, to ca. 40 cm long, branches each subtended by a leafy bracts but usually otherwise leafless, verticils of flowers and fruits usually crowded, sometimes well-spaced. Flowers unisexual, plants monoecious. Outer tepals not reflexed, obovate and similar to the inner tepals in male flowers, slender and oblanceolate in female flowers, valves of mature pistillate flowers 3 to 5.5 mm long, triangular-ovate, the apex rounded to elongate, basally truncate, strongly veined, margins entire or wavy, each valve with an obvious lanceolate to subulate tubercle about 2/3 the length of the valve and extending slightly below the lower margin of the valve, its surface papillate or wrinkled. Fruiting pedicels 2 to 5 times the length of the mature perianth, 10 to 15 mm long, sharply deflexed at ca. 1 mm from the base at the joint, straight thereafter. Achene strongly stipitate, trigonous-ovoid, dark brown, 2.8 to 3.1 mm long. Infrequent in low, moist areas such as wet meadows, swamps, floodplains, etc. SE. TX; Que. to NE, S. to FL and E. TX. Spring.
The leaves are edible if boiled in several changes of water (Mitchell and Dean 1978).
4. R. altissimus Wood Pale Dock, Tall Dock, Peach-leaved Dock. Perennial from a stout root; stems erect or procumbent at base, strongly ribbed, often branched below the inflorescence, 0.5 to 1 m tall, stem and petioles commonly reddish. Leaves lance-oblong to ovate-lanceolate, the lower to ca. 15 to 20 cm long and ca. 5 cm broad, petioles shorter than blades, upper leaves smaller, all leaves acute to acuminate, bases rounded or subcordate to acute, glabrous, margins entire to crenulate but flat and not undulate. Ocreae obvious, thin, lacerate. Inflorescence a panicle of ascending racemes, verticils usually crowded in fruit, ca. 5 to 35 cm long, generally without many leafy bracts; pedicels shorter than the mature perianth in fruit, jointed below the middle and gently recurved. Flowers imperfect, plants monoecious. Outer tepals not reflexed at maturity, valves greenish to reddish brown, broadly ovate, 4 to 5(6) mm long, 3 to 4(5) mm broad, tips acute to obtuse, bases truncate or cordate, reticulate-veined, margins entire, without tubercles or 1 valve with and 2 without (occasionally all 3 with), largest tubercles more than 1/2 the length of the valve, ovate-fusiform, puffy-looking, smooth or wrinkled. Achene ovate-trigonous, the angles sharp or slightly winged, ca. 3 mm long, 2 mm broad, short-acute apically, dark brown. Common in wet areas--flood-plains, wet meadows, ditches, etc. E. and N. Cen. TX, Plains Country, and parts of the Ed. Plat.; NH to MN, S. to CO, TX, AZ, and GA. Spring, fruiting in early summer. [R. ellipticus Greene].
5. R. crispus L. Curly Dock, Yellow Dock, Sour Dock. Coarse perennial from a large root; stems erect, to ca. 1.6 m tall, ribbed, glabrous, unbranched below the inflorescence and without well-developed axillary shoots. Leaves 10 to 30 cm long, to ca. 5 cm wide, blades oblong-lanceolate, apically acute, basally cuneate to subcordate or rounded, glabrous, margins decidedly undulate and crisped, upper leaves smaller and more obtuse basally; petioles well-developed but shorter than the blades, channeled or ribbed on the upper surface, sometimes pubescent. Ocreae thin, striate, becoming brittle and crumbly. Inflorescence an elongate panicle of slender, ascending-erect branches, to 60 cm long, flower whorls usually dense in fruit, bracteal leaves, if any, usually only in the lower 1/2; pedicels slender, 5 to 10 mm long, ca. 1.5 times the length of the mature valves, gently recurved, jointed below the middle. Valves widely ovate to more or less deltoid, 4 to 5 mm long and about as broad, apically obtuse to subacute, reticulate, margins entire to slightly erose, usually all 3 valves bearing tubercles, occasionally only 1 tuberculate or the tubercles unequal, largest tubercles ca. 1/2 the length of the valve or less, not extending below the margin of the valve, elliptic, plump, smooth to slightly wrinkled or pitted. Achene (1.5)2 to 2.5 m long, ovate-trigonous, apically acute, slightly stipitate, reddish-brown, smooth, shiny. Common in seasonally moist, usually disturbed areas--ditches, the margins of ponds, etc. N. Cen., E., and SE. TX; native to Eurasia and widely naturalized in temperate areas; throughout N. America where conditions allow. Spring.
See note at R. chrysocarpus, below. The leaves are edible boiled in 1 or 2 changes of water (Tull 1987).
6. R. chrysocarpus Moris Amamastla. Perennial from a creeping rootstock; stems erect or basally procumbent, usually no taller than 4 to 6 dm, often reddish, commonly with small lateral branches or axillary shoots below the inflorescence. Leaves linear-lanceolate to oblong-linear, usually only 5 to 12 cm long and 1 to 3 cm broad, 3.5 to 5 times longer than wide, upper leaves smaller than lower, apically acute to blunt, basally cuneate or subcordate, margins usually at least somewhat undulate and crisped; petioles well-developed but shorter than the blades. Ocreae to a few cm long, thin, becoming brittle and crumbly. Inflorescence a panicle of slender, ascending racemes, never with bracteal leaves within, verticils of flowers usually well-spaced but sometimes a little crowded in fruit, pedicels gently to strongly recurved, shorter than to about equalling the mature valves. Valves ovate-deltoid to rotund-deltoid, 3.5 to 5 mm long, 3 to 4 mm broad, apically acute to obtuse, reticulate, margins entire to minutely erose, each valve usually with a well-developed oblong tubercle, tubercles at maturity finely alveolate-reticulate. Achene ovate-trigonous, 2.5 to 3 mm long, apically acute, shiny dark brown. Low, seasonally wet places. SE. TX and Rio Grande Plains, less common N. to N. Cen. TX, Ed. Plat., and Trans Pecos; TX, LA, and panhandle FL; also Mex. Spring-summer. [R. berlandieri Meisn.; R. langloisii Small].
NOTE: Young plants of R. crispus and R. chrysocarpus are easily confused. The presence of axillary shoots or branches marks R. chrysocarpus. In fruit, the length of the pedicels and the presence or absence of bracteal leaves in the inflorescence are useful characters.
Annual or perennial herbs or tendril-less herbaceous vines from fibrous roots, taproots, or rhizomes. Nodes often swollen. Leaves alternate, simple, cauline and/or basal; stipules 2-lobed or more commonly tubular and sheathing the stem (ocreae). Ocreae often brittle and shattering or deteriorating into just fibrous veins, margin with or without a ring of bristles or cilia. Petiole present or absent, sometimes incorporated into the ocrea. Flowers pedicellate, in relatively inconspicuous axillary fascicles or else collectively showy in terminal or terminal and axillary spike- or raceme-like panicles. Ocreolae--small bracts--usually present subtending groups of flowers in the inflorescence. Flowers perfect (occasionally functionally unisexual), perianth white, pink, red, green, or some combination of these at anthesis, of undifferentiated tepals, 4- to 6-parted, the members connate below, in fruit the outer tepals often enclosing the inner, sometimes becoming enlarged or winged. Stamens 3 to 9, included or exserted, a nectariferous disk sometimes present near the base of the filaments. Styles 2 to 3, free or connate to some degree, included or exserted. Gynoecium unilocular and uniovulate. Fruit a lenticular or trigonous achene, smooth to variously textured, included or slightly exserted from the persistent perianth.
About 150 species worldwide, especially in the N. temperate region; 18 in TX; 15 here. Those non-viny species with rather showy flowers have been separated as Persicaria (Bauh.) P. Mill. In Texas material, the distinction is usually readily apparent, but the separation breaks down when variation across the entire genus is considered. Other species have at one time or another been treated under Reynoutria, Tovara, Bistorta, Tracaulon, Bilderdykia, and Tiniaria. This treatment is based, in part, on the work of McDonald (1980).
Various species have been used in herbal remedies. Native Americans are known to have eaten the young shoots of some. A few have ornamental uses, particularly Fleece Vine (P. aubertii). Many are troublesome weeds, especially in wet areas (Mabberley 1987; Mitchell & Dean 1978).
Positive identification usually requires a complete plant, including the mature perianth and achene.
1. Stems with numerous recurved prickles .......................................................1. P. sagittatum
1. Stems without prickles ..............................................................................................................2
2(1) Plants twining, trailing, or sprawling vines, usually with cordate or sagittate leaves ..............3
2. Plants erect to ascending or sprawling, but not vinelike; leaves not cordate or sagittate .....4
3(2) Perianth segments strongly winged in fruit .....................................................2. P. scandens
3. Perianth segments smooth or keeled, not winged in fruit .........................3. P. convolvulus
4(2) Styles 2, persistent, usually hook-like; inflorescence a slender, interrupted raceme; leaves ovate to obovate, more than 4 cm long ......................................................4. P. virginianum
4. Styles 2 or 3, deciduous; inflorescence various; leaves proportionately narrower, if ovate or obovate, then less than 4 cm long ...........................................................................................5
5(4) Largest leaves less than 4 cm long; leaves with a joint near the base of the blade; flowers axillary or in raceme-like inflorescences, usually not showy ...................................................6
5. Largest leaves more than 4 cm long; leaves without joints; flowers in spike- or raceme-like inflorescences, collectively conspicuous .................................................................................8
6(5) Leaves with 2 distinct longitudinal pleats, 1 on either side of the midrib ...............5. P. tenue
6. Leaves not pleated ....................................................................................................................7
7(6) Achenes equilaterally trigonous, sharp-angled, shiny, OR achenes greenish, lenticular, and conspicuously exserted; plants generally erect, slender; outer perianth segments often cucullate (hooded) at the apex at maturity ...................................6. P. ramosissimum
7. Achenes inequilaterally trigonous (1 side narrower or wider than the other 2), dull to mildly shiny; plants sprawling or branch ends erect (occasionally the whole plant erect); outer perianth segments not cucullate or only slightly so ........................................7. P. aviculare
8(5) Inflorescence(s) 1 (or 2) per main stem, terminal; plant a rhizomatous or stoloniferous perennial; achene lenticular, strongly biconvex ...........................................8. P. amphibium
8. Inflorescences 2 or usually more per main stem, terminal and axillary; plants perennial or annual; achene trigonous or lenticular .....................................................................................9
9(8) Ocrea without a ring of marginal bristles or with hairs less than 1.5 mm long, though perhaps pubescent on the surface (ocrea can deteriorate into fibrous veins that look like bristles--don't count these) .....................................................................................................10
9. Ocrea with a ring of marginal bristles longer than ca. 1.5 mm (look carefully--these are sometimes broken off as ocrea deteriorate) .........................................................................12
10(9) Peduncle with stalked glands, especially near the base (use a lens) ..9. P. pensylvanicum
10. Peduncle without stalked glands; glands, if present, sessile ................................................11
11(10) Sepals with conspicuous raised inverted-anchor shaped veins near the apex; larger inflorescences nodding; achene flat; plants annual ...............................10. P. lapathifolium
11. Sepals without conspicuous anchor-shaped veins; racemes usually erect; achenes biconvex; plants perennial .........................................................................11. P. densiflorum
12(9) Calyx with yellow to brown glandular dots, easily visible with a lens, especially in dried material .........................................................................................................12. P. punctatum
12. Calyx without glandular dots ...................................................................................................13
13(12) Taprooted annual; inflorescence less than 5(6) and usually less than 3 cm long, blunt and dense; bracts of the inflorescence eciliate or with a few short cilia; ocreal cilia usually less than 3 mm long; achene usually lenticular, sometimes trigonous ..............13. P. persicaria
13. Rhizomatous perennial; larger inflorescences usually more than 4 cm long, blunt or tapered and open; bracts of inflorescence ciliate; ocreal cilia more than 3 mm long; achene usually trigonous .......................................................................................................14
14(13) Ocreal cilia usually shorter than 6 mm; larger leaves less than 1.5 cm wide (or if that broad then ocreal cilia less than 5 mm long); stems usually less than 7 mm broad at the base; annual or perennial; hairs on ocrea often stiff and appressed .......................................
............................................................................................................14. P. hydropiperoides
14. Ocreal cilia usually longer than 6 mm; larger leaves wider than 1.5 cm; stems often more than 7 mm broad at the base; perennial; hairs on ocrea often loosely spreading ...........................
........................................................................................................................15. P. setaceum
1. P. sagittatum L. Tear-thumb, Arrowvine, Arrowleaf, Arrow-leaved Tearthumb. Stoloniferous or sub-rhizomatous annual (perhaps sometimes perennial?); stems simple or branched, to 2 m. long, at first erect, later trailing, twining, scandent, and/or vine-like, weak, divergently branched, 4-angled, channeled, armed with numerous short, sharp, retrorse prickles or barbs. Ocreae entire, oblique, split down one side, 0.5 to 1 cm long, eciliate. Petioles with the same retrorse barbs, petioles of the upper leaves usually less than half the length of the blades; blades sagittate, 1 to 12 cm long, 0.5 to 3 cm broad, the basal lobes rounded to subacute, apices acute, midribs barbed or scabrous below, margins sometimes scabrous beneath, lower surface sometimes lighter than the upper. Inflorescences erect, terminal and axillary, solitary or paired, consisting of clusters of ca. 3 to 12 flowers on slender, barbed or barbless peduncles longer than the leaves and bearing lanceolate bracts with hyaline margins; pedicels ca. 1.5 mm long and jointed just below the perianth. Flowers perfect, white or greenish and tinged with pink or red, subtended by small, chaff-like bracts; perianth parts 5, united below the middle; stamens (6 to)8, included; styles 3, connate below. Perianth in fruit enlarging to ca. 4 mm long; achene ovoid in overall outline, blackish brown, sharply trigonous with 3 subequal sides, 2 sides usually concave, apically acute and basally slightly stipitate, 3 to 3.5 mm long. Infrequent in wet places such as bogs, ditches, and the margins of lakes, marshes, etc. E. TX, but known in our area at least from Leon and Robertson Cos.; Sask. to Newf., S. to FL and TX; also Asia. June-Oct. [Tracaulon sagittatum (L.) Small].
The small barbs are capable of inflicting a lot of damage on skin and clothing.
2. P. scandens L. var. cristatum (Engelm. & Gray) Gleason Thicket Knotweed, False Buckwheat, Climbing False Buckwheat. Perennial vine, flowering the first year and often short-lived; stems climbing, to 5 m long or more, weakly angled to terete, glabrous to occasionally short-pubescent, twining at the tips. Ocreae 1 to 6 mm long, truncate to acute, papillose, eciliate, crumbling with age. Petioles slender, of variable length, from much shorter than to equalling the blades; blades cordate-deltoid to shallowly sagittate-deltoid, basally truncate to cordate, basal lobes rounded, apically acuminate or less often acute, to 12(14) cm long and 8(9) cm broad, reduced upwards, glabrous or else very short-pubescent on the veins below, the midvein above, and on the margins. Flowers greenish-white, arranged along the midstem in axillary clusters of 2 to 3 from small bract-like leaves with ocrea, on upper portions of the plant flowers single or grouped in axillary racemose inflorescences (1)3 to 20(28) cm long with a few reduced leaves below and bracts above; pedicels 4 to 8 mm long, slender, commonly reflexed, jointed near the flower; bracts of inflorescence scarious, sheathing. Flowers pendulous, perianth of 5 parts, free to below the middle, with 3 tepals winged and 2 unwinged, wings in this variety ca. 7 to 9(10) mm long at maturity (measured from the joint in the pedicel) and ca. 1 mm wide, undulate to crinkly, acuminate; stamens 8, included; styles 3, less than 0.5 mm long. Achene included in the perianth, 2.1 to 2.7(3.5) mm long at maturity, tapered to both ends, sharply trigonous, dark brown to black, lustrous and smooth. Wood edges, thickets, fencerows, wet areas. E. and N. Cen. TX (also perhaps Plains Country and occasional elsewhere, possibly adventive); ND E. to Que. and NY, S. to FL and TX; also Eurasia. Aug.-Oct. [P. cristatum Engelm. & Gray; Tiniaria cristata (Engelm. & Gray) Small; Bilderdykia cristata (Engelm. & Gray) Greene; Reynoutria scandens (L.) Shinners var. cristatum (Engelm. & Gray) Shinners.] Outside of TX there are other varieties, including var. dumetorum (L.) Gleason from Europe. There is some debate about the usefulness and circumscription of infraspecific taxa.
3. P. convolvulus L. Black Bindweed, Climbing or Wild Buckwheat, Ivy Bindweed, Nimble Will, Cornbind. Taprooted annual vine; stems to as much as 5 m long, but usually closer to 1 or 2 m, trailing or twining, branched, slightly grooved; herbage glabrous but minutely scabrous or scurfy. Ocreae 2 to 4 mm long, oblique, entire, eciliate, but with scattered, minute, hyaline hairs. Petioles each with a nectar gland at the base, slender, those of the larger leaves about equalling the blades; blades ovate-cordate to deltoid-sagittate, bases cordate, basal lobes acute to rounded, apices acuminate to acute, largest to 6 cm long and 5 cm broad, reduced upwards. Flowers in clusters of 2 to 3(6) at the nodes or, at the ends of the branches, in raceme-like inflorescences produced by shortened internodes and reduced bracts and leaves; pedicels 2 to 3 mm long, shorter than the perianth, jointed above the middle, erect or reflexed. Flowers greenish-white or sometimes blotched with purple; perianth of 5 segments free to below the middle, at maturity 3.5 to 5 mm long, greenish, the angles smooth or keeled but not winged; stamens 8, included. Achene tightly enclosed by the perianth, ovoid, acute at both ends, 3 to 4 mm long, black, dull to somewhat shiny, minutely granular-tuberculate. Usually in disturbed areas such as gardens and fields. Rare to infrequent throughout TX except the far W. Trans-Pecos; widespread in middle N. Amer. and Europe; adventive from Europe. Apr.-Sept. [Bilderdykia convolvulus (L.) Dum.; Tiniaria convolvulus (L.) Webb. & Moq.; Reynoutria convolvulus (L.) Shinners.]
This is a noxious weed in some areas, but not ours. A yellow dye is extractable from the plant. Early Europeans ground the fruits to produce a poor-quality, abrasive flour (Mitchell and Dean 1978).
4. P. virginianum L. Jump-seed. Perennial, rhizomatous herb; stems erect to reclining, single or sometimes several, 3 to 15 dm tall, usually unbranched, nodes only somewhat swollen. Ocreae truncate, with cilia or bristles to ca. 3 mm long. Leaves with relatively short petioles, petioles 10 to 20 mm long on the lower leaves, the upper leaves subsessile; blades ovate to ovate-elliptic or ovate-lanceolate, the largest to 16 cm long and 9 cm broad, upper leaves reduced and more lanceolate, bases acute, apices acuminate to acute, upper surface with scattered appressed hairs (strigose-pilose), lower surface scabrous-pilose or occasionally nearly glabrous, margins ciliate. Inflorescences 1 to several, terminal, elongate, to 50 cm long, loosely flowered below and more dense above, spike- or raceme-like in appearance, but actually panicles of bracteate fascicles, each bract subtending 2 or 3 flowers, bracts sheathing and ciliate; peduncles pubescent; pedicels ca. 3 mm long, jointed just beneath the flower, sometimes deflexed. Flowers greenish-white (occasionally pinkish); perianth with 4 parts free to below the middle, at maturity 3.6 to 4 mm long, inner tepals slightly longer than the outer; stamens 5, about as long as the perianth. Achene with 2 persistent styles, these separate to the base, stiff, springy, deflexed-hooked at the apex, 2 to 4 mm long. Achene lenticular, strongly biconvex, 3.4 to 4 mm long, ovoid-oblong, brown, included in the perianth or with the tip exserted. Local in rich woods, along streambeds, lake margins, etc. E., SE., and N. Cen. TX; MN and NE to Ont., Que., NH, and NY, S. to FL and TX; also Asia and parts of Mex. June-Oct. [Tovara virginiana (L.) Raf.; Antenoron virginianum (L.) Roberty & Vautier].
The persistent styles, if snagged properly, can spring back abruptly, propelling the achene up to several meters--hence the common name.
5. P. tenue Michx. Knotweed, Pleatleaf Knotweed, Slender Knotweed. Annual from a slender taproot; stems erect, simple or usually well-branched, 20 to 30(40) cm tall, stems and branches slender, 1 to 2 mm in diameter, wiry, more or less 4-angled above the base. Ocreae hyaline, 3 to 15 mm long, usually with 2 subulate lobes and often with other bristles or lacerations as well, nodes only slightly swollen. Leaves not crowded, base of blade jointed to petiole, petiole as long as ocrea and adnate to it, leaf blades therefore appearing sessile; blades linear to lance-linear, 5 to 40 mm long, 1 to 3(5) mm broad, firm, basally acute and apically acute to cuspidate, with 2 distinct longitudinal pleats, 1 on either side of the midrib, margins very slightly toothed or sometimes revolute. Flowers axillary, solitary or in bracteate groups of 2 to 3, on the upper part of the plant the leaves reduced and the inflorescence resembling an inter-rupted spike or raceme; pedicels very short, 1 to 1.5 mm long, flowers erect. Perianth segments 5, free nearly to the base, greenish to brownish with white or pinkish margins, the outer tepals cucullate (hooded) and longer than the obtuse-tipped inner tepals; perianth at maturity 2.6 to 4.2 mm long; stamens mostly 8, the inner with dilated filament bases, the outer adnate to the perianth. Achene enclosed by the persistent perianth, sharply trigonous-ovoid, with 3 concave sides, 2.5 to 4 mm long, apex acute and base with a minute stipe, dark brown or black, smooth or commonly dull and striate, especially on the apex and/or angles. Not very common in our area. Dryish open areas, slopes, and open areas in woods. E. 1/2 TX; WY E. to S. Can. and ME, S. to GA, OK, and TX. Aug.-Nov.
6. P. aviculare L. Knotweed, Prostrate Knotweed. Taprooted annual or rarely weak perennial; stems 1 or commonly several from the base, usually quite leafy and well-branched, main stem and branches ascending when growing among supporting vegetation, but more commonly prostrate or sprawling and often rooting at the nodes, sometimes the ends of the branches erect, stems smooth or slightly grooved, nodes only slightly swollen. Ocreae hyaline, 4 to 8 mm long, becoming lacerate or crumbling. Leaves blue-green to gray-green, blades jointed to very short petioles, elliptic to lanceolate or ovate-lanceolate, flat, not pleated, without conspicuous veins, apices acute, plants heterophyllous: early leaves 1.5 to 3(6) cm long, 4 to 15 mm broad, later and upper leaves 1/2 to 1/3 as large and often somewhat narrower, often deciduous late in the season, especially the larger, lower ones. Flowers in axillary fascicles of (1)3 to 6, pedicels filiform, shorter than and hidden by the ocreae so flowers always erect and scarcely exserted from the ocreae and bracts. Perianth of 5 tepals free to below the middle, green with white or pink margins, the 3 outer segments incurved (sometimes cucullate) and positioned over the angles of the achene, the inner 2 segments flat, over 2 faces of the achene; perianth at maturity 2.5 to 3.1 mm long, never opening; stamens (5 to 6) 8, included, filaments dilated basally. Achene usually included, 2.5 to 3.5 mm long, ovoid, apically acute, base very shortly stipitate, inequilaterally trigonous: in cross-section with 1 face broader or narrower than the other 2, 2 or 3 faces concave, dull brown, punctate or striate and usually not very shiny. Widespread in TX (except the extreme S.) but occurring in scattered localities; cosmopolitan in distribution, perhaps originally native to Europe or Eurasia. May-Nov.
This plant is part of a complex of closely-related and very similar species, including P. buxiforme Small and P. arenastrum Ford ex Bor., neither of which occurs here. However, these species may appear in the literature as subspecific taxa under P. aviculare.
The plant is a weed of fields and cultivated areas. The shoots are eaten by birds and livestock. The achenes have been used, cooked, as human food. Herbal lore gives the plant medicinal properties. It has been used to treat bleeding, kidney ailments, and malaria, and has been used as a diuretic (Mitchell and Dean 1978). Tull (1987) suggests using the leaves and fruits as a peppery spice and says that tan to yellow dyes can be obtained from the plant.
7. P. ramosissimum Michx. Knotweed. Annual from a taproot up to 1 cm thick; stem usually 1 from the base, with slender branches above, usually erect to ascending, striate, somewhat woody near the base, (2)3 to 12(20) dm tall, nodes slightly swollen. Ocreae 2-cleft to several-parted, brownish or hyaline, becoming lacerate and fibrous. Leaves with the short petiole jointed to the ocrea, yellowish- to bluish-green, sometimes somewhat glaucous, often falling early, lower, larger leaves lanceolate to linear, or else oblanceolate, (4)5 to 30 mm long, 1 to 5 mm broad, flat, without obvious nerves, apically obtuse to acuminate, upper, bracteal leaves smaller. Flowers solitary or in 2's or 3's at the nodes on the upper parts of branches, pedicels 1 to 3.5 mm long, at least some on the plant well-exserted from the ocreae and usually some reflexed or drooping. Perianth with 5 or 6 parts united at the base, (2)3 to 4 mm long, yellow- or brownish-green or with pinkish margins, inner tepals about as long as the outer, the outer cucullate (hooded) and the inner flat; perianth at maturity commonly slightly keeled or pouch-like at the base; stamens 3 to 6, the inner filaments dilated. Achenes dimorphic: either ovoid-trigonous with 3 concave sides, 2 to 3.5 mm long, brown, shiny, included in the persistent perianth OR lanceolate and lenticular, biconvex, 4 to 6.5 mm long, pale yellow-green, shiny, exserted from the perianth; both types can be found on one plant late in the season. Seasonally wet, low places, sometimes in saline or brackish habitats. Widespread in TX except the Rio Grand Plain, usually localized; B.C. and Newf. S. to CA, VA, TX. Summer-fall. [P. latum Small; P. leptocarpum Robins.].
A very variable species. Much-branched plants with unreduced upper leaves, obtuse to rounded leaf tips, and pedicels less than 2 mm long have been designated as var. prolificum Small [P. prolificum (Small) Robins.], while less-branched plants with reduced upper leaves, obtuse to acuminate leaf tips, and pedicels 2.5 to 3.5 mm long have been designated var. ramosissimum. P. exsertum Small has been described as a salt marsh plant similar to var. ramosissimum but with most of the achenes exserted. It is not clear at present how much of the variation in P. ramosissimum is genetic and how much is seasonal or due to environmental factors.
8. P. amphibium L. Water Smartweed. Rhizomatous perennial, variously aquatic, emergent, or amphibious; stems floating to decumbent, ascending, or erect, depending on conditions, to 2 m long, sometimes rooting at the nodes, nodes swollen. Ocreae and leaves variable (see below.) Inflorescence 1 per main stem (sometimes with a second smaller one), terminal, erect. Perianth 4 to 5 mm long, rose or pink; plants functionally dioecious: stamens 8, included or exserted, always of a different length than the 2 included or exserted styles, styles united to about the middle. Achenes lens-shaped, strongly biconvex, ca. 2 to 4 mm long, shiny and dark. Cosmopolitan.
This plant has an almost unlimited array of forms depending upon habitat. The extremes have been recognized under varietal names; both can be expected here. The first has two forms, and neither can be ruled out for our area.
var. stipulaceum Colem. Natant form: Leaves and stems floating, rooting at the nodes, stems to ca. 1 m long. Ocreae glabrous, oblique, opaque to silvery. Leaves with petioles 10 to 80 cm long, blades glabrous, shape varying from elliptic with obtuse to rounded tips to broadly cordate with acute tips, 2 to 15 cm long, 1 to 5 cm wide. Inflorescences spike-like racemes held above the water, cylindrical and dense, 1.5 to 3(4) cm long, 0.5 to 1.5 cm thick; peduncle shorter than to longer than the inflorescence, glabrous; bracts of inflorescence ovate, entire, pink, glabrous. Summer to fall. Land form: Plants sprawling on sand or muck, to ca. 1 m long, herbage pubescent to hirsute. Ocreae tubular, hirsute, with a flange or collar at the apex, margin ciliate. Leaves short-petiolate to sessile, elliptic, acute, bases tapered-cuneate to acute, sometimes narrowly cordate but never truncate, to 10 cm long and 3 cm wide. Peduncle rarely glandular pubescent if plants stranded away from water. Inflorescence similar to that of the natant form but produced less often. Summer-fall. Less common in our area than the next variety. [Persicaria mesochora Greene; P. nebraskensis Greene; P. psycrophila Greene; Polygonum natans Eat.; P. hartwegii Gray; P. fluitans Eat.].
var. emersum Michx. Swamp Smartweed. Stems erect or decumbent, to 1 m tall; stems and leaves glabrous (especially if rooted in water) to strigose (typical of dry areas), becoming rusty in age. Floating leaves and stems not produced; stipules tubular, glabrous to strigose, without a collar, more or less truncate, becoming brown and shattering. Petioles 1 to 3(7) cm long; upper leaves usually the same size as the lower, ovate-lanceolate to oblong-lanceolate, acuminate, basally acute, truncate, or cordate, to 25 cm long and 6 cm broad, margins often undulate. Inflorescences erect, 2 to 15 cm long, commonly tapering, peduncles glandular-pubescent to strigose; inflorescence bracts ovate, strigose to ciliate, overlapping. Plants of ponds, marshes, streams, lakes, ditches or (usually seasonally) moist upland sites. Summer-fall. Apparently more common in our area than the preceding variety. [Persicaria coccinea (Muhl.) Greene; P. iowense Rydb.; P. mesochora Greene; P. pratincola Greene; P. rigidula (Sheld.) Greene; P. vestita Greene; Polygonum coccineum Muhl.; P. muhlenbergii (Meisn.) Small].
9. P. pensylvanicum L. Pennsylvania Smartweed, Pinkweed, Smartweed. Taprooted annual; stems erect to ascending, usually well-branched, 1 to 23 dm tall, glabrous or sometimes strigose towards the apex, sometimes glaucous, often cherry-red, at least at the swollen nodes. Ocreae longer than broad, ca. 1 to 1.5(2) cm long, truncate, glabrous or sparsely pubescent, eciliate or sometimes with cilia less than 1 mm long, becoming lacerate with age, internodes hollow. Petioles to ca. 4 cm long, lower ones longer, jointed to the ocrea; blades broadly to narrowly lanceolate, to ca. 18 cm long and 4(5) cm broad, apices acute to acuminate, bases acute to obtuse or rounded, glabrous to slightly hispid, strigose, or glandular, visibly punctate, especially below, margins entire, glabrous to strigillose. Inflorescences numerous, terminal and lateral, thick, spike-like, 1 to 1.5 cm broad, 0.5 to 5 cm long (usually ca. 3 cm), blunt; peduncles (at least some on a plant) with stalked glands sometimes mixed with strigillose hairs, occasionally completely glabrous; bracts of inflorescence sheathing, overlapping, glabrous to sparingly glandular, acute, margin entire or with minute cilia, pedicels exserted. Flowers pale to deep pink or rose, rarely white; perianth of 5 parts united below the middle, in fruit 2.7 to 5(6) mm long, only rarely some on a plant glandular; stamens 6 to 8, attached to a glandular disk at the base of the ovary; styles 2 or 3, partially united below, about as long as the stamens in our area, usually stamens and style not exserted (some heterostylous populations are known.) Achene enclosed by the persistent perianth, ovoid-lenticular (rarely trigonous), ovate to subreniform in outline, flattened, 1 or both faces indented, without (or rarely with) humps in the indentions, 2.5 to 3.5 mm long, dark brown to black, minutely roughened but nonetheless shiny. Wet, disturbed areas: ditches, fields, roadsides, etc., sometimes weedy. Throughout TX (but absent from drier areas) and N. Amer. May-Nov.(Jan.) [Persicaria pensylvanica (L.) Small; P. omissa Greene; P. bicornis (Raf.) Nieuw.; Polygonum longistylum Small; P. bicorne Raf.].
The achenes are a source of food for gamebirds and have been planted by some gun clubs. Sometimes planted for erosion control. (Mitchell and Dean 1978).
10. P. lapathifolium L. Pale Smartweed, Willow-weed, Dock-leaved Smartweed. Taprooted annual; stems erect or erect-ascending, occasionally decumbent, well-branched, to ca. 2.5 m tall, nodes swollen, internodes hollow, glabrous. Ocreae tubular, orifice oblique, entire, eciliate, glabrous, strongly nerved, becoming brittle and lacerate. Petioles 0.3 to 3.5 cm long, glabrous to glandular-pubescent to scabrous or strigillose; blades usually lanceolate and acuminate, but varying from lance-ovate to oblong-ovate to nearly linear or even somewhat rhombic, 5 to 25(30) cm long, to ca. 5 cm wide, sometimes glandular punctate, glabrous to sparsely pubescent or even tomentose (in aquatic forms), commonly strigose-scabrous along margin and on veins beneath. Inflorescences numerous, terminal and lateral, nodding or drooping (sometimes erect on small specimens,) dense-cylindrical, 0.5 to 1.1 cm wide but usually less than 1 cm, ca. 3 to ca. 8 cm long; peduncles glabrous to sparingly appressed pubescent, sometimes glandular, the glands sessile or subsessile; bracts oblique, scarcely sheathing, glabrous or with a few tiny bristles. Perianth 2 to 3(4) mm long, constricted above the achene, white to pinkish white or greenish with a silvery sheen, with 4 or 5 parts united below the middle, each part usually with veins branched near the apex and recurved into an inverted anchor-shape; stamens usually 6, included; styles 2, free nearly to the base. Achene included in calyx, oval, lenticular, flattened, both faces plane or 1 or both faces concave, 1.8 to 2.8 mm long, shiny dark brown to black. Wet, disturbed areas--ditches, fields, etc. Throughout TX and N. Amer.; introduced from Eurasia and perhaps also native; circumboreal. Apr.-Dec. [Persicaria lapathifolia (L.) Small; P. tomentosa (Schrank) Bickn.; Polygonum tomentosum Gray; P. scabrum Moench; P. incarnatum Ell.; P. nodosum Pers.].
This plant can be weedy, but the fruits are of some use to waterfowl as food. (Mitchell and Dean 1978).
11. P. densiflorum Meisn. Rhizomatous perennial; stems erect, to ca. 2 m tall, often 7 mm or more wide at the base, well-branched, herbage usually glabrous throughout. Ocreae tubular, longer than wide, truncate or slightly oblique, eciliate or sometimes ciliate, otherwise glabrous, brittle, becoming lacerate or crumbling. Petioles attached to base or middle of ocrea, to ca. 2.5 cm long; blades lanceolate, 5 to 25 cm long, 2 to 5 cm broad, bases obtuse to acuminate, apices acuminate, mostly glabrous, punctate. Inflores-cences numerous, terminal and lateral, erect or slightly curved, usually dense, occasionally appearing compound, to ca. 6 cm long, terminal portion often quite pointed when in bud, more blunt at anthesis; peduncles glabrous or occasionally with essentially sessile glands or pubescent; bracts of inflorescence ovate, from scarcely to definitely sheathing, glabrous or slightly pubescent or glandular. Perianth with 5 parts united below the middle, 2.3 to 3.2(4) mm long, whitish to whitish-pink or pink, margins of tepals lighter on dried specimens, eglandular or often lightly glandular; styles 2. Achene included in the persistent calyx, lenticular, plumply biconvex, oval in cross-section, 1.9-2.5(3) mm long, shortly beaked at the apex, shiny dark brown to black. Wet areas, often growing in water. In TX on the Coastal Plain and N. to our area; throughout E. U.S. on the Coastal Plain and Piedmont and southward, N. to MO and W. VA. (Apr.) Jun.-Nov. [Persicaria densiflora (Meisn.) Moldenke; P. portoricensis (Bertero) Small.].
12. P. punctatum Ell. Water Smartweed, Dotted Smartweed. Annual from a rootstock or perennial from stolons and rhizomes; stems decumbent at base, often rooting at the nodes, thereafter erect or ascending, to ca. 1.2 m tall, slender, simple to well-branched, green glabrous or short-pubescent. Ocreae tubular to obconic or flared in the upper half, 2 to 4 times longer than broad, to 1.8 cm long, glabrous or sparingly short pubescent, margin with cilia 5 to 15 mm long, often lost as the ocrea becomes brown and crumbles. Petioles short, to ca. 2 cm long, dilated where attached to the lower part of the ocrea; blades mostly lanceolate (occasionally long-elliptic or long-rhombic,) to 15 mm long and 2.4 cm broad, tapered and slightly decurrent on the petiole basally, apically acuminate or acute, somewhat reduced near the inflorescence, glabrous or short pubescent or minutely scabrous on the midvein and major lateral veins, usually scabrous-pubescent on or near the margins. Peduncles usually glabrous, or with sessile glands; inflorescences usually numerous, occasionally only few, erect or arching, slender, to 15(20) cm long, usually about 7 cm, 1 to 5 mm wide, flowers usually not very dense, giving the inflorescence an interrupted look; bracts not over-lapping, sheathing, truncate, ciliate-margined, body glabrous. Perianth of 5 parts free to below the middle, greenish near the base and the lobes creamy (rarely with a pink tinge,) 3 to 3.5 mm long, rather densely glandular-dotted on lobes and tube, glands yellowish when fresh, drying brown, easily seen at 10x; stamens 6 to 8, included; styles 2 or 3, united above the middle, included. Achene included in perianth, usually trigonous or sometimes plumply lenticular, oval in outline, 2.5 to 3 mm long, shiny dark brown to black. Wet or seasonally wet areas--ponds, ditches, swamps, moist fields, etc. Throughout TX, more common in E. TX; throughout temperate, subtropical, and tropical N., Cen., and S. Amer. [Persicaria punctata (Ell.) Small.].
Perennial plants with terminal inflorescences of uniform size and trigonous achenes are recognized by some as var. punctatum. Annual plants from taproots or slightly creeping stems, with very long branch-like inflorescences from the lower nodes and smaller, scattered inflorescences, and having lenticular achenes have been called var. confertiflorum (Meisn.) Fassett. There is some doubt as to the benefit of naming varieties, because numerous intermediate forms may be found. Many recent treatments do not recognize varieties.
13. P. persicaria L. Lady's Thumb, Moco de Guajolote, Heart's Ease. Taprooted annual; stems erect-ascending, simple to well-branched, to 1(1.5) m tall, glabrous or sparsely pubescent, green or occasionally with some reddish color. Ocreae loose-tubular, 1 to 2 cm long, strigillose to glabrous, sometimes striate, marginal cilia usually less than 3 mm long, ocreae becoming thin and brittle. Leaves short-petiolate to sessile, petioles to 12 mm long, glabrous; blades lanceolate to linear-lanceolate, variable in size, 3 to 15(20) cm long, 3 to 18(26) mm wide, tapered to base, apex acute-acuminate (rarely blunt or obtuse,) sparsely strigose (especially on the veins) or glabrous, often with purple spot or down-pointing crescent in the middle, this marking not always obvious in dried material. Peduncles usually glabrous, usually glabrous; racemes numerous, more or less erect, short, dense, 1 to 4.5(6) cm long, usually 3 cm or less, 7 to 12 mm thick, sometimes smaller inflorescences produced at the lower nodes; bracts of inflorescence overlapping, nearly truncate, strigose, with short, weak marginal cilia less than half as long as the bract or the margins entire. Perianth glandless or with just a few glands, with 5 parts free to below the middle, pink to greenish, purple, rose, or rarely white, 2 to 3.5 mm long, enclosing the achene at maturity; stamens usually 6, included or slightly exserted; styles 2(3), united at the base. Achene ovoid, lenticular or sometimes trigonous with flat or slightly concave sides, 2 to 3 mm long, shiny brown to black. Moist to wet areas, ditches, river and stream banks, alluvial bars, etc., throughout the state. Naturalized from Eurasia, now in Greenland and N. Amer. from AK to FL and Mex. Jun.-Dec. [Persicaria persicaria (L.) Small; P. maculosa (Schrank) Bickn.; Polygonum puritanorum Fern.; and, of American authors, P. dubium Stein and P. minus Huds.].
There is much variation in leaf shape, stature, percentage of trigonous achenes, and inflorescence length, but the variation seems to be random and most current treatments do not recognize varieties.
The plant is weedy, but the achenes provide food for wild fowl and are sometimes sown purposely. The leaves have a peppery taste and can be used in cooking. Yellow- to brown-gray dyes can be made from this plant (Mitchell and Dean 1978).
14. P. hydropiperoides Michx. Mild Water-pepper. Rhizomatous perennial to ca. 1 m tall; stems erect, or bases decumbent and rooting at the nodes, glabrous to strigose. Ocreae tubular, 1 to 3.5 cm long, at least twice as long as wide, scarious, strigose or scabrous, ocreal hairs appressed-ascending and with their bases flat and expanded, adnate to the ocrea, marginal bristles 3 to 8 mm long. Leaves short-petiolate to sessile, petioles 2 to 20 mm long, pubescence various; blades narrowly to broadly lanceolate, 3 to 25 cm long, 0.4 to 5 cm broad, bases acute to obtuse, apices acuminate, pubescence of stiff, short, appressed hairs, sometimes nearly glabrous or glabrous except near the margins, sometimes with bluish-green to yellow, scale-like glands underneath. Peduncles ca. 1 to 3 cm long, slender, glabrous to strigose or scabrous; inflorescences numerous, elongate, slender, 2 to 5 mm wide, 2 to 8 cm long, erect, usually interrupted near the base and never very dense; bracts perhaps contiguous but little if at all overlapping, pinkish-green, sheathing, with cilia less than half as long as the bract. Perianth oval to spherical in overall outline, of 5 parts united to just below the middle, rosy below, pink to cream at the lobe tips (rarely greenish, purplish, or white,) rarely with a few scale-like glands, 2.5 to 4 mm long, sometimes staminate flowers 1.5 to 2.5 mm long produced; stamens 8, included except in young staminate flowers; styles 3, partially united. Achene trigonous with sharp angles and flat to concave sides, 1.5 to 3 mm long, enclosed in the persistent perianth or with the tip slightly exserted. Wet or seasonally wet areas--shores, clearings, swamps, ditches, etc. Throughout TX; most of N. Amer.:N.S. to B.C. and southward. Jun.-Nov. [Persicaria hydropiperoides (Michx.) Small; Polygonum mite Pers. (not Schrank)]. See notes following:
NOTE #1. The trend in the literature seems to be to describe just one species. This plant has been treated in the past as 1 species with 2 varieties or as 2 separate species. P. hydropiperoides Michx. var. opelousanum (Ridd.) Stone or Polygonum opelousanum Ridd. ex Small was described as having narrower leaves to ca. 1 cm broad. These are the plants with glands on the leaves or spherical calyces; the inflorescences are more dense, the bracts slightly overlapping; achenes 1.5 to 2.5 mm long, slightly exserted from the calyx, staminate flowers not produced. [Persicaria opelousana (Ridd.) Small.].
NOTE #2. There seems to be some disagreement about the relationship of this species to P. setaceum (following.) At least one current treatment (GPFA 1986) combines the two under P. hydropiperoides (along with var. opelousanum or P. opelousanum as described in Note 1, above.) It is clear that further study of this complex is needed.
15. P. setaceum Baldw. Smartweed. Perennial, rather coarse, rhizomatous, sometimes stoloniferous if growing in water, often large at the base, to 7 mm or more in diameter, often decumbent at the base and rooting at the nodes, flowering stems erect, ascending to 2 m tall, greenish, slightly striate, glabrous to sparsely pubescent with spreading hairs. Ocreae tubular, 1 to 2 cm long, variously described as having long appressed hairs or minutely strigose or with loose, spreading hairs expanded at the base but not adnate to the tube, marginal bristles stiff, sometimes twisted, 6 to 22 mm long. Leaves short-petiolate or the upper ones sessile, petioles to ca. 5 mm long; blades lanceolate, the larger 6 to 10(24) cm long, up to 4.8(6) cm broad, generally 1.5 to 2.5 cm broad, basally cuneate, apically acuminate, both surfaces and margins strigose to hispid or with spreading hairs, but vestiture varying with degree of water contact, rarely nearly glabrous. Inflorescences elongate, spike-like, usually uninterrupted, tapered to the apex, 2 to 8 cm long, usually less than 4 cm, 3 to 6 mm wide; bracts contiguous or slightly overlapping, sheathing, with marginal cilia; peduncles strigose to glabrous, often 2 to several from one point on the axis. Perianth greenish white to white, creamy, or tan, occasionally rose-tinted but not wholly pink, of 5 parts 2 to 3.5 mm long, occasionally with a few glands but these faint and restricted to the perianth lobes, never on the tube; stamens 8, included; styles 3. Achene included in the perianth, trigonous, the faces flat, (1.5)2 to 2.5(3) mm long, short-beaked, shiny brown to black. Swamps, shores, wet woods and clearings, ditches, etc. E. part of TX and the Gulf Coast; Coastal plain and Piedmont in N. Amer.: S. NY and NJ to FL, W. to TX, AR, OK. Jun.-Oct. [Persicaria setacea (Baldw.) Small; Polygonum hydropiperoides Michx. var. setaceum (Baldw. ex Ell.) Gleason].
At least one current reference (GPFA 1986) submerges this species in P. hydropiperoides. Many TX specimens seem intermediate between the two. It may in fact be best to treat the two together. More study is needed.