The Aizoaceae and Molluginaceae of the southeastern United States

John R. Boetsch

Draft -- March 2000

This treatment includes original descriptions and keys to those taxa present in the southeastern United States which have been attributed by various authors to the Aizoaceae, sensu lato (see Bittrich and Hartmann 1988). For the purposes of cross-reference, we have incuded keys and descriptions for taxa which are presently attributed to other families (i.e., Gisekia pharnacioides L. and Geocarpon minimum Mackenzie). Included taxa are those documented for the Southeast (AL, AR, DE, FL, GA, KY, LA, MD, MS, NC, SC, TN, VA and WV), and for the neighboring states of NJ, PA, OH, IN, IL, MO, OK and TX. As a matter of convenience, we include all taxa reported for the region regardless of the age of the report. The descriptions and keys were prepared by examining specimens at NCU, DUKE, NCSC and US. Where relevant, synonymy is cited from multi-state and regional floras abutting or overlapping the southeastern states. These are symbolized as follows: S = Small (1933); F = Fernald (1950); G = Gleason 1952; R = Radford et al. (1968); M = McGregor et al. (1986); W = Wofford (1989); and C = Gleason & Cronquist (1991). The format presented follows that of the Vascular Flora of the Southeastern United States (Cronquist 1980; Isely 1990).

AIZOACEAE, sensu lato

Key to families and excluded taxa:

1 Stems glabrous, terete, ribbed with decurrent leaf tissue, branched and radiating from a central axis; gynoecium apocarpous, carpels 5; ovules solitary in each locule; fruit a thin-walled, papillose-hispid achene   i. Gisekia (Gisekiaceae)
1 Stems not as above; gynoecium syncarpous, carpels 2-16; ovules solitary to numerous in each locule; fruit a capsule or fleshy and indehiscent (if indehiscent, sometimes stony and nutlike).
2 Plants diminutive, glabrous; stems erect, less than 5 cm tall; leaves less than 6 mm long; locules 1, not septate, placentation free-central   ii. Geocarpon (Caryophyllaceae)
2 Plants of variable stature and vestiture; stems prostrate, ascending, erect or absent, if present either prostrate or greater than 5 cm tall; leaves mostly greater than 6 mm long (or if less than 6 mm long stems prostrate and mat forming); locules 1-16 (if unilocular, prominent septa present), placentation axile or parietal.
3 Plants generally fleshy or succulent; sepals united basally into a calyx tube; ovary position inferior, half-inferior or superior; stamens fused or distinct, filaments adnate to the calyx tube   I. Aizoaceae, sensu stricto
3 Plants not succulent or only slightly succulent; sepals distinct; ovary superior; stamens free, distinct   II. Molluginaceae


i. Gisekia pharnacioides L.* Annual herbs, somewhat succulent. Stems prostrate to ascending, radiating from a central axis, terete, glabrous, ribbed with decurrent leaf tissue, nodes swollen. Leaves simple, exstipulate, opposite or in whorls, narrowly lanceolate to oblong, margins entire, revolute; petioles with decurrent margins. Dichasia compact, axillary or terminal, on peduncles to 1 cm long. Flowers perfect, actinomorphic; sepals 5, free, distinct, margins hyaline; corolla absent; stamens 5, alternisepalous, filaments swollen at the base; gynoecium apocarpous, carpels 5, pistils superior, subglobose, angled, densely papillose-hispid; styles 1 per pistil, short, erect to spreading; ovules solitary, basal. Achenes thin-walled, angled, papillose-hispid, ca. 5 per flower. Seeds black, lustrous; placentation basal. The taxonomic placement of this species has been problematic, and further complicated by its superficial resemblance to Mollugo verticillata L. (Godfrey 1961; Bogle 1970). It has been variously placed within the Aizoaceae, sensu lato (Jeffrey 1961; Adamson 1961), within the Molluginaceae (Hofmann 1973; Hutchinson 1973), and most recently as a member of the Phytolaccaceae (Takhtajan 1980; Brown and Varadarajan 1985). Following Nakai (1942), Gilbert (1993) has reinstated the family-level distinction of Gisekiaceae, stating that the presence of betalains and the apocarpous gynoecium are sufficient to reject placement within the Molluginaceae, and that the cymose inflorescence is distinct from other (racemose) members of the Phytolaccaceae. Found in disturbed, sandy areas of peninsular FL!. Introduced from Old World tropics.  [specimen photo]

ii. Geocarpon minimum Mackenzie. Diminutive, glabrous winter annuals. Stems erect, branched, less than 5 cm tall. Leaves simple, opposite, basally connate, linear to ovate, less than 6 mm long X 2 mm wide. Flowers few to several, axillary or terminal, sessile. Flowers perfect, actinomorphic; sepals 5, united basally, lobes ovate, 1.5-2.0 mm long, 3-nerved; stamens 5, included, alternating with short, blunt staminodes, adnate to calyx tube; gynoecium syncarpous, carpels 3, stigmata 3, plumose, sessile and recurved between the three apical lobes of the ovary; ovary superior, ovoid-cyllyndric; locules 1. Capsules ca. 4 mm long X 2 mm broad, dehiscence valvular. Seeds numerous, papillose; placentation free-central. This species was originally assigned to the Aizoaceae by Mackenzie (1914). Palmer and Steyermark (1950) transferred this species into Caryophyllaceae based on a suite of morphological evidence, most notably on its unilocular ovary with free-central placentation. Bogle et al. (1971) found further evidence for this placement in the accumulation of anthocyanins rather than betalains. Spring. Endemic to alluvial glades and moist depressions on sandstone outcrops of MO!, AR! and LA. Federally listed. = F, G, Wilson (1932), under Aizoaceae.  [specimen photo]

AIZOACEAE, sensu stricto

Succulent herbs or subshrubs, annual or perennial. Stems repent, prostrate, ascending or erect. Leaves simple, cauline, alternate or opposite, somewhat to highly succulent; stipules present or absent. Inflorescence axillary or terminal, solitary or cymose. Flowers perfect, actinomorphic; perianth uniseriate or biseriate; sepals 4-5, fused below to a calyx tube, petaloid within (in uniseriate taxa), usually persistent in fruit, subapical dorsal mucros present or absent; petals present or absent, where present of staminoid origin and arising from hypanthium, lobes distinct or connate to a short corolla tube; stamens 1-numerous, fused to calyx tube or arising from the summit of hypanthium tissue; gynoecium syncarpous, carpels 2-16, ovary inferior, half-inferior or superior, locules 1-16, partly to completely septate, styles 0-16, linear, sometimes eccentric. Fruit a valvular, loculicidal or circumcissile capsule, or fleshy and indehiscent, sometimes stony. Seeds solitary to numerous in each locule; placentation axile or parietal.

Seven genera and ten species are documented for the Southeast.

Mesembryanthemum crystallinum L. is a South African species prized as an ornamental and naturalized in coastal areas of southern California. It is documented in eastern North America from a single collection from ballast in Philadelphia County, PA in 1879 (Rhoads and Klein 1993). Although we found no documentation of this species for the Southeast, we have included it in the following key given the potential for its presence as a rare introduction in our area. It can be readily distinguished by its fleshy, broad, flat leaves, by its fleshy, petal-bearing flowers, and by being thoroughly invested with large, crystalline papillae. Low, succulent herbs, prostrate or erect, perennial or annual, densely tuberculate with large epidermal protuberances. Leaves fleshy, opposite basally, becoming alternate apically, flattened, deltoid to obovate or broadly spatulate, lower leaves to 12 cm long X 7 cm broad, upper leaves to 4 cm long X 3 cm broad, crystalline-papillose, papillae obscuring the leaf surface, the leaves often appearing pitted when dry; margins undulate, apex acute, base broadly cuneate. Inflorescence solitary or dichasial, terminal or axillary near the summit of erect branches, on stout peduncles with reduced bracts. Flowers ca. 2 cm long X 1.5 mm broad; sepals 4-5, lobes fleshy, unequal; petals numerous, linear, spreading, united below to a short tube, white to cream colored; stamens numerous, arising from the corolla tube; carpels 5, ovary half-inferior, locules 5, styles 5, distinct. Fruit a capsule. Seeds numerous, brown or black, ca. 1 mm in diameter, folded near summit, minutely tuberculate; placentation axile. Native to s. Africa. = F.

Key to the genera of Aizoaceae, sensu stricto:

1 Perianth in two series, corolla present, petals numerous; sepals succulent, not petaloid within; ovary inferior or half-inferior.
2 Plants glabrous; leaves subterete, sharply 3-angled, erect or ascending; placentation parietal; petals not fused into a corolla tube; stamens not adnate to corolla   2. Carpobrotus.
2 Plants moderately to densely papillose or tuberculate; leaves flattened, not 3-angled, spreading; placentation axile; petals fused to a short corolla tube; stamens adnate to corolla tube.
3 Leaves deltoid, obovate or broadly spatulate, sessile, the margins undulate; plants completely covered with large, crystalline globules   x. Mesembryanthemum.
3 Leaves ovate-cordate, petiolate, margins not undulate; plants moderately to densely papillose, the papillae small and nodular   1. Aptenia.
1 Perianth uniseriate, corolla absent; sepals membranaceous or succulent, usually petaloid within; ovary superior or half-inferior.
4 Leaves alternate, flattened, blades ovate-triangular to ovate-rhombic; ovary half-inferior; flowers and fruits with 4-5 conspicuous horns near the summit; fruit fleshy, stony, indehiscent   6. Tetragonia.
4 Leaves opposite or alternate, terete or flattened, variously shaped; ovary superior; flowers and fruits lacking horns; fruit a capsule.
5 Leaves sessile, not clasping the stem; fruit a loculicidal capsule; ovules and seeds one per locule, pendulous   4. Galenia.
5 Leaves with petioles inflated toward the base, clasping the stem; fruit a circumcissile capsule; ovules and seeds few to numerous, not pendulous.
6 Stipules present; leaves distinctly petiolate, petioles often equaling the length of the blade; leaves alternate or opposite and unequal.
7 Plants forming dense, leafy mats to 30 cm across; leaves small, elliptic to obovate, 3-9 mm long; stamens 1-3; ovary and capsule subglobose, the apex rounded   3. Cypselea.
7 Plants sprawling to erect, not forming leafy mats; leaves various, mostly over 1 cm long; stamens 5-10; ovary and capsule cyllindric, the apex truncate or depressed with 1-2 prominent lobes at the summit   7. Trianthema.
6 Stipules absent; petioles much shorter than leaf blades; leaves opposite and subequal   5. Sesuvium.

1. APTENIA N.E. Brown

1. Aptenia cordifolia (L. f.) Schwantes.* Prostrate, succulent perennial herbs. Stems branched, suffruticose, densely papillose to glabrous. Leaves fleshy, opposite, subequal, flattened, ovate-cordate, densely papillose, to 3 cm long X 2.5 cm broad, apex acute, base cordate or rounded, petioles to ca. 1 cm long; stipules absent. Inflorescence fleshy, flowers solitary in axils, pedicels 5-10 mm long. Flowers fleshy; sepals 4, fused below, adnate to ovary, calyx tube 6-7 mm tall, lobes in unequal pairs, the larger pair often foliose, erect to spreading, 4-7 mm long; petals numerous, linear to oblong, united below into a short tube; stamens numerous, adnate to the corolla tube; carpels 4, ovary half-inferior, locules 4, stigmata 4, sessile. Fruit a fleshy, 4-valved capsule, ca. 1.5 cm long X 1 cm broad. Seeds compressed, tuberculate; placentation axile. Summer-fall. Disturbed areas in peninsular FL. Introduced from s. Africa.


1. Carpobrotus edulis (L.) N.E. Brown.* Succulent, glabrous, perennial herbs. Stems trailing, mat-forming, suffruticose, angled. Leaves fleshy, opposite, ascending to erect, subterete, sharply 3-angled, slightly connate at the base, 6-10 cm long, 6-12 mm broad, serrate on the lower edge; stipules absent. Inflorescence solitary at the apex of upright shoots; peduncles fleshy, 2.5-7 cm long. Flowers 8-10 cm in diameter; sepals 5, lobes fleshy, unequal; petals numerous, arising from the summit of the hypanthium, distinct, not forming a corolla tube, linear to oblong, white, pink or magenta; stamens numerous, distinct, antherless filaments (staminodes) present; carpels 10-16, ovary inferior, locules 10-16, stigmata 10-16, plumose, sessile, spreading; hypanthium fleshy. Fruit fleshy, indehiscent, edible. Seeds obovoid; placentation parietal. Summer-fall. Beachfront dunes, escaped from cultivation; FL. Introduced from s. Africa.

3. CYPSELEA Turpin

1. Cypselea humifusa Turpin.* Succulent, annual herbs. Stems succulent, procumbent to ascending, forming leafy mats, glabrous, tawny to orange in color. Root a slender, branching taproot. Leaves cauline, alternate or appearing opposite and unequal, fleshy, elliptic to obovate, 3-9 mm long, ca. 1-3 mm wide, glabrous, apex acute to rounded, margins entire to minutely serrulate, with a narrow band of transparent tissue at the margin, base cuneate, venation pinnate, netted; petioles to 6 mm, often exceeding the blade length, winged with decurrent blade tissue, inflated toward the base and clasping the stem; stipules present, petioles and stipules laciniate to long-fimbriate, with appendages to 1 mm or more long. Inflorescence axillary, flowers 1-3 in axillary cymules; pedicels to 1 mm long; bracts obscure, paired in axils, laciniate-fimbriate, margins scarious, ca. 1.5 mm long. Flowers urceolate, 2-3 mm long; sepals 5, fused below for about 1/5 of their length, lobes deltoid-triangular, erect, more or less imbricate, unequal, persistent, ca. 1.8 mm long X 1-1.6 mm wide, with 1-5 green nerves on the dorsal surface, lacking subapical dorsal mucros, margins hyaline, apex acute or obtuse; corolla absent; stamens 1-3, included, adnate to sepals, filaments ca. 0.8 mm long; carpels 2-3, ovary superior, subglobose, glabrous, locules 2-3 at anthesis (becoming unilocular in fruit by recession of the septa from the columella), styles 2-3, short, erect. Capsules thin-walled, circumcissile toward the base, ca. 2.5 mm long X 2-2.5 mm broad. Seeds numerous, 0.3-0.4 mm long, lustrous, reddish-brown, smooth; placentation axile (appearing free-central in fruit). Spring-summer. Sandy soils, depressions and pinelands; FL!, LA!. Introduced from West Indies. = S.  [specimen photo]


1. Galenia secunda (L. f.) Sonder.* Perennial herbs or subshrubs. Stems prostrate or erect, suffruticose, densely strigose or villose-hispid, becoming glabrate below, branching secund. Leaves somewhat fleshy, cauline, alternate or opposite, crowded in axils and on short branches, blades obovate to spatulate, to 2.0 cm long X ca. 1 cm wide, densely papillose with inflated epidermal cells, appearing pitted with ejected crystaline matter when dry; apex acute, obtuse or rounded, margins with a layer of translucent cells, ciliate with long, stiff, flat trichomes, base attenuate to cuneate, tapering to a broad petiole, venation not apparent; petioles to 1.5 mm long, ciliate with stiff trichomes; stipules absent. Inflorescence axillary, flowers solitary in axils of stem leaves, sessile. Sepals 4-5, fused below, lobes ovate-lanceolate, 2-2.5 mm long X 1.0 mm wide, glabrous and petaloid within, dorsal surface tawny-pubescent with a tuft of long hairs near the summit; corolla absent; stamens 8-10, paired, twice the number of calyx lobes, alternisepalous, fused to the calyx lobes near the base, filaments unequal, shorter than calyx lobes; carpels 2-5, ovary superior, angled, truncate or depressed apically, glabrous, locules 2-5, styles 2-5, distinct, unequal, to ca. 1 mm long. Fruit an angled, membranaceous, loculicidal capsule. Seeds oblong, one per locule; placentation axile, the seeds pendulous. Summer-fall. Disturbed areas and ballast; FL!. [NJ]. Introduced from s. Africa. = S, G.  [specimen photo]


Succulent herbs or subshrubs, annual or perennial. Stems prostrate, ascending or erect, branching from a short taproot, sometimes rooting at the nodes. Leaves fleshy, opposite, subequal, blades flattened to subterete, sessile or narrowed to a short petiole; petioles inflated and clasping the stem; stipules absent. Inflorescence terminal and appearing axillary, flowers solitary or few per axil, sessile or pedicellate. Flowers perfect, sepals 5, united basally into a short, subturbinate tube, connivent with the base of the ovary, petaloid within, lobes with well-developed subapical dorsal mucros; corolla absent; stamens 5 and distinct, or numerous and fascicled, adnate to the calyx tube; carpels 2-5, ovary superior, locules 2-5, styles 2-5, linear, persistent in fruit, stigmata longitudinal, papillate. Capsule membranaceous, circumcissile. Seeds numerous, round to reniform, arillate; placentation axile.

Four species are documented for the Southeast. An additional three species approach our area in Texas: S. erectum Correll, S. sessile Persoon and S. trianthemoides Correll. Of these, S. trianthemoides, is the is best circumscribed. It is endemic to wet depressions and coastal dunes along the southern Gulf coast of Texas, and differs from our taxa by the combination of a densely verrucose vestiture and having 5 stamens. Both S. sessile and S. erectum require further study to determine their validity and segregation from S. verrucosum Raf.; at present they are only separable by geography and coloration upon drying (Correll and Johnston 1970; Correll and Correll 1972). We include these taxa in our key to assist in their elucidation.

Key to the species of Sesuvium (adapted from Correll and Johnston 1970; Correll and Correll 1972):

1 Stamens 5, distinct; flowers sessile or on obscure pedicels to 2 mm long.
2 Plants glabrous; seeds smooth, ca. 0.7 mm long   2. S. maritimum.
2 Plants verrucose, with large crystalline globules; seeds rugulose, ca. 1.5 mm long   x. S. trianthemoides.
1 Stamens numerous, in fascicles; flowers sessile or pedicellate.
3 Plants predominantly glabrous, rooting at the nodes; flowers conspicuously pedicellate, pedicels 3-20 mm long   3. S. portulacastrum.
3 Plants moderately to densely verrucose or papillose, not rooting at the nodes; flowers sessile or with obscure pedicels, these rarely to 5 mm long.
4 Plants erect; leaves linear, 3-6 cm long, 10-20x as long as wide   1. S. crithmoides.
4 Plants prostrate to erect; leaves narrowly elliptic to obovate, less than 3 cm long, typically 3-10x as long as wide.
5 Plants drying light brown   4. S. verrucosum.
5 Plants drying blackish or dark grey.
6 Plants prostrate to decumbent, densely verrucose   x. S. sessile.
6 Plants erect or spreading, sparsely verrucose   x. S. erectum.

1. Sesuvium crithmoides Welwitsch.* Succulent herbs, minutely verrucose above, glabrate below. Roots not observed. Leaves opposite, linear, 3-6 cm long, to 3.5 mm wide (10-20x as long as wide), densely papillose with enlarged epidermal cells or appearing glabrous, smooth, margins entire, apex rounded to obtuse; petioles inflated and scarious at base, clasping the stem. Flowers solitary in axils, sessile, 7-10 mm long, densely verrucose; bracts ovate-triangular, acuminate, 3-3.5 mm long. Sepals 5, fused below, lobes ovate-triangular, imbricate, erect, 5-8 mm long, dorsal surface verrucose, margins hyaline-scarious, with hastate bulges at the junction with calyx tube; stamens numerous, adnate to the calyx tube; gynoecium not observed. Summer-fall. From a single collection series, "waste places among rosin wharves," in Brunswick, Georgia in August, 1902 (coll. R.M. Harper 1524; US 431830!). No further documentation of the status of this species in our area is available. Introduced from Africa. = S.  [specimen photo]

2. Sesuvium maritimum (Walter) B.S.P. Succulent, annual herbs. Stems glabrous, spreading, ascending or erect. Root a thickened taproot, often branching and fibrous. Leaves fleshy, cauline, opposite, flattened to cylindrical, obovate to oblanceolate, rarely linear, 10-35 mm long, 1-10 mm broad (3-10x as long as broad), glabrous or rarely minutely papillose, apex rounded, obtuse or acute, margins entire, base cuneate to rounded; petioles short, scarious, inflated at the base and clasping the stem. Flowers sessile or on obscure pedicels to ca. 1 mm long, 4-5.5 mm long, glabrous or sparingly papillose; bracts paired, ovate-triangular, irregularly toothed, ca. 1 mm long. Sepals 5, fused below to 1/3 the length of the flower, lobes erect to spreading, ovate-triangular, cucullate, imbricate, unequal, 2.5-4.0 mm long, petaloid within, margins hyaline, the dorsal surface green or reddened with 5-9 faint nerves, lobes with obvious subapical dorsal mucros, these divergent, to 1.2 mm long; stamens 5, distinct, alternisepalous, attached near the summit of calyx tube, 2-3 mm long, included; carpels 2-5, ovary ovoid, glabrous, locules 2-5, styles 2-5. Capsules scarcely exceeding the sepals at maturity. Seeds numerous, reniform, reddish-brown, smooth, lustrous, enclosed by a translucent aril, ca. 0.7 mm long. Spring-fall. Coastal dunes and beaches, salt flats and brackish marshes; AL!, DE, FL!, GA!, LA!, MD!, MS, NC!, SC!, VA!. [NJ!, OK, PA, TX]. Native. = S, F, G, R, C.  [specimen photo]

3. Sesuvium portulacastrum (L.) L. Succulent, perennial herbs. Stems thick, fleshy, repent to ascending, rooting profusely at nodes, glabrous, often reddened. Leaves fleshy, glabrous or rarely minutely papillose when young, elliptic to obovate or oblanceolate, to 45 mm long X 8 mm wide, 3-8x as long as wide; petioles inflated toward the base, scarious, clasping the stem. Flowers solitary in axils, conspicuously pedicellate, glabrous, 8-12 mm long; pedicels 3-20 mm long. Sepals 5, fused below to a short tube 1/4 to 1/3 the length of the flower, lobes erect, imbricate, unequal, cucullate, ovate or ovate triangular to somewhat hastate, 5-8 mm long, margins hyaline, petaloid, subapical dorsal mucros to 2.0 mm long; stamens numerous in fascicles, cream to pink or violet in color, included or slightly exserted, filaments flattened, fused at base, adnate to the summit of the calyx tube; carpels 3-5, ovary ovoid-conic, 6-7.5 mm long, glabrous, styles 3-5. Capsules ovoid-conic. Seeds numerous, reniform to pyriform, black, enclosed by a grey, translucent aril, ca. 1.1 mm long. Spring-fall. Beaches, bay margins, salt flats and wet sand or clay along the coast; AL!, FL!, GA!, LA!, MS!, NC!, SC!. [PA, TX!]. Native from the Carolinas to South America. = S, R.  [specimen photo 1]  [specimen photo 2]

4. Sesuvium verrucosum Raf. Succulent, perennial herbs or subshrubs. Stems prostrate, spreading or erect, suffruticose toward the base, densely verrucose with large, crystalline papillae. Roots fibrous, with a contorted taproot. Leaves fleshy, opposite, oblanceolate, obovate, spatulate or linear-elliptic, to 30 mm long X 10 mm wide, mostly 3-10x as long as wide, densely verrucose, apex rounded to acute, margins entire, base attenuate to narrowly cuneate; petioles short, inflated and scarious toward the base, clasping the stem. Flowers solitary or few in axils, sessile or pedicellate, 6-9 mm long, densely verrucose; pedicels to 2 mm long; bracts paired at base of pedicels, lanceolate to triangular, often irregularly toothed, 1.0-3.5 mm long. Sepals 5, fused below to a short tube ca. 1/5 the length of the flower, lobes erect to spreading, imbricate, unequal, cucullate, ovate-triangular, 4.5-7.0 mm long, petaloid within, margins hyaline, dorsal surface green or reddened, densely papillose-verrucose, with longitudinal nerves scarcely visible through warty papillae, subapical dorsal mucros obvious, ca. 1 mm long; stamens numerous, included, filaments fused at base, in fascicles of 5-8, adnate to the calyx tube; carpels 3, styles 3, 3-4 mm long, ovary conic to ovoid-cyllindric, glabrous, locules 3. Capsules membranaceous with longitudinal nerves, to ca. 5 mm, the circumcissile perforation in the lower fourth of the fruit. Seeds numerous, comma-shaped, smooth, black, enclosed by a dull, translucent aril, 0.7-1.0 mm long. Spring-summer. Wet depressions with saline or alkaline soils; native to south-central and southwestern N. America, occurring in our area in AR, LA!. [MO, OK!, TX!]. Native. = M.  [specimen photo]


1. Tetragonia tetragonioides (Pallas) Kuntze.* Succulent, annual herbs, densely papillose throughout. Stems prostrate to ascending, suffruticose toward the base. Leaves fleshy, cauline, alternate, flattened, ovate-triangular to ovate-rhombic, 2.5-7 cm long X 2-5 cm wide, densely papillose, apex acute to obtuse, margins entire, base truncate to attenuate to broad petioles; petioles to 2 cm long; stipules absent. Inflorescence axillary, flowers solitary or paired, to 15 mm long, prominently horned toward the summit, papillose; pedicels to ca. 10 mm long. Flowers fleshy; sepals 4-5, fused below and adnate to ovary for greater than 3/4 the length of the flower, lobes fleshy, inwardly curved, deltoid to narrowly triangular, to ca. 3 mm long, petaloid within; corolla absent; stamens 7-13, often in fascicles, adnate to calyx tube, to 1.3 mm; carpels 5-8, ovary half-inferior or inferior, obconic or obovate to subglobose, lobed and densely papillose at summit, locules 5-8, styles 5-8, distinct, short, erect or spreading; receptacle spongy, black. Fruit fleshy, stony, indehiscent, 4-5 horned, to 10 mm long. Seeds one per locule, pyriform, light brown, smooth, dull, ca. 1.5 mm long; placentation axile, seeds pendulous. Summer-fall. Rarely escaped or persisting after cultivation; FL!, GA!, NC!, WV!. [OH, PA]. Introduced from southeast Asia and the South Pacific. = F, C; T. expansa Murr, per Bogle (1970), S, G, R, Wilson (1932); T. tetragonoides (Pallas) Kuntze, per Taylor (1994).  [specimen photo]


1. Trianthema portulacastrum L.* Somewhat succulent, annual or perennial herbs. Stems prostrate, ascending or erect, sparsely to moderately hispid in longitudinal furrows, branching from axils of the smaller of unequal leaf pairs, nodes obscured by connate, sheathing petioles. Root a contorted taproot. Leaves somewhat fleshy, opposite (or appearing alternate), unequal, highly variable in shape, from round to obovate, elliptic or lanceolate, 1-6.5 cm long, to 3 cm wide, glabrous to sparingly pubescent with simple trichomes, apex cuspedate to obtuse or emarginate, margins translucent, serrulate to papillose, occasionally ciliate toward the base, base truncate, rounded or cuneate; petioles mostly 1-2 cm long, sometimes equaling the blade length, sometimes ciliate-margined, connate at the base, enclosing the stem; stipules present, hyaline, the lobes deltoid to narrowly triangular or attenuate. Inflorescence axillary, flowers 1-3 in axils, glabrous, partly concealed by sheathing petioles; bracts paired, aristate to long-acuminate, 2-5 mm long, often fused to the leaf bases. Calyx subtended by a pair of bracteoles, fused to calyx; sepals 5, fused below to a tube 1-3 mm long, lobes erect, imbricate, lanceolate to aristate or long-acuminate, 3-4 mm long, externally glabrous or hispid, petaloid and papillose within, mucros hispid, to 1.5 mm; corolla absent; stamens 5-10, included, adnate to the calyx tube; carpels 2, ovary superior, cyllindric, glabrous, locule 1, horizontally divided, styles 1, eccentric, to 2 mm long, stigma longitudinal, papillate. Fruit a thin-walled, circumcissile capsule, longitudinally nerved, to 7 mm long, the apex truncate and crested by 1-2 supra-opercular lobes. Seeds few to several, reniform, black, rugulose, enclosed by a dull, membranaceous aril, ca. 1.5 mm long; placentation parietal. Floral tube deciduous above the epicalyx bracts; seeds often evident as bulges in the connate leaf bases. Most Florida specimens were distinct in drying black rather than tawny or brown, and in having only one supra-opercular lobe with obvious ennervations. Spring-fall. Moist, sandy soils of thickets, dunes and wastelands; infrequent in waste areas of the Gulf coast and south-central U.S., a rare introduction to seaports of the mid-Atlantic states; AL!, AR!, FL!, LA!, MD, MS!, SC!, TN, VA?. [MO, NJ, OK, TX!]. A pantropical weed, likely introduced from the Old World tropics. = S, F, G, R, M, C.   [specimen photo 1]  [specimen photo 2]  [specimen photo 3]


Annual or perennial herbs, non-succulent or slightly succulent. Stems procumbent or ascending from a slender taproot, or absent. Leaves simple, basal or cauline, alternate or in false whorls. Inflorescence cymose, axillary or terminal. Flowers perfect, actinomorphic; sepals 5, free, distinct, with petaloid margins, persistent in fruit; corolla absent; stamens 3-5, free, distinct; gynoecium syncarpous, carpels 3-5, ovary superior, stigmata 3-5, generally sessile, persistent. Fruit a glabrous, loculicidal capsule. Seeds numerous, subreniform; placentation axile.

Two genera and four species are documented for the Southeast.

Key to the genera of Molluginaceae:

1 Plants moderately to densely tomentose, especially in axils of inflorescence; trichomes predominantly stellate; seeds with an inflated caruncle and a slender filiform appendage attached below the hilum   1. Glinus.
1 Plants predominantly glabrous; trichomes simple, if present; seeds lacking inflated caruncle or filiform appendage   2. Mollugo.


Annual, procumbent or ascending herbs, stellate-tomentose throughout. Stems sparsely to densely floccose with stellate trichomes, becoming glabrate below with age, usually less than 3 mm in diameter. Leaves simple, predominantly cauline, alternate or in false whorls, blades sparsely to densely tomentose, becoming glabrate; petioles densely to moderately tomentose. Flowers numerous in crowded, axillary cymules, inflorescence densely tomentose. Sepals petaloid within, green with white-hyaline margins, moderately to densely tomentose on external surface; carpels 3 (-5); stigmata 3 (-5), sessile; locules 3 (-5). Capsules turgid. Seeds numerous, visible through the green ovary wall, with an inflated caruncle and a long, contorted, filiform appendage attached below the hilum.

Key to the species of Glinus:

1 Leaf margins entire, or rarely with a few obscure teeth; sepals ovate-elliptic, apices acute, mucros 0.2-0.4 mm long; seeds densely papillose (visible at 10x magnification)   1. G. lotoides.
1 Leaf margins minutely serrulate to coarsely serrulate-lacerate (visible at 10x magnification); sepals lanceolate to ovate-lanceolate, apices long-attenuate, mucros 0.5-1.0 mm long, conspicuous to the unaided eye, often contorted; seeds with minute striations, not papillose   2. G. radiatus.

1. Glinus lotoides L.* Prostrate or ascending herbs. Stems spreading, nodes typically obscured by dense tomentum. Leaves variable, rotund, rhombic, elliptic, obovate or transversely elliptic, to 2 cm long and 2 cm wide, sometimes wider than long but typically 1-1.5x as long as wide, densely tomentose, becoming glabrate, apex rounded to obtuse, margins entire or rarely with a few obscure teeth, base cuneate to attenuate. Inflorescence axillary, flowers typically 6-12 per node, pedicels obscure, 1-2 mm long, densely tomentose. Sepals ovate to elliptic, 3.6-5.0 mm long X 1.5-2.2 mm wide, moderately to densely tomentose, apex ending in a mucro 0.2-0.4 mm long; stamens 3-5, included, to ca. 2.5 mm long; ovary ovoid, 3.3-4.0 mm long X 2.0-2.5 mm broad, glabrous. Seeds chestnut brown, ca. 0.5 mm in diameter, densely papillose (visible at 10x magnification). Summer-fall. Waste areas, mud flats and other moist areas; AR!, LA!. [MO!, OK!, TX]. Introduced from the Old World tropics and subtropics. = F, G, C.  [specimen photo]

2. Glinus radiatus (Ruiz & Pavón) Rohrbach.* Prostrate or ascending herbs. Stems spreading. Leaves variable, broadly obovate, elliptic or rhombic, to 3 cm long and 2 cm wide, typically 1.5-2.0x as long as wide, moderately to sparingly pubescent with stellate trichomes, becoming glabrate, apex acute to cuspidate, rarely obtuse, margins minutely to coarsely serrulate-lacerate (visible at 10x magnification), base cuneate to attenuate. Inflorescence axillary, flowers typically 8-12 per node, pedicels obscure, ca. 1 mm long, densely tomentose. Sepals lanceolate to ovate-lanceolate or elliptic, 4.0-5.2 mm long X 1.0-1.5 mm wide, tomentose, apex long acuminate, ending in a conspicuous, contorted subapical dorsal mucro 0.5-1.0 mm long (visible to the unaided eye); stamens 3-5, included, 2-3 mm long, filaments flattened; stigmata 3, ca. 0.5 mm long; ovary ovoid, 3.1-3.8 mm long X 1.6-2.0 mm broad, glabrous. Seeds numerous, chestnut brown to reddish, lustrous, smooth with minute striations, 0.4-0.5 mm in diameter. Spring-fall. Wet, muddy or sandy soils; LA!. [OK?, TX]. Likely introduced from S. America.  [specimen photo]


Annual or perennial herbs, glabrous or sparingly pubescent with simple trichomes. Leaves simple, basal or cauline and appearing whorled, linear to elliptic or spatulate, predominantly glabrous. Inflorescence either axillary or in terminal dichasia. Sepals glabrous with petaloid-hyaline margins; stamens 3-5; carpels 3 (-5); stigmata 3 (-5); locules 3 (-5). Seeds lacking an inflated caruncle.

M. cerviana (L.) Seringe, an introduced weed of canyons and brushlands in the southwest, approaches our area in TX and PA. This is a smaller, more delicate plant than M. verticillata L., and can be readily distinguished by its slender stems (0.2-0.5 mm in diameter), and by its linear, glaucous cauline leaves. The inflorescence is generally more open than that of M. verticillata, with some terminal dichasia present. The sepals have either a single nerve or an anastomosing network of veins. The capsule is globose in shape and does not exceed the sepals when mature, unlike M. verticillata. The minutely rugulose seeds are smaller than for our other species and lack the knob-like projection above the hilum that is present in our taxa. Not documented for the SE flora. Introduced to sw US from Old World tropics and subtropics, occurring eastward to sw TX; also known from a single occurrence in Philadelphia Co., PA (Wherry et al. 1979).

Key to the species of Mollugo:

1 Leaves predominantly basal; inflorescence a leafless thyrse; seeds verrucose-papillose   1. M. nudicaulis.
1 Leaves predominantly cauline; seeds longitudinally ribbed, smooth or rugulose, not verrucose-papillose.
2 Leaves glaucous, linear or oblanceolate; sepals with either a single, broad dorsal nerve or with an anastomosing network of veins; seeds minutely rugulose, ca. 0.3 mm in diameter   x. M. cerviana.
2 Leaves glabrous, rarely glaucous, elliptic, oblong, obovate or linear; sepals with 3 raised, pigmented nerves; seeds with 3-7 prominent dorsal nerves or turgid and appearing smooth, ca. 0.5 mm in diameter   2. M. verticillata.

1. Mollugo nudicaulis Lam.* Annual or perennial herbs, glabrous, to 30 cm tall, rosette-bearing. Leaves basal, oblanceolate, obovate or spatulate, to 6 cm long X 13 mm wide, glabrous, apex rounded, acute or obtuse, margins entire with a thin layer of translucent cells, base long-attenuate or cuneate. Inflorescence an open thyrse; peduncle slender, elongate, glabrous, leafless; bracts paired, ovate, ca. 1 mm long, scarious. Sepals ovate to elliptic, 2-3 mm long, with 3 slightly raised nerves and hyaline margins; stamens 3-5, 1.2-1.7 mm long; ovary cyllindric, ellipsoidal or ovoid, 2-2.5 mm long. Capsules exceeding sepals when mature. Seeds numerous, dark maroon-chestnut, ca. 0.6 mm in diameter, verrucose-papillose. Reed (1964, p. 323) collected this species once from ore piles in Canton, Maryland in 1958, and no documentation of its persistence or occurrence elsewhere in our area has been found. No specimens from our area seen. Introduced from Old World tropics.  [specimen photo]

2. Mollugo verticillata L. Annual herbs. Stems prostrate or ascending, divergent, glabrous or rarely with a few simple trichomes at the nodes. Leaves appearing whorled, linear, oblanceolate or oblong (those of the basal rosette and lowest nodes sometimes spatulate), to 4 cm long X 0.8 cm wide, glabrous or with a few simple trichomes near the leaf base, midrib conspicuous, translucent in fresh material, apex obtuse or acute, margins entire. Flowers numerous in axillary cymules; pedicels 2-20 mm long, glabrous or sparsely puberulent, spreading in flower, often becoming recurved in fruit. Sepals obovate to elliptic, unequal, slightly inflated or cucullate toward apex, 1.8-2.4 mm long, with three raised, pigmented nerves, spreading at anthesis, becoming erect and appressed to the capsule at maturity; stamens 3 (-5), 1.2-2 mm long; ovary cyllindric to ovoid, 1.8-3.5 mm long, 1.2-2.0 mm broad. Capsules light brown in color, exceeding sepals at maturity. Seeds numerous, deep chestnut brown, lustrous, ca. 0.5 mm in diameter, with 3-7 prominent longitudinal nerves or turgid and appearing smooth. Seeds held in capsule horizontally, readily visible as bulges in the capsule wall. Spring-fall. Waste areas, cultivated grounds and disturbed woodlands with moist soil; ALL!. [IL!, IN!, MO!, NJ!, OH!, OK!, PA!, TX!]. Native to tropical Americas, apparently present at an archaeological site in the Little Tennessee River valley 3000 years ago (Chapman et al. 1974); possibly introduced to temperate N. America by Native Americans and spread incidentally with cultivation. = S, F, G, R, C, W; including M. gracillima Andersson (Reed 1964, p. 323).[specimen photo1]  [specimen photo2]


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