EUPHORBIACEAE 353 Tragia urticifolia Michaux, Nettleleaf Noseburn. Pd (GA, NC, SC, VA), Cp (GA, SC), Mt (SC): dry woodlands and rock outcrops, particularly over mafic or calcareous rocks; common (VA Rare). May-October. Sc. VA west to MO, KS, and CO, south to FL and AZ. [= RAB, F, G, K, W; = T. urticaefolia – S, orthographic variant]

Triadica Loureiro 1790 (Chinese Tallow-tree)

A genus of 2-3 species, native to tropical and subtropical Asia. The most recent monographers of Sapium and related genera (Kruijt 1996; Esser 2002) place our single naturalized species in the genus Triadica, native to Asia; Sapium (excluding Triadica) is a genus of 21 species restricted to the neotropics. This conclusion is corroborated by molecular phylogenetic analysis (Wurdack, Hoffmann, & Chase (2005). References: Kruijt (1996)=Z; Esser (2002)=Y; Govaerts, Frodin, & Radcliffe-Smith (2000)=X.

* Triadica sebifera (Linnaeus) Small, Chinese Tallow-tree, Popcorn Tree. Cp (GA, NC, SC): marsh edges, shell deposits, disturbed areas; uncommon. May-June; August-November, native of e. Asia. With Euphorbia, Chamaesyce, and Cnidoscolus, one of our few Euphorbiaceous genera with milky sap. Triadica has become locally common from Colleton County, SC southward through the tidewater area of GA, and promises to become a serious weed tree (as it is in parts of LA, TX, and FL). [= K, S, X, Y, Z; = Sapium sebiferum (Linnaeus) Roxburgh – RAB, GW]

Vernicia Loureiro 1790 (Tung-oil Tree)

A genus of 3 species, trees, native of se. Asia. References: Govaerts, Frodin, & Radcliffe-Smith (2000)=Z.

* Vernicia fordii (Hemsley) Airy-Shaw, Tung-oil Tree, Tung Tree. Cp (GA, NC): planted for the oil and for ornament, rarely naturalizing; rare, introduced from central and western China. Naturalized in GA, from former plantations; planted and showing a tendency to naturalize in the Coastal Plain of NC (Mount Olive, Wayne Co.). [= K, Z; = Aleurites fordii Hemsley]

FABACEAE Lindley 1836 or LEGUMINOSAE A.L. de Jussieu 1789 (Legume Family)

A family of about 730 genera and 20,000 species, trees, shrubs, and herbs, cosmopolitan. References: Isely (1990)=SE (throughout the family treatment); Wojciechowski, Lavin, & Sanderson (2004); Wilbur (1963); Isely (1998)=I; Robertson & Lee (1976).

1 Trees, shrubs, or woody vines...... Key A 1 Herbs (including herbaceous vines). 2 Leaves 4-many-foliolate. 3 Leaves palmately compound, with 4 or more leaflets ...... Key B 3 Leaves pinnately or bipinnately compound. 4 Leaves bipinnately compound...... Key C 4 Leaves pinnately compound...... Key D 2 Leaves 1-3-foliolate. 5 Leaves unifoliolate...... Key E 5 Leaves trifoliolate. 6 Leaves pinnately trifoliolate...... Key F 6 Leaves palmately trifoliolate...... Key G

Key A – woody legumes (trees, shrubs, or woody vines)

1 Leaves unifoliolate or trifoliolate, or reduced to phyllodial spines. 2 Tree; leaves unifoliolate and > 5 cm wide; [subfamily Caesalpinioideae, tribe Cercideae]...... Cercis 2 Shrubs or woody vines (rarely tree in Erythrina); leaves trifoliolate, unifoliolate, or reduced to phyllodial spines (if unifoliolate, < 2 cm wide); [subfamily Papilionoideae]. 3 Woody vine. 4 Calyx 4.5-6 mm long; leaflets unlobed; [tribe Phaseoleae, subtribe Diocleinae] ...... Dioclea 4 Calyx 10-12 mm long; leaflets generally lobed; [tribe Phaseoleae, subtribe Glycininae] ...... Pueraria 3 Shrub or tree. 5 Shrub or tree with twigs various, but not conspicuously green or flanged; leaves pinnately trifoliolate. 6 Corolla 30-50 mm long, scarlet; legume with several seeds; leaflets lobed; [tribe Phaseoleae, subtribe Erythrininae]...... Erythrina 6 Corolla 8-15 mm long, purplish, pink, or white; legume 1-seeded; leaflets not lobed; [tribe Desmodieae, subtribe Lespedezinae] ...... Lespedeza FABACEAE 354 5 Shrub with angled or flanged green twigs; leaves palmately trifoliolate, unifoliolate, or reduced to spine- tipped phyllodes; flowers bright yellow; [introduced, usually of roadsides or as remnants of cultivation]; [tribe Genisteae, subtribe Genistinae]. 7 Leaves all reduced to phyllodial spines; flowers axillary; calyx 10-15 mm long ...... Ulex 7 Leaves with normal lamina, either unifoliolate or trifoliolate; flowers in terminal racemes; calyx 3-6 mm long. 8 Leaves trifoliolate lower on the stem, often unifoliolate above; corolla 15-22 mm long...... Cytisus 8 Leaves unifoliolate throughout; corolla 10-14 mm long...... [Genista] 1 Leaves pinnate. 9 Woody vines; [subfamily Papilionoideae]. 10 Leaves even-pinnate; legume 3.5-4 cm long; seeds shiny scarlet and black; [tribe Abreae]...... [Abrus] 10 Leaves odd-pinnate; legume 4-15 cm long; seeds brown; [tribe Milletieae]...... Wisteria 9 Trees or shrubs. 11 Leaves 2×-paripinnate; [subfamily Mimosoideae] 12 Stamens connate at the base; inflorescence pink, 2.5-5 cm in diameter; [tribe Ingeae] ...... Albizia 12 Stamens free; inflorescence orange or yellowish-white, 1.0-2.2 cm in diameter. 13 Inflorescence yellowish-white, 1.8-2.2 cm in diameter; stamens 10; [tribe Mimoseae] ...... Leucaena 13 Inflorescence orange, 1.0-1.3 cm in diameter; stamens many; [tribe Acacieae]...... Vachellia 11 Leaves otherwise. 14 [subfamily Papilionoideae]. 15 Leaves glandular-punctate; corolla of only 1 petal; inflorescence a spike; shrubs; [tribe Amorpheae] ...... Amorpha 15 Leaves not glandular-punctate; corolla of 5 petals; inflorescence a raceme or panicle; trees or shrubs. 16 Leaflets alternate on the rachis; leaflets 4-15 (-20) cm long; [tribe Sophoreae]...... Cladrastis 16 Leaflets opposite on the rachis, leaflets (1-) 2-5 (-6) cm long. 17 Leaflets with persistent linear stipels; native and cultivated, collectively widespread in our area; [tribe Robinieae] ...... Robinia 17 Leaflets lacking stipels; cultivated, perhaps not established; [tribe Sophoreae]...... [Styphnolobium] 14 [subfamily Caesalpinioideae]. 18 Leaves all 2-pinnate, or a mixture of 2-pinnate and 1-pinnate on the same plant; shrub or tree; [tribe Caesalpineae]. 19 Leaves a mixture of 1-pinnate and 2-pinnate...... Gleditsia 19 Leaves all 2-pinnate. 20 Leaves petiolate; leaflets 20-70 mm long ...... Gymnocladus 20 Leaves subsessile (the pinnae simulating 1-pinnate leaves); leaflets 1-5 mm long...... Parkinsonia 18 Leaves all 1-pinnate (or appearing so in Parkinsonia); herb, shrub or tree. 21 Shrub with prominent glands on the leafstalk; [tribe Cassieae] ...... Senna 21 Tree or shrub (if a shrub, then lacking prominent glands on the leafstalk); [tribe Caesalpineae]. 22 Leaflets 13-45 mm long; tree; leaves 1-pinnate...... Gleditsia 22 Leaflets 1-5 mm long; shrub; leaves actually 2-pinnate, but subsessile, the pinnae simulating 1-pinnate leaves...... Parkinsonia

Key B – herbaceous legumes with palmate leaves with 4 or more leaflets

Lupinus, Orbexilum, Pediomelum, Psoralidium, Zornia

Key C – herbaceous legumes with bipinnate leaves

1 Petiole with 1-several glands; stems ascending to erect; flowers greenish-white ...... Desmanthus 1 Petiole without glands; stems prostrate to weakly arching; flowers pink-purple, yellow, or greenish-yellow. 2 Flowers pink-purple; legume ribbed, the ribs with prickles ...... Mimosa 2 Flowers yellow to greenish-yellow; legume not ribbed or prickly...... [Neptunia]

Key D – pinnate plus

Aeschynomene, Apios, Arachis, Astragalus, Clitoria, Dalea, Galactia, Glottidium, Glycyrrhiza, Lathyrus, Lotus, Pisum, Securigera, Sesbania, Tephrosia, Vicia

FABACEAE 355

Key E – unifoliolate

Alysicarpus, Baptisia, Crotalaria, Lupinus, Orbexilum, Pediomelum, Rhynchosia

Key F – palmately trifoliolate

Baptisia, Crotalaria, Kummerowia, Lotus, Medicago, Orbexilum, Psoralidium, Thermopsis, Trifolium

Key G – pinnately trifoliolate

Amphicarpaea, Centrosema, Clitoria, Cullen, Dalea, Desmodium, Erythrina, Galactia, Glycine, Indigofera, Lablab, Lespedeza, Lotus, Macroptilium, Medicago, Melilotus, Mucuna, Orbexilum, Pediomelum, Phaseolus, Rhynchosia, Stylosanthes, Strophostyles, Trifolium, Vigna

Abrus Adanson (Precatory Bean)

A genus of about 15 species, woody vines and shrubs, pantropical. References: Isely (1998)=I.

* Abrus precatorius Linnaeus, Precatory Bean, Rosary Pea, Crab's Eye. Apparently reported for GA, AL, and AR by Isely (1998) and Kartesz (1999), but this is actually based on mislabeling in Map 64 in Isely (1998). The species does occur in FL, near the GA border and might be expected in s. GA. [= I, K; = Abrus abrus (Linnaeus) L.F. Wight – S]

Acacia P. Miller (Acacia) [see Vachellia]

Acmispon Rafinesque (American Bird’s-foot-trefoil)

Annual and perennial herbs, of temperate North America. New World taxa often referred to Lotus are not closely related to Lotus, and should be segregated (Degtjareva et al 2006; Allan & Porter 2000). References: Isely (1981)=Z; Isely (1998)=I; Degtjareva et al. (2006); Allan & Porter (2000); Grant & Small (1996).

Acmispon helleri (Britton) A.A. Heller, Carolina Prairie-trefoil. Pd (GA, NC, SC, VA): dry woodlands and openings, originally probably limited to prairie-like sites (fire-maintained, post oak-blackjack oak savannas), generally on clayey soils, now primarily seen on roadbanks, along railroads, and in powerline rights-of-way, where mowing and bush-hogging have replaced fire as the force keeping the habitat open, sunny, and suitable for this plant of prairie affinities; rare (US Species of Concern, GA Special Concern, NC Rare, VA Rare). June-September. A.. helleri is endemic to the Piedmont of extreme sc. VA, NC, SC, and ne. GA. A. helleri is clearly closely related to A. americanus (= Lotus unifoliolatus, = Lotus purshianus, = Lotus americanus), of prairies of the midwestern states and various habitats further west, which ranges east to LA, AR, MO, IL, IN, and WI. Isely (1981) reduced A. helleri to a variety (in Lotus), because it "is but one of many elements within the L. purshianus complex and its differences from the rest are less than among the California races," while also stating "since it has no breeding contact with var. purshianus, it is reasonably maintained as a species." Since A. helleri seems adequately separated from A. americanus by its narrower leaflets, glabrate vestiture, and allopatric distribution, I have chosen to "reasonably maintain it as a species." [= Lotus helleri Britton – RAB; < L. americanus (Nuttall) Bischoff – F; < L. purshianus F.E. & E.G. Clements – G; = L. unifoliolatus (Hooker) Bentham var. helleri (Britton) Kartesz & Gandhi – K; = Acmispon helleri (Britton) Heller – S; = L. purshianus F.E. & E.G. Clements var. helleri (Britton) Isely – C, I, SE, Z]

Aeschynomene Linnaeus (Joint-vetch)

A genus of about 130 species, herbs, pantropical and warm temperate. References: Carulli, Tucker, & Dill (1988)=Z; Rudd (1955)=Y; Isely (1998)=I. Key adapted in part from SE.

1 Prostrate perennial; leaves with 3-7 (-9) leaflets; [of dry sandy habitats] ...... Ae. viscidula 1 Erect or ascending annual; leaves with 20-50 or more leaflets; [of moist to wet habitats]. 2 Leaflets with 2-4 longitudinal nerves; mature fruit stipe 1.5-3 mm long ...... Ae. americana var. americana 2 Leaflets with 1 longitudinal nerve; mature fruit stipe 4-25 mm long. FABACEAE 356 3 Mature fruit stipe 12-25 mm long; corolla (10-) 12-15 mm long; fruit segments 5-7 mm long, 4.5-6.5 mm wide; paired bracts subtending each flower toothed (rarely entire); standard greenish-yellow with distinct dark-red veins; leaflets 6-25 mm long, 2-5 mm wide ...... Ae. virginica 3 Mature fruit stipe 4-8 (-10) mm long; corolla 7-13 (-15) mm long; fruit segments 4-6 mm wide, 3.5-6 mm wide; paired bracts subtending each flower toothed or entire; standard pale orange or reddish-orange, the veins usually indistinct; leaflets 2.5-25 mm long, 1-4 mm wide. 4 Paired bracts subtending each flower entire (rarely toothed); leaflets 2.5-13 mm long, 1-2.5 mm wide; fruit segments 4-5 mm wide, 3.5-5 mm wide...... Ae. indica 4 Paired bracts subtending each flower toothed (rarely entire); leaflets 6-25 mm long, 1.5-4 mm wide; fruit segments 5-6 mm wide, 5-6 mm wide...... Ae. rudis

Aeschynomene americana Linnaeus var. americana, Shyleaf. Cp (GA): moist, disturbed sites; rare (GA Watch List). s. GA southward (Jones & Coile 1988, SE). [= I, SE, Y; < Ae. americana – K, S] Aeschynomene indica Linnaeus, Southern Joint-vetch Cp (GA, NC, SC, VA): marshes, ditches, disturbed wetlands; uncommon. July-October. Apparently native to se. North America, from NC west to AR, south to FL and TX, now widespread in the tropics and subtropics of the Old World and New World. Perry, Ware, & McKenney-Mueller (1998) discuss the occurrence of this species in VA. [= GW, I, K, SE, Y, Z; < Ae. virginica – S, in large part] * Aeschynomene rudis Bentham, Frisolillo. Cp (GA, NC, SC): roadside ditches, rice fields, disturbed wetlands; rare, introduced from South America. July-October. Native to South America, introduced in se. United States, recently becoming a weed. [= I, K, SE, Y, Z] Aeschynomene virginica (Linnaeus) Britton, Sterns, & Poggenburg, Northern Joint-vetch, Sensitive Joint-vetch. Cp (NC, VA): fresh to brackish tidal marshes and adjacent ditches, fields, and disturbed areas; rare (US Threatened, NC Endangered, VA Rare). July-October. NJ to ne. NC. Generally not weedy in most of its range, but in NC (now) found mostly in weedy situations, such as ditches or fields hydrologically connected to tidal waters. See Tyndall, Holt, & Lam (1996) and Belden & Van Alstine (2003) for additional information on habitat, population biology, and survey techniques. See Baskin et al. (1998) for additional information about seed germination and viability. [= RAB, C, F, G, I, K, SE, Y, Z; < Ae. virginica – S (also see Ae. indica)] Aeschynomene viscidula Michaux, Sticky Joint-vetch. Cp (GA): dry sandy areas, such as sandhills, dry pinelands, and barrier islands; rare (GA Special Concern). From s. GA southward (Jones & Coile 1988, SE). [= I, K, SE, Y; = Secula viscidula (Michaux) Small – S]

Albizia Durazzini (Silktree)

A genus of about 100-120 species, trees, shrubs, and vines, of tropical, subtropical, and warm temperate Asia, Africa, and America. References: Isely (1973)=Z; Isely (1998)=I.

1 Leaflets 7-15 mm long; bark of mature trees smoothish, with small wart-like bumps...... A. julibrissin 1 Leaflets 15-30 mm long; bark of mature trees rough, with plates...... A. kalkora

* Albizia julibrissin Durazzini, Mimosa, Silktree. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas, suburban woodlots, escaped and persistent in forests and woodlands; common, native of tropical Asia. May-August; July-November. Becoming a serious weed; "literally almost everywhere in the 'Dixie' south" (Isely 1973). [= RAB, C, I, K, SE, W, Z; = Albizzia julibrissin – F, G, S, orthographic variant] * Albizia kalkora (Roxburgh) Prain, Kalkora Mimosa. Pd (NC): naturalizing in suburban areas; rare, introduced from e. Asia (Japan, Korea, Taiwan). Documented by herbarium specimens at DUKE and NCU.

Alysicarpus Necker ex Desvaux (Alyce Clover)

A genus of about 25 species, herbs, native of the Old World tropics. References: Isely (1998)=I.

* Alysicarpus vaginalis (Linnaeus) Augustin de Candolle, Alyce Clover. Cp (GA), Pd (GA, NC): planted as a forage crop (at least formerly), and rarely naturalized; introduced from the Old World tropics. [= I, K, SE]

Amorpha Linnaeus (Indigo-bush, Leadplant)

References: Wilbur (1964)=Z; Wilbur (1975)=Y; Isely (1998)=I.

1 Short shrubs, usually 0.3-1 (-1.5) m tall; petioles 1-15 (-20) mm long, usually shorter than the width of the contiguous leaflets (except in A. georgiana var. confusa); leaflets usually slightly or conspicuously revolute. 2 Leaflet mucros mostly swollen apically; plant usually evidently and rather densely pubescent or puberulent (except A. herbacea var. floridana, of s. GA and FL). FABACEAE 357 3 Upper portions of the plant (stems and leaves) glabrescent; calyx tube glabrous to sparsely or densely minutely strigillose; fruit glabrous; [of s. GA southward]...... A. herbacea var. floridana 3 Upper portions of the plant (stems and leaves) conspicuously pubescent; calyx tube densely puberulent to short pilose; fruit densely to sparsely puberulent (rarely glabrate); [widespread in our area]....A. herbacea var. herbacea 2 Leaflet mucros mostly tapered apically; plant usually glabrous or sparsely pubescent. 4 Leaflets (10-) 15-25 (-35) mm long, (7-) 9-15 (-18) mm wide; standard intense (rarely light) bright blue; petiole (6-) 8-15 (-20) mm long; racemes mostly panicled, (1-) 3-5 (-8) per flowering branch, 10-20 (-45) cm long; flowering June-July...... A. georgiana var. confusa 4 Leaflets (3-) 6-10 (-15) mm long, (2-) 3-5 (-8) mm wide; standard reddish-purple; petiole 1-3 (-5) mm long; racemes solitary (less commonly panicled), 1 (-4) per flowering branch, (2-) 3-5 (-6) cm long; flowering April- May ...... A. georgiana var. georgiana 1 Taller shrubs, usually 1-3 (-4) m tall, petioles 10-30 mm long, usually exceeding the width of the contiguous leaflets; leaflets not revolute, or slightly so. 5 Calyx lobes (1.2-) 2.0-3.5 mm long (thus approaching, equal to, or exceeding the length of the calyx tube); racemes 3-8 (-15) cm long ...... A. schwerinii 5 Calyx lobes (0-) 0.2-1.2 mm long (thus distinctly shorter than the calyx tube); racemes 5-20 (-25) cm long. 6 Calyx lobes obsolete to very small, (0-) 0.2-0.6 (-0.8) mm long; plants glabrous to glabrate; leaflets usually not mucronate, the midrib commonly terminating in a sessile or shortly exserted (0.2-0.4 mm), slightly enlarged, glandular tip; leaflets relatively few, (9-) 11-15 (-19)...... A. glabra 6 Calyx lobes small, 0.2-1.2 mm long (the abaxial lobe usually 0.8-1.2 mm long); plants pubescent or puberulent, usually conspicuously so; leaflets usually mucronate, the midrib usually slender, exserted, 0.5-1.5 mm long and tapering; leaflets relatively many, 9-23 (-31). 7 Foliage remaining green when dried; leaflets (7-) 9-23 (-31) per leaf, dull to somewhat shiny above; [widespread in our area]...... A. fruticosa 7 Foliage blackening when dried; leaflets (7-) 9-15 (-19) per leaf, usually shiny above; [of s. SC and southward]...... A. nitens

Amorpha fruticosa Linnaeus, Tall Indigo-bush. Pd, Mt, Cp (GA, NC, SC, VA): forests, woodlands, marsh edges, sometimes in disturbed sites; uncommon (rare in VA Mountains). April-June; June-October. [= RAB, C, G, GW, I, K, SE, W, Y; > A. fruticosa var. fruticosa – F; > A. fruticosa var. tennessensis (Shuttleworth) E.J. Palmer – F; > A. curtissii Rydberg – S; > A. fruticosa – S; > A. tennesseensis Shuttleworth – S; > A. virgata Small – S] Amorpha georgiana Wilbur var. confusa Wilbur, Savanna Indigo-bush. Cp (NC, SC): pine savannas; rare (NC Threatened). (May-) June-July; August-October. The varietal epithet is rather unfortunate; it refers to nomenclatural, rather than taxonomic, confusion. In fact, the two varieties of A. georgiana seem so distinct that they warrant specific status. Var. confusa is a narrow endemic of the se. Coastal Plain of NC (Brunswick, Columbus, and Bladen counties) and immediately adjacent SC (Horry County). It is restricted to moist loamy savannas, especially on the Foreston soil series, a habitat now largely destroyed by fire suppression, real estate development, and conversion of savannas to pine tree farms. [= I, K, SE, Y, Z; < A. georgiana – RAB, GW; ? A. cyanostachya auct. non M.A. Curtis – S, in part] Amorpha georgiana Wilbur var. georgiana, Georgia Indigo-bush. Cp (GA, NC, SC): pine savannas, sandy river terraces; rare (GA Special Concern, NC Endangered). Late April-June; July-October. As mentioned above, the two varieties of A. georgiana probably deserve specific recognition. Var. georgiana is endemic to the Coastal Plain of sc. NC, SC, and se. GA, primarily in the fall-line Sandhills region, but rarely found on younger terraces (as far east as Pender County, NC). Much of its habitat has been destroyed. [= I, K, SE, Y, Z; < A. georgiana – RAB, GW] Amorpha glabra Desfontaines ex Poiret, Appalachian Indigo-bush, Mountain Indigo. Mt, Pd (GA, NC, SC): dry to dry- mesic ridgetop and slope forests, primarily in the Blue Ridge escarpment; uncommon (SC Rare). May-July; July-October. Endemic to the Southern Appalachian mountains (and nearby provinces) of n. AL, ne. GA, w. NC, nw. SC, and e. and c. TN. [= RAB, I, K, S, SE, W, Y] Amorpha herbacea Walter var. floridana (Rydberg) Wilbur, Florida Indigo-bush. Cp (GA): pine flatwoods and sandy river terraces; rare (GA Special Concern). Se. GA (Echols County) south into FL (Sorrie 1998b). [= Y, Z; < A. herbacea var. herbacea – I, K, SE; = A. floridana Rydberg – S] Amorpha herbacea Walter var. herbacea, Dwarf Indigo-bush. Cp (GA, NC, SC), Pd (GA, SC), Mt (NC): pine savannas, pine flatwoods, sandhills, other open forests and disturbed sites; common (rare in Mountains). May-July; July-October. Endemic to FL, GA, SC, and NC, mostly limited to the Coastal Plain. [= Y, Z; < A. herbacea – RAB, W; = A. herbacea – S; < A. herbacea var. herbacea – I, K, SE] Amorpha nitens Boynton, Dark Indigo-bush. Cp (SC), Pd, Mt (GA): sandy woodlands, rocky slopes, bottomland forests; rare (GA Special Concern). April-June. S. SC south to GA, west to LA, north in the interior to w. KY, s. IL, AR, and e. OK. First reported for SC by Nelson & Kelly (1997). [= I, K, S, SE, Y] Amorpha schwerinii C. Schneider, Piedmont Indigo-bush. Pd (GA, NC, SC): forests and woodlands, primarily rather xeric and rocky (though not exclusively so); rare (GA Special Concern, NC Rare, SC Rare). April-June; June-October. Endemic to the Piedmont (rarely adjacent provinces) of sc. NC, c. SC, nc. GA, e. AL, and ne. MS. [= RAB, I, K, S, SE, Y]

Amphicarpaea Elliott ex Nuttall (Hog-peanut)

FABACEAE 358 A genus of 3-5 species, of Asia, North America, and Africa. It now appears that 2-3 semi-cryptic taxa should be recognized in what has traditionally been considered a single species of Amphicarpaea (Callahan 1997, Parker 1996). The genus name has been corrected to Amphicarpaea from the frequently used Amphicarpa. References: Callahan (1997)=Y; Parker (1996)=Z; Isely (1998)=I.

1 Petiole 3.5-5.3 cm long; petiolule of the terminal leaflet 1.0-1.4 mm long; terminal leaflet 4.2-5.2 cm long...... A. bracteata var. bracteata 1 Petiole 6.0-6.8 cm long; petiolule of the terminal leaflet 1.7-1.9 mm long; terminal leaflet 5.5-6.1 cm long...... A. bracteata var. comosa

Amphicarpaea bracteata (Linnaeus) Fernald var. bracteata, Hog-peanut. {Mt, Pd, Cp (GA, NC, SC, VA): dry to moist forests, thickets; common. July-September; August-October. Producing inflorescences of 2 types, one with chasmogamous flowers and aerial legumes, the other with cleistogamous flowers and subterranean legumes. The distributions and habitats of the two varieties in our area require herbarium and field investigation}. [= K; = Amphicarpa bracteata var. bracteata – F, G, orthographic variant; < Amphicarpaea bracteata – C, I, SE; < Amphicarpa bracteata – RAB, orthographic variant; < Falcata comosa (Linnaeus) Kuntze – S] Amphicarpaea bracteata (Linnaeus) Fernald var. comosa Fassett, Hog-peanut. {Mt, Pd, Cp (GA, NC, SC, VA): dry to moist forests, thickets; common. July-September; August-October. Producing inflorescences of 2 types, one with chasmogamous flowers and aerial legumes, the other with cleistogamous flowers and subterranean legumes. The distributions and habitats of the two varieties in our area require herbarium and field investigation}. [= K; = Amphicarpa bracteata var. comosa – F, G, orthographic variant; < Amphicarpaea bracteata – C, I, SE; < Amphicarpa bracteata – RAB, orthographic variant; < Falcata comosa (Linnaeus) Kuntze – S]

Apios Fabricius (Groundnut)

A genus of about 10 species, perennial vines, of temperate e. Asia and e. North America. References: Woods (2005)=Z; Isely (1998)=I. Key based on Z.

1 Petiole 20-58 mm long; flower deep maroon to pale maroon and white; style glabrous; legume 6-10 (-12) cm long; seed 5-6 mm long; tubers several in a chain, each 2-10 cm in diameter...... A. americana 1 Petiole 70-75 mm long; flower pale green and rose-purple; style bearded; legume 12-15 (-18) cm long; seed 7.2-11.0 mm long; tuber 1, 15-20 cm in diameter...... [A. priceana]

Apios americana Medikus, Common Groundnut. Cp, Pd, Mt (GA, NC, SC, VA): marshes, wet thickets, streambanks; common. June-August; July-September. Nova Scotia, New Brunswick, and Québec west to MN and SD, south to s. FL and TX. [= RAB, C, GW, I, K, SE, W, Z; > A. americana var. americana – F, G; > A. americana var. turrigera Fernald – F, G; = Glycine apios Linnaeus – S]

Apios priceana B.L. Robinson, Kentucky Groundnut, Price's Potato-bean. Mixed oak woods, especially over limestone. Sw. KY, c. TN, ne. MS, and n. and c. AL. It is a rare species. [= C, F, G, I, K, SE, Z; = Glycine priceana (B.L. Robinson) Britton – S]

Arachis Linnaeus (Peanut)

A genus of 9-60 species, annual and perennial herbs, native of South America (especially Brazil). References: Isely (1998)=I.

* Arachis hypogaea Linnaeus, Peanut. Cp (GA, NC, SC, VA), Pd (NC): fields; commonly cultivated, rarely persistent, introduced from South America. July-October. This remarkable plant bears normal aerial flowers, but following pollination the pedicels elongate and arch downward, the legume soon buried and developing underground. [= RAB, C, F, K, S, SE]

Astragalus Linnaeus (Milkvetch)

Astragalus is a massive genus, usually considered to include over 2000 species, and most diverse in arid regions of w. North America and w. and c. Asia. The habitats of the southeastern species may be characterized as rocky or sandy, "relictual islands" of aridity in the generally moist landscape of eastern North America. References: Barneby (1964)=Z; Isely (1998)=I.

1 Legume pilose with hairs ca. 1 mm or more long; stems conspicuously pubescent, the hairs spreading and simple; plants decumbent, spreading, or ascending, the stems 0.5-4 dm long. 2 Corolla 14-19 mm long; leaflets 17-29; legume bilocular; calyx lobes shorter than the calyx tube; [of calcareous habitats of the interior]...... [A. tennesseensis] FABACEAE 359 2 Corolla 8-12 mm long; leaflets (5-) 7-15; legume unilocular; calyx lobes equaling or longer than the calyx tube [of dry sandy habitats from SC south] ...... A. villosus 1 Legume glabrous; stems glabrous or inconspicuously pubescent, the hairs appressed, simple or dolabriform; plants erect with stems (3-) 4-15 dm long, or decumbent with stems 1-3 dm long (A. distortus var. distortus and A. bibullatus). 3 Plants decumbent or ascending, stems 1-5 dm long; legume either dry and strongly curved (about 90 degrees), or globose and initially fleshy. 4 Legume globose, 1.3-2 cm in diameter, initially fleshy; corolla 18-25 mm long; [of calcareous glades of c. TN]...... [A. bibullatus] 4 Legume lanceolate, 1.2-2.5 cm long, 4-7 mm wide, strongly curved; corolla (7-) 8-15 mm long; [of shaley habitats from w. VA northward or of dry sandy sites in FL and possibly adjacent GA]. 5 Leaflets mostly 2-3.5× as long as wide; mature legume lacking a reticulately textured surface; corolla 10-15 mm long; [of shaley habitats from w. VA northward]...... A. distortus var. distortus 5 Leaflets mostly 1-2× as long as wide; mature legume reticulately textured; corolla 8-11 mm long; [of dry sandy sites in FL and possibly adjacent GA]...... [A. obcordatus] 3 Plants erect, stems (3-) 4-15 dm long; legume straight to moderately curved. 6 Lower stipules connate; pubescence dolabriform; legumes 1-1.5 cm long, 4-5 mm in diameter; [typically of dry to mesic soils]...... A. canadensis var. canadensis 6 Lower stipules free; pubescence simple; legumes either longer or wider (2-3 cm long and 4-6 mm in diameter in A. michauxii, 1.5-2.0 cm long and 8-18 mm in diameter in A. neglectus); [typically of notably dry, either rocky or sandy, soils]. 7 Leaves with 21-31 coriaceous to somewhat fleshy leaflets, many of the leaflets alternate or subopposite; legumes 2-3 cm long, 4-6 mm in diameter; [of dry sandy habitats from NC south]...... A. michauxii 7 Leaves with 11-23 thin-textured leaflets, all of the leaflets usually opposite; legumes 1.5-2 cm long, 8-18 mm in diameter; [of rocky calcareous habitats from VA north]...... A. neglectus

Astragalus canadensis Linnaeus var. canadensis, Canada Milkvetch. Mt (GA, NC, SC, VA), Pd, Cp (NC, SC, VA): forests, woodlands, streambanks, rocky slopes and bluffs; uncommon, rare in VA Piedmont, rare in Coastal Plain (GA Special Concern). June-August; July-October. Ranging through much of North America, from Québec and Hudson Bay west to British Columbia, south to GA, TX, CO, and Utah; also apparently in Siberia. The other varieties occur further west. See Barneby (1964) for a detailed discussion of taxonomic and nomenclatural problems involving A. canadensis. Barneby comments that "the eastern mountain race [in the Southern Appalachians] is commonly distinguished from var. canadensis of the Mississippi Valley and northward by a narrower and more open flowering and fruiting raceme, and the flowers at the same time are relatively small. There is something to be said in favor of recognizing an eastern montane variety, so long as we confine its distinguishing characteristic to a loose raceme." The distribution, as mapped by Barneby, is suggestive of a composite map of 2 (or more) different taxa, one of them being a Southern (and Central) Appalachian endemic. F and G separate var. carolinianus, basing the distinction, however, on a different set of characters, and considering var. canadensis to range south to VA (at least). Further study is needed; it seems we may have in our area 2 taxa worthy of distinction at the varietal level. [= I, K, SE, Z; < A. canadensis – RAB, C, W; > A. canadensis var. canadensis – F, G; > A. canadensis var. carolinianus (Linnaeus) M.E. Jones – F, G; > A. carolinianus Linnaeus – S] Astragalus distortus Torrey & A. Gray var. distortus, Ozark Milkvetch, Bent Milkvetch. Mt, Pd (VA): shale barrens and other dry, shaley places; rare (VA Rare). May-July. A. distortus is interpreted by Z (and followed by C and SE) to consist of 2 varieties: var. distortus, occurring in the s. Midwest from IL, MO, and OK south to MS, LA, and AR, and disjunct in n. and sc. VA, e. WV, and w. MD, and var. engelmannii (Sheldon) M.E. Jones, of TX and ne. LA. The two varieties seem fairly readily distinguishable morphologically in the Midwest. Appalachian var. distortus complicates the issue, since it approaches var. engelmannii in flower size and matches it in ovule number. The Appalachian plant, with a combination of morphologic characters not matching the two named varieties and far allopatric from them might better be considered a distinct variety. Further study is needed. [= C, I, K, SE, Z; < A. distortus – F, G; = Holcophacos distortus (Torrey & A. Gray) Rydberg – S] Astragalus michauxii (Kuntze) F.J. Hermann, Sandhills Milkvetch, Michaux's Milkvetch. Cp (GA, NC, SC): sandhills; rare (GA Special Concern, NC Proposed Threatened). Late April-June; June-October (and persisting). Sc. NC south through SC to GA, a Southeastern Coastal Plain endemic. "The Michaux milk-vetch is greatly isolated from any member of the genus morphologically similar" (Barneby 1964). [= RAB, I, K, SE, Z; = Tium michauxii (Kuntze) Rydberg – S] Astragalus neglectus (Torrey & A. Gray) Sheldon, Cooper Milkvetch. Mt (VA): dry calcareous woodlands and barren, over dolostone and limestone; rare (US Species of Concern, VA Rare). June-September. Se. Ontario west to se. Saskatchewan and ne. ND, south to w. NY, ne. PA, c. PA, n. OH, s. MI, se. WI, and e. SD; disjunct in w. VA and e. WV (Wieboldt et al. 1998). [= C, F, G, I, K, SE, Z] Astragalus villosus Michaux, Bearded Milkvetch, Southern Milkvetch. Cp (GA, SC): sandhills and other dry, sandy places; rare (SC Rare). May-June; June-August. A Southeastern Coastal Plain endemic: s. SC south to Panhandle FL, west to s. MS. This species is described by Barneby as "a lowly but delightful little astragalus." [= RAB, I, K, SE, Z; = Phaca intonsa (Sheldon) Rydberg ex Small – S]

Astragalus bibullatus Barneby & E.L. Bridges, Pyne's Ground-plum. Endemic to calcareous glades of c. TN (Barneby & Bridges 1987). [= I, K, SE; = Geoprumnon crassicarpum (Nuttall) Rydberg ex Small – S, misapplied; = A. crassicarpus Nuttall, misapplied] FABACEAE 360 Astragalus obcordatus Elliott, Florida Milk-vetch. Cp (GA?): sandhills; rare. S. MS south to c. peninsular FL. Reported for s. GA, but no specimen documentation is known (Barneby 1964). [= I, K, SE, Z; = Phaca obcordata (Elliott) Rydberg ex Small – S] Astragalus tennesseensis A. Gray ex Chapman. Calcareous glades. C. TN, n. AL, s. KY (and formerly IL and IN, and possibly MO). [= I, K, SE, Z; >< A. tennessensis – F; >< A. plattensis Nuttall – F; > Geoprumnon tennesseense (A. Gray ex Chapman) Rydberg – S; >< Geoprumnon plattense (Nuttall) Rydberg – S]

Baptisia Ventenat 1808 (Wild Indigo)

A genus of about 15 species, perennial herbs, of temperate e. and c. North America. References: Isely (1981)=Y; Larisey (1940a)=Z; Mendenhall (1994a, 1994b)=X; Turner (2006)=Q; Isely (1998)=I.

1 Leaves 1-foliolate, sessile or perfoliate. 2 Leaves perfoliate; plant glabrous or nearly so; [widespread from s. SC southward]...... B. perfoliata 2 Leaves sessile; plant glabrous or densely cobwebby pubescent. 3 Plant cobwebby-pubescent; leaves ca. 1× as long as wide, cordate at base; corolla 9-11 mm long, yellow; [of e. GA (Brantley and Wayne counties)] ...... B. arachnifera 3 Plant glabrous; leaves 1.3-1.6× as long as wide, rounded to broadly cuneate at base; corolla 12-15 mm long, pale yellow to greenish; [of the FL Panhandle] ...... [B. simplicifolia] 1 Leaves 3-foliolate, petiolate or sessile. 4 Flowering or fruiting pedicels bracteolate; corolla 11-14 mm long 5 Calyx lobes about as long as the calyx tube ...... B. lecontei 5 Calyx lobes much > the calyx tube. 6 Plant glabrous; [of ne. FL] ...... [B. calycosa] 6 Plant tomentose to hirsute; [of FL Panhandle] ...... [B. hirsuta] 4 Flowering or fruiting pedicels lacking bracteoles; corolla larger (except B. tinctoria). 7 Plants in flower ...... Key A 7 Plants in fruit...... Key B

Key A – flowering Baptisia

1 Flowers lavender or blue. 2 Leaflets 2-4 (-5) cm long, mostly < 10 mm wide (if wider, then < 4 cm long); leaflets mostly oriented in a vertical plane; fertile stems usually 0.4-1.0 m tall, the leafy branches horizontally spreading; racemes 1-2.5 (-4) dm long, rather densely flowered; petioles 0-4 (-12) mm long; [of diabase glades and barrens] ...... B. australis var. aberrans 2 Leaflets 4-6 (-9) cm long, mostly > 12 mm wide; leaflets not oriented in a vertical plane; fertile stems usually 1-1.5 m tall, the leafy branches ascending; racemes 2-4 (-5) dm long, rather sparsely flowered; petioles 5-20 (-40) mm long; [of flood-scoured riverside cobblebars and rock outcrops, also frequently cultivated and sometimes persistent or escaped] ...... B. australis var. australis 1 Flowers yellow, cream-white, or white. 3 Flowers white or cream-white. 4 Flowering pedicels 10-18 (-30) mm long, subtended by persistent bracts 10-25 mm long and 7-10 mm wide; flowers cream-white (to pale-yellow). 5 Petioles of median leaves 4-10 mm long...... B. bracteata 5 Petioles of median leaves 2-4 mm long...... [B. leucophaea] 4 Flowering pedicels 3-10 mm long, subtended by caducous bracts 4-7 mm long and 1-2 mm wide; flowers white. 6 Calyx 4.5-6.5 mm long; corolla 13-16 (-18) mm long; petioles 5-10 (-20) mm long...... B. albescens 6 Calyx 7-8 mm long; corolla 20-25 mm long; petioles (of the lower leaves at least) 10-20 mm long. 7 Legume usually 15-20 (-30) mm in diameter, thin-walled and brittle; [of NC south through GA to FL and AL] ...... B. alba 7 Legume usually 10-12 (-15) mm in diameter, rigid and tough; [of c. TN, c. KY, and MS westward] ...... [B. leucantha] 3 Flowers yellow. 8 Flowering pedicels 14-18 (-30) mm long, subtended by persistent bracts 10-25 mm long and 7-10 mm wide; flowers pale-yellow (to cream-white). 9 Petioles of median leaves 4-10 mm long...... B. bracteata 9 Petioles of median leaves 2-4 mm long...... [B. leucophaea] 8 Flowering pedicels 2-10 mm long, subtended by caducous bracts 2-10 mm long and 1-2 mm wide; flowers bright yellow. 10 Leaflets mostly 1-2.5 (-4) cm long, 1-2.5× as long as wide, the petiolules 0-1 mm long; corolla 12-16 mm long; racemes numerous, terminating most of the branches...... B. tinctoria FABACEAE 361 10 Leaflets mostly 4-9 cm long, 1.5-4× as long as wide, the petiolules 2-10 mm long; corolla 20-28 mm long; racemes solitary (-3) (B. cinerea) or numerous (B. lanceolata). 11 Inflorescences of many-flowered cylindrical racemes; stipules persistent or caducous. 12 Plant persistently cinereous-pubescent stipules (some of them at least) persistent...... B. cinerea 12 Plant puberulent when young, soon glabrate to glabrous; stipules caducous; [of MS westward]...... [B. sphaerocarpa] 11 Inflorescence of solitary axillary flowers or flowers in clusters of 2-4 in axils or terminal racemes; stipules caducous. 13 Petiolules 2-3 mm long; leaflets 1-2.5× as long as wide; [of LA, AR, TX, and OK] ...... [B. nuttalliana] 13 Petiolules 4-10 mm long; leaflets 1.7-5× as long as wide; [of SC, GA, FL, and AL]. 14 Leaflets 3-5× as long as wide, usually < 1.5 cm wide; flowers usually solitary or in clusters of 2-3; fruits broadly ellipsoid or subspheroidal, < 2× as long as wide; [se. SC south through GA Coastal Plain to ne. FL] ...... B. lanceolata var. lanceolata 14 Leaflets 1.7-3.2 (-5)× as long as wide, the larger typically > 2 cm wide; flowers in racemes of (1-) 3-10 flowers; fruits usually ellipsoid, often > 2× as long as wide; [FL Panhandle, s. AL, and c. peninsular FL]...... [B. lanceolata var. tomentosa]

{add B. megacarpa to key}

Key B – fruiting Baptisia

1 Legume 5-11 mm in diameter. 2 Legume cylindric, 20-30 (-35) mm long, 7-9 mm in diameter, yellow-brown, leathery in txture...... B. albescens 2 Legume globose or subspheroidal, 7-25 mm long, 5-11 mm in diameter, black, woody in texture. 3 Leaflets mostly 1-2.5 (-4) cm long...... B. tinctoria 3 Leaflets 3.5-10 cm long. 4 Leaflets 3-5× as long as wide, usually < 1.5 cm wide; infructescence nodes (fruits or aborted fruits) usually 1- 3; fruits broadly ellipsoid or subspheroidal, <2× as long as wide; [se. SC south through GA Coastal Plain to ne. FL]...... B. lanceolata var. lanceolata 4 Leaflets 1.7-3.2 (-5)× as long as wide, the larger typically > 2 cm wide; infructescence nodes (1-) 3-10; fruits usually ellipsoid, often > 2× as long as wide; [FL Panhandle, s. AL, and c. peninsular FL]...... [B. lanceolata var. tomentosa] 1 Legume 8-25 mm in diameter. 5 Pod drying tan, thin-walled and brittle...... B. megacarpa 5 Pod drying black to blackish-brown, leathery or tough. 6 Stems puberulent (sometimes inconspicuously so), not glaucous. 7 Legume (20-) 30-40 (-50) mm long, 15-25 mm in diameter; pedicels 14-18 (-30) mm long, subtended by persistent bracts10-25 mm long and 7-10 mm wide...... B. bracteata, [B. leucophaea] 7 Legume 10-35 mm long, 8-15 mm in diameter; pedicels 2-10 mm long, subtended by caducous bracts 2-10 mm long and 1-2 mm wide. 8 Petiolules 2-5 mm long; stipules (some of them at least) persistent ...... B. cinerea 8 Petiolules 4-10 mm long; stipules caducous...... B. lanceolata var. lanceolata 6 Stems glabrous and generally glaucous as well...... B. alba, [B. leucantha],[B. leucophaea], B. australis var. australis, B. australis var. aberrans

{add B. nuttalliana to fruiting key}

Baptisia alba (Linnaeus) Ventenat, Thick-pod White Wild Indigo. Pd, Cp (GA, NC, SC): dry woodlands, roadsides; rare (NC Rare). May-July; June-October. AL, FL, GA, SC, and NC. B. leucantha (see below) is a western sibling, treated as either a species or a variety. In fruit, it is easily separated from B. albescens and other Baptisia by its nearly spheroidal legume. B. alba and B. albescens have been nomenclaturally confused; Isely (1986a) corrects the application of the epithet "alba." [= S; = B. alba var. alba – I, K, SE; = B. pendula Larisey – RAB; = B. lactea (Rafinesque) Thieret var. obovata (Larisey) Isely – C (by implication), X, Y; = B. lactea var. pendula (Larisey) B.L. Turner – Q; > B. pendula var. pendula – Z; > B. pendula var. obovata Larisey – Z] Baptisia albescens Small, Narrow-pod White Wild Indigo, Spiked Wild Indigo. Pd, Mt (GA, NC, SC), Cp (GA, NC, SC, VA): dry woodlands, roadsides; uncommon (NC Rare, VA Rare). May-July; June-October. Se. VA south through NC, SC, and GA to n. FL, e. AL and e. TN. The fruits are unlike any of our other species in being cylindric, about 3× as long as the diameter, and yellowish-brown (rather than black) when mature. [= I, K, S, SE; = B. alba – RAB, C, F, G, Q, W, X, Y, misapplied; > B. alba – Z; > B. albescens – Z] Baptisia arachnifera Duncan, Hairy Rattleweed, Hairy Wild Indigo. Cp (GA): sandhills; rare (US Endangered, GA Endangered). With simple leaves and unmistakable for its dense "cobwebby" pubescence, this species is endemic to GA (Wayne and Brantley counties). [= I, K, Q, SE, X, Y] FABACEAE 362 Baptisia australis (Linnaeus) R. Brown var. aberrans (Larisey) M. Mendenhall, Eastern Prairie Blue Wild Indigo, Glade Wild Indigo. Pd (NC), Mt (GA): glades, barrens, and open woodlands over limestone (or other calcareous rocks) and diabase (or other mafic rocks), in areas that were formerly prairies, barrens, glades, or oak savannas; rare (GA Special Concern, NC Rare). April-May; June-August. C. and se. TN, nw. GA, c. NC, and (possibly) s. KY and sc. VA. Blue-flowered Baptisia from mafic glades, barrens, and former prairies and oak savannas in NC has proven problematic to taxonomists. Larisey (1940a) treated B. australis and B. minor as separate species, and placed eastern plants resembling B. minor in B. minor var. aberrans Larisey, but without providing very satisfying characters for separating it from typical B. minor of midwestern North America. RAB apparently (though tacitly) include B. minor within B. australis. Isely (1981, 1990) treated blue-flowered Baptisia as B. australis var. australis and var. minor, regarding var. minor as reaching its eastern limit in MO (the two varieties thus allopatric), and stating that "sporadic collections within the range of var. australis have the pods and some of the vegetative characters of var. minor... most of these collections are from dry or sterile habitats, e.g., cedar glades, that var. australis typically does not inhabit" (Isely 1990). His treatment of australis and minor at the varietal level seems largely based on the existence of minor-like plants within his concept of the range of australis. NC plants from glade-like sites are morphologically more similar to midwestern prairie B. minor, occur in similar habitats, and grow with a large number of other plants with midwestern phytogeographic affinities, such as Eryngium yuccifolium var. yuccifolium, Echinacea laevigata (an eastern sibling of E. purpurea), Oligoneuron album, Oligoneuron rigidum ssp. glabratum (an eastern sibling of O. rigidum ssp. rigidum), Silphium terebinthinaceum, and others. The affinities of these plants seem to be with B. minor; "shoehorning" them into the more eastern B. australis, which they do not resemble in morphology, habitat, or (indeed) range is not a desirable disposition. Eastern plants referrable to B. minor do, however, as noted by Larisey and Isely, differ from midwestern plants in leaflet size and shape, branching, and pod shape; they are best treated as an eastern, relictual variety, var. aberrans Larisey. Mendenhall (1994a, 1994b) found that B. minor var. aberrans warranted taxonomic recognition, and indeed that it is less closely related to B. australis s.s. and B. minor than they are to one another; she chose to treat the three entities as varieties under B. australis. For now, the best treatment seems to be to follow Mendenhall, and acknowledge the existence of three varietal entities, with the phylogenetic affinities uncertain. The range of B. australis var. minor is thus largely midwestern, from se. NE, s. MO, and e. and c. KS south to w. AR, e. and c. OK, and ne. TX. [= K, X; = Baptisia minor Lehmann var. aberrans Larisey – Z; < B. australis (Linnaeus) R. Brown – RAB, S; < B. australis var. australis – I, Q, SE; < B. australis var. minor (Lehmann) Fernald – C, G; < B. minor – F] Baptisia australis (Linnaeus) R. Brown var. australis, Tall Blue Wild Indigo, Streamside Blue Indigo. Mt (GA, NC*?, VA), Pd (NC*?, VA): riverbank scour areas, gravel bars, and disturbed areas (where persisting from cultivation); rare (GA Special Concern, VA Watch List). April-May; June-August. Native to w. and n. VA, w. MD, WV, w. PA, e. and c. KY, ne. TN, se. IN, and s. OH, and possibly native to other states, the original range somewhat obscured by its frequent cultivation. [= C, G, K, X; = B. australis – F, W, Z; < B. australis – RAB, S (also see B. minor); < B. australis var. australis – I, Q, SE (also see B. australis var. aberrans)] Baptisia bracteata Elliott, Creamy Wild Indigo. Pd (GA, NC, SC), Mt (GA, SC), Cp (SC): sandhills, other dry woodlands; uncommon (NC Rare). March-April; May-June. Ne. AL northwest through n. GA and n. SC to w. NC. The more western B. leucophaea Nuttall is better treated as a species than as B. bracteata var. leucophaea (Nuttall) Kartesz & Gandhi (Mendenhall 1994b). [= RAB, Q, S, W, X, Z; = B. bracteata var. bracteata – C, I, K, SE] Baptisia cinerea (Rafinesque) Fernald & Schubert, Carolina Wild Indigo. Cp (NC, SC, VA), Pd (VA): sandhills, other dry sandy woods; common, rare in VA (VA Rare). Late April-June; June-July. Though common in the Coastal Plain of the Carolinas, B. cinerea is a narrow endemic, ranging only from s. VA south to s. SC. The large, yellow flowers are very showy. In fall, the leaves do not drop, but stay attached to the stems, the whole plant turning an ashy gray; these dried plants are conspicuous through the following winter. The report in Jones & Coile (1988) of B. cinerea in GA is in error; the specimen is of B. lanceolata. [= RAB, C, F, G, I, K, Q, SE, X; = B. villosa auct. non (Walter) Nuttall – S, Z] Baptisia lanceolata (Walter) Elliott var. lanceolata, Gopherweed. Cp (GA, SC): sandhills; rare (SC Rare). April-May; June-November. Var. lanceolata ranges from s. SC south to ne. FL and sw. GA, a Southeastern Coastal Plain endemic. Small (1933) alleges that B. lanceolata ranges north to NC, but no documentation is known. The plant is reminiscent of B. cinerea, but forms larger, bushier plants and is separable by characters in the key. [= I, K, Q, SE, X; < B. lanceolata – RAB, S; = B. lanceolata – Z] Baptisia lecontei Torrey & A. Gray, Leconte's Wild Indigo. Cp (GA): sandhills; rare (GA Special Concern). Sc. GA south to e. Panhandle FL and s. peninsular FL. [= I, K, Q, S, SE, X, Y, Z] Baptisia megacarpa Chapman ex Torrey & A. Gray, Apalachicola Wild Indigo, Bigpod Wild Indigo. Cp (GA): moist florests of floodplains and lower slopes; rare (GA Special Concern). Late April-early June; June-July. E. Panhandle FL and sw. GA west to se. AL. [= I, K, Q, S, SE, X, Y; > B. megacarpa – Z; > B. riparia Larisey var. riparia – Z; > B. riparia var. minima – Z] Baptisia perfoliata (Linnaeus) R. Brown ex Aiton f., Catbells, Gopherweed. Cp (GA, SC): sandhills; uncommon. April- May; May-July. S. SC through e. GA to peninsular FL, a Southeastern Coastal Plain endemic. [= RAB, I, K, Q, S, SE, X, Y, Z] Baptisia tinctoria (Linnaeus) Ventenat, Honesty-weed, Rattleweed. Cp, Pd, Mt (GA, NC, SC, VA): sandhills, pine flatwoods, xeric woodlands, ridges, woodland edges, and roadbanks; common. April-August; July-November. Widespread in eastern United States, from NY and MN south to GA. The most widespread and common of our species of Baptisia, B. tinctoria is readily recognizable from its small, yellow flowers, small leaflets, and small fruits. The taxa synonymized need further investigation. [= RAB, C, I, K, Q, S, SE, W, X; > B. tinctoria var. projecta Fernald – F, G, Z; > B. tinctoria var. tinctoria – F, G, Z; > B. tinctoria var. crebra Fernald – F, Z; > B. tinctoria – S; > B. gibbesii Small – S]

Baptisia calycosa Canby, Florida Wild Indigo. Dry pinelands. Endemic to ne. FL (Clay and St. Johns counties). [= Q, S, Z; = B. calycosa var. calycosa – I, K, SE, Y] {synonymy incomplete} FABACEAE 363 Baptisia hirsuta Small, Hairy Wild Indigo, Panhandle Wild Indigo. Dry pinelands. Endemic to FL Panhandle (Escambia, Holmes, Okaloosa, Santa Rosa, and Walton counties). May; June-September. [= Q, S, Z; = B. calycosa Canby var. villosa Canby – I, K, SE, Y] {synonymy incomplete} Baptisia lanceolata (Walter) Elliott var. tomentosa (Larisey) Isely. Var. tomentosa (Larisey) Isely occurs in panhandle FL and adjacent s. AL; disjunct in c. peninsular FL. Two forms have been recognized, the "narrow-leaved form," endemic to the Apalachicola Lowlands portion of the FL Panhandle, and the "typical form", occupying the FL Panhandle, s. AL, and disjunct in c. peninsular FL (Isely 1981). Mendenhall (1994b) included broad-leaved and narrow-leaved forms of var. tomentosa in her study, which provided some support for the taxonomic recognition of these unnamed entities. [= I, K, SE, Y; = B. lanceolata var. elliptica (Small) B.L. Turner – Q; = B. elliptica Small – S; > B. elliptica var. elliptica – Z; > B. elliptica var. tomentosa Larisey – Z] Baptisia leucantha Torrey & A. Gray. East to s. MS, ne. MS, c. TN, c. KY, and s. OH; alleged by S to occur in NC, probably based on misinterpreted material of B. alba. [= S, X; = Baptisia alba var. macrophylla (Larisey) Isely – I, K, SE; = B. lactea (Rafinesque) Thieret var. lactea – C, Q, Y; > B. leucantha var. leucantha – Z; > B. pendula Larisey var. macrophylla Larisey – Z] {keyed} Baptisia leucophaea Nuttall. Nw. IN west to s. MN and e. NE, south to w. KY, c. MS, c. LA, se. LA (Turner 2006), and e. TX. [= F, G, Q; = B. bracteata Muhlenberg ex Elliott var. leucophaea (Nuttall) Kartesz & Gandhi – K; = B. bracteata var. glabrescens (Larisey) Isely – C, I, SE, Y; = B. leucophaea var. glabrescens Larisey – Z] Baptisia nuttalliana Small. Woodlands and prairies. S. AR and se. OK south to se. LA (Florida parishes) and se. TX. [= I, K, Q, S, SE, Y, Z] {not yet fully keyed; synonymy incomplete} Baptisia simplicifolia Croom. Flatwoods. Endemic to Panhandle FL. [= I, K, Q, S, X, Y, Z] {synonymy incomplete} Baptisia sphaerocarpa Nuttall. Woodlands and prairies. S. MS west to se. MO, e. OK, and e. TX. [= I, K, SE; > B. sphaerocarpa – Z; > B. viridis Larisey – Z] {synonymy incomplete}

Many of our species hybridize; hybrids known in our area include the following, listed in alphabetic order by hybrid formulae. Others may certainly occur. Additional hybrids have been created by plant breeders and may be found in cultivation.

B. albescens × cinerea. Known from SC. B. albescens × perfoliata [B. ×fulva Larisey]. Known from GA and SC. [B. ×fulva Larisey – Z] B. albescens × tinctoria [B. ×serenae M.A. Curtis (pro sp.); B. ×pinetorum Larisey]. Known from GA, NC, SC, and VA. B. cinerea × tinctoria. Known from NC (Brunswick County). B. perfoliata × tinctoria [B. ×microphylla Nuttall (pro sp.) – Z]. Known from SC.

Cassia (see Chamaecrista, Senna)

Centrosema (Augustin de Candolle) Bentham (Spurred Butterfly Pea)

A genus of about 40 species, perennial vining herbs, of tropical and warm temperate regions of the Western Hemisphere. References: Isely (1998)=I; Fantz (2002a).

Identification notes: Centrosema virginianum is easily confused with Clitoria mariana. The following key summarizes the differences:

1 Calyx 10-16 mm long, the calyx lobes 7-11 mm long, distinctly longer than the calyx tube; legume 7-12 cm long, 3-4 mm broad ...... Centrosema virginianum 1 Calyx 15-20 mm long, the calyx lobes 5-7 mm long, distinctly shorter than the calyx tube; legume 3-5 cm long, 5-7 mm broad ...... Clitoria mariana var. mariana

Centrosema virginianum (Linnaeus) Bentham, Spurred Butterfly Pea. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): dry woodlands and openings; common. June-August; July-October. S. NJ south to FL, west to KY, AR, and TX. [= RAB, C, G, K, SE, W; > C. virginianum var. virginianum – F; > C. virginianum var. ellipticum Fernald – F; = Bradburya virginiana (Linnaeus) Kuntze – S]

Cercis Linnaeus (Redbud)

A genus of about 6 species, trees of north temperate areas. References: Isely (1975)=Z; Robertson & Lee (1976)=Y; Isely (1998)=I.

1 Flowering pedicels 6-8 mm long; flowers 8-11 mm long ...... C. canadensis var. canadensis 1 Flowering pedicels 10-20 mm long; flowers (11-) 12-14 mm long ...... [C. chinensis]

FABACEAE 364 Cercis canadensis Linnaeus var. canadensis, Eastern Redbud. Pd, Mt, Cp (GA, NC, SC, VA): moist to dry forests and woodlands, especially over calcareous or mafic rocks, also commonly planted as an ornamental; common (uncommon in Coastal Plain south of VA, uncommon in Mountains). March-May; June-November. This spectacular small tree is showy in bud or flower. The smooth, medium gray bark is distinctive in winter. Other varieties occur in TX and Mexico. [= C, G, I, K, SE, Y, Z; < C. canadensis – RAB, F, S, W]

* Cercis chinensis Bunge, native to China, is sometimes cultivated. [= I]

Chamaecrista (Linnaeus) Moench (Partridge-pea)

A genus of about 265 species, shrubs and herbs, of primarily tropical and warm temperate areas. References: Isely (1975)=Z; Irwin & Barneby (1982)=Y; Robertson & Lee (1976)=X; Isely (1998)=I.

1 Corolla 0.8-1.0 cm in diameter, the larger petals 4-7 (-8) mm long; functional stamens 5-8. 2 Petiole pilose with hairs 1-2 (-3) mm long; petiolar gland cylindric or clavate; functional stamens 5-8; leaflets 5-6× as long as wide ...... Ch. nictitans var. aspera 2 Petiole incurved-puberulent with hairs 0-0.8 mm long; petiolar gland stalked-cupuliform or stalked-discoid; functional stamens 5; leaflets 3-5× as long as wide ...... Ch. nictitans var. nictitans 1 Corolla 2.5-3.5 cm in diameter, the larger petals 15-20 mm long; functional stamens 10. 3 Perennial from a horizontal woody root or crown; stems clustered ...... Ch. deeringiana 3 Annual from a taproot; stems solitary. 4 Pods 6.5-10 mm wide; seeds 4.7-6.3 mm across; [of tidal marshes in e. VA] ..... Ch. fasciculata var. macrosperma 4 Pods 3-5 (-6.5) mm wide; seeds (2.8-) 3.2-4.8 mm across; [widespread geographically and ecologically]. 5 Surface of leaflets pubescent; [from w. Panhandle FL and s. AL westward] .... [Ch. fasciculata var. puberula] 5 Surface of leaflets glabrous; [collectively widespread in our area]. 6 Petiolar gland depressed, 1.5-2.5 mm wide, raised at both ends; pods 6-10 cm long; plant usually glabrous or glabrescent, to 24 dm tall...... Ch. fasciculata var. brachiata 6 Petiolar gland not depressed, <1.5 mm wide; pods 4-6 cm long; plant usually pubescent, to 10 dm tall ...... Ch. fasciculata var. fasciculata

Chamaecrista deeringiana Small & Pennell, Florida Senna. Cp (GA): sandhills, dry longleaf pine woodlands, disturbed sandy areas; rare (GA Special Concern). Sw. and wc. GA (Jones & Coile 1988) south to Panhandle FL and west to s. MS (Sorrie & Leonard 1999); disjunct in s. FL. [= I, K, S, SE, Y, Z] Chamaecrista fasciculata (Michaux) Greene var. brachiata (Pollard) Isely. Cp (GA): fields, disturbed areas; uncommon. E. GA south to s. FL, west to w. panhandle FL. [= I, SE; < Chamaecrista fasciculata var. fasciculata – K; = Cassia fasciculata var. brachiata (Pollard) Pullen ex Isely – X, Z; = Chamaecrista brachiata Pollard – S; < Ch. fasciculata – Y] Chamaecrista fasciculata (Michaux) Greene var. fasciculata, Common Partridge-pea. Cp, Pd, Mt (GA, NC, SC, VA): fields, disturbed areas, fencerows, and a wide range of other habitats; common. June-September; July-November. MA west to MN, south to FL and Mexico. See discussion of the Chamaecrista fasciculata complex under var. macrosperma. [< Cassia fasciculata Michaux – RAB, W; < Chamaecrista fasciculata – C, Y; > Cassia fasciculata var. fasciculata – F, G, X; > Cassia fasciculata var. robusta (Pollard) J.F. Macbride – F, G, X; > Chamaecrista fasciculata – S; > Chamaecrista robusta Pollard – S; = Chamaecrista fasciculata var. fasciculata (variant 1, variant 2, and typical variant) – Z; < Chamaecrista fasciculata var. fasciculata – I, SE (also see var. macrosperma); < Chamaecrista fasciculata var. fasciculata – K (also see var. brachiata); > Cassia fasciculata var. littoralis (Pollard) J.F. MacBride – X] Chamaecrista fasciculata (Michaux) Greene var. macrosperma (Fernald) C.F. Reed, Tidal-marsh Partridge-pea. Cp (VA): freshwater tidal marshes; rare (US Species of Concern, VA Rare). Endemic to e. VA and MD. Isely (1975) did not recognize it formally, but treated it informally as "variant 1," commenting (incorrectly) that it is "apparently a local, saline-adapted ecotype." Irwin & Barneby (1982) treated Ch. fasciculata as a very polymorphic species in which it was impractical to recognize infraspecific taxa, concluding "a student of tropical Chamaecrista has the choice of accepting as a fact of life that sort of infraspecific variability that inspired the dissection of Ch. fasciculata or of retreating to the position of Britton & Urban who, driven by logic rather than sense, found a species in every particular combination of gland and hairiness that fell in their way. But in passing over the observed variation as taxonomically insignificant, it is well to bear in mind that its cause and its biological significance remain a mystery." Botanists familiar with var. macrosperma in the field contend that it shows a cohesion in morphologic characters, ecology, and distribution that is biologically and taxonomically significant; it seems clearly to warrant varietal status. [= K; < Chamaecrista fasciculata – C, S, Y; = Cassia fasciculata var. macrosperma Fernald – F, G; < Chamaecrista fasciculata var. fasciculata – I, SE; = Cassia fasciculata var. fasciculata "variant 1" – Z] Chamaecrista nictitans (Linnaeus) Moench var. aspera (Muhlenberg ex Elliott) Irwin & Barneby, Southern Sensitive-plant. Cp (GA, SC): savannas, pinelands, disturbed sandy soils; uncommon. June-October; July-November. Var. aspera ranges from se. SC south to s. FL. [= I; = Cassia aspera Muhlenberg ex Elliott – RAB, X, Z; = Chamaecrista nictitans ssp. nictitans var. aspera (Muhlenberg ex Elliott) Irwin & Barneby – K, SE, Y; = Chamaecrista aspera (Muhlenberg ex Elliott) Greene – S] Chamaecrista nictitans (Linnaeus) Moench var. nictitans, Common Sensitive-plant. Cp, Pd, Mt (GA, NC, SC, VA): forests, woodlands, disturbed areas, pine savannas, and a wide variety of other habitats; common. June-October; July-November. Ch. nictitans is widely distributed in e. North America, and (depending on the scope of what is included in it) south into South FABACEAE 365 America. Var. nictitans ranges throughout se. United States, north to MA, NY, OH, and KA. [= I; < Cassia nictitans Linnaeus – RAB, W, X, Z; < Chamaecrista nictitans – C; > Cassia nictitans var. nictitans – F, G; > Cassia nictitans var. hebecarpa Fernald – F, G; = Chamaecrista nictitans ssp. nictitans var. nictitans – K, SE, Y; > Chamaecrista procumbens (Linnaeus) Greene – S; > Chamaecrista multipinnata Pollard – S]

Chamaecrista fasciculata (Michaux) Greene var. 1. Dunes, sandy disturbed areas. s. AL west to e. and s. TX. [= I; < Ch. fasciculata var. fasciculata – K; > Chamaecrista littoralis Pollard – S; > Chamaecrista mississipiensis (Pollard) Pollard ex Heller – S; < Ch. fasciculata – Y; = Cassia fasciculata Michaux var. puberula (Greene) J.F. Macbride (variants 1, 2, and 3) – SE, Z; > Chamaecrista puberula Greene] {synonymy incomplete}

Cladrastis Rafinesque (Yellow-wood)

A genus of about 6 species, trees, of the se. United States and montane regions of Japan and China. References: Duley & Vincent (2003)=X; Isely (1981)=Z; Isely (1998)=I; Rudd (1972)=Y.

Cladrastis kentukea (Dumont de Courset) Rudd, Yellow-wood. Mt (GA, NC), Pd (NC*, SC): mountain forests, Piedmont bluffs, especially on calcareous or mafic rocks (introduced only in the Piedmont of NC); uncommon (NC Watch List, SC Rare). April-May; July-August. This small to large tree has a native range primarily in the Southern Appalachians (mostly on the west side), the Ozarks, and limestone regions in-between (such as c. TN), ranging from s. OH, s. IN, and s. MO south to sw. NC, sc. SC, n. GA, AL, c. AR, and e. OK, but is cultivated more widely. As discussed by Wyatt (1985), the SC occurrence on Fall Line bluffs of the Savannah River is an interesting disjunction, apparently relictual. Yellow-wood is a distinctive tree, distinguished by its smooth silvery-gray bark, peculiar leaves with alternate leaflets, and pendent panicles of white flowers. The genus Cladrastis has 4 other species, all of temperate e. Asia. Cladrastis is the only member of the tribe Sophoreae in our area, with the exception of the cultivated (and weakly, if at all, established) Sophora. [= K, W, X, Y; = C. lutea (Michaux f.) K. Koch – RAB, C, F, G, I, S, SE, Z]

Clitoria Linnaeus (Butterfly Pea)

A genus of about 60 species, of tropical and warm temperate regions of the New and Old World. References: Isely (1998)=I; Fantz (2000, 2002b)=Z.

Identification notes: Centrosema virginianum is easily confused with Clitoria mariana. See key under Centrosema.

1 Leaves 3-foliolate ...... C. mariana var. mariana 1 Leaves 5-7-foliolate ...... C. ternatea var. ternatea

Clitoria mariana Linnaeus var. mariana, Butterfly Pea. Cp, Pd, Mt (GA, NC, SC, VA): dry woodlands and openings; common. June-August; July-October. NY (Long Island), NJ west to s. OH, s. IL, MO, and OK, south to FL, TX, and South America; disjunct in s. AZ; also in se. Asia. [= Z; < C. mariana – RAB, C, F, G, I, K, SE, W; < Martiusia mariana (Linnaeus) Small – S] * Clitoria ternatea Linnaeus var. ternatea, Blue-pea. Cp (GA): disturbed areas; rare, introduced and weakly naturalized. In s. GA (Isely 1998). [= Z; < C. ternatea – I, K, S, SE]

Coronilla Linnaeus (Crown-vetch) (see Securigera)

Crotalaria Linnaeus 1753 (Rattlebox)

A genus of about 600 species, annual and perennial herbs, nearly cosmopolitan in tropical and temperate regions (especially diverse in Africa). References: Windler (1974)=Z; Isely (1986b)=Y; Isely (1998)=I. Key adapted in part from SE.

1 Leaves trifoliolate; erect annual herb, typically 1-2 m tall. 2 Leaflets obovate to elliptic-oblong, 1.5-3.5× as long as wide; legume conspicuously curved (or straight in C. incana). 3 Legume 10-15 mm in diameter, pilose; stem pubescence spreading...... C. incana 3 Legume 5-6 mm in diameter, minutely puberulent; stem pubescence appressed...... C. pallida var. obovata 2 Leaflets lanceolate, often narrowly so, 3-15× as long as wide; legume straight or nearly so (or upcurved at the tip). 4 Corolla 8-10 mm long; legume 4-6 mm in diameter, upcurved at tip ...... C. lanceolata 4 Corolla 18-20 mm long; legume 15 mm in diameter, not upcurved...... C. ochroleuca 1 Leaves unifoliolate; plants of various habits, mostly either perennial, smaller, or both. FABACEAE 366 5 Corolla 1.7-3.0 cm long; leaflets 4-15 cm long; stipules not decurrent on the stem and not conspicuously foliose; [exotic annual herbs, in disturbed habitats]. 6 Bracts of the inflorescence 2-3 mm long, caducous; leaflets 4-8 cm long ...... C. retusa 6 Bracts of the inflorescence 5-8 mm long, persistent; leaflets 5-15 cm long ...... C. spectabilis 5 Corolla 0.7-1.4 cm long; leaflets 1-8 cm long; stipules of at least the upper leaves conspicuously decurrent on the stem, giving the impression of a downward-pointing arrowhead (this feature sometimes inconspicuous or essentially absent in C. rotundifolia); [native perennial or annual herbs, in natural or disturbed habitats]. 7 Plant an erect annual; stems with spreading pubescence, the longer hairs 1-2 mm long; leaflets of the upper portion of the plant (4-) avg. 6 (-8)× as long as wide; [mostly of the Piedmont and Mountains (and Coastal Plain of VA)]...... C. sagittalis 7 Plant a decumbent, sprawling, or erect perennial; stems with appressed or spreading pubescence, the longer hairs <1.2 mm long; leaflets of the upper portion of the plant averaging either (1-) avg. 1-2 (-4)× or (5-) avg. 8-10 (- 15)× as long as wide; [mostly of the Coastal Plain]. 8 Leaflets glabrous above; leaflets of the upper portion of the plant usually (5-) 10 (-15)× as long as wide; plant erect or ascending ...... C. purshii 8 Leaflets pubescent above (the hairs sometimes sparse – check with hand lens); leaflets of the upper portion of the plant usually (1-) 2 (-4)× as long as wide; plant decumbent to low-ascending. 9 Stem pubescence appressed...... C. rotundifolia var. rotundifolia 9 Stem pubescence spreading...... C. rotundifolia var. vulgaris

* Crotalaria incana Linnaeus, Shake-shake. Cp (SC): disturbed areas; rare, introduced from Africa. [= I, K, S, SE] * Crotalaria lanceolata E. Meyer, Lanceleaf Rattlebox. Cp (GA, NC, SC): sandy fields, roadsides, other disturbed areas; rare, introduced from Africa. July-October; August-November. [= RAB, I, K, SE] * Crotalaria ochroleuca G. Don, Slenderleaf Rattlebox. Cp (GA, NC, SC): roadsides and sandy fields; rare, introduced from Africa. July-August; August-October. [= I, K, SE; ? C. intermedia – RAB, misapplied; ? C. brevidens Bentham var. intermedia (Kotschy) Polhill, misapplied] * Crotalaria pallida Aiton var. obovata (G. Don) Polhill, Smooth Rattlebox. Cp, Pd (GA, NC, SC): roadsides and fields; common, introduced from Africa. July-September; August-October. [= I, K, SE; ? C. mucronata – RAB; ? C. striata Augustin de Candolle – S] Crotalaria purshii Augustin de Candolle, Coastal Plain Rattlebox, Pursh's Rattlebox. Cp (GA, NC, SC, VA), Pd?, Mt? (GA): mesic to dry pinelands, sandy openings, roadsides; common (VA Watch List). May-July; July-September. A Southeastern Coastal Plain endemic: se. VA south to n. FL, c. peninsular FL, and west to e. LA, with scattered locations inland. [= RAB, C, G, I, K, S, SE, W; > C. purshii var. purshii – F; > C. purshii var. bracteolifera Fernald – F] * Crotalaria retusa Linnaeus, Rattleweed. Cp (GA, NC, SC), Pd (NC): disturbed areas; rare, introduced from the Old World tropics. July-September; August-October. [= RAB, F, G, I, K, S, SE] Crotalaria rotundifolia Walter ex J.F. Gmelin var. rotundifolia, Low Rattlebox, Rabbitbells. Cp (GA, SC): sandy forests and woodlands, roadsides; rare. E. SC south to s. FL, and west to e. LA, endemic to the Southeastern Coastal Plain. [= Z; < C. rotundifolia – C, I, K, SE, Y; < C. angulata – RAB, F, G, apparently misapplied; = C. maritima Chapman – S] Crotalaria rotundifolia Walter ex J.F. Gmelin var. vulgaris Windler, Low Rattlebox, Rabbitbells. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (GA): sandy forests and woodlands, roadsides; common (VA Rare). Se. VA south to c. peninsular FL, west to se. LA; also widespread in Mexico. [= Z; < C. rotundifolia – C, I, K, SE, Y; < C. angulata – RAB, F, G, misapplied; = C. rotundifolia – S] Crotalaria sagittalis Linnaeus, Common Rattlebox. Cp (NC, SC, VA), Pd, Mt (GA, NC, SC, VA): woodlands, woodland edges, openings, fields; common. June-August; July-September. MA and VT west to s. MI, s. WI, and c. MN, south to c. SC, s. AL, s. MS, TX, Mexico and Central America; West Indies. [= RAB, C, G, I, K, S, SE, W; > C. sagittalis var. sagittalis – F; > C. sagittalis var. oblonga Michaux – F] * Crotalaria spectabilis Roth, Showy Rattlebox. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): fields, roadsides, disturbed areas; common, introduced from s. Asia. July-September; August-October. [= RAB, C, F, G, I, K, SE; ? C. retzii A. Hitchcock – S]

Cullen Medikus

A genus of ca. 35 species, herbs, of the Old World. References: Isely (1998)=I.

* Cullen americanum (Linnaeus) Rydberg, Scurf-pea. Cp (SC): waste areas around wool-combing mill; rare, perhaps only a waif, native of the western Mediterranean region (a misnomer). There are other (older) reports from other southeastern states, including FL and MS. [= I, S; = C. americana – K, SE, orthographic variant]

Cytisus Linnaeus (Broom)

References: Isely (1998)=I.

FABACEAE 367 * Cytisus scoparius (Linnaeus) Link, Broom, Scotch Broom, Besom, Ginster. Cp (NC, SC, VA), Pd, Mt (GA, NC, SC, VA): roadbanks, woodland borders, disturbed areas; common (rare in Mountains), introduced from Europe. April-May; May-July. [= RAB, C, F, G, I, S, SE, W; > C. scoparius var. scoparius – K]

Dalea Linnaeus 1758 (Prairie-clover)

A genus of about 160 species, of temperate and tropical America, especially dry areas. The inclusion of Petalostemon in Dalea is controversial; recent evidence suggests that the recognition of Petalostemon may be warranted. If Petalostemon is recognized, only D. leporina is retained in Dalea. References: Barneby (1977)=Z; Ward (2004c)=Y; Isely (1998)=I. Key adapted from SE.

1 Spikes corymbosely aggregated, capitate, surrounded by an involucre of 3-4 series of sterile bracts; [subgenus Dalea, section Kuhnistera]. 2 Leaflets 5-9 (-15); petals (other than the standard) mostly 3.7-4.5 mm long ...... D. pinnata var. pinnata 2 Leaflets usually 3; petals (other than the standard) mostly 4.8-6.8 mm long ...... D. pinnata var. trifoliata 1 Spikes not corymbosely disposed, ovoid to cylindric, with or without a few subtending, sterile bracts. 3 Corolla subpapilionaceous, with apparent, differentiated wings and keel; stamens 9-10; annual herb; [alien, of disturbed habitats]; [subgenus Dalea, section Dalea]...... D. leporina 3 Corolla not papilionaceous, the wings and keel not differentiated; stamens 5; perennial herb; [native, primarily of calcareous glades and Coastal Plain pinelands]; [subgenus Dalea, section Kuhnistera]. 4 Leaflets 15-25; leaflets 2.5-3.5× as long as wide ...... [D. foliosa] 4 Leaflets 3-9; leaflets 3-10 (or more)× as long as wide. 5 Plants slightly to obviously pubescent (at least the spikes obviously pubescent); leaflets commonly involute or tubular, and > 10× as long as wide; corolla purple or pink. 6 Leaflets 5-7 (-9); spikes lengthening and loosening in fruit, often becoming sinuous; plants decumbent to ascending, stems normally branching only below the middle ...... D. gattingeri 6 Leaflets 3-5 (-7); spikes remaining compact; plants decumbent or ascending, stems normally branching only below the middle (D. cahaba), or ascending to erect, the stems branching above the middle (D. purpurea var. purpurea). 7 Interfloral bracts with pucescence along the keel and margins; plants decumbent to ascending, stems normally branching only below the middle ...... [D. cahaba] 7 Interfloral bracts with pubescence in a transverse band only; plants ascending to erect, the stems branching above the middle...... [D. purpurea var. purpurea] 5 Plants glabrous (except that the calyx lobes may be pubescent); leaflets broad and flat or narrow and involute; corolla pink-purple or white. 7 Calyx tube not incised on the ventral (upper) side; blade of the standard cordate; corolla white; [of calcareous habitats of inland provinces of GA, AL, TN, WV and westward]...... D. candida 7 Calyx tube deeply incised on the ventral (upper side; blade of the standard not cordate; corolla pink- purple or white; [of the Coastal Plain of GA southward and westward]. 8 Leaflets linear, folded, or involute and terete-filiform; spikes globose ca. 6-12 mm long and in diameter; bracts much shorter than the calyx; corolla usually bright pink-purple (less commonly white or lavender)...... D. feayi 8 Leaflets elliptic to oblanceolate, flat or folded; spikes ovoid to cylindric, 7-40 mm long; bracts as long as or longer than the calyx; corolla pink or white. 9 Plants spreading or decumbent; leaves widely spaced, generally lacking axillary fascicles; bract tips recurved in bud; calyx 2.7-3.3 mm long; flowers white; [of sc. and sw. GA west to se. LA]...... D. gracilis 9 Plants erect-ascending to sprawling; leaves more densely spaced, generally with well- developed axillary fascicles; bract tips not recurved in bud; flowers pink or white; [of the GA Coastal Plain, se. AL, and south through e. FL Panhandle to the s. FL peninsula]. 10 Leaflets of primary stem leaves mostly 5; corolla white...... D. albida 10 Leaflets of primary stem leaves mostly 7-9; corolla pink (rarely white) ...... D. carnea

Dalea albida (Torrey & A. Gray) D.B. Ward, White-tassels. Cp (GA): pinelands; uncommon. July-November. E. GA (near the SC border) west to se. AL, south ne. FL, n. peninsular FL, and e. FL Panhandle. [= Y; = Dalea carnea (Michaux) Poiret var. albida (Torrey & A. Gray) Barneby – I, K, SE, Z; = Petalostemon albidus (Torrey & A. Gray) Small – S] Dalea candida Michaux ex Willdenow, White Prairie-clover. Mt (GA): limestone glades and barrens; rare (GA Special Concern). Late May-August. WV, KY, IN, WI, MN, and Saskatchewan south to nw. GA, e. TN, w. AL, sc. MS, s. LA, and ne. TX. [= I, SE (excluding D. occidentalis); = D. candida var. candida – C, K, Z; = Petalostemum candidum (Michaux ex Willdenow) Michaux – F, G; = Petalostemon candidus (Michaux ex Willdenow) Michaux – S] Dalea carnea (Michaux) Poiret, Pink-tassels. Cp (GA): dry sandy pinelands; rare (GA Special Concern). June-November. Se. GA south to s. peninsular FL. [= Y; = Dalea carnea (Michaux) Poiret var. carnea – I, K, SE, Z; = Petalostemon carneus Michaux – S] FABACEAE 368 Dalea feayi (Chapman) Barneby, Feay's Prairie-clover. Cp (GA): sandhills; rare (GA Special Concern). June-October. E. GA (vicinity of the Altamaha River); FL peninsula; Panhandle FL (vicinity of the Altamaha River). [= I, K, SE, Z; = Petalostemon feayi Chapman – S] Dalea gattingeri (A. Heller) Barneby, Gattinger's Prairie-clover. Mt (GA): limestone glades and barrens; rare (GA Special Concern). May-August. C. TN, nw. GA, n. AL, s. MO, and n. AR (Sundell et al. 1999). [= I, K, SE; = Petalostemon gattingeri (A. Heller) A. Heller – S] Dalea gracilis (Nuttall) D.B. Ward, Sprawling White-tassels. Cp (GA): wet pine savannas; rare (GA Special Concern). August -September. Sc. and sw. GA west to se. LA. [= Y; = Dalea carnea (Michaux) Poiret var. gracilis (Nuttall) Barneby – I, K, SE, Z; = Petalostemon gracilis Nuttall – S] * Dalea leporina (Aiton) Bullock, Hare's-foot Dalea. Mt (VA): {habitat not known}; rare, introduced from farther west. [= I, K, SE, Z; ? Parosela alopecuroides (Willdenow) Rydberg – S] Dalea pinnata (J.F. Gmelin) Barneby var. pinnata, Summer Farewell, Eastern Prairie-clover. Cp (GA, NC, SC): sandhills and other dryish pinelands, especially in loamy sands; uncommon (NC Watch List). August-November. Sc. and se. NC south through SC and GA to c. peninsular FL and e. panhandle FL. [= I, K, SE, Z; < Petalostemum pinnatum (J.F. Gmelin) Blake – RAB; < Kuhnistera pinnata (J.F. Gmelin) Kuntze – S] Dalea pinnata (J.F. Gmelin) Barneby var. trifoliata (Chapman) Barneby. Cp (GA): sandhills, dry to moist longleaf pine flatwoods; uncommon. September-November. E. GA (near the Savannah River) south and west to w. Panhandle FL, s. AL, and s. MS. [= I, K, SE, Z; < Kuhnistera pinnata (J.F. Gmelin) Kuntze – S; = Petalostemon pinnatus (J.F. Gmelin) Blake ssp. trifoliatus (Chapman) Wemple]

Dalea cahaba J. Allison, Cahaba Prairie-clover. Dolomitic Ketona glades. May-June; June-September. C. AL (Bibb County) (Allison & Stevens 2001). Dalea foliosa (A. Gray) Barneby, Cedar Glade Prairie-clover. Calcareous glades. Late June-September. C. TN, n. AL, IL, and OH (?). [= C, I, K, SE, Z; = Petalostemum foliosum A. Gray – F, G; = Petalostemon foliosus A. Gray – S] Dalea purpurea Ventenat var. purpurea, Purple Prairie-clover. Prairies, glades, and open woodlands. NY and Ontario west to British Columbia, south to KY, TN, n. AL, c. MS, TX, and NM. [= C, I, K, SE, Z; < Petalostemum purpureum (Ventenat) Rydberg – F, G; < Petalostemon purpureus (Ventenat) Rydberg – S]

Daubentonia (see Sesbania)

Desmanthus Willdenow (Bundleflower)

A genus of about 25 species, mostly herbs, of warm temperate and subtropical America. References: Isely (1973)=Z; Isely (1998)=I.

* Desmanthus illinoensis (Michaux) MacMillan ex B.L. Robinson & Fernald, Bundleflower, Prairie Mimosa. Pd (NC, SC), Cp (GA, SC, VA), Mt (GA): disturbed areas; rare, introduced from west of the Appalachians. June-July; August-November. Native primarily west of the Mississippi River (but probably also on limestone through much of TN), with scattered occurrences to the east, most of these adventive. The species is probably not native in our primary area. [= RAB, C, F, G, I, K, SE, W, Z; = Acuan illinoense (Michaux) Kuntze – S; = Mimosa illinoensis Michaux]

Desmodium Desvaux 1813 (Tick-trefoil, Tick-clover, Beggar's-ticks, Stick-tights)

A genus of about 300 species, annual herbs, perennial herbs, and shrubs, nearly cosmopolitan (but lacking from Europe). In our area, Desmodium is a complex genus. Some of the species in our area are confusing and can be identified only with difficulty. The three species in Key A form a group quite distinct from the others in our area. It may be that these two groups warrant recognition as two genera, in which case D. nudiflorum, D. glutinosum, and D. pauciflorum are retained in Desmodium, and the rest of our taxa would be in Meibomia (which is the older genus name, but Desmodium is conserved against it when the two are united). References: Isely (1998)=I. Key based on SE, C, RAB, and F. Some parts adapted with little change from SE. Some parts, especially Key E, will likely be substantially revised, based on additional herbarium and field testing.

1 Calyx lobes shorter than the calyx tube; stipe of the loment 4-20 mm long, about 3× or more as long as the calyx; mature leaves without stipels at the base of the petiolules of the leaflets; leaves subverticillate at the top of the stem (alternate in D. pauciflorum); stamens monadelphous; lower margin of the loment incised to the upper suture...... Key A 1 Calyx lobes equal to or longer than the calyx tube (at least the lower calyx lobes longer than the calyx tube); stipe of the loment absent or nearly so, included within the calyx; mature leaves retaining stipels at the base of the petiolules of the leaflets; leaves alternate; stamens diadelphous; lower margin of the loment not incised to the upper suture. 2 Leaflets narrow, the terminal leaflet < 10 mm wide, and also 4-12× as long as wide, typically thick and strongly reticulate; petioles of midstem leaves 1-10 (-15) mm long; [primarily of the Coastal Plain and lower Piedmont (rarely Mountains) in our area]...... Key B FABACEAE 369 2 Leaflets broader, the terminal leaflet > 15 mm wide, or < 4× as long as wide, typically thin and not reticulate; petioles of midstem leaves various, but > 15 mm long if leaflet proportions are narrow; [collectively widespread in our area]. 3 Stems trailing vinelike along the ground...... Key C 3 Stems erect or ascending, not vinelike. 4 Stipules persistent (most or all of the stipules persisting through the year), 4-20 mm long, ovate to amplexicaul-clasping (to lance-attenuate, and if so, generally longer than 8 mm long, except in D. floridanum); leaflets 1.5-3× as long as wide ...... Key D 4 Stipules caducous (most or all of the stipules falling soon after expansion of the leaves), 2-6 (-8) mm long, mostly linear-subulate or lance-attenuate (in some species narrowly ovate to triangular); leaflets 1-8 (-10)× as long as wide ...... Key E

Key A

1 Stems dimorphic, the flowering stem normally lacking leaves (rarely with leaves), the sterile stem with a subverticillate cluster of 3-7 leaves near the top; pedicels 10-20 mm long ...... D. nudiflorum 1 Stems monomorphic, bearing both leaves and flowers, the leaves either subverticillate or not; pedicels 2-10 mm long. 2 Leaves subverticillate, clustered; leaflets conspicuously and strongly acuminate, 5-10 cm long; flowers usually distinctly pink or pink-purple; inflorescence 3-8 dm long, elongate, large, and conspicuous, much exceeding the leaves...... D. glutinosum 2 Leaves alternate, scattered; leaflets acute to slightly acuminate, 3-7 cm long; flowers white; inflorescence 1-2 dm long, small and inconspicuous, often partly obscured by the leaves ...... D. pauciflorum

Key B (Desmodium with very narrow leaflets)

1 Petioles (0-) 1-3 (-4) mm long, the leaves thus subsessile; leaflets 5-10 mm wide, strongly pubescent on the lower surface ...... D. sessilifolium 1 Petioles 3-15 mm long, the leaves thus obviously petiolate; leaflets 2-5 (-8) mm wide, glabrate or inconspicuously puberulent on the lower surface. 2 Loment segments flat to distinctly concave along the upper (suture) margin; [of dry to mesic habitats]...... D. strictum 2 Loment segments slightly convex along the upper (suture) margin; [of boggy, wet, or mesic habitats]..... D. tenuifolium

Key C (Desmodium with trailing stems)

1 Stipules ovate, persistent, slightly to strongly clasping at the base, 6-12 mm long. 2 Leaflets ovate, 1.2-1.9× as long as wide; flowers white to yellowish; loment uncinate-puberulent only along the sutures ...... D. ochroleucum 2 Leaflets ovate, 0.8-1.1× as long as wide; flowers blue-purple; loment uncinate-puberulent over the surface ...... D. rotundifolium 1 Stipules lanceolate to linear (or deltate in D. humifusum), usually quickly deciduous, not clasping at the base, 2-8 mm long. 3 Terminal leaflet 1.4-2.0× as long as wide, 3.0-7.0 cm long; loment segments 6-8 mm long; stipules 4-8 mm long, ovate to lance-acuminate ...... [D. humifusum] 3 Terminal leaflet 0.9-1.2× as long as wide, 1.5-2.3 cm long; loment segments 4-5 mm long; stipules lanceolate, 1-5 mm long ...... D. lineatum

Key D

1 Loment segments nearly symmetrical along the axis of the loment (the isthmi more or less equal above and below, thus each segment diamond-shaped, rounded-diamond-shaped, or essentialy elliptical), each segment 3-3.5 mm long; annual from a taproot; [of the Coastal Plain of NC and SC]...... D. tortuosum 1 Loment segments asymmetrical along the axis of the loment (the isthmi deeper below than above, thus each segment triangular, rounded-triangular, or semi-circular), each segment 5-11 mm long; perennial; [collectively widespread in our area]. 2 Corolla 6-7 mm long; loment with 2-4 segments, each 5-7 mm long; lower leaves often 1-foliolate; [of se. SC and southward] ...... D. floridanum 2 Corolla 8-13 mm long; loment with 4-6 segments, each 6.5-11 mm long; lower leaves usually 3-foliolate; [collectively widespread in our area]. 3 Stem densely spreading pilose (at least the upper stem) and also uncinate-puberulent; loment segments 6.5-10 mm long ...... D. canescens 3 Stem glabrous or uncinate-puberulent; loment segments 9-11 mm long...... D. cuspidatum var. cuspidatum FABACEAE 370

Key E

1 Corolla 8-11 mm long; petioles mostly 1-8 mm long; [plants of the Mountains of VA and possibly NC]...... D. canadense 1 Corolla 3-8 (-9) mm long (or 8-10 mm long in D. laevigatum); petioles mostly longer, mostly 10-60 mm long (except D. ciliare and D. obtusum); [plants collectively widespread in our area]. 2 Loment with 1-3 segments, rounded below. 3 Leaflets cinereous on the lower surface; corolla 6-7 mm long; loment with 3 (-4) segments ...... D. nuttallii 3 Leaflets not cinereous on the lower surface; corolla 3.5-6 mm long; loment with 1-2 (-3) segments; ["Desmodium ciliare group"]. 4 Leaflets 3-5.5× as long as wide...... D. ciliare 4 Leaflets 1.2-3-5× as long as wide. 5 Terminal leaflet usually distinctly longer and narrower than the lateral leaflets; stem (near the middle) sparsely to densely uncinate-pubescent...... D. obtusum 5 Terminal leaflet similar to the lateral leaflets; stem (near the middle) glabrous to pilose, or also with some uncinate-pubescence. 6 Stem usually pilose; leaflets sub-appressed pubescent (to glabrate)...... D. ciliare 6 Stem glabrous (to sparsely uncinate-puberulent); leaflets glabrous or with only a few scattered hairs ...... D. marilandicum 2 Loment with 3-5 segments, mostly obtusely angled below. 7 Leaves densely villous on the lower surface; stem densely pubescent with uncinate or non-uncinate hairs. 8 Leaflets 1.5-2.0 (-2.2)× as long as wide; loment usually curved (the upper margin convex); loment with 2-4 segments; loment segments 4-5 mm long...... D. nuttallii 8 Leaflets 1.0-1.5 (-1.9)× as long as wide; loment straight; loment with (3-) 4-5 (-6) segments; loment segments (4-) 5-8 (-9) mm long ...... D. viridiflorum 7 Leaves glabrous to moderately appressed-villous on the lower surface; stem glabrate, pilose or uncinate pubescent. 9 Bracts (subtending clusters of 2-3 flowers) usually villous; plants moderately to densely villous; loment usually incurved (the upper margin convex); loment with 2-4 segments, each segment 4-5 mm long...... D. nuttallii 9 Bracts (subtending clusters of 2-3 flowers) not villous; plants glabrous or slightly to moderately villous or pilose; loment usually nearly straight; loment with 3-5 segments, each segment 4-8.5 mm long. 10 Corolla 8-10 mm long; pedicels usually 10-15 (-20) mm long; stems and leaves glabrous; leaflets distinctly pale on the lower surface ...... D. laevigatum 10 Corolla 6-8 (-9) mm long; pedicels 3-12 mm long; stems and leaves pubescent or glabrate (but pubescent at least on the leaves); leaflets green or slightly pale on the lower surface; ["Desmodium paniculatum group"]. 11 Leaflet lower surface glabrous, except for the conspicuous uncinate puberulence on the veins; stems and petioles glabrous or uncinate-puberulent; [plant of the Coastal Plain and possibly lower Piedmont] ...... D. fernaldii 11 Leaflet lower surface strigose to conspicuously sub-appressed-villous, and sometimes also uncinate-puberulent; stems and petioles glabrate to conspicuously pilose or uncinate-puberulent; [plants collectively widespread in our area]. 12 Leaflets (2.5-) 3-8 (-10)× as long as wide; leaflet pubescence usually sparse, of straight, appressed hairs < 0.5 mm long (or sometimes of longer spreading hairs); leaflets usually lacking uncinate pubescence on either surface; mid-stems glabrous or glabrate, the pubescence usually uncinate puberulence. 13 Loment segments rounded on the lower margin (thus semicircular to gibbous); leaves subsessile to short-petiolate; [plant restricted to Coastal Plain]...... D. paniculatum var. epetiolatum 13 Loment segments angled on the lower margin (thus triangular to sub-rhombic); leaves long-petiolate; [plant widespread in our area] ...... D. paniculatum var. paniculatum 12 Leaflets 1.5-3 (-4)× as long as wide; leaflet pubescence usually evident, of spreading hairs > 0.5 mm long; leaflets usually with uncinate pubescence on the veins of the upper surface; mid- stems pubescent, either pilose or with uncinate pubescence (if not, evidently pubescent on the petioles). 14 Stem and petiole pubescence sparsely to densely uncinate-puberulent; upper surface of leaflets commonly uncinate-puberulet on the veins...... D. glabellum 14 Stem and petiole pubescence pilose; upper surface of leaflets occasionally uncinate- puberulent on the veins...... D. perplexum

Desmodium canadense (Linnaeus) Augustin de Candolle, Showy Tick-trefoil, Canadian Tick-trefoil. Mt (NC?, VA), Pd (VA): {insert habitat}; rare (VA Rare). July-September; August-October. Québec and Nova Scotia west to Alberta, south to n. FABACEAE 371 VA, sw. VA, NC (?), MO, and OK. Small (1933) reports this species for NC; the documentation is not known. [= C, F, G, I, K, SE, W; = Meibomia canadensis (Linnaeus) Kuntze – S] Desmodium canescens (Linnaeus) Augustin de Candolle, Hoary Tick-trefoil. Pd, Mt, Cp (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-August; August-October. MA west to WI and NE, south to FL and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia canescens (Linnaeus) Kuntze – S] Desmodium ciliare (Muhlenberg ex Willdenow) Augustin de Candolle. Cp, Pd, Mt (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. MA west to IN, MO, and se. KS, south to FL and TX; also in Cuba. [= RAB, C, G, I, SE, W; > D. ciliare var. ciliare – F, K; > D. ciliare var. lancifolium Fernald – F, K; = Meibomia ciliaris (Muhlenberg ex Willdenow) Blake – S] Desmodium cuspidatum (Muhlenberg ex Willdenow) Augustin de Candolle ex Loudon var. cuspidatum, Toothed Tick- trefoil. Mt, Pd, Cp (NC, SC, VA): fields, woodland borders, disturbed areas; uncommon (VA Rare). June-August; August- October. VT and MA west to MI and WI, south to GA and OK. [= C, F, G, K, SE; < D. cuspidatum – RAB, I, W; = Meibomia grandiflora (Augustin de Candolle) Kuntze – S] Desmodium fernaldii Schubert, Fernald's Tick-trefoil. Cp (NC, SC, VA), Pd? (SC): sandhills, dry flatwoods, woodland borders; common (VA Watch List). June-September; August-October. Se. VA south to s. SC (and maybe e. GA); Isely (1998) states that reports from the Gulf Coast are based on "glabrate forms of D. glabellum," and also suggests that D. fernaldii is only weakly differentiated from D. glabellum. [= RAB, C, F, G, I, K, SE, W; < Meibomia rhombifolia Vail – S (also see D. floridanum)] Desmodium floridanum Chapman, Florida Tick-trefoil. Cp (GA, SC): sandhills, other dry sandy habitats; uncommon. June-September; August-October. Se. SC south to FL. [= RAB, I, K, SE; < Meibomia rhombifolia Vail – S (also see D. fernaldii)] Desmodium glabellum (Michaux) Augustin de Candolle. Cp, Pd, Mt (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. ME west to WI and NE, south to FL and TX. [= RAB, F, I, K, SE; < D. glabellum – C (also see D. perplexum); ? Meibomia paniculata (Linnaeus) Kuntze – S, in part; ? Meibomia pubens (Torrey & A. Gray) Rydberg – S (also see D. paniculatum var. paniculatum); < D. paniculatum var. dillenii (Darlington) Isely – W] Desmodium glutinosum (Muhlenberg ex Willdenow) A. Wood, Heartleaf Tick-trefoil, Clusterleaf Tick-trefoil. Mt, Pd, Cp (NC, SC, VA): moist forests, especially nutrient-rich; uncommon. June-August; August-October. Nova Scotia west to Saskatchewan, south to FL and Mexico. [= RAB, C, F, G, I, K, SE, W; = Meibomia acuminata (Michaux) Blake – S] {Desmodium humifusum (Muhlenberg ex Bigelow) Beck. MA (Nova Scotia?) south to MD and DC (and possibly VA). Perhaps only a hybrid. [= C, F, I, K, SE; D. glabellum – G, misapplied; Meibomia glabella – S, misapplied]} Desmodium laevigatum (Nuttall) Augustin de Candolle. Cp, Pd, Mt (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. S. NY west to IN and MO, south to FL and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia laevigata (Nuttall) Kuntze – S] Desmodium lineatum Augustin de Candolle, Matted Tick-trefoil. Cp (NC, SC, VA), Pd (NC): sandhills and other dry forests and woodlands; common (VA Watch List). June-August; August-October. Se. MD south to FL, west to TX. [= RAB, C, F, G, I, K, SE, W; > Meibomia arenicola Vail – S; > Meibomia polymorpha (A. Gray) Small – S] Desmodium marilandicum (Linnaeus) Augustin de Candolle. Mt, Pd, Cp (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. MA west to MI and MO, south to FL and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia marilandica (Linnaeus) Kuntze – S] Desmodium nudiflorum (Linnaeus) Augustin de Candolle, Naked Tick-trefoil. Cp, Pd, Mt (NC, SC, VA): moist to dry forests; common. July-August; August-October. ME west to MN, south to FL and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia nudiflora (Linnaeus) Kuntze – S] Desmodium nuttallii (Schindler) Schubert. Cp, Pd, Mt (NC, SC, VA): fields, woodland borders, disturbed areas; common. July-September; August-October. NY west to IN, south to n. FL, AL, and AR. [= RAB, F, I, K, SE, W; < D. viridiflorum – C, G; < Meibomia viridiflora (Linnaeus) Kuntze – S (also see D. viridiflorum)] Desmodium obtusum (Muhlenberg ex Willdenow) Augustin de Candolle. Mt, Pd, Cp (NC, SC, VA): dry pine woodlands, fields, woodland borders, disturbed areas; common. June-September; August-October. MA west to s. MI, south to FL and TX. [= RAB, I, K, SE, W; = D. rigidum (Elliott) Augustin de Candolle – C, F, G; = Meibomia rigida (Elliott) Kuntze – S] Desmodium ochroleucum M.A. Curtis ex Canby, White Tick-trefoil, Creamflower Tick-trefoil. Pd (NC, VA), Cp (VA), Mt (GA, NC): dry woodlands, especially over calcareous soils; rare (GA Special Concern, NC Rare, VA Rare). June-August; August-October. NJ (?), DE, and MD south to sc. and sw. NC, GA, TN, AL, FL, MS, and MO. [= RAB, C, F, G, I, K, SE, W; = Meibomia ochroleuca (M.A. Curtis ex Canby) Kuntze – S] Desmodium paniculatum (Linnaeus) Augustin de Candolle var. epetiolatum Schubert. Cp (NC, SC?, VA): pine savannas and flatwoods, bogs; uncommon? (VA Watch List). June-September; August-October. Var. epetiolatum ranges from se. VA south to se. NC or e. SC. It may reflect hybridization between D. paniculatum var. paniculatum and another species. Further study is needed. [= F, I, K, SE; < D. paniculatum – RAB, C; < D. paniculatum var. paniculatum – W] Desmodium paniculatum (Linnaeus) Augustin de Candolle var. paniculatum. Mt, Pd, Cp (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. S. ME west to s. Ontario, MI, and NE, south to FL and TX. [= F, I, K, SE; < D. paniculatum – RAB, C; > Meibomia chapmanii (Britton) Small – S; = D. paniculatum var. pubens Torrey & A. Gray – G; > Meibomia paniculata (Linnaeus) Kuntze – S; >< Meibomia pubens (Torrey & A. Gray) Rydberg – S (also see D. glabellum); < D. paniculatum var. paniculatum – W] Desmodium pauciflorum (Nuttall) Augustin de Candolle, Few-flowered Tick-trefoil. Cp, Pd (NC, SC, VA), Mt (VA): moist forests; common (NC Watch List). June-August; August-October. NY west to OH and IA, south to FL and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia pauciflora (Nuttall) Kuntze – S] FABACEAE 372 Desmodium perplexum Schubert. Mt, Pd, Cp (NC, SC, VA): fields, woodland borders, disturbed areas; common. July- September; August-October. [= RAB, F, I, K, SE; < D. glabellum – C; ? Meibomia dillenii (Darlington) Kuntze – S; < D. paniculatum var. dillenii (Darlington) Isely – W] Desmodium rotundifolium Augustin de Candolle, Roundleaf Tick-trefoil. Mt, Pd, Cp (NC, SC, VA): dry forests and woodlands; common. June-August; August-October. VT and MA west to s. MI, south to GA, LA, and MO. [= RAB, C, F, G, I, K, SE, W; ? Meibomia michauxii Vail – S] Desmodium sessilifolium (Torrey) Torrey & A. Gray, Sessile-leaf Tick-trefoil. Pd (NC, SC, VA), Mt (VA), Cp (GA, SC?): dry woodlands; rare (NC Rare, VA Rare). July-August; August-October. RI west to s. MI and KS, south to NC, MS, and TX. [= RAB, C, F, G, I, K, SE, W; = Meibomia sessilifolia (Torrey) Kuntze – S] Desmodium strictum (Pursh) Augustin de Candolle, Pineland Tick-trefoil, Pinebarren Tick-trefoil. Cp (NC, SC, VA), Pd (NC, SC): sandhills, other dry wodlands; common (VA Rare). July-August; August-October. NJ south to FL, west to LA. [= RAB, C, F, G, I, K, SE, W; = Meibomia stricta (Pursh) Kuntze – S] Desmodium tenuifolium Torrey & A. Gray, Slimleaf Tick-trefoil. Cp (NC, SC, VA), Pd? (NC?): bogs, pine savannas, wet pine flatwoods; common (VA Rare). July-August; August-October. Se. VA south to FL, west to LA. [= RAB, C, F, G, I, K, SE; = Meibomia tenuifolia (Torrey & A. Gray) Kuntze – S] Desmodium tortuosum (Swartz) Augustin de Candolle. Cp (NC, SC): fields, woodland borders, disturbed areas; common. July-August; August-October. E. NC south to FL, west to TX. [= RAB, I, K, SE; = Meibomia purpurea (P. Miller) Vail – S] Desmodium viridiflorum (Linnaeus) Augustin de Candolle. Pd, Cp, Mt (NC, SC, VA): fields, woodland borders, disturbed areas; common. June-September; August-October. DE south to FL, west to TX, and inland to w. VA, w. NC, n. TN, and AR. [= RAB, F, I, K, SE, W; < D. viridiflorum – C, G (also see D. nuttallii); < Meibomia viridiflora (Linnaeus) Kuntze – S (also see D. nuttallii)]

Desmodium cuspidatum (Muhlenberg ex Willdenow) Augustin de Candolle ex Loudon var. longifolium (Torrey & A. Gray) Schubert, in GA. Var. longifolium (Torrey & A. Gray) Schubert, differing in its pubescent stem, leaves, stipules, and calyx (vs. nearly glabrous), is generally more western, ranging from OH west to MN and NE, south to GA and ne. TX. [= C, F, G, K, SE; < D. cuspidatum – RAB, I] {not keyed at this time} Desmodium incanum Augustin de Candolle. Cp (GA): lawns, disturbed areas; rare, presumably introduced or adventive from further south. A pantropical weedy species. [= I, SE; > D. incanum var. incanum – K; = Meibomia cana (J.F. Gmelin) Blake – S, illegitimate basionym; = D. canum (J.F. Gmelin) Schinz & Thellung, illegitimate basionym] {not keyed at this time}

Dioclea Kunth

A genus of about 30 species, perennial to woody vines, of tropical (rarely temperate) regions of the Old and New World. References: Isely (1998)=I.

Dioclea multiflora (Torrey & A. Gray) C. Mohr. Cp (GA): alluvial forests; rare. S. GA and FL Panhandle west to e. TX, north in the interior to w. TN and w. KY. [= C, G, I, K, S, SE; = Galactia mohlenbrockii R.H. Maxwell]

Erythrina Linnaeus (Coral Bean)

A genus of about 100 species, trees, shrubs, and perennial herbs, of tropical and subtropical regions of the Old and New World. References: Isely (1998)=I.

1 Leaflets not lobed; [cultivated tree, persistent] ...... E. crista-galli 1 Leaflets hastately lobed; [native herb or shrub] ...... E. herbacea

* Erythrina crista-galli Linnaeus, Coraltree. Cp (GA): cultivated, disturbed areas, roadside ditches; rare, introduced from South America. [= I, K, SE; = Micropteryx crista-galli (Linnaeus) Walpers – S] Erythrina herbacea Linnaeus, Coral Bean, Cardinal-spear. Cp (GA, NC, SC): maritime forests, dry sandy woodlands, sandhills in the outer Coastal Plain; uncommon (rare in NC). May-July; July-September. Se. NC south to FL, west to se. TX, and south to e. Mexico (Tamaulipas and e. San Luis Potosi). [= RAB, I, K, SE; > E. herbacea – S; > E. arborea (Chapman) Small – S]

Galactia P. Browne (Milkpea) (also see Dioclea)

A genus of about 50 species, herbs, of tropical and warm temperate regions, primarily American. References: Duncan (1979)=Z; Isely (1998)=I; Ward & Hall (2004)=Y.

Identification notes: Definite identification of the taxa from key lead 4 on is problematic. Note also that the traditional application of G. regularis and G. volubilis is reversed. FABACEAE 373

1 Leaves with (5-) 7 (-9) leaflets; flowers white...... G. elliottii 1 Leaves with 3 leaflets; flowers white, pink, red, or purple. 2 Plant erect, with 4-6 (-8) leaves ...... G. erecta 2 Plant prostrate, trailing, or twining, generally with numerous leaves. 3 Legumes villous with hairs 1-1.5 mm long; corolla when fresh bright reddish purple and white (drying dark on herbarium specimens); stems villous with hairs > 0.5 mm long...... G. mollis 3 Legumes glabrous, or pubescent with hairs < 1 mm long; corolla pink to pink-purple (drying pale or the petals dropping on herbarium specimens); stems glabrate to villosulous with hairs < 0.5 mm long. 4 Inflorescences generally exserted, (2-) 4-15 (-20) cm long, the flowers well distributed along half or more of the length of the inflorescence axis; flower buds generally 5-8 mm long; corolla 8-12 (-14) mm long; plants twining (rarely trailing) ...... G. regularis 4 Inflorescences short or exserted, if exserted then the flowers generally from nodes crowded into the upper half of the axis; flower buds 5-10 mm long; corolla (11-) 12-16 (-18) mm long; plants trailing or twining. 5 Internodes short, 1-2 cm long, the stem thus appearing leafy; inflorescences with 1-3 flowers (or also with solitary axillary flowers); [plants of Coastal Plain of SC southward]...... G. minor 5 Internodes generally longer; inflorescences with 1-3 or more flowers; [plants collectively widespread in our area]. 6 Stems glabrescent to conspicuously pubescent with spreading hairs < 0.5 mm long; leaflets 1.5-3.5 (-4) cm long, the undersurface with hairs 0.4-0.7 mm long; [plants of e. GA southward] ...... G. floridana 6 Stems antrorse- or retrorse-strigose; leaflets 1.5-5 (-7) cm long, the undersurface strigose with hairs < 0.3 mm long; [plants widespread in our area] ...... G. volubilis var. volubilis

Galactia elliottii Nuttall, Elliott's Milkpea. Cp (GA, SC): moist forests; uncommon. July-September; August-October. S. SC south to s. FL. [= RAB, I, K, S, SE, Y, Z] Galactia erecta (Walter) Vail, Erect Milkpea. Cp (GA, NC, SC): sandhills; common. May-July; July-September. Se. NC south to Panhandle FL, west to e. TX. [= RAB, I, K, S, SE, Y, Z] Galactia floridana Torrey & A. Gray, Florida Milkpea. Cp (GA): sandhills and other xeric sands; rare (GA Special Concern). S. GA south to s. FL, west to s. MS. [= Y; < G. floridana – I, K, SE, Z (also see G. volubilis var. fasciculata); = G. floridana var. floridana – S] Galactia minor Duncan, Little Milkpea. Cp (GA, NC, SC): sandhills; uncommon. June-August; July-October. Sc. NC south to Panhandle FL, west to s. MS. [= Y, Z; < G. regularis (Linnaeus) Britton, Sterns, & Poggenburg – RAB, misapplied; = G. microphylla (Chapman) H.J. Rogers ex Isely – I, K, SE; = G. floridana Torrey & A. Gray var. microphylla Chapman – S] Galactia mollis Michaux. Cp (GA, NC, SC): sandhills; uncommon (rare in NC and SC). May-July; July- September. Se. NC south to c. peninsular FL, west to Panhandle FL and se. AL. [= RAB, I, K, S, SE, Y, Z] Galactia regularis (Linnaeus) Britton, Sterns, & Poggenburg. Cp, Pd, Mt (GA, NC, SC, VA): dry forests and woodlands; common. July-September; August-October. Se. PA west to MO and OK, south to s. FL and se. TX. [= Y, Z; > G. volubilis (Linnaeus) Britton – RAB, C, F, G, misapplied; > G. macreei M.A. Curtis – RAB, C, F, G; = G. volubilis – I, K, S, SE, misapplied] Galactia volubilis (Linnaeus) Britton var. volubilis. Cp (GA, NC, SC, VA), Pd?, Mt? (VA): sandhills, other dry forests and openings; common. June-August; July-October. NJ and s. PA west to c. AR, south to s. FL and LA. Var. baltzelliana D.B. Ward & D.W. Hall and var. fasciculata (Vail) D.B. Ward & D.W. Hall are endemics of the FL Peninsula. Duncan (1979) describes additional forms of this taxon (which he treated under the name G. glabella) that he considered to potentially warrant description as varieties or species; they need further study. [= Y; < G. regularis (Linnaeus) Britton, Sterns, & Poggenburg – RAB, C, F, G, I, K, SE, misapplied; > G. regularis – S, misapplied; > G. brevipes Small – S; < G. glabella Michaux – Z]

Genista Linnaeus (Dyer's Greenweed)

A genus of about 80 species, shrubs and small trees, native to Eurasia. References: Isely (1998)=I.

* Genista tinctoria Linnaeus, Dyer's Greenweed. Cp (VA): disturbed areas; rare, introduced from Europe. June-September. Not cited in Harvill et al. (1992), but described as naturalized in sterile soils south to VA in C, F, and G. [= C, F, G, I, K]

Gleditsia Linnaeus (Honey Locust, Water Locust)

A genus of 13-14 species, scattered in the Old and New Worlds. References: Isely (1975)=Z; Robertson & Lee (1976)=Y; Isely (1998)=I; Schnabel & Wendel (1998).

1 Legume ovate, 3-5 (-8) cm long and 1-3-seeded; foliage glabrous (or slightly puberulent when young); [trees of frquently flooded swamps, often with Taxodium, rarely planted and escaped] ...... G. aquatica FABACEAE 374 1 Legume elongate, 20-40 cm long and multi-seeded; foliage puberulent (even in age); [trees of moist to dry forests, frequently planted and escaped in disturbed areas]...... G. triacanthos

Gleditsia aquatica Marshall, Water Locust. Cp (GA, SC): swamp forests; common. April-May; July-November. E. SC south to c. peninsular FL, west to TX, and north in the interior to IN, IL, and MO; occasionally cultivated north of its native range. [= RAB, C, F, G, GW, I, K, S, SE, Y, Z] Gleditsia triacanthos Linnaeus, Honey Locust. Pd, Mt, Cp (GA, NC, SC, VA): woodlands, forests (generally bottomland), fencerows, often planted as a street tree; common (uncommon in Mountains). April-May; July-November. NY west to SD, south to panhandle FL and TX. Its occurrence over much of our region appears to be as an adventive; the native range is poorly known. G. triacanthos is more likely to be native in the western part of our region, particularly in the Mississippi drainage. The trunks are normally beset with lengthy, branched thorns, but thornless trees are encountered (and are usually favored for horticultural planting). [= RAB, C, G, GW, I, K, S, SE, W, Y, Z]

The hybrid Gleditsia ×texana Sargent (pro sp.) [G. aquatica × triacanthos] occurs occasionally in the area of range overlap of its parents. Isely (1975) reports its occurrence in SC. It is intermediate between its parents. [= I, K]

Glottidium Desvaux (Bladderpod)

A genus of a single species, a coarse annual herb, of se. North America. References: Isely (1998)=I.

Identification notes: See key under Sesbania for distinctions.

Glottidium vesicarium (Jacquin) Harper, Bladderpod, Bagpod. Cp (GA, NC, SC), Pd (GA): ditches, marshes, disturbed wet areas; common. The native status of G. vesicarium is uncertain; its distribution is from ne. NC south to s. FL, west to e. OK and se. TX, and Isely (1998) states that it is unknown from outside the United States. Its weedy habit suggests that it may have expanded its distribution northward into our area from the deeper South. July-September; August-November. [= RAB, I, K, S, SE; = Sesbania vesicaria (Jacquin) Elliott – GW]

Glycine Linnaeus (Soybean, Soya)

A genus of about 10 species, annual and perennial herbs, of Asia and Australia. References: Isely (1998)=I.

* Glycine max (Linnaeus) Merrill, Soybean. Cp, Pd, Mt (GA, NC, SC, VA): abundantly cultivated, rarely persisting as a waif, introduced from e. Asia. July-October. One of the most important legume crops in the world. [= RAB, F, I, K, SE]

Glycyrrhiza Linnaeus (Licorice)

References: Isely (1998)=I.

* Glycyrrhiza lepidota Pursh, Wild Licorice, American Licorice. Cp (VA): disturbed areas; rare, introduced from w. North America. Not cited in Harvill et al. (1992), but described as naturalized in old fields in e. VA in C, F, and G. [= K; > G. lepidota Pursh var. glutinosa (Nuttall) S. Watson – C, F, G, I]

Gymnocladus Lamarck (Kentucky Coffee-tree)

A genus of 5 species, all trees, ours in e. North America and four species in e. Asia. References: Isely (1975)=Z; Robertson & Lee (1976)=Y; Lee (1976)=X; Isely (1998)=I.

Gymnocladus dioicus (Linnaeus) K. Koch, Kentucky Coffee-tree. Mt (GA*, VA), Pd* (GA*, NC*): native in rich bottomland and slope forests, also in disturbed areas, persistent and weakly spreading from horticultural plantings; rare (VA Watch List), also rare east of the Appalachians as an introduction. April-May; August-November (and persistent). The original native range has been obscured, perhaps PA west to se. SD, south to w. VA, TN, n. AL, and OK. [= RAB, C, F, G, I, K, S, SE, X, Y, Z]

Indigofera Linnaeus (Indigo)

A genus of about 700 species, annual herbs, perennial herbs, and shrubs, nearly cosmopolitan in tropical and warm temperate regions. References: Isely (1998)=I.

FABACEAE 375 1 Stem pubescence hirsute or pilose with long brownish hairs...... I. hirsuta 1 Stem pubescence strigose-appressed. 2 Legume 7-9 mm long, ovoid, not falcate, indehiscent, with 2-3 seeds; corolla 6-9 mm long; [native species] ...... I. caroliniana 2 Legume 15-36 mm long, linear-cylindric, slightly to strongly falcate, dehiscent, with 3-12 or more seeds; corolla either 5-6 mm long or 15-18 mm long; [introduced species]. 3 Corolla 15-18 mm long; legume 30-40 mm long, straight; leaflets 2.5-4 cm long...... I. decora 3 Corolla 5-6 mm long; legume 15-36 mm long, slightly to strongly falcate; leaflets (0.5-) 1-3 cm long. 4 Legume 15-20 mm long, strongly falcate...... I. suffruticosa 4 Legume 28-36 mm long, slightly falcate...... I. tinctoria

Indigofera caroliniana P. Miller, Wild Indigo, Carolina Indigo. Cp (GA, NC, SC): sandy forests and woodlands, including sandhills and sandy maritime forests; common (uncommon in NC and SC). June-August; July-October. E. NC south to FL, west to se. LA, a Southeastern Coastal Plain endemic. [= RAB, I, K, S, SE] * Indigofera decora Lindley, Chinese Indigo. Pd (GA): planted horticulturally and spreading to nearby roadbanks; rare (but potentially invasive), native of China. June-July (-September). In GA (Oglethorpe County). * Indigofera hirsuta Linnaeus, Hairy Indigo. Cp (GA, SC): sandy disturbed areas, such as wildlife "food fields"; rare, introduced from the Old World tropics. First reported for SC by Nelson & Kelly (1997). Also known from other scattered locations in the Southeast, such as s. MS (Leonard, 2006, pers.comm.). [= I, K, SE] * Indigofera suffruticosa P. Miller, West Indian Indigo. Cp (GA, NC, SC): disturbed areas, dry sandy woodlands; rare, formerly commonly cultivated, locally established as a weed at that time, perhaps no longer present in our area, introduced from native range in the New World tropics, including s. FL. [= I, K, S, SE] * Indigofera tinctoria Linnaeus, African Indigo. Cp (GA, NC, SC): formerly commonly cultivated, locally established as a weed at that time, perhaps no longer present in our area, introduced from Africa. Both this species and I. suffruticosa were cultivated as an important export crop in the Coastal Plain of GA, SC, and (less so) NC in the seventeenth and eighteenth centuries. [= I, K, S, SE]

Kummerowia Schindler (Korean-clover, Japanese-clover)

A genus of 2 species, annual herbs, native to temperate e. Asia. Kummerowia differs from Lespedeza in its annual habit (vs. perennial), conspicuous stipules (vs. not conspicuous), inflorescence branching pattern (see Akiyama & Ohba 1985), and leaflets with striate, parallel, lateral veins (vs. with reticulate lateral veins). It is distinct at the generic level. References: Akiyama & Ohba (1985)=Z; Isely (1998)=I. Key based closely on SE.

1 Mid-stem leaves with petioles 4-10 mm long; leaflets emarginate at the apex; leaflets conspicuously spreading-ciliate; stems antrorsely appressed-strigose; calyx covering 1/3-1/2 of the legume...... K. stipulacea 1 Mid-stem leaves with petioles 1-2 (-4) mm long; leaflets not emarginate at the apex; leaflets inconspicuously appressed- ciliate; stems retrorsely appressed-strigose; calyx covering 1/2-4/5 of the legume...... K. striata

* Kummerowia stipulacea (Maximowicz) Makino, Korean Lespedeza, Korean-clover. Mt, Pd, Cp (GA, NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from e. Asia. July-September; August-November. [= I, K, SE, Z; = Lespedeza stipulacea Maximowicz – RAB, C, F, G, W] * Kummerowia striata (Thunberg) Schindler, Japanese-clover, Common Lespedeza. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disurbed areas; common, introduced from e. Asia. July-September; August-November. [= I, K, SE, Z; = Lespedeza striata (Thunberg) Hooker & Arnott – RAB, C, F, S, G, W]

Lablab Adanson (Hyacinth-bean)

A genus of a single species, an annual to perennial herb, native of the Old World tropics. References: Isely (1998)=I.

* Lablab purpureus (Linnaeus) Sweet, Hyacinth-bean, is cultivated and rarely may escape or persist as a waif in disturbed areas; it is reported from se. PA (Rhoads & Klein 1993). [= I, K, SE; = Dolichos lablab Linnaeus]

Lathyrus Linnaeus 1753 (Wild-pea, Vetchling)

A genus of about 150 species, annual and perennial herbs, of nearly cosmopolitan distribution. References: Isely (1998)=I. Key adapted in part from C.

1 Leaflets > 2, generally 4-12; [native species of various habitats]. 2 Foliaceous stipules laterally symmetrical, with 2 approximately equal basal lobes; leaves somewhat fleshy; [plants of ocean beaches and dunes] ...... [L. maritimus var. pellitus] FABACEAE 376 2 Foliaceous stipules asymmetrical, oblique at the base, the basal lobe well-developed only on one side. 3 Basal lobe of stipule broadly rounded at tip; flowers whitish ...... [L. ochroleucus] 3 Basal lobe of stipule acute to acuminate at tip; flowers purplish (rarely whitish, or faded in old specimens). 4 Racemes with 2-6 (-9) flowers; leaflets 4-8 (-10) per leaf; [plants of marshes, bottomlands, and other wet habitats]...... L. palustris 4 Racemes with (5-) 10-20 flowers; leaflets (8-) 10-14 per leaf; [plants of dry to mesic forests] ...... L. venosus 1 Leaflets 0-2; [alien species, except L. pusillus]. 5 Leaflets absent (but with foliaceous stipules) ...... [L. aphaca] 5 Leaflets 2. 6 Stems with wings 0-1 (-2) mm wide; corolla 6-14 mm long; flowers 1-3 (-4) per raceme. 7 Legume (in fruit) and ovary (in flower) hirsute with swollen-based hairs; corolla 10-14 mm long...... L. hirsutus 7 Legume (in fruit) and ovary (in flower) glabrous; corolla 6-9 mm long ...... L. pusillus 6 Stems with wings 1-3 mm wide; corolla 13-30 mm long; flowers 2-12 per raceme. 8 Stems hirsute with swollen-based hairs; plant an annual; flowers 2-4 per raceme ...... L. odoratus 8 Stems glabrate; plant a perennial; flowers (3-) 4-12 per raceme. 9 Stipules 4-10 mm wide; leaflets 2-5× as long as wide...... L. latifolius 9 Stipules 2-3 mm wide; leaflets 6-15× as long as wide...... L. sylvestris

* Lathyrus hirsutus Linnaeus, Caley Pea, Singletary Pea. Cp, Pd (GA, NC, SC, VA), Mt (GA, VA): roadsides, fields, disturbed areas; common, introduced from Europe. April-July. [= RAB, C, F, G, I, K, S, SE, W] * Lathyrus latifolius Linnaeus, Everlasting Pea, Perennial Sweet Pea. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): roadsides, fencerows, disturbed areas; common, introduced from Europe. May-September. [= RAB, C, F, G, I, K, SE, W] * Lathyrus odoratus Linnaeus, Sweet Pea. Cp, Pd, Mt (GA, NC, SC, VA): cultivated, and occasionally persisting; rare, introduced from s. Europe. [= C, F, G, I, K, SE] Lathyrus palustris Linnaeus, Marsh Pea, Marsh Vetchling. Pd (GA, VA), Mt (VA), Cp (NC): bottomland forests, marshes, streambanks; rare (GA Special Concern). May-June; July-September. Circumboreal, ranging in North America south to NJ, VA, ne. NC, ec. GA, OH, IN, MO, CO, and CA. [= RAB, I, K, SE; > L. palustris var. palustris – C, F, G; > L. palustris var. myrtifolius (Muhlenberg ex Willdenow) A. Gray – F, G; > L. palustris var. linearifolius Seringe – G; > L. myrtifolius Muhlenberg ex Willdenow – S] Lathyrus pusillus Elliott. Cp (NC, VA): open areas in bottomlands; rare. April-July. E. VA, MO and KS south to FL and TX. [= RAB, F, G, I, K, S, SE] * Lathyrus sylvestris Linnaeus, Perennial Pea. Pd (GA) (NC?, SC?, VA): cultivated, and occasionally persisting; rare, introduced from Europe. [= C, F, G, I, K, SE] Lathyrus venosus Muhlenberg ex Willdenow, Forest Pea, Bush Vetch. Mt, Pd (GA, NC, VA), Cp (VA): dry to mesic slope and bottomlands forests and woodlands, especially in base-rich soils; common. July-September. S. Ontario west to MN and Saskatchewan, south to c. NC, wc. GA, and MO. [= RAB, I, K, S, SE, W; > L. venosus var. venosus – C, F, G; > L. venosus var. intonsus Butters & St. John – C, F, G; > L. venosus var. meridionalis Butters & St. John – F]

* Lathyrus aphaca Linnaeus, Yellow Vetchling, introduced from Eurasia, is scattered in occurrence in the Southeast, including AL, TN, and KY (Kartesz 1999). [= G, I, K, SE] Lathyrus maritimus (Linnaeus) Bigelow var. pellitus (Fernald) Gleason, Beach Pea, has been reported from ocean beaches in Dare County (NC). Documentation of the occurrence of this species in our area is needed. [= C, G; = Lathyrus japonicus Willdenow var. pellitus Fernald – F, K; < L. japonicus – I] Lathyrus ochroleucus Hooker, White Pea. Forests over limestone. May-July. South sc. PA (Rhoads & Klein 1993) and NJ (Kartesz 1999). [= C, F, G, I, K] * Lathyrus pratensis Linnaeus, Meadow Pea. Reported for VA on the basis of "personal communication" (Kartesz 1999). {investigate} [= C, F, I, K] {not keyed at this time} * Lathyrus tuberosus Linnaeus. Introduced in e. TN (Chester, Wofford, & Kral 1997), WV (Kartesz 1999), and at scattered locations in PA (Rhoads & Klein 1993). [= C, F, G, I, K, SE] {not keyed at this time}

Lespedeza Michaux 1803 (Lespedeza) (also see Kummerowia)

A genus of about 40 species, perennial herbs and shrubs, of temperate regions of e. Asia and e. North America. References: Clewell (1966a)=Z; Clewell (1966b)=Y; Isely (1986b)=X; Akiyama (1988)=Q; Clewell & Stickel (1990); Isely (1998)=I. Key based primarily on Z and SE.

1 Plants annual; stipules ovate to ovate-lanceolate, conspicuous; leaflets with striate, parallel, lateral veins...... [Kummerowia] 1 Plants perennial, stipules subulate, setaceous, or lanceolate, not conspicuous; leaflets with reticulate lateral veins, joining before reaching the margin. 2 Plant a shrub, usually 1-3 m tall, bushy-branched, the woody stems over-wintering; corolla 8-15 mm long; [plants alien, planted in "wildlife food plots" and persisting or spreading]; [section Macrolespedeza). FABACEAE 377 3 Calyx lobes equal to or shorter than the calyx tube; corolla 8-11 mm long; leaflets 1.5-2× as long as wide; racemes erect or strongly ascending; stems 1-several per crown, brown when young...... L. bicolor 3 Calyx lobes longer than the calyx tube (especially or at least the lowest lobe); corolla (10-) 12-15 mm long; leaflets 2-3× as long as wide; racemes lax and drooping; stems many per crown, purplish when young ...... L. thunbergii 2 Plant an herb, 0.1-2 m tall, not bushy-branched (the taller species often wand-like), the stems in some species somewhat woody but dying back; corolla to 10 mm long; [plants native, except L. cuneata and L. virgata). 4 Leaflets distinctly widest toward the tip, 3-5× as long as wide, the base and apex very differently shaped (the base narrowly cuneate, the tip rounded, truncate or even retuse); racemes reduced, with 2-3 flowers, shorter than the subtending leaves; [plants alien] ...... L. cuneata 4 Leaflets generally widest near the middle, 1-8 (-10)× as long as wide, the base and apex shaped similarly (i.e. both rounded, or both cuneate); racemes with 3-many flowers, shorter or longer than the subtending leaves; [plants native, except L. virgata]. 5 Midrib of leaflets distinctly excurrent as a spinose bristle 0.5-1.5 mm long; [plant a rare introduction] ...... L. virgata 5 Midrib of leaflets not excurrent, or only as an obscure mucro, not at all spinose; [plants native]. 6 Plants trailing at maturity (young stems erect to arching-ascending up to 2 dm tall, then lopping over); stems slender, wiry; corolla pink to purple. 7 Pubescence of the stem spreading (pilose) ...... L. procumbens 7 Pubescence of the stem appressed (strigose). 8 Calyx of legumes produced from cleistogamous flowers 1/4-1/3 as long as the pod; stems usually lacking axillary leaves; keel subequal to the wings, or shorter; stipules 2-4 (-5) mm long...... L. repens 8 Calyx of legumes produced from cleistogamous flowers ca. 1/5 as long as the pod; stems often with axillary leaves distinctly smaller than the primary leaves; keel usually longer than the wings; stipules 3-5 (-6) mm long...... L. frutescens 6 Plants erect at maturity; stems generally stout, stiff; corolla pink, purple, white, cream, or mixed. 9 Plants in flower. 10 Corolla primarily white or cream (often with a purplish throat). 11 Raceme peduncles short (shorter than the subtending leaf), the inflorescence itself barely if at all exceeding the subtending leaf; calyx lobes 6-10 mm long; leaflets (2-) 2.5-5 (-8)× as long as wide...... L. capitata 11 Raceme peduncles elongate (often longer than the subtending leaf), the inflorescence itself well-exserted beyond the subtending leaf; calyx lobes 3-7 mm long; leaflets either narrower or wider (see below). 12 Leaflets 4-8 (-10)× as long as wide...... L. angustifolia 12 Leaflets 1.3-1.8× as long as wide. 13 Leaves closely strigose on both surfaces with hairs 0.2-0.5 mm long, silvery when fresh; leaflets 1-2 cm long; petiole of midstem leaves not generally > 1 cm long, about the same length as the rachis; [plants of the Coastal Plain and, in NC and SC, the lower Piedmont]...... L. hirta var. curtissii 13 Leaves glabrate, or strigose above only, at least some of the hairs (especially those on the veins below) > 0.5 mm long, green or grey (to somewhat silvery) when fresh; leaflets 1.5-4 (-5) cm long; petiole of midstem leaves 1-1.5 (-2) cm long, much exceeding the rachis; [plants widespread in our area] ...... L. hirta var. hirta 10 Corolla primarily pink or purple. 14 Peduncles of the racemes of chasmogamous (petaliferous) flowers longer than the subtending leaves; keel 1-2 mm longer than the wings ...... L. frutescens 14 Peduncles of the racemes of chasmogamous (petaliferous) flowers shorter than the subtending leaves; keel about as long as or shorter than the wings. 15 Upper surface of the leaflets glabrous (sometimes strigose along the midrib only); pubescence of the stem appressed; leaflets 1.5-3× as long as wide ...... L. violacea 15 Upper surface of the leaflets pubescent; pubescence of the stem appressed or spreading; leaflets 1.3-7× as long as wide. 16 Leaflets 1.3-3 (-3.5)× as long as wide...... L. stuevei 16 Leaflets (4-) 5-7× as long as wide...... L. virginica 9 Plants not in flower. 17 Leaflets of average, mid-stem leaves > 4× as long as wide (L. capitata keyed here and below). 18 Petioles of mid-stem leaves ca. 10 mm long...... L. virginica 18 Petioles of mid-stem leaves 1-3 mm long. 19 Leaflets 4-8(-10)× as long as wide; pubescence of the stems and leaves usually not silvery-cinereous...... L. angustifolia FABACEAE 378 19 Leaflets (2-) 2.5-5 (-8)× as long as wide; pubescence of stems and leaves usually silvery-cinereous...... L. capitata 17 Leaflets of average, mid-stem leaves < 3.5× as long as wide (L. capitata keyed here and above). 20 Leaflets (2-) 2.5-5 (-8)× as long as wide; leaf rachis (the apparent petiolule of the terminal leaflet) longer than the petiole ...... L. capitata 20 Leaflets 1.3-3 (-3.5)× as long as wide; leaf rachis shorter than the petiole (or about equal in L. hirta var. curtissii). 21 Central axis not strongly dominant, branches divaricate, irregular; stems slender, wiry ...... L. frutescens 21 Central axis strongly dominant, branches ascending, mostly on the upper stem; stems stout, stiff. 22 Leaflets (1.3-) 1.8-3 (-3.5)× as long as wide. 23 Upper surface of the leaflets glabrous (sometimes strigose along the midrib only); pubescence of the stem appressed...... L. violacea 23 Upper surface of the leaflets pubescent; pubescence of the stem appressed or spreading ...... L. stuevei 22 Leaflets 1.3-1.8× as long as wide. 24 Leaves closely strigose, the hairs 0.2-0.5 mm long, silvery when fresh; leaflets 1-2 cm long; [plants of the Coastal Plain and, in NC and SC, the lower Piedmont]...... L. hirta var. curtissii 24 Leaves glabrate, or strigose above only, at least some of the hairs > 0.5 mm long, green or grey when fresh; leaflets 1.5-4 (-5) cm long; [plants widespread in our area]...... L. hirta var. hirta

Lespedeza angustifolia (Pursh) Elliott, Narrow-leaved Lespedeza. Cp (GA, NC, SC, VA), Pd (GA), Mt (NC): sandhill- pocosin ecotones and dry to moist savannas, mountain bogs; common. August-October; September-November. MA south to c. FL, west to s. MS, essentially a Southeastern Coastal Plain endemic, rarely disjunct inland to w. NC, c. GA, and ec. TN (Chester, Wofford, & Kral 1997). [= RAB, C, G, I, K, S, SE, W, Y, Z; > L. angustifolia – F; > L. hirta var. intercursa Fernald – F] * Lespedeza bicolor Turczaninow, Bicolor Lespedeza. Pd, Cp, Mt (GA, NC, SC, VA): "wildlife food plots", roadsides; common, introduced from e. Asia. June-September; August-November. [= RAB, C, I, K, Q, S, SE, W] Lespedeza capitata Michaux, Bush-clover. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): woodlands and woodland borders; common. August-October; September-November. ME and s. Ontario west to MN, SD, and NE, south to FL and TX. [= RAB, C, I, K, S, SE, W, Y, Z; > L. capitata var. capitata – F, G; > L. capitata var. stenophylla Bissell & Fernald – F, G; > L. capitata var. velutina (Bicknell) Fernald – F, G; > L. capitata var. vulgaris Torrey & A. Gray – F] * Lespedeza cuneata (Dumont-Cours.) G. Don, Sericea Lespedeza, Chinese Lespedeza. Cp, Pd, Mt (GA, NC, SC, VA): roadbanks, "wildlife food plots", disturbed areas; common, introduced from e. Asia. July-September; October-November. [= RAB, C, F, G, I, K, SE, W] Lespedeza frutescens (Linnaeus) Elliott, Violet Lespedeza. Mt, Pd (GA, NC, VA), Cp (GA, SC, VA): woodlands and woodland borders; common. July-September; October-November. MA and NY west to MI, WI, IA, and KS, south to NC, SC, GA, AL, MS, AR, and TX. [= K, S; = L. violacea (Linnaeus) Persoon – RAB, C, F, G, I, S, SE, W, Y, Z, misapplied] Lespedeza hirta (Linnaeus) Hornemann var. curtissii (Clewell) Isely, Silvery Lespedeza. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): sandhills and dry to moist savannas; common. August-October; September-November. Se. VA south to s. FL, west to panhandle FL and se. AL, barely extending onto the Piedmont in NC, SC, and GA. Clewell (1966a) discusses apparent intergrades between the two varieties in s. NJ. [= C, I, SE, X; < L. hirta – RAB, G, S; = L. hirta var. appressipilis Blake – F (as to intent, but not the type); = L. hirta ssp. curtissii Clewell – K, Y, Z] Lespedeza hirta (Linnaeus) Hornemann var. hirta, Hairy Lespedeza. Mt, Pd, Cp (GA, NC, SC, VA): woodlands and woodland borders; common. August-October; September-November. S. ME and s. Ontario west to MI, n. IL, c. MO, and OK, south to FL and TX. [= C, I, SE, X; < L. hirta – RAB, G, S, W; > L. hirta var. hirta – F; > L. capitata var. calycina (Schindler) Fernald – F; = L. hirta ssp. hirta – K, Y, Z] Lespedeza procumbens Michaux, Downy Trailing Lespedeza. Pd, Cp, Mt (GA, NC, SC, VA): woodlands and woodland borders; common. July-September; August-November. MA, NH, and NY west to IL, MO, and KS, south to panhandle FL and TX. [= RAB, C, G, I, K, S, SE, W, Y, Z; > L. procumbens var. procumbens – F; > L. procumbens var. elliptica Blake – F] Lespedeza repens (Linnaeus) W. Barton, Smooth Trailing Lespedeza. Cp, Pd, Mt (GA, NC, SC, VA): woodlands and woodland borders; common. July-September; August-November. CT and NY west to n. OH, s. WI, MO, and KS, south to panhandle FL and c. TX. [= RAB, C, F, G, I, K, S, SE, W, Y, Z] Lespedeza stuevei Nuttall, Velvety Lespedeza. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC): woodlands and woodland borders; common. July-September; August-November. MA south to n. FL, west to c. and n. TX, north in the interior to. NC, TN, s. IN, s. IL, c. MO, and nc. KS. [= RAB, C, F, G, I, K, SE, W, Y, Z; = L. stuvei – S, orthographic variant] * Lespedeza thunbergii (Augustin de Candolle) Nakai. Pd (GA, NC, SC), Cp (NC, SC), Mt (NC): "wildlife food plots"; rare, introduced from e. Asia. Reported for Macon County, NC by Pittillo & Brown (1988). [= C, F, G, I, K, Q, SE] Lespedeza violacea (Linnaeus) Persoon, Wand Lespedeza. Mt, Pd, Cp (GA, NC, SC, VA): woodlands and woodland borders; common. July-September; August-November. S. ME and s. Ontario west to MI and se. MN, south to panhandle FL and e. TX. [= K, S; = L. intermedia (S. Watson) Britton – RAB, C, F, G, I, SE, W, Y, Z] FABACEAE 379 * Lespedeza virgata (Thunberg) Augustin de Candolle. Mt (NC): roadbanks; rare, introduced from e. Asia. Clewell & Stickel (1990) report the occurrence of this species in NC. [= I, K] Lespedeza virginica (Linnaeus) Britton, Virginia Lespedeza. Cp, Pd, Mt (GA, NC, SC, VA): woodlands and woodland borders; common. July-September; August-November. MA and NH west to MI, WI, IA, and KS, south to n. FL and c. TX. [= RAB, C, F, G, I, K, S, SE, W, Y, Z]

Many species of Lespedeza hybridize, and most combinations may occur in our area. Some of the hybrids have been named in the past as varieties or species. Hybrids generally occur in mixed populations with both parents and can usually be identified by their intermediate morphology (identification much easier in the field where context is apparent than in the herbarium). See Isely (1990) and Clewell (1966a) for additional hints about identification of hybrids. The following hybrids combinations are known to occur in our area.

L. angustifolia × capitata. L. angustifolia × hirta [= L. ×oblongifolia (Britton) W. Stone (pro sp.) – K; L. hirta var. appressipilis Blake – F (as to type)]. L. angustifolia ×repens. L. angustifolia ×stuevei. L. capitata × hirta. [= L. ×longifolia Augustin de Candolle (pro sp.) – K; L. hirta var. longifolia (Augustin de Candolle) Fernald – F; L. longifolia Augustin de Candolle – S] L. capitata × repens. L. capitata × stuevei. L. capitata × virginica. [= L. ×simulata Mackenzie & Bush (pro sp.) – F, K; L. simulata Mackenzie & Bush – S] L. hirta × procumbens. L. hirta × repens. L. hirta × stuevei. L. hirta × violacea. [= L. ×nuttallii Darlington (pro sp.) – K; L. nuttallii Darlington – RAB, F, S; L. nuttallii var. nuttallii – G]. L. hirta × virginica. L. intermedia × procumbens. L. intermedia × repens. L. intermedia × stuevei. L. procumbens × repens. L. procumbens × virginica. [= L. ×brittonii Bicknell (pro sp.) – K; L. brittonii Bicknell – F; L. procumbens var. elliptica Blake – F]. L. repens × stuevei. L. repens × virginica. L. stuevei × violacea. L. stuevei × virginica [= L. ×neglecta Mackenzie & Bush (pro sp.) – K; L. stuevei var. angustifolia Britton – F; L. neglecta Mackenzie & Bush – S]. L. violacea × virginica [= L. ×acuticarpa Mackenzie & Bush (pro sp.) – K]. Documented from Northampton Co. VA.

Leucaena Bentham (Leadtree)

A genus of about 22 species, of tropical and warm temperate America. References: Hughes (1998)=Z; Isely (1998)=I.

* Leucaena leucocephala (Lamarck) de Wit ssp. leucocephala, Leadtree. Cp (GA): disturbed areas; rare, introduced from the New World tropics. E. GA (Kartesz 1999, voucher at UGA), south into FL and the New World tropics. [= Z; < L. leucocephala – I, K, SE; < L. glauca (Linnaeus) Bentham – S, misapplied]

Lotus Linnaeus (Birdsfoot-trefoil) (also see Acmispon)

A genus of about 120-130 species, annual and perennial herbs, of temperate Eurasia. New World taxa often referred to Lotus are not closely related to Lotus, and should be segregated (Degtjareva et al 2006; Allan & Porter 2000). References: Isely (1981)=Z; Isely (1998)=I; Degtjareva et al. (2006); Allan & Porter (2000); Grant & Small (1996).

1 Leaves 3-foliolate, the upper commonly 1-foliolate; flowers solitary in leaf axils; [native annual herbs]...... [see Acmispon helleri] 1 Leaves 5-foliolate; flowers in umbels; [alien perennial herbs]. 2 Calyx tube 2.8-3.5 mm long; corolla usually 10-14 mm long; leaflets of the medial leaves mostly 1.5-2.5 (-5)× as long as wide ...... L. corniculatus 2 Calyx tube 1.8-2.8 mm long; corolla usually 8-10 mm long; leaflets of the medial leaves 3-4 (-6)× as long as wide ...... L. tenuis

FABACEAE 380 * Lotus corniculatus Linnaeus, Birdsfoot-trefoil, Eggs-and-Bacon. Cp (NC, VA), Pd (VA), Mt (GA, VA): fields, roadsides, and waste places; uncommon, introduced from Eurasia. June-September. First reported for GA (Rabun County) by Stiles & Howel (1998). [= RAB, C, F, G, K, S, SE, W, Z; < L. corniculatus Linnaeus – I (also see L. tenuis)] * Lotus tenuis Waldstein & Kitaibel ex Willdenow, Slender Birdsfoot-trefoil. Pd (NC): fields, roadsides, and waste places; rare, introduced from Eurasia. June-September. [= C, K, SE, Z; < L. corniculatus Linnaeus – I]

Lupinus Linnaeus (Lupine)

A genus of about 150-200 species, annual herbs, perennial herbs, and shrubs, of temperate and tropical regions in North America, Mediterranean Europe, South America, and Africa (especially diverse in w. North America and South America). References: Isely (1998)=I.

1 Leaves palmately compound; leaves and stems deciduous, dying back in winter; plant inconspicuously pubescent. 2 Stem short; leaves clustered, nearly whorled; leaflets narrow; racemes long exserted; flowers small; [plants of e. GA southward] ...... L. perennis ssp. gracilis 2 Stem elongate; leaves alternate; leaflets broad; racemes only moderately exserted; flowers large; [plants of n. SC northward]...... L. perennis ssp. perennis 1 Leaves unifoliolate; leaves and stems evergreen, overwintering (absent in midsummer); plant conspicuously pubescent. 3 Standard blue, with a white to creamy eyespot; hairs on legume ca. 2 mm long; petioles with short and appressed pubescence...... L. diffusus 3 Standard reddish to purplish, with a red or deep purple eyespot; hairs on legume 3-5 mm long; petioles with long and spreading (shaggy) pubescence...... L. villosus

Lupinus diffusus Nuttall, Blue Sandhill Lupine. Cp (GA, NC, SC): sandhills, sandy roadsides; common. March-May; June-July. Se. NC south to s. FL, west to s. MS. I concur with Duncan & McCartney (1992) in recognizing L. cumulicola Small of peninsular FL as distinct from L. diffusus. [= RAB, K, S; < L. diffusus – I, SE] Lupinus perennis Linnaeus ssp. gracilis (Nuttall) Dunn, Southern Sundial Lupine. Cp, Pd (GA): sandhills and sandy or dry rocky roadsides; uncommon. E. GA (immediately across the Savannah River from SC), south to n. FL and west to s. AL. The validity of this taxon is uncertain; the differences may be only clinal. [= K, SE; < L. perennis – RAB, C, G; = L. perennis var. gracilis (Nuttall) Chapman – I; = L. nuttallii S. Watson – S] Lupinus perennis Linnaeus ssp. perennis, Northern Sundial Lupine. Mt (VA), Pd (NC, SC, VA), Cp (NC, SC, VA): sandhills, sandy roadsides, other dry habitats; uncommon. April-May; June-July. ME west to MN, south to n. SC, w. VA, e. WV, IN, and IL. [= SE; < L. perennis – RAB, C, G, W; > L. perennis var. perennis – F, I; > L. perennis var. occidentalis S. Watson – F; > L. perennis ssp. perennis var. perennis – K; > L. perennis ssp. perennis var. occidentalis – K; = L. perennis – S] Lupinus villosus Willdenow, Pink Sandhill Lupine. Cp (GA, NC, SC): sandhills, sandy roadsides; uncommon (NC Watch List). April-May; June-August. Se. NC south to n. FL, west to se. LA. [= RAB, I, K, S, SE]

Macroptilium (Bentham) Urban

A genus of about 20 species, annual and perennial herbs, of tropical and subtropical America. References: Isely (1998)=I.

* Macroptilium lathyroides (Linnaeus) Urban. Cp (GA, SC): disturbed areas; rare, naturalized from a native range in tropical America. [= I, K, SE; = Phaseolus lathyroides Linnaeus]

Medicago Linnaeus (Medick, Bur-clover)

A genus of about 80 species, annual and perennial herbs, of Eurasia and Africa. References: Isely (1998)=I. Key based largely on SE.

1 Legume 1-seeded, reniform, black at maturity; corolla 2-3 mm long ...... M. lupulina 1 Legume several-seeded, spirally coiled or falcate, tan to dark brown; corolla 3-11 mm long. 2 Plants perennial, mostly erect or ascending, 2-8 (-10) dm tall; corolla 6-11 mm long, violet, yellow, or varicolored; legumes spineless. 3 Legume falcate; corolla yellow...... M. falcata 3 Legume spiral; corolla violet or varicolored (rarely yellow)...... M. sativa 2 Plant annual, mostly prostrate or ascending, 1-6 dm tall; corolla 3-6 mm long, yellow; legumes spiny (except lacking spines in M. orbicularis). 4 Stipules entire or slightly dentate (M. minima) or the base only of the stipule lacerate (M. laciniata); plants pilose (M. minima) or glabrous (M. laciniata). 5 Stipules lacerate at the base; plant glabrous ...... M. laciniata 5 Stipules entire or slightly dentate; plant pilose...... M. minima FABACEAE 381 4 Stipules lacerate; plants glabrous or sparsely pubescent. 6 Legume lacking spines; stipules deeply lacerate, the sinuses extending nearly to the base...... M. orbicularis 6 Legume spiny; stipules either deeply lacerate (M. polymorpha) or shallowly lacerate (M. arabica). 7 Leaflets 0.7-1.1× as long as wide, usually marked with a central dark spot; leaflet tip usually strongly notched; stipules shallowly lacerate, the sinuses extending < ½ way to the base...... M. arabica 7 Leaflets 1-2× as long as wide, not marked with a central dark spot; leaflet tip not strongly notched; stipules deeply lacerate, the sinuses extending > ½ way to the base ...... M. polymorpha

* Medicago arabica (Linnaeus) Hudson, Spotted Medick, Spotted Bur-clover. Cp, Pd (GA, NC, SC), Mt (GA, VA): fields, roadsides, disturbed areas; uncommon, introduced from Mediterranean Europe. April-August. [= RAB, F, G, I, K, S, SE] * Medicago falcata Linnaeus, Yellow Alfalfa, Sickle Medick. Mt? (VA?): disturbed areas; rare, introduced from n. Eurasia. April-July. The occurrence of this taxon in our area requires verification. [= F, G, I, S, SE; = M. sativa Linnaeus ssp. falcata (Linnaeus) Arcangeli – C, K] * Medicago laciniata (Linnaeus) P. Miller. Cp (SC): waste areas aropund wool-combing mills; rare, native of Europe, perhaps merely a waif. [= F, I, K] * Medicago lupulina Linnaeus, Black Medick, Yellow Trefoil. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from Europe. April-August. [= RAB, C, G, I, K, S, SE, W; > M. lupulina var. lupulina – F; > M. lupulina var. glandulosa Neilreich – F] * Medicago minima (Linnaeus) Linnaeus, Downy Bur-clover, Bur Medick. Cp (NC, SC, VA): fields, roadsides, disturbed areas; rare, introduced from Eurasia. April-August. [= RAB, C, G, I, K, S, SE; > M. minima var. minima – F; > M. minima var. compacta Neyraut – F; > M. minima var. longiseta Augustin de Candolle – F] * Medicago orbicularis (Linnaeus) Bartalini. Pd (GA, NC): lawns, disturbed areas; rare, introduced from Mediterranean Europe and n. Africa. April-July. [= RAB, G, I, K, SE] * Medicago polymorpha Linnaeus, Smooth Bur-clover, Toothed Medick. Cp (GA, NC, SC), Pd (SC): fields, roadsides, lawns, disturbed areas; uncommon, introduced from Mediterranean Europe. March-April. [= RAB, C, I, K, SE; = M. hispida Gaertner – F, G, S] * Medicago sativa Linnaeus, Alfalfa, Lucerne, Blue Alfalfa. Mt, Pd, Cp (GA, NC, SC, VA): roadsides, fields, disturbed areas; common, introduced from se. Europe. April-July. [= RAB, F, G, I, S, SE, W; = M. sativa Linnaeus ssp. sativa – C, K]

Melilotus P. Miller (Melilot, Sweetclover, Sourclover)

A genus of about 20 species, annual and perennial herbs, of temperate Eurasia. References: Stace (1997)=Z; Isely (1998)=I. Key based in part on Stace (1997).

1 Flowers white...... M. albus 1 Flowers yellow. 2 Flowers 2-3.5 mm long; fruits < 3 mm long ...... M. indicus 2 Flowers > 4 mm long; fruits > 3 mm long. 3 Fruits > 5 mm long, mostly 2-seeded, black when ripe, pubescent (at 10× magnification); keel about equalling the wings...... [M. altissimus] 3 Fruits < 5 mm long, mostly 1-seeded, brown when ripe, glabrous; keel shorter than the wings ...... M. officinalis

* Melilotus albus Medikus, White Melilot, White Sweetclover. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from Eurasia. April-October. M. albus and M. officinalis, nearly identical except in flower color, are apparently incompatible (Isely 1998); they should not be synonymized, as was done by K. Other differences useful in the determination of faded herbarium specimens are given but Isely (1998): corolla 3.5-5 mm long, the wing petals about as long as the keel (M. albus) vs. corolla 5-7 mm long, the wing petals generally longer than the keel (M. officinalis). [= I, Z; = M. alba – RAB, C, F, G, S, SE, W, orthographic variant; < M. officinalis – K] * Melilotus indicus (Linnaeus) Allioni, Small Melilot, Sourclover. Cp (GA, NC, SC, VA), Pd (GA, SC): roadsides, disturbed areas; uncommon, introduced from Mediterranean Europe. April-October. [= I, K, Z; = M. indica – RAB, C, F, G, S, SE, orthographic variant] * Melilotus officinalis (Linnaeus) Pallas, Yellow Melilot, Yellow Sweetclover, Ribbed Melilot. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from Eurasia. April-October. [= RAB, C, F, G, I, S, SE, W, Z; < M. officinalis – K (also see M. albus)]

* Melilotus altissimus Thuiller, Tall Melilot, another Eurasian weed, is known to be naturalized as far south as e. PA (Rhoads & Klein 1993). It likely occurs in our area. It superficially resembles M. officinalis; see key for distinguishing characteristics. [= I, Z; = Melilotus altissima – C, F, G, orthographic variant]

Mimosa Linnaeus (Mimosa)

FABACEAE 382 A genus of about 500 species, herbs, shrubs, trees, and vines, of tropical, subtropical, and warm temperate areas, especially America. Barneby (1991) and Beard (1963) argue that there are no characters which serve to separate Schrankia from Mimosa. References: Barneby (1991)=Y; Isely (1973)=Z; Isely (1998)=I.

Identification notes: Unmistakable in our flora for its bipinnate leaves, with tiny (2-4 mm long) leaflets, responding to touch by closing.

1 Plant unarmed ...... M. strigillosa 1 Plant armed. 2 Leaflets without apparent secondary veins; pinnae 4-8 (-11) pairs ...... M. microphylla 2 Leaflets with evident (sometimes weakly so) secondary veins; pinnae 3-5 (-6) pairs...... M. quadrivalvis var. floridana

Mimosa microphylla Dryander, Eastern Sensitive-brier. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): dry woodlands and forests, especially sandhills, disturbed areas; common (VA Rare). June-September; August-November. DE, WV, and MO south to FL and e. TX. A form with smaller fruits (3-5 cm long vs. 5-12 cm long) has been variously treated as a species [Leptoglottis chapmanii – S, Schrankia chapmanii] or a "recurrent fruit-form genotype" [phase brachycarpa of Isely (1973)]. [= K; = Mimosa quadrivalvis Linnaeus var. angustata (Torrey & A. Gray) Barneby – C, I, Y; = Schrankia microphylla (Dryander) J.F. Macbride – RAB, F, G, W; = Schrankia microphylla (Dryander) J.F. Macbride var. microphylla – SE; > Leptoglottis microphylla (Dryander) Britton & Rose – S; > Leptoglottis chapmanii Small ex Britton & Rose – S; > Schrankia microphylla "phase brachycarpa" – Z; > Schrankia chapmanii (Small ex Britton & Rose) F.J. Hermann] Mimosa quadrivalvis Linnaeus var. floridana (Chapman) Barneby, Florida Sensitive-briar. Cp (GA): xeric sandhills and other dry, sandy habitats; rare. A Southeastern Coastal Plain endemic: GA south into FL. This taxon is distinct at the specific level from M. quadrivalvis and nomenclatural adjustments are forthcoming (Flores-Cruz et al. 2004). [= I, K, Y; = Leptoglottis floridana (Chapman) Small ex Britton & Rose – S; = Schrankia microphylla (Dryander) J.F. MacBride var. floridana (Chapman) Isely – SE] Mimosa strigillosa Torrey & A. Gray, Powderpuff Mimosa. Cp (GA): floodplain forests, open wet areas; uncommon. A Southeastern Coastal Plain endemic: e. GA south to FL, west to TX. It might be expected in se. SC (see SE, Y, Z). [= I, K, S, SE, Y, Z]

Mucuna Adanson (Velvetbean)

A genus of about 100 species, perennial herbs, annual herbs, and woody vines, of tropical regions of Old World and New World. References: Isely (1998)=I.

* Mucuna pruriens (Linnaeus) Augustin de Candolle. Velvetbean, Bengal Bean, Florida Bean. Cp (NC, SC): disturbed areas, fields, cultivated and sporadically established in disturbed areas; rare, introduced from se. Asia. [= I, SE; > M. pruriens var. pruriens – K; > Stizolobium deeringianum Bort – S; > M. deeringiana (Bort) Merrill]

Neptunia Loureiro (Neptunia)

A genus of ca. 10-11 species, herbs, of the tropics and subtropics of America and Eurasia. References: Isely (1998)=I; Windler (1966)=Z.

1 Leaflets 9-15 pairs per pinna; stipules 2-4 mm long; all flowers perfect, with functional stamens; stipe of fruit 4-14 mm long ...... [N. lutea] 1 Leaflets (12-) 15-25 pairs per pinna; stipules 4-10 mm long; flowers in the lower part of the inflorescence with flattened staminodes; stipe of fruit 2-5 mm long ...... [N. pubescens var. pubescens]

Neptunia lutea (Leavenworth) Bentham, Yellow Neptunia. Savannas, prairies, roadsides. AL west to OK and TX. [= I, K, S, SE, Z] Neptunia pubescens Bentham var. pubescens, Tropical Neptunia. Savannas, sandhills, scrub, prairies, roadsides. AL and FL west to TX and south to Argentina. [= I, K, SE, Z; > N. floridana Small – S]

Orbexilum Rafinesque (Scurfpea, Sampson's-snakeroot)

A genus of about 9 species, perennial herbs, of s. North America and Mexico. References: Grimes (1988, 1990)=Z; Isely (1998)=I.

1 Leaves unifoliolate; [subgenus Poikadenia] ...... O. virgatum 1 Leaves with 3-7 leaflets. FABACEAE 383 2 Leaves palmately (3-) 5-7 foliolate, the leaflets linear to very narrowly oblanceolate, 2-7 cm long, 0.5-2.0 (-3.5) mm wide, > 10× as long as wide; [subgenus Orbexilum]...... O. lupinellum 2 Leaves pinnately 3-foliolate, the leaflets orbicular, ovate, elliptic or lanceolate, > 8 mm wide, 1-8× as long as wide. 3 Leaflets 1.5-7 cm wide, 1-2.5× as long as wide; [subgenus Orbexilum]. 4 Upper leaf surfaces lacking glands; leaflets 3.7-5.5 cm long; [endemic to Jefferson Co. KY and now extinct] ...... [O. stipulatum] 4 Upper leaf surfaces glandular; leaflets 4-12 cm long. 5 Leaflets subcordate, 4-7 cm wide, 1-1.5× as long as wide, glandular-punctate above and below, the apex obtuse; calyx stipitate-glandular; petals 8-10 mm long; [endemic to Polk Co. NC]...... O. macrophyllum 5 Leaflets rounded at base, 2-4 cm wide, 1.5-2.5× as long as wide, eglandular or sparsely glandular above, the apex acute; calyx lacking stipitate glands; petals 5-7 mm long; [widely scattered from w. VA and w. NC westward]...... O. onobrychis 3 Leaflets 0.8-2 cm wide, 2.5-7.5× as long as wide; [subgenus Poikadenia]. 6 Flowers 8-10 mm long; [of s. AL westward] ...... [O. simplex] 6 Flowers 5-7 mm long; [collectively widespread in our area]. 7 Calyx tube, fruits, and bracts of the inflorescence eglandular (rarely slightly glandular-punctate); leaflets eglandular below (rarely slightly punctate); hairs on calyx 0.7-1.0 mm long; upper 2 calyx teeth 1.0-1.5 mm long, lateral teeth 1.5-2.0 mm long, lower calyx tooth 2.0-3.0 mm long...... O. pedunculatum var. pedunculatum 7 Calyx tube, fruits, and bracts of the inflorescence conspicuously glandular-punctate; leaflets conspicuously glandular-punctate below; hairs on calyx 0.3-0.5 mm long; upper 2 calyx teeth 0.7-1.0 mm long, lateral teeth 1.2-1.5 mm long, lower calyx tooth 1.7-2.0 mm long...... O. pedunculatum var. psoralioides

Orbexilum lupinellum (Michaux) Isely, Lupine Scurfpea. Cp (GA, NC, SC): sandhills; uncommon (NC Watch List). May-July; July-October. This peculiar species is a Southeastern Coastal Plain endemic, ranging from sc. and se. NC, south to c. peninsular FL, s. AL, and e. GA. The very peculiar leaves, palmately 5-7-foliolate with "oblinear" leaflets, make the species unmistakable. First reported for SC by McMillan et al. 2002). [= K; = Psoralea lupinellus Michaux - RAB; = Orbexilum lupinellus – I, SE, Z, orthographic variant; = Rhytidomene lupinellus (Michaux) Rydberg – S] Orbexilum macrophyllum (Rowlee in Small) Rydberg, Bigleaf Scurfpea. Mt (NC): wooded slopes of mountain on Blue Ridge escarpment, precise habitat not known (probably nutrient-rich dry woodlands); rare (US Species of Concern, NC Endangered). June; July-August? This species was discovered on 18 June 1897 and subsequently collected on 8 June 1899 by E.C. Townsend, somewhere on the double peak of Tryon Mountain and White Oak Mountain, Polk County, NC, a phytogeographically interesting area with disjunct, endemic, and relictual species largely of midwestern affinities. It is currently presumed to be extinct, following a number of unsuccessful attempts to relocate it. Isely (1990) is correct in stating that the assignment of "this distinctive species" to Orbexilum is "reasonably assumptive," since fruits have never been seen. [= I, K, S, SE, Z; = Psoralea macrophylla Rowlee in Small – RAB, W] Orbexilum onobrychis (Nuttall) Rydberg, Lanceleaf Scurfpea. Mt (NC, VA): habitat in our area not known, elsewhere usually in nutrient-rich, open or semi-open areas; rare (NC Watch List, VA Rare). June-July; August-October. Primarily a species of prairies and prairie-like areas of OH and KY west to se. IA and e. MO, O. onobrychis also occurs (at least formerly) as a rare disjunct in the mountains of w. NC, nw. SC, w. VA, and e. TN. The only report for NC was in the 1800's. [= C, I, K, S, SE, Z; = Psoralea onobrychis Nuttall – RAB, F, G, W] Orbexilum pedunculatum (P. Miller) Rydberg var. pedunculatum, Western Sampson's-snakeroot. Mt (GA, NC, SC), Pd, Cp (GA, SC): open woodlands; rare (NC Watch List). May-July; July-September. Var. pedunculatum, the western and more widespread variety, occurs primarily west of the Blue Ridge, with scattered occurrences in and east of the Blue Ridge. Its range is s. OH, s. IN, s. IL, c. MO, and se. KS, south to sw. NC, sc. SC, sw. GA, s. AL, s. LA, and e. TX. [= C, I, K, SE, Z; = Psoralea psoralioides (Walter) Cory var. eglandulosa (Elliott) F.L. Freeman – RAB, F, G, GW, W; = Orbexilum pedunculatum – S] Orbexilum pedunculatum (P. Miller) Rydberg var. psoralioides (Walter) Isely, Eastern Sampson's-snakeroot. Cp (GA, NC, SC, VA), Pd (NC, SC, VA): savannas, open woodlands; uncommon. May-July; July-September. Var. psoralioides, the eastern variety, occurs primarily on the Atlantic Coastal Plain, ranging from e. VA to n. FL, inland to the Piedmont of NC and SC. [= K; = Psoralea psoralioides (Walter) Cory var. psoralioides – RAB, F, G, GW; = Orbexilum pedunculatum var. gracile (Torrey & A. Gray) Grimes – C, I, SE, Z; = Orbexilum gracile (Torrey & A. Gray) Rydberg – S] Orbexilum virgatum (Nuttall) Rydberg, Slender Leather-root. Cp (GA, SC?): sandhills; rare (GA Special Concern). Se. GA (or SC?) south to ne. FL. A collection by Curtiss is labeled as from South Carolina. [= I, K, SE, Z; = Psoralea virgata Nuttall]

Orbexilum simplex (Nuttall ex Torrey & Gray) Rydberg. Prairies, open woodlands. AR and OK south to s. AL and e. TX; perhaps disjunct in IL. [= I, K, S, SE, Z] Orbexilum stipulatum (Torrey & Gray) Rydberg. Rocky limestone glade. So far as is known, once endemic to Rock Island, Jefferson Co., KY, and now extinct. [= C, I, K, SE, Z; = Psoralea stipulata Torrey & Gray – F, G]

Parkinsonia Linnaeus (Jerusalem Thorn)

FABACEAE 384 A genus of about 30 species (if circumscribed to include Cercidium), shrubs and trees of sw. North America, Central America, and Africa. References: Isely (1975)=Z; Robertson & Lee (1976)=Y; Isely (1998)=I.

* Parkinsonia aculeata Linnaeus, Jerusalem Thorn, Retama, Horse-bean, Mexican Palo Verde. Cp (GA, SC): disturbed areas; rare, introduced from sw. North America. May. Rarely established or spread from cultivation in our area, more commonly so in much of FL. [= I, K, S, SE, Y, Z]

Pediomelum Rydberg (Buckroot, Prairie-turnip)

A genus of about 21 species, perennial herbs, of North America. References: Allison, Morris, & Egan (2006)=Y; Grimes (1988, 1990)=Z; Isely (1998)=I.

1 Plants acaulescent, 1-2 dm tall; leaves (4-) 5-7-foliolate; [plants of calcareous glades of the Interior (nw. GA, TN, n. AL)]; [subgenus Disarticulatum]...... P. subacaule 1 Plants caulescent, 3-10 dm tall; leaves 3-foliolate; [plants of sandhills of the Coastal Plain and rocky woodlands of the lower Coastal Plain]; [subgenus Pediomelum]. 2 Inflorescence loose (much of the axis exposed); leaflets (1-) 3; leaflets < 2× as long as wide, petiolules 5-9 mm long; [of longleaf pine sandhills of the Coastal Plain] ...... P. canescens 2 Inflorescence congested (the axis usually concealed); leaflets 3 (-5); leaflets > 2× as long as wide, petiolules 1.8–3 mm; [of rocky woodlands of the lower Piedmont]...... P. piedmontanum

Pediomelum canescens (Michaux) Rydberg, Buckroot, Eastern Prairie-turnip, Hoary Scurfpea. Cp (GA, NC, SC, VA): sandhills; rare (NC Watch List, VA Rare). May-July; July-October. A Southeastern Coastal Plain endemic: se. VA south to c. peninsular FL, Panhandle FL, and s. AL. This uncommon species tends to occur as very widely scattered individuals in sandhill habitats, rarely with more than a few seen at a time. It is related to P. esculentum (Pursh) Rydberg, the "prairie potato," prized by early travelers across the prairies for its edible tubers. An interesting collection label (by R.E. Wicker, collected in 1942, the specimen at NCU) mentions both the edible tubers and the characteristically sparse population structure of the species. "Not uncommon near Pinehurst in ... open places in sandy pine woods..., but usually only one plant at a time. Tuber hard, dark brown, about size of a medium-sized Irish potato, somewhat ventral-elongated with roots coming from pointed base. Internal pure white, apparently almost entirely starch... Mr. Wicker says that he rather likes to take a bit of it and chew when fresh, has a rather condiment taste, but does not think it well to eat..." Because of its rarity, P. canescens should not (of course) be eaten. Because of its habit, that of a very bushy, tumbleweed-like plant, it superficially most closely resembles various Baptisia species, but it is easily separated by its rather dense and soft pubescence (our Baptisia are all glabrous or rather inconspicuously puberulent, except the very unifoliolate B. arachnifera). [= C, I, K, S, SE, Z; = Psoralea canescens Michaux – RAB, F, G] Pediomelum piedmontanum J.R. Allison, M.W. Morris, & A.N. Egan, Piedmont Buckroot. Pd (GA, SC): open, rocky woodlands in the lower Piedmont; rare. Late May-late June (-late July); July-August (-September). Apparently endemic to the lower Piedmont of c. SC and e. GA. See Allison, Morris & Egan (2006) for additional details. [= P. piedmontanum J.R. Allison, M.W. Morris, & A.N. Egan – Y] Pediomelum subacaule (Torrey & A. Gray) Rydberg, Nashville Breadroot. Mt (GA): limestone glades; rare (GA Special Concern). E. TN and nw. GA west to c. TN and n. nw. AL. [= I, K, SE, Z; = P. subacaulis – S, orthographic variant]

Petalostemon (see Dalea)

Phaseolus Linnaeus (Bean)

A genus of about 50 species, annual and perennial herbs, of tropical and warm temperate America (now widely distributed worldwide in cultivation). References: Isely (1998)=I; Maréchal, Mascherpa, & Stainier (1978)=Z; Freytag & Debouck (2002)=Y. Key based on SE.

1 Raceme axes slender, flexuous; [plants native perennials]; [section Paniculati; subsection Volubili]. 2 Stems climbing and twining on other vegetation; leaflets 3-10 cm long, usually not lobed and only slightly reticulate; [plants of various habitats]...... Ph. polystachios 2 Stems trailing across ground; leaflets 1-4 cm long, usually strongly 3-lobed and strongly reticulate; [plants of sandhill habitats of the Coastal Plain]...... Ph. sinuatus 1 Raceme axes stout, stiff; [plants alien annuals, only weakly naturalized]. 3 Corolla 1.5-2 cm long, scarlet red or bright lavender; racemes exserted; plants twining; [section Coccinei] ...... Ph. coccineus ssp. coccineus 3 Corolla ca. 1 cm long, pink-purple, greenish white, or bicolored pink and white; racemes short or exserted; plants bushy-erect (rarely twining). FABACEAE 385 4 Legumes distinctly flattened at maturity, 15-20 mm wide; corolla usually greenish white; [section Paniculati; subsection Volubili]...... Ph. lunatus 4 Legumes nearly terete at maturity, about 8 mm in diameter; corolla usually pink-purple, at least in part; [section Phaseoli] ...... Ph. vulgaris

* Phaseolus coccineus Linnaeus ssp. coccineus, Scarlet Runner Bean. Cp, Pd, Mt (GA, NC, SC, VA): infrequently cultivated, mostly as an ornamental in home gardens, rarely found as a waif, introduced from tropical America. [= Z; < Ph. coccineus – C, F, G, I, K, SE; > Ph. coccineus ssp. coccineus var. coccineus – Y] * Phaseolus lunatus Linnatus, Lima Bean. Cp, Pd, Mt (GA, NC, SC, VA): frequently cultivated (both commercially and in home gardens), rarely found as a waif, introduced from tropical America. [= I, K, S, SE, Y, Z; = Ph. limensis Macfadyen – F] Phaseolus polystachios (Linnaeus) Britton, Sterns, & Poggenburg, Wild Bean, Wild Kidney Bean. Mt, Pd, Cp (GA, NC, SC, VA): thickets, woodlands; uncommon. July-September; August-October. S. ME west to OH, IL, and MO, south to FL and TX. [= RAB, C, G, I, SE, W; > Ph. polystachios var. polystachios – F; > Ph. polystachios var. aquilonius Fernald – F; = Ph. polystachios var. polystachios – K; = Ph. polystachyus – S, orthographic variant; = Ph. polystachyus ssp. polystachyus – Y; = Ph. polystachyus var. polystachyus – Z] Phaseolus sinuatus (Nuttall) Torrey & A. Gray, Sandhills Bean. Cp (GA, NC, SC): sandhills; rare (GA Special Concern). July-September; August-October. Sc. NC south to s. peninsular FL, west to s. MS, a Southeastern Coastal Plain endemic. Freytag & DeBouck (2002) describe Ph. sinuatus and Ph. polystachios as being "very distinct and there seems to be no intergradation;" I choose to recognize them as species. Not easy to distinguish in sterile condition from Strophostyles. [= RAB, I, S, SE, W; = Ph. polystachios (Linnaeus) Britton, Sterns, & Poggenburg var. sinuatus (Nuttall) R. Marechal, J.M. Mascherpa, & F. Stainier – K; = Ph. polystachyus ssp. sinuatus (Nuttall) Freytag – Y; = Ph. polystachyus var. sinuatus (Nuttall) R. Marechal, J.M. Mascherpa, & F. Stainier – Z] * Phaseolus vulgaris Linnaeus, Garden Bean, Green Bean, Snap Bean, String Bean, Kidney Bean, Pole Bean, Bush Bean. Cp, Pd, Mt (GA, NC, SC, VA): frequently cultivated (both commercially and in home gardens), rarely found as a waif, introduced from tropical America. [= C, F, G, I, K, S, SE; > Ph. vulgaris var. vulgaris – F; > Ph. vulgaris var. humilis Alefeld – F; < Ph. vulgaris var. vulgaris – Z]

Pisum Linnaeus (Pea)

A genus of 2 species, annual herbs, native to the Mediterranean region. References: Isely (1998)=I.

* Pisum sativum Linnaeus, Pea, Garden Pea, English Pea. Mt, Pd, Cp (GA, NC, SC, VA): commonly cultivated in home gardens, rarely found as a waif. March-May. [= I, K, SE; > P. sativum var. sativum – F; > P. sativum var. arvense (Linnaeus) Poiret – RAB, F]

Psoralea (see Orbexilum, Pediomelum)

Psoralidium Rydberg

A genus of 3 species, herbs, of c. and w. North America. References: Grimes (1988, 1990)=Z; Isely (1998)=I.

Psoralidium tenuiflorum (Pursh) Rydberg, Gray Scurf-pea. Prairies. KY to MT, south to TX and n. Mexico; disjunct in MS. [= C, I, K, SE, Z; = Psoralea tenuiflora Pursh – F, G]

Pueraria Augustin de Candolle (Kudzu)

A genus of about 15 species, perennial vining herbs, of tropical and subtropical Asia. References: Isely (1998)=I; Ward (1998)=Z.

* Pueraria montana (Loureiro) Merritt var. lobata (Willdenow) van der Maesen & S. Almeida, Kudzu. Mt, Pd, Cp (GA, NC, SC, VA): roadsides, waste areas; common, introduced from e. Asia. July-October. Kudzu was strongly promoted in the 1920's and 1930's in the Southeastern United States as a stabilizer of eroded areas. Hundreds of Kudzu Clubs formed, and Kudzu Songbooks were published. It is now notorious as a weed and symbol of the South. Despite its notoriety in the popular press, kudzu is an ecologically relatively trivial (though conspicuous) weed, since it rarely produces viable seeds in our area, and generally does not invade high quality natural areas. The thickened rhizome can weigh as much as 100 kg, and is the source of a high quality cooking starch prized in the Orient. The purple flowers smell like artificial grape flavoring. The leaves are very frost-sensitive. [= I, K, Z; = P. lobata (Willdenow) Ohwi – RAB, C, F, G, SE, W; = P. thunbergiana (Siebold & Zuccarini) Bentham – S]

FABACEAE 386

Rhynchosia Loureiro (Snoutbean)

A genus of about 200 species, perennial herbs, of tropical and warm temperate regions, nearly cosmopolitan. References: Grear (1978)=Z; Isely (1998)=I.

1 Leaves unifoliolate (rarely with a few upper leaves trifoliolate). 2 Plant prostrate, trailing, usually with many leaves; stipels absent; [plant very rare in our area, probably introduced] ...... Rh. michauxii 2 Plants erect or ascending, usually with fewer than 6 leaves; stipels present; [plant common in the Coastal Plain in our area] ...... Rh. reniformis 1 Leaves trifoliolate (rarely with a few lowermost leaves unifoliolate, these generally withering before flowering and fruiting). 3 Plant trailing or twining; pubescence of the lower leaf surface mostly restricted to the veins. 4 Calyx 8-10 (-12) mm long, about as long as the corolla; [plants widespread in our area]...... Rh. difformis 4 Calyx 2.5-3 (-4) mm long, clearly shorter than the corolla; [plants of e. GA southward] ...... Rh. minima 3 Plant erect; pubescence of the lower leaf surface not restricted to the veins, grayish tomentose and velvety to the touch. 5 Inflorescence a single exserted terminal raceme, 5-20 cm long; stipules caducous; [plants of s. SC southward] ...... Rh. tomentosa var. mollissima 5 Inflorescences several and axillary, 1-3 cm long; stipules persistent; [plants widespread in our area] ...... Rh. tomentosa var. tomentosa

Rhynchosia difformis (Elliott) Augustin de Candolle. Cp (GA, NC, SC, VA), Pd (SC): sandhills; common. June-August; July-October. Se. VA south to s. FL, west to e. TX. [= RAB, C, F, G, I, K, SE; = Rh. tomentosa – S, misapplied] *? Rhynchosia michauxii Vail. Cp (GA, NC): disturbed areas; rare. June-August; August-October. Se. NC (one record) and e. GA (one record) south to s. peninsular FL, west to Panhandle FL. The disjunct sites are of uncertain origin. [= I, K, S, SE; = Rh. americana (Houston ex P. Miller) M.C. Metz – RAB (based on misidentification of specimen)] Rhynchosia minima (Linnaeus) Augustin de Candolle. Cp (GA): coastal sands; uncommon. Along the coast in e. GA, south to FL, west to s. TX. The species also occurs in the Old World, and the New World distribution is sometimes considered a result of introduction. [= I, K, SE; = Dolicholus minimus (Linnaeus) Medikus – S] Rhynchosia reniformis Augustin de Candolle, Dollarweed. Cp (GA, NC, SC): sandhills; common. June-September; August-October. Se. NC south to s. FL, west to e. TX; disjunct (introduced?) in e. TN (Chester, Wofford, & Kral 1997). [= RAB, K, SE; = Rh. simplicifolia (Walter) Wood – S] Rhynchosia tomentosa (Linnaeus) Hooker & Arnott var. mollissima (Elliott) Torrey & A. Gray. Cp (GA, SC): sandhills, rare. June-August; August-October. Peninsular FL, e. GA, and se. SC (Beaufort County, documented by an old specimen [GH] by Mellichamp from the vicinity of Bluffton, where it was probably native). [= I, K, SE; = Rh. mollissima (Elliott) S. Watson – S] Rhynchosia tomentosa (Linnaeus) Hooker & Arnott var. tomentosa. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): xeric woodlands and forests, sandhills, edges, open areas; common. June-August; August-October. DE south to FL, west to LA, and north in the interior to e. and c. TN. [= C, I, K, SE; < Rh. tomentosa – RAB, F, G, W; > Rh. erecta (Walter) Augustin de Candolle – S; > Rh. intermedia (Torrey & Gray) Small – S]

Rhynchosia cytisoides (Bertol.) Wilbur. AL and Panhandle FL west to MS. [= I, K, SE; = Pitcheria galactioides Nuttall – S] {not keyed at this time}

Robinia Linnaeus (Locust)

A genus of 5-8 species, shrubs and trees, of e. and sw. North America. The Southern Appalachians are a center of diversity of Robinia, with active hybridization, introgression, and formation of local (sterile) races involved; a fully satisfying taxonomic treatment of such a situation is not possible. Isely & Peabody's (1984) treatment seems a reasonable approach, and I have largely followed it here, differing in the rank of some of the taxa. The key is differently structured than that in RAB or SE; it is presented as an alternative. {NOTE: This treatment may be altered substantially prior to publication.} References: Isely & Peabody (1984)=Z; Ashe (1922)=Y; Isely (1998)=I.

1 Corolla white, 1.5-2.0 cm long; peduncles, pedicels, and calyces velvety-puberulent, the hairs neither glandular nor hispid; plant a small to large tree...... R. pseudoacacia 1 Corolla pink to pink-purple (rarely white or nearly so), (1.5-) 2.0-2.5 cm long; peduncles, pedicels, and calyces glandular- pubescent, hispid, or with short-stalked to sessile glands; plant a shrub to small tree. 2 Twigs and leafstalks conspicuously hispid with hairs 1-5 mm long, these stiff, thick-based, and typically persistent several years. 3 Plants fruiting abundantly; shrubs 0.6-2 (-3) m tall; leaflets relatively broad, mostly 1.2-1.8× as long as wide...... R. hispida var. fertilis 3 Plants sterile (rarely fruiting scantily); shrubs 0.5-1.5 m tall; leaflets relatively narrow, mostly 1.8-2.5× as long as wide...... R. hispida var. hispida FABACEAE 387 2 Twigs and leafstalks either viscid with sessile or short-stalked glands, or densely glandular-pubescent (the hairs 0.5-2 mm long), or tomentulose, or sparsely hispid with weak, non-persistent hairs. 4 Leaflets usually 13-21, permanently but inconspicuously appressed-pubescent beneath; bracts (evident only before anthesis) aristate; plants never with long, hispid pubescence. 5 Twigs and peduncles finely glandular-pubescent with hairs 1 (-2) mm long; plants abundantly pod-forming ...... R. hartwigii 5 Twigs and peduncles viscid with sessile or short-stalked glands (the racemes sometimes with some glandular pubescence); plants sterile or forming pods ...... R. viscosa 4 Leaflets usually 9-13, initially appressed-silky but later glabrate beneath; bracts (evident only before anthesis) not aristate; plants with or without sparse long, hispid pubescence. 6 Plants fruiting abundantly; shrubs 1-3 m tall...... R. hispida var. kelseyi 6 Plants sterile (rarely fruiting scantily); shrubs or small trees 0.4-3 (-8) m tall. 7 Shrubs or small trees, 1-3 (-8) m tall, much branched, the stems and branches relatively straight, the nodes usually lacking spines; leaflets 1.5-3 cm long...... R. hispida var. rosea 7 Shrubs, 0.4-1 m tall, little branched, the stems and branches typically zigzag (bent at each node), the nodes usually with spines; leaflets 1-2 (-3) cm long...... R. nana

Robinia hartwigii Koehne, Granite Dome Locust, Highlands Locust, Hartwig's Locust. Mt (NC, SC): forests and outcrop edges on high elevation granitic domes, also clearings; rare (NC Rare). June-July; August-September. Apparently endemic to several mountains within a 5 km radius of Highlands, NC. While certainly related to and apparently hybridizing with R. viscosa, R. hartwigii seems worthy of recognition as a species. The original spelling (in Koehne 1913) is "hartwigii;" it is not clear why the variants (see synonymy) arose. [= RAB, S; = R. viscosa var. hartwegii (Koehne) Ashe – K, orthographic variant; = R. viscosa var. hartwigii – SE, Z; < R. viscosa – W; = R. viscosa var. hardwegii – Y, orthographic variant] Robinia hispida Linnaeus var. fertilis (Ashe) Clausen, Arnot Bristly Locust. Mt (NC): woodlands and forests; rare (NC Rare). May-June; July-August. Apparently endemic to the Southern Appalachians of w. NC and e. TN. A horticultural selection of var. fertilis, the Arnot Bristly Locust, is used as a soil binder. [= C, F, K, SE, Z; < R. hispida – RAB; > R. fertilis Ashe – S; > R. grandiflora Ashe – S, Y; > R. pedunculata Ashe – S; < R. hispida – W] Robinia hispida Linnaeus var. hispida, Common Bristly Locust. Mt, Pd*, Cp* (GA, NC, SC, VA); woodlands and forests, and as an escape in disturbed areas and roadsides; common (uncommon in Piedmont and Coastal Plain, where mostly or entirely introduced). May-June. Probably originally endemic to the Southern Appalachians (and perhaps adjacent provinces) of NC, SC, GA, and VA, now widely distributed in e. North America as an escape from cultivation. [= C, F, K, SE, Z; < R. hispida – RAB (also see R. hispida var. fertilis); = R. hispida – G, S, Y; > R. hispida – S; > R. pallida Ashe – S; > R. speciosa Ashe – S; < R. hispida – W] Robinia hispida Linnaeus var. kelseyi (Cowell ex Hutchinson) Isely, Kelsey's Locust. Mt (NC, SC), Pd* (VA*): mountain woodlands, introduced elsewhere; rare (NC Rare). April-July; July-October. Traditionally considered an endemic originally restricted to w. NC, but SE and Z suggest that var. kelseyi may have been only of horticultural origin. [= K, SE, Z; = R. kelseyi Cowell ex Hutchinson – RAB, G, S, Y; < R. hispida – W] Robinia hispida Linnaeus var. rosea Pursh, Boynton's Locust. Mt (GA, NC, SC, VA): mountain woodlands; rare (NC Watch List). April-July. Originally distributed from w. NC and e. TN south to nw. SC, n. GA, and ne. AL, now ocasionally found outside that range as an escape from cultivation. [= C, K, SE, Z; = R. boyntonii Ashe – RAB, G, S, Y; < R. hispida – W] Robinia nana Elliott, Dwarf Bristly Locust. Cp, Pd, Mt (GA, NC, SC): sandhills, dry rocky forests (especially associated with chestnut oak); uncommon (rare in Piedmont and escarpment region of Mountains). April-June; July-October. Se. and nc. NC south through SC to GA and AL. This species flowers infrequently. [= RAB, S, Y; = R. hispida Linnaeus var. nana (Elliott) Augustin de Candolle – K, SE, Z; = R. elliottii (Chapman) Ashe ex Small – RAB, F, G, S; < R. hispida – W; > R. nana – Y; > R. elliottii – Y] Robinia pseudoacacia Linnaeus, Black Locust. Mt, Pd, Cp (GA, NC, SC, VA): forests, woodlands, disturbed areas, roadcuts; common. April-June; July-November. Native in the s. and c. Appalachians, from PA south to GA and AL, now much more widespread, throughout e. and c. North America, also widely cultivated and escaped in Europe. Generally considered a weed tree. [= C, K, SE, Z; = R. pseudo-acacia – RAB, F, S, orthographic variant; > R. pseudo-acacia var. pseudo-acacia – G, orthographic variant; > R. pseudo-acacia var. rectissima (Linnaeus) Raber – G] Robinia viscosa Ventenat, Clammy Locust. Mt (GA, NC, SC), Pd* (NC), Cp* (NC): mountain forests and woodlands, roadsides, disturbed areas; rare in wild, uncommon as an escape (NC Watch List). May-July; July-August. Originally a Southern and Central Appalachian endemic, ranging from PA south through w. MD, w. VA, e. WV, w. NC, and e. TN, to n. GA and n. AL, now much more widespread as an escape from cultivation. [= RAB, F, G, S; = R. viscosa var. viscosa – C, K, SE, Y, Z; < R. viscosa – W (also see R. hartwigii)]

A variety of hybrids (including some cultivars) are known, including the following:

Robinia ×longiloba Ashe (pro sp.) [R. hispida × viscosa]. Known from NC and SC. Robinia ×margaretta Ashe (pro sp.) [R. hispida × pseudoacacia]. Known from NC, SC, and GA. Robinia ×ambigua Poiret (pro sp.) [R. pseudoacacia × viscosa]. Known from NC. Robinia hartwigii × hispida. Known from Whiteside Mountain, Jackson County, NC. Robinia hartwigii × viscosa. Known from Whiteside Mountain, Jackson County, NC.

FABACEAE 388

Securigera Augustin de Candolle (Crown-vetch)

A genus of about 12 species, annual and perennial herbs, of Eurasia. This genus is sometimes included in Coronilla, but is apparently better separated (Isely 1998). References: Isely (1998)=I.

1 Corolla yellow; annual; [rare waif]...... S. securidaca 1 Corolla white and pink; perennial; [common alien, planted and established]...... S. varia

* Securigera securidaca (Linnaeus) Degen & Dörfler. Cp (SC): disturbed areas; rare, native of Europe. Reported by Small (1933); rejected by Isely (1990) on the basis of no material seen to document the occurrence. This taxon’s status as part of our flora is uncertain. [= I, SE; = Bonaveria securidaca (Linnaeus) Reichenbach – S; = Coronilla securidaca Linnaeus] * Securigera varia (Linnaeus) Lassen, Crown-vetch. Mt, Pd (GA, NC, SC, VA), Cp (VA): roadbanks, woodland borders; common (rare in Piedmont south of NC), introduced from Europe. This species, generally known as Coronilla varia, is now widely used to stabilize road-cuts. [= I; = Coronilla varia Linnaeus – RAB, C, F, G, K, SE, W]

Senna P. Miller (Senna, Sicklepod, Wild Coffee)

A genus of about 350 species, trees, shrubs, and herbs, of tropical and warm temperate areas. References: Isely (1975)=Z; Irwin & Barneby (1982)=Y; Robertson & Lee (1976)=X; Isely (1998)=I; Marazzi et al. (2006).

1 Racemes spike-like, 3-6 (-10) dm long; legume winged; [section Senna, series Pictae] ...... [S. alata] 1 Racemes not spike-like, <3 dm long; legume not winged; [section Chamaefistula]. 2 Plant a shrub, 1-3 m tall; gland between the lowest pair of the acute or acuminate leaflets; [plants aliens, barely established in the vicinity of cultivation]; [section Chamaefistula, series Coluteoideae]. 3 Gland between the lowest pair of leaflets only; leaflets 3-5× as long as wide ...... S. corymbosa 3 Glands between each pair of leaflets; leaflets 2-3× as long as wide, acuminate...... S. septemtrionalis 2 Plant an herb, 0.1-1.5 m tall; gland near the base of the petiole (except in S. occidentalis which has leaflets rounded to emarginate at the apex); [plants natives, or aliens generally well-established and weedy]. 4 Leaflets obovate, the apex rounded to emarginate, 1.3-2× as long as wide; gland between the lowest pair of leaflets; [section Chamaefistula, series Trigonelloideae]...... S. obtusifolia 4 Leaflets ovate or narrowly elliptic, the apex acute or acuminate, 2-3.5× as long as wide; gland near the base of the petiole. 5 Leaflets 1.5-3.0 cm wide, in 3-6 pairs; racemes with 1-5 flowers; [section Chamaefistula, series Basiglandulosae]...... S. occidentalis 5 Leaflets 0.7-2.0 cm wide, in 6-10 pairs; racemes with 5-10 (-25) flowers; [series Temperatae]. 6 Legume 5.5-8 mm wide, with broad, nearly square segments, usually pilose initially, the hairs up to 0.8- 2 mm long (sometimes glabrate); ovary lanate with hairs to 1 mm long; ovules 10-15 (-18); petiolar gland broadest above the middle ...... S. hebecarpa 6 Legume 8-11 mm wide, with narrow segments (much shorter than broad), glabrous (or with a few hairs, these < 0.6 mm long); ovary strigulose with hairs to 0.5 mm long; ovules 20-25 (-30); petiolar gland usually broadest at or below the middle ...... S. marilandica

* Senna corymbosa (Lamarck) Irwin & Barneby. Cp (GA, SC), Pd (GA): cultivated as an ornamental, rarely persistent or spreading to disturbed areas; rare, introduced from South America. August-September. [= I, K, SE, Y; = Adipera corymbosa (Lamarck) Britton & Rose – S; = Cassia corymbosa – X, Z] Senna hebecarpa (Fernald) Irwin & Barneby, Northern Wild Senna. Pd (GA?, NC, VA), Mt (NC, VA), Cp (VA): open wet habitats, moist forests; uncommon in VA Mountains and VA Piedmont, rare in VA Coastal Plain and NC (NC Watch List). July-August; August-November. MA and s. NH west to s. WI, south to sc. NC, e. TN, s. IN, and c. IL. [= C, I, K, SE, Y; = Cassia hebecarpa Fernald – RAB, G, W, X, Z; > C. hebecarpa var. hebecarpa – F; > C. hebecarpa var. longipila E.L. Braun – F; = Ditremexa marilandica (Linnaeus) Britton & Rose – S, misapplied] Senna marilandica (Linnaeus) Link, Maryland Wild Senna. Pd (GA, NC, SC, VA), Mt, Cp (NC, GA, VA): dry to moist forests, especially on greenstone and diabase barrens and rocky woodlands, thickets, woodland borders, sometimes somewhat weedy; uncommon (rare in NC). July-August; August-November. S. MA and s. NY west to e. NE, south to c. peninsular FL and c. TX. [= C, I, K, SE, Y; = Cassia marilandica Linnaeus – RAB, F, G, W, X, Z; = Ditremexa medsgeri (Shafer) Britton & Rose – S] * Senna obtusifolia (Linnaeus) Irwin & Barneby, Sicklepod, Coffeeweed. Cp, Pd (GA, NC, VA), Mt (GA): fields (especially soybean fields), disturbed areas; common (uncommon in VA), probably introduced from the New World Tropics. July-September; August-November. The species is now pantropical. [= C, I, K, SE, X, Y; = Cassia obtusifolia Linnaeus – RAB, W, Z; < Cassia tora Linnaeus – F, G, misapplied; < Emelista tora (Linnaeus) Britton & Rose – S, misapplied] * Senna occidentalis (Linnaeus) Link, Coffee Senna. Cp, Pd (GA, NC, SC), Mt (VA): disturbed places; rare, native of the Old World Tropics. July-August; August-November. The species is now pantropical. [= C, I, K, SE, Y; = Cassia occidentalis Linnaeus – RAB, F, G, X, Z; = Ditremexa occidentalis (Linnaeus) Britton & Rose ex Britton & Wilson – S] FABACEAE 389 * Senna septemtrionalis (Viviani) Irwin & Barneby. Cp (NC): disturbed areas; rare, introduced from the tropics, where it is widespread, probably originally from tropical America, perhaps not truly established, though Isely (SE) states that "the weedy nature of this species suggests that it is almost certainly somewhat established." [= I, K, SE, Y; = Cassia laevigata Willdenow – Z]

* Senna alata (Linnaeus) Roxburgh, Emperor's Candlesticks, Candlestick Plant. September-November. Native to the American tropics, planted and slightly naturalized from s. AL and FL west to OK and TX. [= I, K, SE, Y; = Cassia alata Linnaeus – Z]

Sesbania Scopoli (Rattlebox, Sesban) (also see Glottidium)

A genus of about 50 species, annual herbs, perennial herbs, shrubs, and trees, of tropical, subtropical, and less commonly warm temperate regions of the Old and New World. References: Isely (1998)=I.

1 Corolla 8-9 mm long; legume flat; leaves with 8-13 pairs of leaflets ...... [see Glottidium vesicarium] 1 Corolla 10-25 mm long; legume quadrangular or 4-winged; leaves with 10-35 pairs of leaflets. 2 Legume quadrangular in cross-section, the corners not winged, 15-20 cm long, 0.3-0.5 cm wide; corolla 10-15 (-20) mm long, yellow, often marked with purple; leaves with 15-35 pairs of leaflets; [plant a robust herb, to 4 m tall]...... S. herbacea 2 Legume conspicuously 4-winged longitudinally, 3-8 cm long, 1-1.5 cm wide; corolla 13-25 mm long, yellow, orange, or scarlet; leaves with 10-20 pairs of leaflets; [plant a shrub, to 4 m tall]. 3 Corolla yellow; pedicels 0.5-1.0 cm long; legume blunt or short-acuminate to a beak...... S. drummondii 3 Corolla orange or red; pedicels 0.5-1.2 (-1.5) cm long; legume acuminate or tapering to beak...... S. punicea

* Sesbania drummondii (Rydberg) Cory, Rattlebox, Poison-bean. Cp (GA, SC): disturbed areas, spoil, marsh edges, ditches; rare, introduced. First reported for GA and SC by Townsend et al. (2000). [= GW, I, K, SE; = Daubentonia drummondii Rydberg – S] *? Sesbania herbacea (P. Miller) McVaugh, Sesban, Coffee-weed, Indigo-weed, Peatree. Cp (GA, NC, SC, VA), Pd (NC): ditches, wet fields; common (rare in Piedmont), perhaps native only in the deeper South. July-September; August-November. [= K; = S. exaltata (Rafinesque) Cory – RAB, C, F, G, I, SE; = Sesbania macrocarpa Muhlenberg ex Rafinesque – GW; = Sesban exaltatus (Rafinesque) Rydberg – S] * Sesbania punicea (Cavanilles) Bentham, Rattlebox, Scarlet Wisteria-tree, Purple Sesban. Cp (GA, NC, SC, VA), Pd (GA): ditches, wet fields, marshes, ponded wetlands, wet pinelands; common, presumably introduced from South America. June- October; August-November. [= GW, I, K, SE; = Daubentonia punicea (Cavanilles) Augustin de Candolle – RAB, S]

Sophora Linnaeus [see Styphnolobium]

Strophostyles Elliott (Sand Bean, Woolly Bean, Wild Bean)

A genus of 3 species, annual and perennial herbs, of North America. References: Pelotto & Martínez (1998)=Z; Isely (1998)=I. Key adapted from SE.

1 Legumes 2-4 cm long, permanently pubescent; corolla 5-8 mm long; leaves permanently pubescent on the upper surface; seeds glabrous...... S. leiosperma 1 Legumes 3-8 cm long, glabrate at maturity; corolla 8-15 mm long; leaves usually glabrate on the upper surface; seeds pubescent. 2 Bracteoles (immediately subtending the calyx) 2-3 mm long, equalling or exceeding the calyx tube; leaflets usually prominently 3-lobed; terminal leaflet 2.5-3.5 cm wide; plant an annual...... S. helvula 2 Bracteoles (immediately subtending the calyx) 0.5-1.0 (-1.5) mm long, shorter than the calyx tube; leaflets not lobed; terminal leaflet 0.3-2.0 cm wide; plant a perennial ...... S. umbellata

Strophostyles helvula (Linnaeus) Elliott, Annual Sand Bean. Cp, Pd, Mt (GA, NC, SC, VA): coastal dunes, beaches, dry sandy woodlands, disturbed areas; common. June-September; August-October. Québec west to MN and SD, south to n. peninsular FL and e. TX. See Isely (1986b) for a discussion of the orthography of the epithet. [= K, SE, W, Z; = S. helvola – RAB, C, G, S, orthographic variant; > S. helvola var. helvola – F; > S. helvola var. missouriensis (S. Watson) Britton – F] * Strophostyles leiosperma (Torrey & A. Gray) Piper, Small-flowered Sand Bean. Cp (VA): disturbed areas; rare, presumably introduced but possibly native (VA Watch List). This species is native east to KY and TN; it should be sought in our area in prairies, glades, and barrens with midwestern affinities. [= C, F, G, K, SE, Z; S. pauciflora (Bentham) S. Watson – S] FABACEAE 390 Strophostyles umbellata (Muhlenberg ex Willdenow) Britton, Perennial Sand Bean. Cp, Pd, Mt (GA, NC, SC, VA): dry sandy or rocky woodlands, disturbed areas; common. June-September; August-October. S. NY west to s. IN, s. MO, and KS, south to c. peninsular FL and s. TX. [= RAB, C, F, G, K, S, SE, W, Z; > S. umbellata var. umbellata – F; > S. umbellata var. paludigena Fernald – F]

Stylosanthes Swartz (Pencil-flower)

A genus of about 25 species, annual and perennial herbs, pantropical and less commonly temperate. References: Isely (1998)=I.

Stylosanthes biflora (Linnaeus) Britton, Sterns, & Poggenburg, Pencil-flower. Cp, Pd, Mt (GA, NC, SC, VA): sandhills, dry to moist (but not wet) pine savannas and flatwoods, dry forests, woodlands, woodland borders, glades, barrens, rock outcrops; common. June-August; July-October. S. NY west to OH, s. IL, and KS, south to c. peninsular FL and e. TX. The large, adnate stipules are distinctive. Variation in this species (see synonymy) needs additional study. [= RAB, C, I, K, SE, W; > S. biflora var. biflora – F, G; > S. biflora var. hispidissima (Michaux) Pollard & Ball – F, G; > S. riparia Kearney – G, S; > S. riparia var. riparia – F; > S. riparia var. setifera Fernald – F; > S. biflora – S]

Styphnolobium Schott (Pagoda Tree)

A genus of about 9 species, trees, shrubs, of central and South America and e. Asia. References: Isely (1998)=I; Isely (1981)=Z; Sousa S. & Rudd (1993)=Y; Palomino et al. (1993).

* Styphnolobium japonicum (Linnaeus) Schott, Pagoda Tree. Pd (GA, NC, SC, VA), Mt (VA): cultivated ornamental, native of China. Reported as "slightly escaped" in the United States by Isely (1981), but all specimens seen are from cultivated plants. Also reported for VA, MD, PA, and OH (Kartesz 1999). [= Y; = Sophora japonica Linnaeus – I, K, Z]

Tephrosia Persoon (Goat's-rue)

A genus of about 400 species, perennial herbs, of tropical and warm temperate regions of the Old World and New World. References: Isely (1998)=I; Ward (2004c)=Z; Wood (1949)=Y. Key adapted from SE.

1 Corolla bicolored, the standard yellow and the wings pink; racemes terminal; stems erect; stamens monadelphous; leaves with (9-) 13-23 (-37) leaflets. 2 Inflorescence reduced, foliose, flowers solitary or in small clusters overtopped by leaves; plants < 25 cm tall; leaflets generally < 10 mm long and < 5 mm wide; [restricted to the West Gulf Coastal Plain of sw. GA, adjacent FL and westward] ...... T. mohrii 2 Inflorescence terminal, not foliose and overtopped by leaves; plants > 25 cm tall; leaflets generally > 10 mm long and > 5 mm wide; [widespread in our area]...... T. virginiana 1 Corolla unicolored, initially white or pink, darkening in age to a dark maroon or purple; racemes opposite the leaves (the uppermost appearing terminal); stems decumbent or ascending; stamens diadelphous; leaves with (3-) 5-23 (-27) leaflets. 3 Upper stamen fused with the staminal sheath for part or most of its length (submonadelphous); leaves with (9-) 13-23 (- 27) leaflets; [plants from s. AL westward]...... [T. onobrychoides] 3 Upper stamen completely separate from the staminal sheath (diadelphous); leaves with (3-) 5-17 (-19) leaflets; [plants collectively widespread in our area]. 4 Petiole 1-4× as long as the lowest leaflets of the leaf; peduncle and rachis of inflorescence strongly flattened (2- angled, or rarely, 3-angled) in cross-section; leaflets averaging 25 mm long and 12 mm wide ...... T. florida 4 Petiole 1/3-1× as long as the lowest leaflets of the leaf; peduncle and rachis of inflorescence terete or inconpicuously 2-4-angled in cross-section; leaflets averaging smaller. 5 Leaves with (3-) 5-7 leaflets; petiole 0-5 mm long; stem and fruit hairs < 0.5 mm long ...... T. chrysophylla 5 Leaves with (7-) 9-17 (-19) leaflets; petiole 2-15 mm long; some stem and fruit hairs > 0.5 mm long. 6 Inflorescence with 1-3 (-5) nodes; plants inconpicuously pubescent with gray hairs (the hairs appressed or spreading, short to fairly long); leaflets (3-) avg. 5-6 (-7) mm wide, mostly acute; [plants of the Coastal Plain of NC and SC]...... T. hispidula 6 Inflorescence with 2-20 nodes; plants conspicuously tawny long-pilose with rusty brown hairs; leaflets (6-) avg. 8 (-12) mm wide, mostly obtuse; [plants widespread in our area] ...... T. spicata

Tephrosia chrysophylla Pursh, Sprawling Goat's-rue. Cp (GA): sandhills; rare (GA Rare). E. GA s. to s. FL, and west to s. MS. [= I, K, SE, Y; = Cracca chrysophylla (Pursh) Kuntze – S] Tephrosia florida (F.G. Dietrich) C.E. Wood. Cp (GA, NC, SC): pine savannas and other pinelands; common. May-July; June-September. E. NC south to s. FL, west to se. LA, a Southeastern Coastal Plain endemic. [= RAB, I, K, SE, Y; = Cracca ambigua (M.A. Curtis) Kuntze – S] FABACEAE 391 Tephrosia hispidula (Michaux) Persoon. Cp (GA, NC, SC, VA?): pine savannas and other pinelands; common. May- August; July-October. E. NC (se. VA?) south to c. peninsular FL, west to se. LA, a Southeastern Coastal Plain endemic. Fernald (1950) reports this species from se. VA. [= RAB, F, I, K, SE, Y; = Cracca hispidula (Michaux) Kuntze – S] Tephrosia mohrii (Rydberg) Godfrey, Dwarf Goat's-rue. Cp (GA): sandhills, dry savannas; rare (GA Rare). GA and westward in the East Gulf Coastal Plain. Perhaps not distinct from T. virginiana, but not easily dismissed as "little more than a freak" (Wood 1949); see Godfrey & Kral (1958). [= K; < T. virginiana – I, SE, Y; = Cracca mohrii Rydberg – S; = T. virginiana var. mohrii (Rydberg) D.B. Ward – Z] Tephrosia spicata (Walter) Torrey & A. Gray. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): woodlands; common. June- August; July-October. S. DE south to s. FL, west to w. LA, north in the interior to se., sc., and sw. TN and se. KY. [= RAB, C, G, K, SE, W, Y; > T. spicata var. semitonsa Fernald – F; > T. spicata var. spicata – F; = Cracca spicata (Walter) Kuntze – S] Tephrosia virginiana (Linnaeus) Persoon, Virginia Goat's-rue. Cp, Pd, Mt (GA, NC, SC, VA): sandhills, other pinelands, xeric and rocky woodlands and forests, outcrops, barrens; common. May-June; July-October. S. NH west to WI, se. MN, and c. KS, south to FL, c. TX, and nw. TX. [= RAB, C, I, K, SE, W; > T. virginiana var. glabra Nuttall – F, G; > T. virginiana var. virginiana – F, G; < T. virginiana – I, SE, Y (also see T. mohrii); = Cracca virginiana Linnaeus – S; = T. virginiana var. virginiana – Z]

Tephrosia onobrychoides Nuttall. Dry pinelands, s. AL west to c. TX. [= I, K, SE, Y; = Cracca onobrychoides (Nuttall) Kuntze – S]

Thermopsis R. Brown ex Aiton & Aiton f. (Golden-banner)

A genus of 8-10 species, perennial herbs, of temperate e. North America, w. North America, and e. Asia. References: Larisey (1940b); Chen, Mendenhall, & Turner (1994)=Y; Isely (1981)=Z; Isely (1998)=I.

1 Legumes erect or strongly ascending, densely villous; stipules clasping, those of the principal leaves (20-) 35-65 mm long, 10-30 mm wide; pedicels 2-3 mm long; plants mostly 6-20 dm tall, strict or few-branched ...... Th. villosa 1 Legumes spreading to ascending, glabrate or pubescent; stipules not clasping, those of the principal leaves 12-25 (-32) mm long, 1-5 mm wide; pedicels 4-20 mm long; plants mostly 3-10 dm tall, branched. 2 Plants from a single woody rootstock, mostly 5-10 dm tall; calyx glabrous or very sparsely pubescent, often also glaucous, the lobes often only 1-1.5 mm long; pedicels glabrate, (4-) 7-20 mm long (as long as or longer than the bracts); racemes terminal or lateral; plants flowering (late May-) early June-July; [plants of moderate to high elevations, (300-) 700-2000 m]...... Th. fraxinifolia 2 Plants from extensive rhizomes, mostly 3-6 dm tall; calyx pubescent, the lobes 2-2.5 mm long; pedicels villosulous, 2-6 (-10) mm long (shorter than the bracts); racemes terminal; plants flowering late April-early May (-June); [plants of low to moderate elevations, 200-800 m]...... Th. mollis

Thermopsis fraxinifolia (Nuttall ex Torrey & A. Gray) M.A. Curtis, Ash-leaf Golden-banner. Mt, Pd (GA, NC, SC): dry slopes and ridges; rare (GA Rare, NC Rare). Late May-July; July-October. A Southern Appalachian endemic: w. NC and e. TN south to nw. SC and n. GA. In addition to the key characters above, Th. fraxinifolia tends to have thinner stems than Th. mollis, to average taller, and to have the inflorescence generally arching to reclining (vs. erect to sometimes arching). The phenologic separation (peak flowering times separated by about 6-7 weeks, generally with a 2 week period between the last flowering of Th. mollis and the first flowering of Th. fraxinifolia) provides strong support to the recognition of Th. fraxinifolia and Th. mollis at the species level. [= RAB, K, S, W, Y; = Th. mollis var. fraxinifolia (Nuttall ex Torrey & A. Gray) Isely – I, SE, Z] Thermopsis mollis (Michaux) M.A. Curtis ex A. Gray, Appalachian Golden-banner. Pd, Mt (GA, NC, SC, VA): dry slopes and ridges; rare (GA Rare, NC Rare, SC Rare, VA Watch List). April-May; June-August. Centered in the Southern Appalachians, but mostly in the Piedmont and lower elevation periphery of the mountains, ranging from sc. VA south through w. and c. NC and e. TN to nw. SC, n. GA, and ne. AL. See comments under Th. fraxinifolia. [= RAB, C, F, G, K, W, Y; = Th. mollis var. mollis – I, SE, Z; > Th. hugeri Small – S; > Th. mollis – S] Thermopsis villosa (Walter) Fernald & Schubert, Aaron's-rod, Blue Ridge Golden-banner. Mt (GA, NC, VA*): floodplains, mesic disturbed areas, woodland edges, roadbanks; rare (GA Rare, NC Watch List). May-June; July-September. A Southern Appalachian endemic: w. NC and e. TN to n. GA, and escaped from cultivation more widely, as in w. VA, s. MD, c. TN, and WV probably representing escapes from cultivation. Th. villosa is a more erect and unbranched plant than our other 2 species. It is generally found in disturbed sites, its natural habitat somewhat of a mystery. [= RAB, C, I, K, SE, W, Y, Z; = Th. caroliniana M.A. Curtis – S]

Trifolium Linnaeus 1753 (Clover)

A genus of abut 240 species, annual and perennial herbs, nearly cosmopolitan. References: Isely (1998)=I. Draft key adapted from various published sources, including SE and C.

1 Flowers bright yellow (fading brown); [section Chronosemium]. 2 Leaves palmately trifoliolate (all leaflets essentially sessile); heads 10-13 mm in diameter; flowers 5-7 mm long ...... FABACEAE 392

...... T. aureum 2 Leaves pinnately trifoliolate (the lateral leaflets essentially sessile, the terminal leaflet with a petiolule 0.8-3 mm long; heads 5-13 mm in diameter; flowers 2.5-5 mm long. 3 Standard with 5 obvious diagonal veins (striations); heads 8-13 mm in diameter, generally with 20-30 flowers; flowers 3.5-5 mm long; petiolule of the terminal leaflet 1-3 mm long...... T. campestre 3 Standard inconspicuously veined; heads 5-8 mm in diameter, generally with 5-15 (-20) flowers; flowers 2.5-3.5 m long; petiolule of the terminal leaflet ca. 1 mm long...... T. dubium 1 Flowers not bright yellow. 4 Flowers borne on distinct pedicels, (1-) 2-10 mm long, these often curving or reflexing in age; flowers white, fading pink with age in most species; [native and alien species]; [section Lotoidea]. 5 Plants stoloniferous, all or some of the leaves alternate from ground level and long petioled. 6 Calyx lobes narrowly triangular, about as long as the calyx tube; peduncles axillary along the stolons; stipules scarious-membranaceous; [plant an abundant introduced weed]...... T. repens 6 Calyx lobes subulate, distinctly longer than the calyx tube; peduncles terminal, either at tips of the stolons, or at tips of erect flowering branches; stipules green, foliaceous; [plants rare natives]. 7 Peduncle terminal, at the tip of the stolon, lacking leaves; pedicels 2-3 mm long...... T. calcaricum 7 Peduncle terminal at tip of erect flowering branches, subtended by a pair of opposite or subopposite, short-petioled leaves; pedicels (2-) 4-8 mm long ...... [T. stoloniferum] 5 Plants not stoloniferous, clumped (though sometimes with prostrate or lax stems). 8 Calyx lobes narrowly triangular, about as long as the calyx tube (or longer in T. hybridum); stipules scarious- membranaceous; [plants introduced]. 9 Calyx lobes not scarious-margined, straight, equal to or longer than the tube ...... T. hybridum 9 Calyx lobes scarious-margined, becoming divergent and twisted, about equal to the tube ...... [T. nigrescens] 8 Calyx lobes subulate to lanceolate, distinctly longer than the calyx tube; stipules green, foliaceous; [plants rare natives]. 10 Flowers 4-6 mm long; calyx lobes lanceolate, foliaceous, 3-nerved, 0.4-0.8 mm wide ....T. carolinianum 10 Flowers 8-12 mm long; calyx lobes subulate, setaceous, 1-nerved, < 0.4 mm wide. 11 Leaflets 1-2.8× as long as wide; stems erect or ascending; flowers purplish; plant an annual or biennial; [plants of a variety of natural woodlands, widespread in our area]...... T. reflexum 11 Leaflets 3-7× as long as wide; stems prostrate; flowers creamy white and purple-veined; plant a perennial; [plants of shale barrens and other rock outcrops, from VA northward]...... T. virginicum 4 Flowers sessile or on very short pedicels (usually < 1 mm long); flowers pink, purplish, white, or scarlet; [alien species]. 12 Plants stoloniferous, all or some of the leaves alternate from ground level and long petioled. 13 All flowers with petals; fruiting heads enlarging, becoming a reddish brown, pubescent ball ca. 2 cm in diameter, remaining aerial; [section Vesicaria]...... T. fragiferum 13 Only 2-5 outer flowers of the head with petals, the others lacking petals and sterile; fruiting heads becoming a subterranean bur, buried by curvature and growth of the peduncle; [section Trichocephalum] ...... T. subterraneum 12 Plants not stoloniferous, the leaves clustered at or near ground level and/or produced on aerial stems. 14 Heads subtended by a pseudo-involucre of 2 (-3) enlarged stipules and/or opposite or subopposite leaves; [section Trifolium]. 15 Flowers white (fading pink), 7-8 mm long; calyx tube both externally glabrous and 20-nerved ...... T. lappaceum 15 Flowers red, pink-purple, or bicolored, either 11-20 mm long or 4-6 mm long; calyx tube not both externally glabrous and 20-nerved (externally pubescent, or 10-nerved, or both). 16 Flowers 4-6 mm long ...... T. striatum 16 Flowers 11-20 mm long. 17 Stipules gradually tapering to a long slender tip, longer than the fused part; calyx densely hirsute; stem soft pubescent with deflexed to spreading hairs...... T. hirtum 17 Stipules abruptly narrowed to a short awn; calyx glabrous to pilose; stem appressed pubescent ...... T. pratense 14 Heads not subtended by a pseudo-involucre of leaves or expanded stipules. 18 Heads axillary, sessile, in the axils of subtending leaves; calyx tube glabrous (except for a few hairs at apex); [section Lotoidea]...... T. glomeratum 18 Heads terminal or axillary; calyx tube pubescent. 19 Calyx bladdery-inflated in fruit; corolla resupinate (inverted 180 degrees, such that the standard is lowermost); [section Vesicaria]. 20 Inflorescence with a prominent peduncle; head lobed in fruit ...... T. resupinatum 20 Inflorescence subsessile to shortly peduncled; head spherical in fruit...... T. tomentosum 19 Calyx not bladdery-inflated in fruit; corolla orientation normal (standard uppermost). 21 Corolla 3-6 mm long; section Trifolium]...... T. arvense 21 Corolla 10-18 mm long. FABACEAE 393 22 Corolla crimson, 10-13 (-15) mm long; floral bracts absent; heads 1-1.5 (-2) cm in diameter; [section Trifolium] ...... T. incarnatum 22 Corolla white, 15-18 mm long; floral bracts present; heads 2.5-3 cm in diameter; [section Mistyllus]...... T. vesiculosum

* Trifolium angustifolium Linnaeus, Narrowleaf Clover. (SC). SC according to Kartesz (1999), based on specimen at NCU {check}. [= I, K] {not keyed at this time} * Trifolium arvense Linnaeus, Rabbitfoot Clover. Pd (GA, NC, SC, VA), Cp, Mt (NC, SC, VA): disturbed areas; common, introduced from the Mediterranean region. April-August. [= RAB, C, F, G, I, K, S, SE, W] * Trifolium aureum Pollich, Large Hop Clover, Yellow Clover. Mt, Pd, Cp (NC, VA): fields, roadsides, disturbed areas; uncommon, introduced from Eurasia. May-August. [= C, I, K, SE, W; = T. agrarium Linnaeus – RAB, F, G, S, misapplied] Trifolium calcaricum J.L. Collins & Wieboldt. Mt (VA): limestone glades; rare. In c. TN (Chester, Wofford, & Kral 1997). For additional information, see Collins & Wieboldt (1992). [= I, K] * Trifolium campestre Schreber, Hop Clover. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, fields, lawns, disturbed areas; common, introduced from Eurasia. April-October. [= RAB, C, I, K, SE, W; ? T. procumbens Linnaeus – F, G, S, misapplied] Trifolium carolinianum Michaux, Wild White Clover, Carolina Clover. Cp (GA, NC, SC), Pd (GA, SC): open woodlands, woodland edges, pine savannas, thin soils around rock outcrops; rare. April-July. [= RAB, C, F, G, I, K, S, SE, W; > T. carolinianum – S; > T. saxicola Small – S] * Trifolium cernuum Brot., Nodding-head Clover. (SC). (Kartesz 1999). [= K] {not keyed at this time} * Trifolium depauperatum Desvaux var. depauperatum. (SC). (Kartesz 1999). [= I, K] {not keyed at this time} * Trifolium dubium Sibthorp, Low Hop Clover. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, lawns, disturbed areas; common, introduced from Europe. April-September. [= RAB, C, F, G, I, K, S, SE, W] * Trifolium fragiferum Linnaeus, Strawberry Clover. Pd (GA): disturbed areas; rare, introduced from Middle East. Introduced in c. GA (Jones & Coile 1988) and reported from an old collection from se. PA (Rhoads & Klein 1993). [= C, F, G, I, K, SE] * Trifolium glomeratum Linnaeus, Cluster Clover. {prov} (SC): {habitat}; rare, introduced from Mediterranean region. [= I, K, S, SE] * Trifolium hirtum Allioni, Rose Clover. Pd (NC, VA): roadsides, disturbed areas; rare, introduced from Eurasia and n. Africa. April-July. [= RAB, C, G, I, K, SE] * Trifolium hybridum Linnaeus, Alsike Clover. Cp, Pd, Mt (GA, NC, SC, VA): lawns, fields, roadsides, disturbed areas; common, introduced from Europe. April-September. [= RAB, C, G, I, K, S, SE, W; > T. hybridum var. hybridum – F; > T. hybridum var. elegans (Savi) Boiss. – F] * Trifolium incarnatum Linnaeus, Crimson Clover. Cp, Pd, Mt (GA, NC, SC, VA): fields, disturbed areas; uncommon, introduced from Europe. April-June; June-August. [= RAB, C, F, G, I, K, S, SE, W] * Trifolium lappaceum Linnaeus, Lappa Clover. Cp (NC): disturbed areas; rare, introduced from Mediterranean Eurasia and Africa. April-August. [= RAB, I, K, S, SE] * Trifolium pratense Linnaeus, Red Clover. Mt, Pd, Cp (GA, NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from Europe. April-September. [= RAB, C, G, I, K, S, SE, W; > T. pratense var. pratense – F; > T. pratense var. sativum (P. Miller) Schreber – F] Trifolium reflexum Linnaeus, Buffalo Clover. Pd, Cp (GA, NC, SC, VA), Mt (GA, SC): open woodlands, woodland edges; rare. April-August. [= RAB, C, I, K, S, SE, W; > T. reflexum var. reflexum – F, G; > T. reflexum var. glabrum Lojacono – F, G] * Trifolium repens Linnaeus, White Clover, Dutch Clover, Ladino Clover. Cp, Pd, Mt (GA, NC, SC, VA): lawns, roadsides, disturbed areas; common, introduced from Eurasia. April-September. [= RAB, C, F, G, I, K, S, SE, W] * Trifolium resupinatum Linnaeus, Persian Clover, Reversed Clover. Cp (NC, SC), Pd (GA): lawns and disturbed areas; rare, introduced Mediterranean region and w. Asia. April-July. [= RAB, C, F, G, I, K, S, SE] * Trifolium striatum Linnaeus, Knotted Clover. Pd (GA, NC): roadsides, disturbed areas; rare, introduced from Europe. April-August. [= RAB, C, F, G, I, K, S, SE] * Trifolium subterraneum Linnaeus, Subterranean Clover. Pd (GA), {prov.} (NC, SC): disturbed areas; rare, introduced from Europe, Asia, and n. Africa. Reported for NC and SC by Isely (1990); reported for Piedmont of GA by Jones & Coile (1988). [= I, K, SE] * Trifolium tomentosum Linnaeus. {prov.} (NC): disturbed areas; rare, introduced from Mediterranean region. Reported for NC by Isely (1998). [= I, K] * Trifolium vesiculosum Savi, Arrowleaf Clover. Pd (GA, SC), Cp (GA): roadsides, disturbed areas; uncommon, introduced from s. Europe. First reported for South Carolina by Hill & Horn (1997). [= I, K, SE] Trifolium virginicum Small, Kates Mountain Clover, Shale-barren Clover. Mt, Pd (VA): shale barrens, other rock outcrops; uncommon (rare in Piedmont). May-June. Ranges from sc. PA through w. MD south to w. VA and e. WV. [= C, F, G, I, K, SE, W]

* Trifolium medium Linnaeus, Zigzag Clover. Introduced in MD (Kartesz 1999). [= K] {not keyed} * Trifolium nigrescens Viviani, Ball Clover. Introduced in c. TN (Chester, Wofford, & Kral 1997). [= I, K, S, SE] Trifolium stoloniferum Muhlenberg ex Eaton, Running Buffalo-clover, a rare native, occurs (at least formerly) in dry upland woodlands and prairies from WV, OH, n. IN, IL, MO, and e. KS, south to KY and AR. [= C, F, G, I, K, S, SE]

FABACEAE 394

Ulex Linnaeus (Gorse)

References: Isely (1998)=I.

* Ulex europaeus Linnaeus, Gorse, Furze. Cp (VA): disturbed areas; rare, introduced from Europe. June. Not cited in Harvill et al. (1992), but naturalized in sandy soils in York County, VA. Also reported from WV and PA. [= C, F, G, I, K, SE]

Vachellia Wight & Arnott 1834 (Acacia)

A genus of about 163 species, of tropoical and subtropical America, Africa, Asia, and Australia. Formerly considered part of Acacia. References: Isely (1998)=I; Isely (1969)=Z; Ebinger, Seigler, & Clarke (2002)=Y; Seigler & Ebinger (2005)=X; Maslin, Miller, & Seigler (2003).

1 Leaves with 2-4 (-6) pairs of pinnae; each pinna with 10-20 pairs of leaflets...... A. farnesiana var. farnesiana 1 Leaves with 10-15 (-20) pairs of pinnae; each pinna with 20-30 pairs of leaflets...... A. macracantha

Vachellia farnesiana (Linnaeus) Wight & Arnott var. farnesiana, Sweet Acacia. Cp (GA): sandy flats on barrier islands, maritime scrub; rare, apparently native but possibly adventive from further south (GA Special Concern). E. GA, along the coast, south to s. FL. See Duncan (1985). [= X; < Acacia farnesiana (Linnaeus) Willdenow – I, K, SE, Z; < Vachellia farnesiana (Linnaeus) Wight & Arnott – S; = Acacia farnesiana ssp. farnesiana – Y] * Vachellia macracantha (Humboldt & Bonplandt ex Willdenow) Seigler & Ebinger, Apopanax. Cp (GA): planted as an ornamental and rarely naturalized; rare, introduced from further south. [= X; = Acacia macracantha Humboldt & Bonplandt ex Willdenow – I, K, SE, Z; Vachellia]

Vicia Linnaeus 1753 (Vetch, Tare)

A genus of about 150 species, annual and perennial herbs, of temperate Eurasia and North America. References: Isely (1998)=I; van de Wouw, Maxted, & Ford-Lloyd (2003)=Y. Key adapted from I.

1 Inflorescence nearly sessile, of 1-4 flowers clustered in the leaf axil; [alien species]. 2 Leaves with 2-6 leaflets, succulent; leaflets 3-7 cm long; legume with pectinate sutures...... [V. narbonensis] 2 Leaves with 4-20 leaflets, not succulent; leaflets 0.3-3.5 cm long; legume not pectinate (except V. lutea). 3 Corolla 5-6 mm long; leaves with 4-6 (-8) leaflets...... V. lathyroides 3 Corolla 10-30 mm long; leaflets 6-16 (-20). 4 Calyx lobes conspicuously unequal; legumes pilose with pustulate-based hairs...... V. lutea 4 Calyx lobes more or less equal; legumes glabrous at maturity, or very finely pubescent with non-pustulate- based hairs. 5 Calyx lobes all shorter than the calyx tube; corolla yellow, often streaked with purple, 25-30 mm long...... V. grandiflora 5 Calyx lobes (at least the longer) about as long as the calyx tube; corolla pink, purple, lavender, white, or creamy yellow, 10-25 (-30) mm long. 6 Standard pubescent dorsally; corolla 15-25 (-30) mm long, creamy yellow to purple; legume pubescent, with a basal stipe...... V. pannonica 6 Standard glabrous; corolla 10-25 (-30) mm long, pink, purple, lavender, or whitish; legume glabrous, sessile. 7 Calyx 7-11 (-12) mm long; corolla pink-purple to whitish, 10-18 mm long; leaflets 4-10× as long as wide...... V. sativa ssp. nigra 7 Calyx 10-15 mm long; corolla generally pink-purple, 18-25 (-30) mm long; leaflets 2-5 (-7)× as long as wide ...... V. sativa ssp. sativa 1 Inflorescence pedunculate, of 2-many flowers along a well-developed raceme; [alien and native species]. 8 Peduncles 1-10 mm long; raceme axis 2-10 mm long, with 2-7 (-10) flowers. 9 Plant a robust annual, 10-20 dm tall; tendrils absent; leaves with (2-) 4-6 leaflets; leaflets 5-10 cm long; corolla 20-30 mm long...... [V. faba] 9 Plant a trailing perennial, 3-10 dm tall; tendrils present; leaves with 8-16 leaflets; leaflets 2-3.5 cm long; corolla 10-15 mm long...... [V. sepium] 8 Peduncles usually >10 mm long; raceme axis usually >10 mm long, with (1-) 2-many flowers. 10 Corolla 10-25 mm long. 11 Stipules dimorphic, one of each pair entire, the other palmately lacerate; flowers 1 (-2) per inflorescence...... [V. articulata] 11 Stipules of a pair alike; flowers 1-numerous per inflorescence. FABACEAE 395 12 Flowers 15-22 (-25) mm long; legumes with a basal stipe 2-5 mm long; leaves with 8-16 leaflets ...... V. americana var. americana 12 Flowers 8-16 (-18) mm long; legumes with a basal stipe 1-3 mm long; leaves with 8-22 leaflets. 13 Calyx swollen on one side; plant an annual; inflorescence not secund. 14 Plant glabrate or with pubescence of hairs < 1 mm long; lower calyx lobe lanceolate to linear- lanceolate, 1-2 (-2.4) mm long ...... V. villosa ssp. varia 14 Plant conspicuously villous, the hairs 1-2 mm long; lower calyx lobe acicular or weak, 2-4 mm long...... V. villosa ssp. villosa 13 Calyx not swollen on one side; plant a rhizomatous perennial; inflorescence not secund. 16 Flowers white to lavender, the keel spotted; legumes 4-5 mm wide; inflorescence not second ...... V. caroliniana 16 Flowers blue-violet or purple; legumes 6-8 mm wide; inflorescence generally second...... V. cracca 10 Corolla 2.5-8 (-10) mm long. 17 Plant an annual. 18 Legume symmetrically rounded at the apex; inflorescence with 1-2 (-4) flowers...... V. tetrasperma 18 Legume asymmetrically acute at the apex; inflorescence with 1-15 flowers. 19 Legume finely hirsute; leaves with (8-) 10-16 leaflets ...... V. hirsuta 19 Legume glabrous to inconspicuously puberulent; leaves with 2-4 leaflets...... V. minutiflora 17 Plant a rhizomatous perennial. 18 Leaves with 2-4 (-6) leaflets; [plants of s. SC southward, native, of the Coastal Plain]. 19 Legumes 2.5-3.0 cm long; leaflets 1.5-4.5 cm long, oblong to linear, 8-20× as long as wide...... V. acutifolia 19 Legumes 0.8-1.5 cm long; leaflets 1-1.5 cm long, usually elliptic, 2-4 (-10)× as long as wide...... V. floridana 18 Leaves with 10-25 leaflets; [plants collectively widespread in our area, native or alien]. 20 Flowers white to lavender, the keel spotted; legumes 4-5 mm wide; inflorescence not second ...... V. caroliniana 20 Flowers blue-violet or purple; legumes 6-8 mm wide; inflorescence generally secund ...... V. cracca

Vicia acutifolia Elliott, Fourleaf Vetch. Cp (GA, SC): pond margins, pine flatwoods, ditches; rare. April-May; May-June. Se. SC south to s. FL, west to e. Panhandle FL. [= RAB, GW, I, K, S, SE] Vicia americana Willdenow var. americana, American Vetch, Purple Vetch, Tare. Mt (VA): {habitat}; rare. May-June. Var. americana ranges from Québec west to AK, south to w. VA, MO, OK, TX, Mexico. Var. minor Hooker occurs in w. North America. [= C, F, G, I, SE; = V. americana ssp. americana – K; < V. americana – W] Vicia caroliniana Walter, Pale Vetch, Wood Vetch, Carolina Vetch. Mt, Pd, Cp (GA, NC, SC, VA): forests, woodlands, and disturbed areas; common. April-June; May-July. NY west to WI, south to s. GA, s. MS, and c. TX. [= C, F, G, I, K, SE, W; > V. caroliniana – RAB, S; > V. hugeri Small – RAB, S] * Vicia cracca Linnaeus, Tufted Vetch, Cow Vetch, Canada-pea. Cp, Pd (VA), Mt (NC): disturbed areas; rare, introduced from Europe. May-July; June-August. [= RAB, C, G, S, SE; > V. cracca var. cracca – F, I; > V. cracca ssp. cracca – K] Vicia floridana S. Watson, Florida Vetch. Cp (GA): moist soils of hammocks, ditches, roadbanks; rare. S. FL north to McIntosh Co., GA (Isely 1998). [= GW, I, K, S, SE] * Vicia grandiflora Scopoli, Large Yellow Vetch. Cp, Pd (GA, NC, SC, VA): disturbed areas; uncommon, introduced from Europe. April-June; May-July. [= C, I, F, G, K, SE, W; > V. grandiflora var. kitaibeliana W.D.J. Koch – RAB] * Vicia hirsuta (Linnaeus) S.F. Gray, Tiny Vetch, Hairy Tare. Cp, Pd (GA, NC, SC, VA), Mt (SC, VA): disturbed areas, uncommon, introduced from Europe. April-June; May-July. [= RAB, C, F, G, I, K, S, SE] * Vicia lathyroides Linnaeus, Spring Vetch. Cp (NC, SC, VA), Pd (GA, NC): lawns, disturbed areas; rare, introduced from Europe. April-June; May-July. [= RAB, C, I, F, G, K, SE] * Vicia lutea Linnaeus, Yellow Vetch. Pd (NC): disturbed areas; rare, introduced from Europe. [= I, K, SE] Vicia minutiflora F.G. Dietrich, Smallflower Vetch. Cp (GA): woodlands; rare. TN, w. FL, and sw. GA west to OK and TX. [= GW, I, K, SE, V; = V. micrantha Nuttall ex Torrey & A. Gray – F, G, S] * Vicia pannonica Crantz, Hungarian Vetch. Pd (GA): disturbed areas; rare, introduced. Introduced in c. GA. Reported for NC (Kartesz 1999). {investigate} [= I, K, SE] * Vicia sativa Linnaeus ssp. nigra (Linnaeus) Ehrhart, Narrowleaf Vetch. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas; common, introduced from Mediterranean Europe. March-June; May-July. [= I, K, SE; = V. angustifolia Linnaeus – RAB, C, S, W; V. sativa var. angustifolia (Linnaeus) Ehrhart; > V. angustifolia var. angustifolia – F, G; > V. angustifolia var. segetalis (Thuill.) Ser. – F, G; > V. angustifolia var. uncinata (Desv.) Rouy – F] * Vicia sativa Linnaeus ssp. sativa, Common Vetch. Pd (GA, NC, VA), Mt (NC), Cp (VA): disturbed areas; rare, introduced from Mediterranean Europe. April-June; May-July. [= I, K, SE; = V. sativa – RAB, C, G, S; > V. sativa var. sativa – F; > V. sativa var. linearis Lange – F] * Vicia tetrasperma (Linnaeus) Schreber, Slender Vetch, Smooth Tare, Lentil Vetch. Cp, Pd (GA, NC, SC, VA): disturbed areas; uncommon, introduced from Europe. April-June; May-July. [= RAB, C, G, I, K, S, SE; > V. tetrasperma var. tetrasperma – F; > V. tetrasperma var. tenuissima Druce – F] FABACEAE 396 * Vicia villosa Roth ssp. varia (Host) Corbière, Winter Vetch. Mt, Pd, Cp (GA, NC, SC, VA): disturbed areas; common, introduced from Europe. May-September. [= I, K, SE; = V. dasycarpa Tenore – RAB, C, F, G, W] * Vicia villosa Roth ssp. villosa, Hairy Vetch, Fodder Vetch. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): disturbed areas; common, introduced from Europe. May-September. [= I, K, SE; = V. villosa – RAB, C, F, G, W]

* Vicia articulata Hornemann, Monantha Vetch. Probably only cultivated. [=I, K, SE] * Vicia faba Linnaeus, Horse Bean, Faba Bean, Broad Bean. Introduced in se. PA (Rhoads & Klein 1993). Also reported for VA (Kartesz 1999). {investigate} [= C, F, G, I, K, SE] Vicia ludoviciana Nuttall ssp. ludoviciana, Louisiana Vetch. AL and FL west to OR and CA. [= K] * Vicia narbonensis Linnaeus, Narbonne Vetch. Introduced in MD and DC (Fernald 1950). [= C, F, G, I, K, SE] * Vicia sepium Linnaeus, Bush Vetch, Wild Tare. Introduced south to WV and in e. PA (Rhoads & Klein 1993). [= C, G, I, SE; > L. sepium var. sepium – F, K]

Vigna Savi (Cow Pea)

A genus of about 150 species, annual and perennial herbs, pantropical, rarely extending into warm temperate regions. References: Isely (1998)=I; Maréchal, Mascherpa, & Stainier (1978)=Z.

1 Corolla yellow, 1.5-1.7 cm long; leaves somewhat fleshy-thickened; [plant native or introduced in maritime situations]; [section Vigna] ...... V. luteola 1 Corolla pink to purple, 1.5-2.5 cm long; leaves herbaceous; [plant a cultivated introduction]; [section Catiang]...... V. unguiculata

Vigna luteola (Jacquin) Bentham, Wild Cow Pea. Cp (GA, NC, SC): edges of freshwater tidal marshes, beaches, disturbed areas, railroad embankments, low fields, in the outer Coastal Plain; rare (GA Special Concern). July-September; August- October. Se. NC south to FL, west to se. TX, and in the New World tropics. Often weedy in appearance, and its nativity in our area perhaps uncertain. [= RAB, GW, I, K, Z; ? V. repens (Linnaeus) Kuntze – S; = V. marina (Burmann) Merrill (the correct name according to some authors, based on uncertain typification)] * Vigna unguiculata (Linnaeus) Walpers, Black-eyed Pea, Field Pea, Cow Pea. Cp, Pd (GA, NC, SC, VA): commonly cultivated in commercial and home gardens, rarely persistent or occurring as a waif, introduced from tropical Africa or Asia. June-August; July-September. [= RAB, I, K; ? V. sinensis (Linnaeus) Savi – F, S; > V. unguiculata var. unguiculata – Z]

Wisteria Nuttall (Wisteria)

A genus of about 6 species, woody vines, shrubs, and small trees, of temperate e. Asia and e. North America. Some research suggests that the Asian species should be placed in a separate genus (see Isely 1998). References: Isely (1998)=I; Valder (1995)=Z; Stritch (1984)=Y.

Identification notes: Twining direction can be determined by looking at (or imagining) the vine twining around a branch or pole. Look at the pole or branch from the base (from the direction from which the vine is growing). If the vine is circling the branch or pole in a clockwise direction, that is dextrorse; if counterclockwise, that is sinistrorse.

1 Legume and ovary glabrous; pedicels 5-10 (-15) mm long; standard reflexed near the middle; seeds reniform; [native species of swamps and bottomland forests and thickets]...... W. frutescens 1 Legume and ovary velvety pubescent; pedicels 15-20 mm long; standard reflexed at the base; seeds lenticular; [introduced species, naturalized in a wide variety of situations]. 2 Vine twining counter-clockwise (sinistrorse; from lower right ascending to upper left); leaflets 11-19 per leaf; racemes 2-5 (-7) dm long, the 50-170 flowers opening successively from base to tip of the inflorescence; corolla 1.5-2.0 cm long ...... W. floribunda 2 Vine twining clockwise (dextrorse; from lower left ascending to upper right); leaflets 7-13 per leaf; racemes 1-2 (-3.5) dm long, the 25-95 flowers opening nearly simultaneously; corolla 2.0-2.5 (-2.7) cm long...... W. sinensis

2 Standard 20-23.5 mm long, 21-23 mm wide; leaflets (7-) 9-11 (-13) per leaf; raceme to 33 cm long, with 25-95 flowers opening nearly simultaneously...... W. sinensis 2 Standard 16-18 mm long, 16-18 mm wide; leaflets 7-17 (-19) per leaf; raceme to 132 cm long, with 25-170 flowers opening nearly simultaneously or sequentially. 3 Auricles of the standard’s callosity 1.1-1.2 mm long; leaflets (11-) 13-17 (-19) per leaf; raceme to 132 cm long, with the 50-170 flowers opening successively from base to the tip of the inflorescence, those at the base withering before those at the tip have opened ...... W. floribunda 3 Auricles of the standard’s callosity 0.7-0.8 mm long; leaflets 7-17 per leaf; racemes to 36 cm long .... W. ×formosa

FABACEAE 397 * Wisteria floribunda (Willdenow) Augustin de Candolle, Japanese Wisteria. Cp, Pd, Mt (GA, NC, SC, VA): commonly cultivated, sometimes escaped to urban, suburban, and rural forests and woodlands, introduced from Japan. April-July; July- November. [= RAB, C, F, G, I, K, SE, Z; = Kraunhia floribunda (Willdenow) Taubert – S; = Rehsonia floribunda (Willdenow) Stritch – Y] * Wisteria ×formosa Rehder, Hybrid Asian Wisteria. Cp, Pd, Mt (GA, NC, SC, VA): commonly cultivated, sometimes escaped to urban, suburban, and rural forests and woodlands, introduced from Japan. April-July; July-November. [= Rehsonia ×formosa (Rehder) Stritch – Y] Wisteria frutescens (Linnaeus) Poiret, American Wisteria, Swamp Wisteria, Atlantic Wisteria. Cp (GA, NC, SC, VA), Pd (GA, NC), Mt (GA, VA): swamp forests, wet thickets; common. April-May; June-September. E. VA south to FL, west to TX, north in the interior to AR, s. IN, and s. MO. The issue of the distinctiveness of W. frutescens and W. macrostachya needs further study. Harvill et al. (1992) reports W. macrostachya from Northumberland and Shenandoah counties, VA. [= RAB, GW, I, K, SE, W; > W. frutescens – C, F, G, Z; > W. macrostachya (Torrey & A. Gray) Nuttall ex B.L. Robinson & Fernald – C, F, G, Z; > Kraunhia frutescens (Linnaeus) Greene – S; > Kraunhia macrostachya (Torrey & A. Gray) Small – S] * Wisteria sinensis (Sims) Augustin de Candolle, Chinese Wisteria. Cp, Pd (GA, NC, VA), Mt (GA): commonly cultivated, commonly escaped to urban, suburban, and rural forests and woodlands, introduced from China. April-July; July-November. [= RAB, C, F, I, K, SE; = Rehsonia sinensis (Sims) Stritch – Y]

Zornia J. F. Gmelin (Zornia)

A genus of about 50-90 species, perennial herbs, of tropical and warm temperate regions. References: Isely (1998)=I.

Identification notes: The palmately 4-foliolate leaves are unique in the flora of our area.

Zornia bracteata Walter ex J.F. Gmelin. Cp (GA, NC, SC, VA): sandy roadsides, woodlands, and sandhills; common. June-August; July-October. Se. VA south to FL, west to TX and e. Mexico, endemic to the Southeastern Coastal Plain. [= RAB, C, F, G, K, S, SE]

FAGACEAE Dumortier 1829 (Beech Family)

A family of about 8 genera and 620-1050 species, trees and shrubs, mostly of the Northern Hemisphere, but extending into se. and Australia. References: Nixon in FNA (1997); Govaerts & Frodin (1998); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993); Elias (1971a).

1 Fruits partially enclosed in a scaly cup; leaves lobed, toothed, crenate, or entire ...... Quercus 1 Fruits enclosed in a spiny or prickly bur; leaves toothed. 2 Nuts rounded or flattened on one or two sides; bur with long, straight spines; winter buds < 1 cm long; leaves elliptic or oblanceolate, some of them usually > 12 cm long ...... Castanea 2 Nuts sharply triangular; bur with short, recurved prickles; winter buds 1.5-2.5 cm long; leaves ovate, 6-12 cm long ...... Fagus

Castanea P. Miller 1754 (Chestnut, Chinquapin)

A genus of 8-10 species, trees and shrubs, of temperate regions of the Northern Hemisphere. References: Johnson (1988)=Z; Nixon in FNA (1997); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993); Stanford (1998). Key adapted from Z.

1 Leaves elliptic to oblanceolate, mostly < 15 cm long, the apices acute to obtuse; twigs puberulent; spine-covered husk of fruit splitting into 2 sections, enclosing 1 nut; nut circular in cross-section, 7-19 mm in diameter; pistillate dichasia of 1 flower; leaves with stellate trichomes, with few bulbous-based trichomes when young, puberulent, pilose, tomentulose, or tomentose in age (usually rather densely so). 2 Longest spines of the fruit husk usually >10 mm long; young twigs glabrous; petiole 8-10 (-15) mm long; [plants of n. AL and westward]...... [C. ozarkensis] 2 Longest spines of the fruit husk usually <10 mm long; young twigs puberulent; petiole 3-7 (-10) mm long; [plants widespread in our area]...... C. pumila 1 Leaves elliptic, oblanceolate or lanceolate, 8-30 cm long, the apices acuminate, sometimes only shortly so; spine-covered husk of fruit splitting into 4 sections, enclosing (2-) 3 (-5) nuts; nut flattened on at least one side, 18-25 mm in diameter; pistillate dichasia of 3 flowers; leaves usually without stellate trichomes; twigs puberulent or glabrous. 3 Undersurface of leaves densely covered with bulbous-based trichomes when young, essentially glabrous in age; leaves mostly > 15 cm long, generally long-acuminate; twigs glabrous...... C. dentata 3 Undersurface of leaves persistently and densely tomentose beneath; leaves mostly < 15 cm long, generally short- acuminate; twigs puberulent ...... C. mollissima

FAGACEAE 398 Castanea dentata (Marshall) Borkhausen, American Chestnut. Mt (GA, NC, SC, VA), Pd (GA, NC, VA), Cp (GA, VA): mesic and xeric forests; common (uncommon in Piedmont, rare in Coastal Plain). June-July; September-October. Formerly one of the most important, largest, and most abundant forest trees in the Mountains of our area, C. dentata was severely affected by chestnut blight, Cryphonectria parasitica (Murrill) Barr, introduced at New York City in 1904 on nursery stock of C. mollissima. Blight spread steadily southward, reaching our area in the 1920's and 1930's. C. dentata remains rather abundant, but now occurs only as stump sprouts and small trees, usually reinfected by blight persisting on oaks and killed at about the size of first fruit production. The accidental introduction of chestnut blight and the subsequent profound alteration of the role of chestnut is one of the most tragic ecological disasters to have affected our area. [= RAB, C, F, FNA, G, K, S, W, Z] * Castanea mollissima Blume, Chinese Chestnut. Pd, Mt (NC): forests; rare, introduced and naturalized from China. June; September. This species is relatively resistant to chestnut blight and has been planted widely as an ornamental and nut tree; it sometimes naturalizes and appears nearly native. Reported for NC (Macon County) by Pittillo & Brown (1988). [= C, FNA, K] Castanea pumila (Linnaeus) P. Miller, Common Chinquapin. Cp, Pd, Mt (GA, NC, SC, VA): xeric forests and woodlands, generally in fire-maintained habitats; uncommon. May-July; September-October. NJ, s. PA, s. OH, n. KY, and s. MO, south to c. FL and se. TX. It is relatively resistant to chestnut blight. [= FNA; = C. pumila var. pumila – C, K, Z (in a broad sense, in relation to var. ozarkensis); > C. pumila var. pumila – RAB, F (in a narrow sense); > C. pumila var. ashei Sudworth – RAB, F; > C. alnifolia Nuttall var. alnifolia – RAB; > C. alnifolia var. floridana Sargent – RAB; > C. pumila – G, S, W; > C. ashei (Sudworth) Sudworth – S; > C. floridana (Sargent) Ashe – S; > C. alnifolia – S]

Castanea ×neglecta Dode (pro sp.) [C. dentata × pumila], occurs in our area; "the leaves of the hybrid resemble those of C. dentata in size and shape but have the vestiture and stellate trichomes of C. pumila" (Johnson 1988). [= K, Z] {not keyed} Castanea ozarkensis W.W. Ashe, Ozark Chinquapin, is mainly distributed in the Ozarks and Ouachitas of AR, MO, and OK, but also occurs (at least formerly) as a disjunct in nc. AL. It is affected by the chestnut blight. Related to C. pumila, though showing some relation as well to C. dentata, is C. ozarkensis Ashe [sometimes treated as C. pumila var. ozarkensis (Ashe) Tucker]. C. ozarkensis is more susceptible to blight, and occurs in s. MO, e. OK, and w. AR, and disjunct in c. AL, where now apparently extirpated by blight. [= FNA, S; = C. pumila P. Miller var. ozarkensis (W.W. Ashe) G.E. Tucker – K, Z] * Castanea sativa P. Miller, Spanish Chestnut. Reported as naturalized in KY, AL, PA, and elsewhere in e. North America (Clark et al. 2005). [= K] {not keyed at this time; synonymy incomplete}

Fagus Linnaeus 1753 (Beech)

A genus of about 10 species, trees, of temperate regions of the Northern Hemisphere. Our native trees belong to subgenus Fagus, section Grandifolia (Shen 1992). References: Cooper & Mercer (1977)=Z; Nixon in FNA (1997); Shen (1992)=X; Kubitzki in Kubitzki, Rohwer, & Bittrich (1993); Elias (1971a)=Y; Stanford (1998); Govaerts & Frodin (1998)=V.

1 Leaves denticulate; cupule prickles 1-2.5 (-4) mm long, slightly to strongly recurved; cupule valves generally ovate, the apex obtuse, reddish in color; leaves with fairly dense white acicular trichomes on the lower leaf surface at maturity; leaf base sometimes clearly cordate; [plants of the Coastal Plain, Piedmont, and low to moderate elevations (mostly below 1050 m or 3500 feet) in the Mountains]...... F. grandifolia var. caroliniana 1 Leaves sharply serrate; cupule prickles (3-) 4-10 mm long, projecting forward; cupule valves generally narrowly triangular, the apex acute, grayish-yellow in color; leaves usually lacking white acicular trichomes on the lower surface at maturity, instead with yellowish glandular hairs (acicular trichomes often present on the veins); leaf base not clearly cordate; [plants of moderate to high elevations (mostly above 1050 m or 3500 feet) in the Mountains] ...... F. grandifolia var. grandifolia

Fagus grandifolia Ehrhart var. caroliniana (Loudon) Fernald & Rehder, White Beech, American Beech. Cp, Pd, Mt (GA, NC, SC, VA): moist forests, from near sea level to low elevations in the Mountains, mostly below 1050 meters (3500 feet); common (uncommon in Coastal Plain south of VA). March-April; September-October. Se. MA, OH, IN, s. IL, s. MI (?), and MO south to Panhandle FL and e. TX. Several subspecies, varieties, or phases of Fagus grandifolia have been described, and their taxonomic recognition is controversial. The most recent monographer, Shen (1992), recognizes three subspecies, one of which is limited to Mexico, the other two as treated here but at the subspecific level. I have here chosen to recognize 2 intergradient varieties in our area. A third variety, var. mexicana (Martínez) Little, of the mountains of México, is apparently most closely related to var. grandifolia. Cooper & Mercer (1977) studied variation in NC, concluding that two genetic races or varieties were present, the montane var. grandifolia and the Piedmont and Coastal Plain var. caroliniana, but that patterns of variation were complicated. Hardin & Johnson (1985) and Hardin (1992, 1985) note that variation is "more-or-less" clinal, variation within populations is great, and they do not favor recognition of infraspecific taxa. Depending on one's tolerance or intolerance for intergradational varieties, one may choose to recognize one or two taxa in our area. [= C, F, G, Y; < F. grandifolia – RAB, FNA, K, S, W, Z; < F. grandifolia ssp. grandifolia – V; = F. grandifolia ssp. caroliniana (Loudon) Camp ex Shen – X, nomen nudum; = F. ferruginea Aiton] Fagus grandifolia Ehrhart var. grandifolia, Gray Beech, Red Beech, American Beech. Mt (NC, VA): moderate to high elevation forests, especially on high elevation ridges, gaps, and open slopes, often forming clonal dwarfed thickets in the most exposed situations; common. April; September-October. Nova Scotia, New Brunswick, and s. Québec west to s. Ontario and n. MI, south to VA, w. NC, n. GA, e. TN, and n. OH. "Red beech" is alleged to differ from "gray beech" in having the involucral segments not covering the nutlets at maturity. Hardin & Johnson (1985), Hardin (1985), and Shen (1992) point out that var. mexicana (Martínez) Little, of the mountains of México, is more closely related to our montane variety or phase than to the lower FAGACEAE 399 elevation variety or phase. [= C, F, G, Y; < F. grandifolia – RAB, FNA, K, S, W, Z; < F. grandifolia ssp. grandifolia – V; = F. grandifolia ssp. grandifolia – X]

* Fagus sylvatica Linnaeus, European Beech, Copper Beech, of subgenus Fagus, section Fagus, is sometimes cultivated, but is not known to escape in our area. It has only 5-10 (-12) lateral veins, as opposed to (12-) 15-18 (-20) in F. grandifolia. {not keyed} [= V; ? F. sylvatica ssp. sylvatica – X]

Quercus Linnaeus 1753 (Oak)

A genus of about 350-530 species, trees and shrubs, of temperate, subtropical, and rarely tropical regions of the Northern Hemisphere. Oaks are the predominant tree of our area, with a variety of species dominating much of the landscape in nearly every ecological situation. Only in a few specialized (and usually in some sense edaphically extreme) communities are oaks generally entirely absent: deepest Coastal Plain swamps, some Coastal Plain depression ponds, wettest pine savannas, pocosins, spruce-fir forests, highest elevation northern hardwood forests, and mountain bogs. Our oaks are divided into two well-marked sections; other sections occur outside our area. Red oaks (section Lobatae, sometimes treated as subgenus Erythrobalanus) are characterized by acorns maturing in two years (in one year in Qu. elliottii), styles elongate, abortive ovules are at the top of the seed, leaves and leaf lobes bristle-tipped, inner surface of the acorn shell velvety-pubescent, and acorns rooting in spring. Twenty-one of our Quercus species are in this group: Qu. arkansana, Qu. coccinea, Qu. elliottii, Qu. falcata, Qu. georgiana, Qu. hemisphaerica, Qu. ilicifolia, Qu. imbricaria, Qu. incana, Qu. laevis, Qu. laurifolia, Qu. marilandica var. marilandica, Qu. myrtifolia, Qu. nigra, Qu. pagoda, Qu. palustris, Qu. phellos, Qu. rubra var. ambigua, Qu. rubra var. rubra, Qu. shumardii var. shumardii, and Qu. velutina.

White oaks (section Quercus, sometimes treated as subgenus Quercus) are characterized by acorns maturing in a single year, styles short or absent, abortive ovules at the base of the seed, leaves and leaf lobes not bristle-tipped, inner surface of the acorn shell smooth, and acorns rooting in autumn. Twenty of our Quercus species are in this group: Qu. alba, Qu. austrina, Qu. bicolor, Qu. boyntonii, Qu. chapmanii, Qu. geminata, Qu. lyrata, Qu. macrocarpa, Qu. margaretta, Qu. michauxii, Qu. minima, Qu. montana, Qu. muehlenbergii, Qu. oglethorpensis, Qu. prinoides, Qu. robur, Qu. similis, Qu. sinuata var. sinuata, Qu. stellata, and Qu. virginiana. Hybrids within each section are frequent and diverse; hybrids do not naturally occur between the two sections. The live oaks of the southeastern Coastal Plain and Central America are subsection Virentes; other white oaks are subsection Quercus. References: Nixon in FNA (1997) (overall treatment); Jensen in FNA (1997) (red oaks); Nixon & Muller in FNA (1997) (white oaks); Godfrey (1988); Stein, Binion, & Acciavatti (2003); Cronquist (1991); Duncan & Duncan (1988); Hunt (1990)=Z; Hunt (1994); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

Many oak species are well-adapted to ecological situations in which fires frequently burn the ground layer. Fire-maintained communities of the Piedmont and Mountains typically have oaks such as Qu. stellata, Qu. marilandica var. marilandica, Qu. ilicifolia, and Qu. prinoides. The two latter species are normally shrubby, and have become rarer because of fire suppression (they require fire to prevent larger trees from outcompeting them). In contrast, Qu. stellata and Qu. marilandica var. marilandica become larger and more frequent in fire-suppressed conditions.

Fifteen oak species in our area are typical of upland Coastal Plain communities with at least occasional fire: Qu. arkansana, Qu. chapmanii, Qu. geminata, Qu. hemisphaerica, Qu. incana, Qu. laevis, Qu, margerettiae, Qu. marilandica var. marilandica, Qu. minima, Qu. myrtifolia, Qu. stellata, and less typically Qu. falcata, Qu. nigra, Qu. velutina, and Qu. virginiana. Fire suppression of Coastal Plain communities, especially of sandhills, leads to an unnatural increase in the stature and abundance of oaks present. In frequent fire conditions, most oaks will persist as short, shrubby fire sprouts. Additional suggestions of how to recognize fire sprouts of hese species are given below. In general, leaves of fire sprouts are larger and more deeply lobed than normal leaves. In species of the red oak group, the bristle tips are larger and more pronounced. Increased size in leaves is particularly noticeable when an area previously long fire-suppressed is burned (the large underground root system and nutritional resources of a small tree destroyed by fire being devoted to a few very vigorous sprouts). Fire sprouts are often in sunny conditions, which tend to make oak leaves more deeply lobed and more coriaceous than shaded leaves.

White oaks with lobed leaves: Qu. margaretta – Often forms dense clonal, stoloniferous patches in frequent fire conditions. Tends to retain standard leaf characteristics. Qu. stellata – Less prone to formal clonal patches. Sprout leaves often very large, with exaggerated lobing.

Red oaks normally with deeply lobed leaves: Qu. laevis – Not clonal. Vigorous sprouter, leaves more deeply lobed than any other fire red oak. Small sprouts often have vertical leaf orientation characteristic of adults, though vigorous fire sprouts sometimes have more normally disposed leaves. Sprout leaves sometimes very large, with very long, curving lobes. Qu. falcata – Not clonal. Sprout leaves generally less lobed than typical adult leaves, more like forma triloba, but larger and coarser in texture, difficult to distinguish in shape from Qu. marilandica var. marilandica and Qu. velutina. See pubescence differences in main key. FAGACEAE 400 Qu. velutina – Not clonal. Leaves variable, sometimes minimally lobed and closely resembling Qu. marilandica var. marilandica and Qu. falcata. See pubescence differences in main key.

Red oaks normally with unlobed leaves: Qu. marilandica var. marilandica – Sprout leaves sometimes coarsely (though never deeply) lobed. Texture often very coriaceous, shiny, and very stiff. See pubescence characters in main key. Qu. nigra – Not very typically in fire-prone situations, but sometimes so. Young saplings, as well as fire sprouts, often with wildly different leaves than the typical adult form, frequently deeply lobed (for excellent illustrations showing variability in leaf shapes, see p. 329 of Godfrey, 1988 and pp. 51-52 of Godfrey & Wooten, 1981). Leaves always smaller and more glabrous than those of other fire oaks (except Qu. hemisphaerica). Qu. incana – Generally not strongly clonal and stoloniferous even in frequently burned situations. Fire sprouts and vigorous shoots more prone to lobing than adult trees. Even fire shoots, though, usually with only one to several lobes, and the characteristic bluish-green characteristic holds. See comments above on Qu. elliottii. Qu. elliottii – Strongly clonal via a stoloniferous "runner", never tree-like. Leaves never lobed, even on fire sprouts, though fire sprout leaves can be larger (to 15 cm long and 5 cm wide). Very difficult to tell from fire sprouts or seedlings of Qu. incana, best separated by leaf pubescence (white in Qu. elliottii, gray in Qu. incana), margin (slightly revolute in Qu. elliottii, flat in Qu. incana), leaf vernation (planate in Qu. incana, rolled in Qu. elliottii), and acorn maturation (1 year in Qu. elliottii, with acorns often on small plants, 2 years in Qu. incana, with small plants rarely produing acorns). Qu. hemisphaerica – Not clonal. Leaves of vigorous shoots and fire sprouts often shallowly lobed, the lobing usually fairly neat and regular, triangular-ascending, and with bristle tips.

Live oaks: Qu. geminata – Sometimes clonal. Leaves, even of sprouts, not normally with lobes or teeth. Qu. virginiana – Sometimes clonal. Leaves of vigorous summer shoots (but apparently not spring shoots) often coarsely toothed, very similar to similar leaves of Qu. hemisphaerica, but lacking bristle tips (instead the translucent margin with a darker, thickened callus at the tip of the tooth). Qu. minima – Always clonal. Leaves often with teeth or lobes.

Identification notes: Some oaks with ambiguous leaves are keyed in both Key A and Key D or in both Key B and Key C. The leaves of juvenile (seedling or sapling) branches, fire-sprout shoots, or other vigorous shoots (resulting from similar stimuli such as insect damage) are often much different than typical leaves and are not accounted for in these keys (see discussion at end of generic treatment). Hybrids are frequently encountered; they, too, are not keyed here, but can usually be identified (with difficulty) by their intermediate morphology and by parental context. Trichome types are useful in making and confirming identifications of oaks, since certain types are restricted to various groups of species. Hand lenses of 10× or 20× can be useful, but a compound microscope is far preferable. See Hardin (1992, 1976, 1979), and Thomson & Mohlenbrock (1979).

1 Most of the leaves on a relatively mature tree entire and unlobed (some species frequently with some leaves on a tree, especially those on young or vigorous growth, that are toothed or shallowly lobed, the teeth or lobes generally few and irregular in size or location); [primarily either "laurel oaks" of section Lobatae or "live oaks'"of section Quercus]...... Key A 1 Most of the leaves on a relatively mature tree lobed or toothed. 2 Venation of the leaves neatly and evenly pinnate, the 3-17 (or more) main lateral veins on each side distinctly parallel to each other, each ending in a tooth or shallow, crenate lobe extending < ¼ of the way to the midrib; ["chestnut oaks" of sections Quercus and Cerris]...... Key B 2 Venation of the leaves pinnate, but more branched and irregular, the 1-7 main lateral veins on each side rebranching into prominent side veins, the leaf usually distinctly lobed, at least some of the lobes of some of the leaves of a tree extending > ¼ of the way to the midrib 3 Apices of the lobes or teeth obtuse (rarely acute), lacking bristle tips; ["white oaks" of section Quercus]...... Key C 3 Apices of the lobes or teeth acuminate (rarely acute), and with bristle tips; ["red oaks" of section Lobatae]...Key D

Key A – Leaves (most of them) entire and unlobed (Laurel Oaks and Live Oaks)

1 Leaves broadly obovate or spatulate, 1-2.5 (-3)× as long as wide. 2 Leaves 10-30 cm long, with rounded, subcordate, truncate, or oblique bases; lower leaf surfaces thinly to densely pubescent with tawny to orange glandlike hairs; [section Lobatae]...... Qu. marilandica var. marilandica 2 Leaves 2-10 (-15) cm long, mostly with cuneate or rounded bases (in some species sometimes subcordate, truncate, or oblique); lower leaf surfaces glabrous, glabrescent, or pubescent, but the pubescence not orange and glandlike. 3 Twigs of the current year densely and finely hairy, obscuring the surface; [scrubby trees of sandhills from se. SC southward]; [section Quercus] ...... Qu. chapmanii 3 Twigs of the year glabrous or sparsely pubescent; [shrubs, scrubby small trees, or large trees of various habitats]. 4 Leaves grayish beneath; [section Quercus] ...... Qu. sinuata var. sinuata 4 Leaves bright green beneath; [section Lobatae]. FAGACEAE 401 5 Leaves evergreen, (including the petiole) usually < 4 cm long (sometimes to 9 cm long) and < 2 cm wide (to 6 cm wide); lower leaf surface usually entirely glabrous at maturity (rarely with pubescence in the vein axils); leaf blades rarely lobed; [shrub to scrubby tree of sandhills in se. SC and southward]Qu. myrtifolia 5 Leaves deciduous, (including the petiole) usually > 5.5 cm long (rarely smaller) and usually 3-5 cm wide; lower leaf surface usually with tufts of hairs in the main vein axils beneath; leaf blades often lobed. 6 Leaves with broadly cuneate to rounded leaf bases, the blades 5-15 cm long; lower leaf surfaces generally pubescent across the surface, and also with tufts in the axils; [of sw. GA westward]...... Qu. arkansana 6 Leaves with cuneate bases, the blades 5-10 (-15) cm long; lower leaf surfaces glabrous, except for tufts of hairs in the vein axils; [widespread in our area]...... Qu. nigra 1 Leaves linear, elliptic, or narrowly obovate, 2-10× as long as wide. 7 Leaves (at maturity) glabrous or at most sparsely pubescent on the surface below, though often with tufts of hairs in the main vein axils. 8 Twigs of the year densely and finely hairy, obscuring the surface; leaves (at maturity) sparsely pubescent beneath; [scrubby trees of sandhills from se. SC south]; [section Quercus]...... Qu. chapmanii 8 Twigs of the year glabrous or sparsely pubescent; leaves (at maturity) bright green and glabrous beneath, though often with tufts of hairs in the main vein axils; [medium to large trees, more widespread, mostly of moist habitats, except Qu. hemisphaerica]; [section Lobatae]. 9 Leaves predominantly lanceolate, mostly 6-12 cm long and 0.7-2 cm wide, most of them 5-8× as long as wide, the apex acute; mature leaves with tufts of hairs in the vein axils below, and sometimes also some pubescence on the blade surface near the midrib; blades never with lobes or teeth; leaves deciduous in autumn; young leaves bronze red, emerging tightly rolled lengthwise and appearing linear; [trees of bottomlands and upland depression swamps, mesic uplands, and also weedy and frequent in disturbed successional habitats]...... Qu. phellos 9 Leaves predominantly oblanceolate, obovate, or rhombic, mostly 2.5-10 cm long and 1.5-4 cm wide, most of them 2-5× as long as wide, the apex acute, obtuse, or rounded; mature leaves with or without tufts of hairs in the vein axils below, lacking pubescence on the blade surface; blades sometimes with 1-5 lateral lobes or teeth; leaves persisting until spring, or tardily and irregularly deciduous in winter; young leaves red, yellow, or green, not emerging tightly rolled lengthwise; [trees primarily either of swamp forests, maritime forests, or sandhills, not typically weedy]. 10 Mature leaves entirely glabrous below; leaves mostly with acute apices and bristle tips (rarely a few rounded), mostly 2.5-8 cm long and 1-2 (-3) cm wide, the upper surface shiny, the vein network not readily visible when backlit; leaves evergreen (persisting until spring); petiole 0.5-2 mm long; leaves of vigorous growth often with dentate lobes; [trees of dry sandy habitats, such as sandhills and maritime forests]...... Qu. hemisphaerica 10 Mature leaves with tufts of stellate trichomes in the vein axils; leaves mostly with rounded apices (rarely a few acute and then bristle-tipped), mostly 5-10 cm long and (1.8-) 2-4 cm wide, the upper surface dull, the vein network readily visible when backlit; leaves tardily deciduous; petiole 2-6 mm long; leaves of vigorous growth rarely lobed, and then not dentate; [trees of moist habitats, such as floodplain forests]...... Qu. laurifolia 7 Leaves (at maturity) persistently and densely pubescent on the surface below, the pubescence in some species so dense and tight as to be difficult to perceive without at least 10× magnification. 11 Leaves bristle-tipped (sometimes the bristle fallen or broken off, but leaving a truncate scar), deciduous in autumn; multi-armed trichomes of the rosulate or multiradiate types, many of the arms ascending or erect (never with the stellate or fused-stellate trichomes characteristic of the live oaks); acorns maturing in 2 years (immature acorns present through the winter on fruiting trees); [section Lobatae]. 12 Leaves (including petiole) mostly 10-17 cm long, 3.5-7 cm wide; lower leaf surface (at maturity) sparsely to moderately densely pubescent with soft hairs; leaves lustrous dark-green above; [trees of the Mountains, Piedmont, and rarely Coastal Plain] ...... Qu. imbricaria 12 Leaves (including petiole) mostly 4-11 cm long, 0.5-3.0 cm wide; lower leaf surface densely covered with soft hairs; leaves lustrous dark-green or bluish-green above; [stoloniferous shrubs and small to medium trees of the Coastal Plain]. 13 Leaves 1.5-3.0 cm wide, mostly 2-4× as long as wide, dull bluish-green above; acorns 10-15 mm long; petioles 4-15 mm long; [plant a small to medium tree] ...... Qu. incana 13 Leaves 0.5-1.5 cm wide, mostly 4-8× as long as wide, lustrous dark-green above; acorns 8-12 mm long; petioles1-3 mm long; [plant a stoloniferous shrub, to 1 m tall (or to 2 m in fire-suppressed pinelands)]...... Qu. elliottii 11 Leaves not bristle-tipped, evergreen (overwintering, falling with the expansion of new leaves in the spring) or deciduous (in Qu. oglethorpensis); multi-armed trichomes of the fused-stellate and stellate types, the arms parallel to the leaf surface, radiating from a well developed disc that appears as a white eye or dot at 20-40× magnification (or rosulate or multiradiate in Qu. oglethorpensis); acorns maturing in 1 year (immature acorns not present through the winter, unless aborted); [section Quercus]. FAGACEAE 402 14 Leaves deciduous in autumn; bark gray, resembling Qu. alba; [trees of bottomlands and upland clay flats of GA and SC]...... Qu. oglethorpensis 14 Leaves evergreen (overwintering, falling with the expansion of new leaves in the spring); bark (on the tree species) brownish, deeply furrowed; [trees and stoloniferous shrubs of sandy habitats of the Coastal Plain of GA, NC, SC, and VA]. 15 Plant a stoloniferous shrub, to 1 m tall (or to 2 m in fire-suppressed pinelands)...... Qu. minima 15 Plant a small to large tree. 16 Leaf blades with the margins strongly revolute, and also the sides of the blades generally rolled downward and obscuring part of the lower surface, the leaf appearing boatlike (the depth of the "boat" often approaching the width of the leaf); midvein and major lateral veins impressed on the upper surface and raised on the lower surface (the lower surface therefore appearing rugose); pubescence of the lower surface stellate, both appressed and erect, the individual stellae readily visible at 20× magnification (sometimes at 10× magnification); acorns (1-) 2 (-6) per stalk; [typically a small tree of sandhills near the coast]...... Qu. geminata 16 Leaf blades flat, or the margins slightly to strongly revolute, the sides of the blade sometimes rolled downward, usually not obscuring part of the lower surface, the leaf not boatlike (the leaf much wider than deep); midvein and major lateral veins not impressed (or very slightly so) on the upper surface and only very slightly, if at all, raised on the lower surface (the lower surface therefore not appearing notably rugose); pubescence of the lower surface stellate, all of it tightly appressed, the individual stellae readily visible only at 30× magnification (sometimes barely distinguishable at 20× magnification); acorns 1-2 per stalk; [typically a salt-pruned shrub to large tree of dunes, estuarine shorelines, and southward of upland flats and slopes]...... Qu. virginiana

Key B – Leaves with even crenations or teeth (Chestnut Oaks)

1 Scales of the acorn cup prolonged and long tapered; lateral veins terminating in a well-developed bristle; [species planted, rarely escaped]; [section Cerris] ...... Qu. acutissima 1 Scales of the acorn cup acute to obtuse; lateral veins terminating in a minute mucro or hardened projection; [species native]; [section Quercus].× 2 Acorns on peduncles (2-) 4-7 cm long; acorns 1.5-2.5 cm long; veins ending in crenations usually 6-10 on each side of leaf ...... Qu. bicolor 2 Acorns sessile or on peduncles 0-1 cm long; acorns 1-2 cm or 2.5-3.5 cm long; veins ending in crenations usually 8-15 or 3-7 (if 3-7, then a stoloniferous shrub). 3 Leaves mostly obovate, with rounded teeth (crenations), the teeth sometimes with a minute mucro; hairs of the leaf undersurface clustered in sessile, stellate-appearing clusters of 2-8 hairs; acorns 2.5-3.5 cm long; large trees. 4 Hairs of the leaf undersurface in clusters with a diameter of 0.15-0.5 mm, dense to sparse; bark of mature trees light gray, loose, breaking into plates or scales...... Qu. michauxii 4 Hairs of the leaf undersurface in clusters with a diameter of 0.1-0.25 mm, sparse; bark of mature trees dark gray, tight, deeply furrowed ...... Qu. montana 3 Leaves mostly narrowly elliptic, narrowly ovate, or narrowly obovate, with sharp ascending, often incurved teeth, the teeth ending in a hardened projection; hairs of the leaf undersurface tiny and stellate, with 6-16 rays; acorns 1- 2 cm long; medium to large trees or stoloniferous shrubs. 5 Medium to large tree; veins ending in teeth usually 7-13 on each side of the leaf; leaves 8-20 cm long and 4- 10 cm wide...... Qu. muehlenbergii 5 Stoloniferous shrub to 5 m tall; veins ending in teeth usually 3-7 on each side of the leaf; leaves 4-10 cm long and 2-6 cm wide...... Qu. prinoides

Key C – Leaves with lobes not bristle-tipped (White Oaks)

1 Lower surfaces of mature leaves glabrous. 2 Leaf lobes with acute apices; sinuses often both broad and "flat-bottomed" (with portions parallel to the midrib); acorn cup covering 2/3 to 3/4 of acorn...... Qu. lyrata 2 Leaf lobes with obtuse apices; sinuses narrow (often notch-like), narrowly to broadly rounded or triangular (lacking portions parallel to the midrib); acorn cup covering 1/4 to 1/2 of acorn. 3 Leaves mostly 4-10 (-17) cm long, 2-5 (-9) cm wide, with 1-5 shallow lobes or undulations, extending 1/8 to 1/2 of the way to the midrib; acorn cup flat at the base, covering < ¼ of the acorn...... Qu. sinuata var. sinuata 3 Leaves mostly 7-20 cm long, 3-10 cm wide, with 3-11 lobes, extending 1/4 to 5/6 of the way to the midrib (if the lobing < ½ of the way to the midrib, then the acorn cup rounded at the base and covering 1/4 to 1/2 of the acorn). 4 Leaf base deeply cordate; [alien, sometimes planted and persistent] ...... [Qu. robur] 4 Leaf base cuneate; [native] FAGACEAE 403 5 Leaves with 7-11 lobes (the sinuses usually deep, those of the larger leaves usually about 2/3 to 5/6 of the way to the midrib), 10-20 cm long, 5-10 cm wide; terminal bud rounded or globose; basal scales of acorn cup thickened, the thickening giving the cup a knobby texture ...... Qu. alba 5 Leaves with 3-7 lobes (the sinuses usually shallow, those of the larger leaves usually ranging from 1/4 to 1/2 of the way to the midrib), 7-15 cm long, 3-8 cm wide; basal scales of the acorn cup thin, appressed, the cup having a rough but not knobby texture ...... Qu. austrina 1 Lower surfaces of mature leaves pubescent, the pubescence varying from dense to sparse (sometimes minute and requiring 10× magnification to be readily visible). 6 Lower surfaces of mature leaves whitish to pale green, with a mixture of minute, sessile, stellate hairs with horizontal tips and longer stellate hairs with erect ascending tips; leaves shallowly lobed (if so, the lobes 9-19) to deeply lobed (if so, the lobes with acute apices), the sinuses extending 1/4 to 4/5 of the way to the midrib. 7 Leaves mostly shallowly lobed at the base, the sinuses extending 1/4 to 1/2 of the way to midrib, grading into mere crenations toward the tip of the leaf, the total number of lobes/crenations usually 9-19; acorns borne on peduncles 2-10 cm long; acorn cup covering 1/3 to 1/2 of acorn, the upper scales with long-acuminate apices ...... Qu. bicolor 7 Leaves mostly relatively deeply lobed throughout the length of the leaf, the sinuses extending 1/2 to 4/5 of the way to the midrib, the total number of lobes 3-13; acorns sessile or borne on peduncles up to 1 cm long; acorn cup covering 1/3 to 3/4 of acorn, the upper scales with acute, long-acuminate, to long-awned apices. 8 Upper scales of the acorn cups thin and acute; acorn cup covering ½ to 3/4 of the acorn; [swamps in the Coastal Plain and lower Piedmont of GA, NC, SC, and VA]...... Qu. lyrata 8 Upper scales of the acorn cups long-attenuate into nearly terete awns; acorn cup covering 1/3 to 1/2 of the acorn; [Mountains of VA] ...... Qu. macrocarpa var. macrocarpa 6 Lower surfaces of mature leaves gray, green, pale green, or yellowish, glabrescent or densely pubescent, the hairs few- branched and erect; leaves mostly relatively deeply and obtusely lobed, rarely shallowly lobed (if so, the lobes 3-7), the sinuses extending 1/2 to 4/5 of the way to the midrib, the total number of lobes 3-7; acorns sessile or nearly so. 9 Leaf lobes with acute apices; acorn cup covering 2/3 to 3/4 of acorn ...... Qu. lyrata 9 Leaf lobes with obtuse to rounded apices; acorn cup covering 1/3 to 1/2 of acorn. 10 Woody twigs of the season glabrous or with scattered, deciduous 2-forked hairs; petioles of mature leaves 3- 10 (-15) mm long; leaf blades (2.5-) 4-8 (-13.5) cm long, irregularly and often rather shallowly 3-5 (-7) lobed, the overall form of the leaf only rarely cruciform; largest lateral lobes usually at the midpoint of the blade (or even below it), the lobes usually not sublobed, tapering from base to tip; [xeric sandy sites in the Coastal Plain from se. VA southward]...... Qu. margaretta 10 Woody twigs of the season densely and persistently stellate-pubescent, especially toward the tip of the twig; petioles of mature leaves 15-20 mm long (Qu. stellata) or 3-10 (-15) mm long (Qu. boyntonii and Qu. similis); leaf blades (5-) 7.5-15 (-20) cm long, usually 5-lobed, the overall form of the leaf typically cruciform (Qu. stellata) or not (Qu. boyntonii and Qu. similis); largest lateral lobes of the leaves usually above the midpoint of the blade, these lobes often sublobed or squarish in shape, usually wider near their tips than at their bases (Qu. stellata) or not sublobed, tapering from base to tip (Qu. boyntonii and Qu. similis); [collectively widespread in our area]. 11 Leaves usually cruciform, the largest lateral lobes often sublobed or squarish in shape, usually wider near their tips than at their bases, and borne at right angles to the midrib; [usually of dry to dry-mesic upland situations, widespread in our area] ...... Qu. stellata 11 Leaves not cruciform, the largest lateral lobes usually not sublobed, tapering from base to tip, and borne at ascending angles relative to the midrib; [of temporarily flooded calcareous swamps of the Coastal Plain, from SC (NC?) southward in our area (Qu. similis) or localized on sandstone in nc. AL (Qu. boyntonii)]. 12 Rhizomatous shrubs to small trees, generally < 2 m tall; [of sandstone outcrops in nc. AL] ...... [Qu. boyntonii] 12 Single-trunked large trees; [usually of temporarily flooded calcareous swamps of the Coastal Plain, from SC (NC?) southward in our area]...... Qu. similis

Key D – Leaves with lobes or teeth bristle-tipped (Red Oaks)

1 Leaves shallowly 3-lobed near the broad apex (some leaves of sprout or juvenile shoots may be more lobed). 2 Leaf blades 5-15 cm long; lower leaf surfaces glabrous, except for tufts of hairs in the vein axils (or pubescent across the surface in Qu. arkansana]. 3 Leaves with broadly cuneate to rounded leaf bases, the blades 5-15 cm long; lower leaf surfaces generally pubescent across the surface, and also with tufts in the axils; [of sw. GA westward] ...... Qu. arkansana 3 Leaves with cuneate bases, the blades 5-10 (-15) cm long; lower leaf surfaces glabrous, except for tufts of hairs in the vein axils; [widespread in our area]...... Qu.nigra 2 Leaf blades 10-30 cm long; lower leaf surfaces pubescent across the surface (and often also with denser tufts of hairs in the vein axils). FAGACEAE 404 4 Petioles short and stout, 5-15 mm long; lower leaf surfaces thinly to densely pubescent with a mixture of tawny or orange glandlike hairs and stellate hairs whose structure is easily visible at 10× magnification...... Qu. marilandica var. marilandica 4 Petioles long and slender, (14-) 20-50 mm long; lower leaf surfaces densely puberulent with tawny stellate hairs whose structure is barely visible at 10× magnification ...... Qu. falcata 1 Leaves shallowly to deeply 5-12-lobed (some of the leaves of Qu. georgiana only 3-lobed), the lobes primarily lateral. 5 Mature leaves pubescent beneath on the surface with stellate hairs. 6 Leaves 5-10 (-12) cm long, 5-lobed; shrub or small tree; [w. NC northward] ...... Qu. ilicifolia 6 Leaves (8-) 10-20 cm long, 5-12-lobed; small to large trees[collectively widespread in our area]. 7 Petioles 0.5-1.0 (-1.8) cm long, generally twisted such that the blade is oriented in a vertical plane; leaves all deeply lobed, some of the sinuses extending > 4/5 of the way to the midrib; pubescence of the lower leaf surface greenish yellow, matted, and glandlike, usually sloughing off by late in the year...... Qu. laevis 7 Petioles 2-5 cm long, not twisted so that the blade is oriented in a vertical plane; leaves shallowly to deeply lobed, some of the leaves on a tree generally shallowly lobed, none of the sinuses extending > 2/3 of the way to the midrib; pubescence of the lower leaf surface tawny or gray, stellate, not glandlike, persistent or sloughing off by late in the year. 8 Acorns 12-20 mm long, in a cup 15-25 mm across and 10-12 mm deep; mature leaves loosely and rather coarsely pubescent (the stellate hairs conspicuous and readily distinguishable at 10× magnification), often becoming nearly or entirely glabrous by late in the year (except for tufts of hairs in the vein axils); terminal bud 4-angled, 7-10 mm long, densely gray-tomentose...... Qu. velutina 8 Acorns 10-15 mm long, in a cup 12-14 mm across and 4-5 mm deep; mature leaves densely and finely pubescent (the stellate hairs minute and scarcely distinguishable at 10× magnification), the pubescence permanent; terminal bud only obscurely angled (if at all), 5-8 mm long, brown-puberulent. 9 Base of blades of sun-leaves typically rounded, thus forming a U-shape (some leaves cuneate, angled, or oblique); terminal lobe of leaves generally long-attenuated, narrow (its sides nearly parallel for much of its length), and curved to one side (falcate) (note that trees with the trilobed leaf form will key out above); leaves with 3-7 well-developed lobes, these often very irregular in size, shape, spacing, and orientation; pubescence of lower leaf surface normally tawny (when fresh) ...... Qu. falcata 9 Base of blades of sun-leaves typically cuneate or angled, thus forming a V-shape (some leaves somewhat U-shaped or oblique); terminal lobe of leaves generally short, broadly triangular (its sides normally tapering toward the tip for most of their length), not strongly curved to one side; leaves with 5-9 well-developed lobes, these generally rather uniform in size, shape, spacing, and orientation; pubescence of leaf surface gray...... Qu. pagoda 4 Mature leaves glabrous beneath on the surface, with tufts of hairs in the main vein axils beneath. 10 Petioles 0.5-1.0 (-1.8) cm long, generally twisted such that the blade is oriented in a vertical plane; inner cup- scales of the acorn cup inflexed, thus the cup appearing to have a broadly rounded rim ...... Qu. laevis 10 Petioles 2.5-7 cm long, not twisted so that the blade is oriented in a vertical plane; inner cup-scales of the acorn cup not inflexed, thus the cup appearing to have a sharp rim appressed against the acorn. 11 Terminal buds 4-angled, 7-10 mm long, the bud scales densely gray-tomentose...... Qu. velutina 11 Terminal buds not 4-angled, 3-5 (-7) mm long, the bud scales glabrous or with ciliate margins. 12 Leaves relatively shallowly lobed, the sinuses extending up to 2/3 of the way to the midrib; upper leaf surface dull, not lustrous. 13 Acorn cup covering about 1/4 of acorn; leaf sinuses extending about 1/4 of the way to the midrib; bark of mature trees dark gray to black; [widespread in our area, at low to medium elevations] ...... Qu. rubra var. rubra 13 Acorn cup covering about 1/3 of acorn; leaf sinuses extending about 1/3 of the way to the midrib; bark of mature trees medium gray; [of the Mountains, mostly at 1000 m and above]...... Qu. rubra var. ambigua 12 Leaves relatively deeply lobed, the sinuses extending 2/3 to 9/10 of the way to the midrib; upper leaf surface lustrous. 14 Larger lateral lobes of most leaves with 1 bristle per lobe (-2 on some lobes)...... Qu. georgiana 14 Larger lateral lobes of most leaves with 2 or more bristles. 15 Mature leaves mostly 7-12 cm long, 5-11 cm wide (averaging about 9 cm long and 8 cm wide), with 5-7 lobes; acorns (8-) 10-13 (-15) mm long; acorn cup nearly flat at base, covering about 1/4 of the acorn...... Qu. palustris 15 Mature leaves mostly 10-20 cm long, 8-15 cm wide (averaging about 12-15 cm long and 10- 12 cm wide), with 5-11 lobes; acorns (12-) 15-25 (-37) mm long; acorn cup nearly flat, turbinate, or rounded, covering 1/4 to 1/2 of the acorn. 16 Acorn cup turbinate, covering about 1/2 of the acorn; acorn (12-) 15-20 mm long, with 1- 3 concentric grooves near the tip; upper surface of leaves bright green ...... Qu. coccinea 16 Acorn cup nearly flat at base, covering about 1/4 to 1/3 of the acorn; acorn 15-37 mm long, lacking concentric grooves near the tip; upper surface of leaves dark green ...... Qu. shumardii var. shumardii FAGACEAE 405

* Quercus acutissima Carruthers, Sawtooth Oak. Pd (NC, VA): commonly cultivated as a suburban street tree and also widely planted in "wildlife food plots"; commonly cultivated, rarely (at this time) persistent and spreading, introduced from Japan. This species has been a popular recommendation for "wildlife plantings" in the recent past, and entire stands can be encountered in relatively remote areas, planted by federal and state land management agencies; why "wildlife" species in our area need more oak trees is somewhat mystifying! See Whittemore (2004) for additional information. [= K; ? Qu. acutissima ssp. acutissima] Quercus alba Linnaeus, White Oak. Pd, Mt, Cp (GA, NC, SC, VA): mesic to xeric forests; common. April; September- November (of the same year). Widespread in e. North America. Historically, one of the most valuable timber trees of eastern North America. Qu. alba is probably the most abundant native plant in our area, and in eastern North America, based on biomass, leaf area, and ubiquity. Hardin (1975) discusses introgression between Qu. alba and many other species of Quercus subgenus Quercus. [= RAB, C, F, FNA, G, K, W; < Qu. alba – S (also see Qu. austrina)] Quercus arkansana Sargent, Arkansas Oak. Cp (GA): dry bluffs; rare (GA Special Concern). Sw. and wc. GA west in a fragmented distribution to sw. AR and e. TX. [= FNA, K, S; > Q. caput-rivuli W.W. Ashe] Quercus austrina Small, Bluff Oak. Cp (GA, NC, SC), Pd (NC): river bluffs, natural levees of brownwater rivers, over mafic rocks in the Piedmont of NC, on shell or calcareous sediments on the Coastal Plain of SC (Charleston and Beuafort counties); rare (GA Special Concern, NC Rare). April; October (of the same year). Essentially a Southeastern Coastal Plain endemic, ranging from sc. NC south to n. FL and west to MS, nowhere common. [= RAB, FNA, K; < Qu. alba – S (apparently)] Quercus bicolor Willdenow, Swamp White Oak. Mt (VA), Pd (NC, SC, VA), Cp (NC, VA): upland depression swamp forests over mafic rocks such as gabbro or diabase, bottomland swamps with calcareous sediments; uncommon (rare south of VA) (NC Watch List, SC Rare). April; September (of the same year). Widespread in ne. North America, south to NC, SC (Nelson 1993), TN, n. AL, and MO. [= RAB, C, F, FNA, G, GW, K, S, W] Quercus chapmanii Sargent, Chapman Oak. Cp (GA, SC): dry pinelands; rare (GA Special Concern). February-March; September-November (of the same year). A Southeastern Coastal Plain endemic: se. SC south to FL, west to sw. AL. [= RAB, FNA, K, S] Quercus coccinea Muenchhausen, Scarlet Oak. Mt, Pd, Cp (GA, NC, SC, VA): xeric upland forests; common (uncommon in Coastal Plain south of VA). April; September-November (of the second year). Centered in the Appalachians, from s. ME south to c. AL, but ranging west to MS, ne. AR, s. IL, and s. MI. [= RAB, C, F, FNA, G, S, W; > Qu. coccinea var. coccinea – K; > Qu. coccinea var. tuberculata Sargent – K] Quercus elliottii Wilbur, Running Oak. Cp (GA, NC, SC): pine flatwoods, especially on loamy soils in the Middle Coastal Plain; uncommon. March-April; September (of the first year). A Southeastern Coastal Plain endemic: se. NC south to s. FL and west to s. MS. Wilbur (2002) discusses the reasons for rejecting the traditional use of Qu. pumila for this species; Walter's diagnosis states that Qu. pumila has leaves that are glabrous and glaucous below, ruling out application to this species. [= Q. pumila Walter – RAB, FNA, K, S, Z, apparently misapplied] Quercus falcata Michaux, Spanish Oak, Southern Red Oak. Cp, Pd, Mt (GA, NC, SC, VA): upland forests, usually xeric; common (uncommon in Mountains). April; September-November (of the second year). Widespread in se. North America, north to e. OK, s. MO, s. IL, s. IN, s. OH, WV, se. PA, NJ, and reported (apparently without specimen documentation) from Long Island, NY. "Qu. triloba Michaux", the form with the leaves only shallowly trilobed at the apex, causes much confusion. Though even medium-sized trees sometimes have leaves only of this form (rather than the typical form, deeply 5-7-lobed, the terminal lobe long-attenuate and falcate), it has no taxonomic merit. [= C, FNA, K, W; = Qu. falcata var. falcata – RAB, G, GW; > Qu. falcata var. falcata – F; > Qu. falcata var. triloba (Michaux) Nuttall – F (the juvenile form); = Qu. rubra – S, misapplied; ? Qu. digitata Sudworth; > Qu. triloba Michaux] Quercus geminata Small, Sand Live Oak. Cp (GA, NC, SC): xeric sandhills near the coast; uncommon (VA Watch List). April; September-November (of the same year). A Southeastern Coastal Plain endemic: se. NC south to FL (where common) and west to s. MS. The alleged occurrence of Qu. geminata as far north as se. VA is apparently based on ambiguous specimens that probably are only Qu. virginiana (the so-called var. maritima). The relative ranges, habitats, and abundance of this species and Qu. virginiana in NC are poorly understood. Apparently flowering about 2-3 weeks later than Qu. virginiana when growing in close proximity and in similar habitats. [= C, FNA, GW, K, S; < Qu. virginiana – RAB; ? Qu. virginiana var. maritima (Michaux) Sargent – F, misapplied] Quercus georgiana M.A. Curtis, Georgia Oak. Pd (GA, SC): dry slopes and bluffs over granite; rare (SC Rare). April; September-October (of the second year). W. SC south and west through GA to c. AL. [= RAB, FNA, K, S] Quercus hemisphaerica Bartram ex Willdenow, Sand Laurel Oak, Darlington Oak. Cp (GA, NC, SC, VA): sandhills and other dry, sandy soils, an abundant component of maritime forests with Qu. virginiana, and widely planted as a street tree in most parts of our region; common (VA Rare). March-April; September-November (of the second year). Essentially a Southeastern Coastal Plain endemic: se. VA south to s. FL and west to s. TX, north uncommonly in the interior to nc. AL, n. MS, and s. AR. Often confused with Qu. laurifolia (see the key for distinctions). Qu. hemisphaerica is the semi-evergreen laurel oak planted widely as a street tree in southern cities, often intermixed with the strictly deciduous Qu. phellos. [= C, F, FNA, Z; < Qu. laurifolia – RAB; = Qu. laurifolia – S, misapplied; > Qu. hemisphaerica var. hemisphaerica – K; > Qu. hemisphaerica var. maritima (Michaux) Muller – K] Quercus ilicifolia Wangenheim, Bear Oak, Scrub Oak. Mt (VA), Pd (NC, VA): xeric soils in ridges in the Mountains and monadnocks in the upper Piedmont; common (rare in Piedmont and south of VA) (NC Rare). Late April-June; August (of the second year). Primarily Appalachian: s. ME south to w. VA, w. NC, and e. KY. This scrubby oak is limited in NC to dry summits and upper slopes of Piedmont monadnocks; it is rare and probably declining because of fire suppression (Barden 1985), FAGACEAE 406 though recent ice storms have opened the tree canopy at several of its NC sites. The occurrence of Q. ilicifolia in KY was confirmed at the Devil's Teatable, Floyd County (Clark et al. 1997). [= RAB, C, F, FNA, G, K, S, W] Quercus imbricaria Michaux, Shingle Oak. Mt, Pd (NC, VA), Cp (VA): rich soils of upper floodplains of rivers and creeks, often at the base of the slope into the upland, also on lower slopes, and in forests over diabase in the Piedmont of VA and n. NC; uncommon (GA Special Concern). May; October (of the second year). Primarily midwestern, ranging from NJ, PA, n. OH, s. MI, n. IL, and c. IA, south to e. VA, nc. and w. NC, sc. TN, n. AL, and n. AR. [= RAB, C, F, FNA, G, K, S, W, Z] Quercus incana Bartram, Bluejack Oak. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): sandhills, primarily in somewhat loamier textured, submesic soils; common (rare in Piedmont) (VA Rare). April; September-November (of the second year). Primarily a species of the Southeastern Coastal Plain, but rarely extending inland into the Piedmont (especially on coarse sandy alluvium): se. VA south to c. peninsular FL and west to e. TX, sw. AR, and se. OK. This oak is recognizable even at a distance by its bluish color. [= RAB, F, FNA, K, Z; = Qu. cinerea Michaux – C, G, S; ? Qu. humilis Walter] Quercus laevis Walter, Turkey Oak. Cp (GA, NC, SC, VA): sandhills, primarily in very xeric soils of deep sandy deposits (Carolina bay rims, old beach dunes, early Cenozoic deposits of the Sandhills Province); common (VA Rare). April; September- October (of the second year). Essentially a Southeastern Coastal Plain endemic: se. VA south to s. FL and west to e. LA. The leaves turn an intense orange-red in the autumn (November). [= RAB, C, F, FNA, G, K, S, Z; = Qu. catesbaei Michaux] Quercus laurifolia Michaux, Laurel Oak. Cp (GA, NC, SC, VA): mesic to seasonally flooded soils of floodplains, also (rarely) mesic slopes and swamps in maritime forests; common. March-April; September-November (of the second year). A Southeastern Coastal Plain endemic: se. VA south to s. FL and west to e. TX and s. AR. Sometimes confused with Qu. hemispherica, but (in addition to the key characters above) Qu. laurifolia has blunter leaf tips, flowers about 2 weeks earlier, and generally occupies much moister habitats. [= C, F, FNA, G, GW, K, Z; < Qu. laurifolia – RAB (also see Qu. hemisphaerica); = Qu. obtusa (Willdenow) Ashe – S] Quercus lyrata Walter, Overcup Oak. Cp, Pd (GA, NC, SC, VA), Mt (GA): common in seasonally rather deeply and frequently flooded soils of floodplains of the Coastal Plain, less commonly in seasonally flooded swamps in Triassic basins in the lower Piedmont, and rarely in upland depression swamps of the Piedmont (developed over clays weathered from mafic rocks) and Coastal Plain; common (rare in Piedmont and Mountains). March-April; September-October (of the same year). Primarily a species of the Southeastern Coastal Plain: DE south to nw. FL, west to e. TX and se. OK, north in the inland to w. TN, s. IN, s. IL, and se. MO. Of our oaks, Qu. lyrata tolerates the wettest habitats, both in terms of depth and duration of flooding. [= RAB, C, F, FNA, G, GW, K, S] Quercus macrocarpa Michaux var. macrocarpa, Bur Oak, Mossycup Oak. Mt (VA): bottomland forests; rare (VA Rare). New Brunswick and Québec west to s. Manitoba, south to nw. VA, KY, TN, LA, and TX. Variation in this species needs additional study; Qu. macrocarpa in our area is the typic variety or subspecies if other taxa are recognized. [= K; < Qu. macrocarpa – C, F, FNA, G, GW, S, W] Quercus margaretta Ashe ex Small, Sand Post Oak. Cp (GA, NC, SC, VA), Pd (GA): sandhills, typically in slightly loamy or clayey soils, not usual in the deepest and most xeric sands; common (VA Watch List). April; September-November (of the same year). Primarily a species of the Southeastern Coastal Plain: se. VA south to FL and west to TX and se. OK. As stated by F, this oak was "chivalrously named [by W.W. Ashe] in 1903 for Margaret Henry Wilcox, who two years later became Mrs. Ashe." There has been controversy, however, over the spelling of the specific epithet. Some nomenclaturists believe that the original spelling, "margaretta" should be treated as a correctable error, according to Articles 73.10 and Recommendation 73C.1 of the International Code, and replaced with "margarettiae." [= RAB, C, FNA, G, S; = Qu. margarettiae Ashe ex Small – K, orthographic variant; = Qu. stellata var. margaretta (Ashe ex Small) Sargent – F] Quercus marilandica Muenchhausen var. marilandica, Blackjack Oak. Pd, Cp, Mt (GA, NC, SC, VA): uplands forests and woodlands, usually on periodically droughty soils, as over shrink-swell clays, sandstones, deep sands, and sands with clay lenses; common (uncommon in Mountains). April; September-November (of the second year). NY (Long Island), NJ, se. PA, w. VA, s. OH, s. IN, c. IL, s. IA, and se. NE south to s. GA, Panhandle FL, and sc. TX (west to the Prairie border). There are historical accounts of the existence of prairies or barrens in the vicinity of Charlotte in the late eighteenth century, known as the "the blackjack lands." These areas were described as open and prairie-like, until the early nineteenth century, when they became dominated by dense forests of blackjack oak. The previously open condition was almost certainly maintained by fire, perhaps set by the Waxhaw Indians. Blackjack oak has long been considered an indicator of poor soil, as in Guthrie (1820), who states in his discussion of NC, "the Black Jack land is generally poor, though it has sometimes a black appearance, it is wet and loose, and is avoided by farmers, as unproductive." Var. marilandica is the widespread taxon; var. ashei Sudworth [= Qu. neoashei Bush] is worthy of recognition at the varietal level at least, and occurs from s. MO and s. KS south to c. AR, e. TX, and sc. TX, especially on the Edwards Plateau (Hunt 1990). [= FNA, K, Z; < Qu. marilandica – RAB, C, F, G, S, W] Quercus michauxii Nuttall, Basket Oak, Swamp Chestnut Oak. Cp, Pd (GA, NC, SC, VA), Mt (GA): bottomland forests, especially in fertile soils of upper terraces where flooded only infrequently and for short periods; common (uncommon in Piedmont). April; September-October (of the same year). NJ south to n. FL and west to e. TX and se. OK, north in the interior to s. IL and s. IN. See discussion under Qu. montana of the application of the name Qu. prinus Linnaeus. [= RAB, C, F, FNA, G, GW, K, W; = Qu. prinus Linnaeus – S, possibly misapplied] Quercus minima (Sargent) Small, Dwarf Live Oak. Cp (GA, NC, SC): pine flatwoods, coastal fringe sandhills; uncommon (NC Watch List). April; September-November (of the same year). A Southeastern Coastal Plain endemic: se. NC (New Hanover County) south to FL, west to s. MS. [= FNA, K, S] Quercus montana Willdenow, Rock Chestnut Oak. Mt, Pd (GA, NC, SC, VA), Cp (GA, NC, VA): xeric forests of ridges, slopes; common. April; September-November (of the same year). Primarily Appalachian but broadly distributed in e. North America: s. ME, NY, MI, s. UN, s. IL, and se. MO (Smith & Parker 2005) south to c. GA, c. AL, ne. MS (and LA?). The proper application of the Linnaean "Qu. prinus" is controversial and unclear, having been debated and variously applied for well over a FAGACEAE 407 century. I have here decided to err on the side of clarity. The name "Qu. prinus" has nomenclatural priority over either "Qu. montana" or "Qu. michauxii", but it is not clear which species was intended; Whittemore & Nixon (2005) have proposed its formal rejection. [= FNA, S, W; = Qu. prinus Linnaeus – RAB, C, F, G, K, probably misapplied] Quercus muehlenbergii Engelmann, Yellow Oak, Chinquapin Oak. Mt, Pd (GA, NC, SC, VA), Cp (GA, SC, VA): slopes and bluffs, on soils derived from calcareous or mafic rocks; common in VA Mountains (rare elsewhere) (NC Watch List). April; October-November (of the same year). S. New England and Ontario west to WI, se. MN, and IA, south to nw. FL, TX, and n. Mexico. The similar Qu. montana sometimes has a few leaves with somewhat sharply lobed leaves, but these are minutely mucronate and lack the well-developed callus of Qu. muehlenbergii. Additionally, Qu. muehlenbergii has a flaky, light gray bark, very different from the dark gray, deeply furrowed bark of Qu. montana. [= RAB, C, F, K; = Qu. muhlenbergii – FNA, S, W, orthographic variant; = Qu. prinoides Willdenow var. acuminata (Michaux) Gleason – G] Quercus myrtifolia Willdenow, Myrtle Oak. Cp (GA, SC): dry pinelands; rare (SC Rare). February-March; September (of the second year). A Southeastern Coastal Plain endemic: se. SC south to s. peninsular FL, west to se. MS. [= RAB, FNA, K, S, Z] Quercus nigra Linnaeus, Water Oak, Paddle Oak. Cp, Pd (GA, NC, SC, VA), Mt (GA): bottomland forests, especially on levees or second terraces where flooded infrequently and for short periods, less commonly on mesic slopes; common (uncommon in Piedmont). April; September-November (of the second year). Primarily a species of the Southeastern Coastal Plain: s. NJ south to s. FL and west to e. TX and se. OK, north in the interior to se. TN, c. TN, w. and sc. KY (Clark et al. 2005), se. MO, and e. OK. Seedlings and fire sprouts of this species are highly variable; see discussion at end of generic treatment. [= RAB, C, FNA, G, GW, K, S, W, Z; > Qu. nigra var. nigra – F; > Qu. nigra var. heterophylla (Aiton) Ashe – F: = Qu. aquatica Walter] Quercus oglethorpensis Duncan, Oglethorpe Oak. Pd (GA, SC): bottomland forests, upland oak flats over clays (Iredell and Enon soils); rare (GA Threatened, SC Rare). April; September-October (of the same year). Ranging (in a widely scattered pattern) from w. SC, to adjacent e. GA, nw. AL (Sorrie pers. comm. 2002), MS, and LA. [= RAB, FNA, GW, K] Quercus pagoda Rafinesque, Cherrybark Oak, Swamp Spanish Oak. Cp, Pd (GA, NC, SC, VA): bottomland forests, especially on second terraces; common (rare in Piedmont). April; September-November (of the second year). A Southeastern Coastal Plain endemic: e. and c. VA south to nw. FL and west to se. TX and north in the interior to e. TN, s. IL, and s. IN. [= C, FNA, K, S; = Qu. falcata var. pagodifolia Elliott – RAB, F, G, GW] Quercus palustris Muenchhausen, Pin Oak. Pd (NC, VA), Cp (VA), Mt (GA, VA): swamps and bottomlands, especially the broader swamps developed in the sedimentary rocks of Triassic basins of the lower Piedmont, isolated upland sag ponds, also widely planted as a street tree in towns and cities; uncommon (rare south of VA) (GA Special Concern, NC Watch List). March- April; October-November (of the second year). MA and NY west to se. IA and e. KS, south to c. NC, nw. GA, sc. TN, n. AR, and e. OK. [= RAB, C, F, FNA, G, GW, K, S, W] Quercus phellos Linnaeus, Willow Oak. Cp, Pd (GA, NC, SC, VA), Mt (GA): bottomland forests, especially on natural levees and second terraces, also in upland depression swamps developed on clay soils, weedy and successional on slopes and upland sites following disturbance, and widely planted as a street tree in towns and cities; common (uncommon in Piedmont). March-April; September-November (of the second year). Primarily a species of the Southeastern Coastal plain: NY (Long Island), s. NJ, and se. PA south to s. GA and panhandle FL, west to e. TX and se. OK, north in the interior to e. TN, s. KY, w. KY, s. IL, and se. MO, and e. OK. [= RAB, C, F, FNA, G, GW, K, S, W, Z] Quercus prinoides Willdenow, Dwarf Chinquapin Oak. Pd, Mt (GA, NC, VA), Cp (VA): xeric uplands, especially on clay soils derived from mafic rocks, and probably in sites which naturally burned rather frequently; rare (GA Special Concern, NC Rare, VA Rare). April; August-September (of the same year). MA and s. MI south to NC, OK, and TX. Fire suppression in the Piedmont sites where this rare oak occurs has nearly or entirely extirpated it from much of our area. [= RAB, C, FNA, K, S, W; > Qu. prinoides var. prinoides – F; > Qu. prinoides var. rufescens Rehder – F; = Qu. prinoides var. prinoides – G] Quercus rubra Linnaeus var. ambigua (A. Gray) Fernald, Gray Oak. Mt (NC, SC, VA), Pd (VA): forests on ridges, slopes, and coves, mostly at over 1000 meters elevation; common. May; September-October (of the second year). Fairly widespread in ne. North America south to PA, and in the Appalachians to w. NC, nw. SC, and n. GA. This and var. rubra tend to intergrade and their distinction as even varieties may not be warranted. For discussion of the two varieties, see McDougal & Parks (1984) and Jensen (1977). [= K; = Qu. rubra var. borealis (Michaux f.) Farwell – RAB, F, FNA; < Qu. rubra – C, W; = Qu. borealis Michaux f. var. borealis – G; = Qu. borealis – S] Quercus rubra Linnaeus var. rubra, Red Oak. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): moist to fairly dry forests of slopes, coves, and ravines, below 1000 meters elevation; common (rare in Coastal Plain). April; August-September (of the second year). Widespread in e. North America, south to e. VA, GA, AL, MS, AR, and OK. [= RAB, F, FNA, K; < Qu. rubra – C, W; = Qu. borealis Michaux f. var. maxima (Marshall) Ashe – G; = Qu. maxima (Marshall) Ashe – S] Quercus shumardii Buckley var. shumardii, Shumard Oak. Pd, Cp, Mt (GA, NC, SC, VA): moist and fertile soils of bottomlands and lower slopes, also in xeric sites over calcareous rocks (such as limestone); uncommon (rare in Coastal Plain and Mountains). April; September-October (of the second year). Sc. PA, OH, s. MI, IN, s. IL, MO, and e. KS south to FL and TX. A number of varieties have been recognized in Qu. shumardii, and the morphological and habitat variation needs additional study. Var. schneckii (Britton) Sargent is apparently more midwestern, but should be looked for in our area, especially on dry limestone slopes, its preferred habitat. It is allegedly distinguished by the acorn cups rounded to turbinate below (vs. flattened and saucer-shaped in var. shumardii). Hess & Stoynoff (1998) tentatively concluded that no varieties should be recognized within Qu. shumardii, but they plan additional studies. Qu. acerifolia (E.J. Palmer) Stoynoff & W.J. Hess (Qu. shumardii var. acerifolia Palmer] is an endemic of Magazine Mountain in n. AR and scattered sites in OK; it is best treated as a distinct species, though there has been much debate about its taxonomic status, with opinions ranging from full species to mere form (Smith 1988, Stoynoff & Hess 1990, Johnson 1992, Johnson 1994, Hess & Stoynoff 1998). [= F, FNA, G, K; < Qu. shumardii – RAB, C, S, W] FAGACEAE 408 Quercus similis Ashe, Swamp Post Oak, Delta Oak. Cp (GA?, NC?, SC): calcareous stream flats; rare. Qu. similis resembles Qu. stellata, differing in its less definitely cross-shaped leaves and its distinctly wetland habitat. [= FNA, K; = Qu. stellata Wangenheim var. paludosa Sargent; = Qu. ashei Sterret] Quercus sinuata Walter var. sinuata, Bastard Oak. Cp (GA, SC): alluvial and slope forests; rare (GA Special Concern, SC Rare). April-May; September-November (of the same year). Se. SC west to TX. [= FNA, K; > Qu. durandii Buckley – RAB, S] Quercus stellata Wangenheim, Post Oak. Pd, Cp, Mt (GA, NC, SC, VA): upland forests and woodlands, especially in clay or rocky soils and in fire communities; common (uncommon in Mountains). April; September-November (of the same year). Se. MA, s. NY, s. PA, s. OH, s. IN, s. IA, and e. KS south to FL and TX. In KS, OK, and TX, post oak is one of the trees that forms the Prairie boundary. There is no question of the distinctness of Qu. margaretta from Qu. stellata. See Qu. similis. [= RAB, C, FNA, G, K, S, W; = Qu. stellata var. stellata – F; = Qu. villosa Walter] Quercus velutina Lamarck, Black Oak. Mt, Pd, Cp (GA, NC, SC, VA): upland forests and woodlands, especially in fairly xeric and sandy soils; common. April; September-October (of the second year). Widespread in e. North America. [= RAB, C, F, FNA, G, K, S, W] Quercus virginiana P. Miller, Live Oak. Cp (GA, NC, SC, VA): locally common to abundant in maritime forests and maritime scrub on barrier islands, more rarely inland (though regularly on the mainland from se. NC south, and extending substantially inland from s. SC south), sometimes in dry, fire-maintained habitats more usually occupied by Qu. geminata, also planted (especially in the outer Coastal Plain); uncommon (VA Watch List). April; September-November (of the same year). A Southeastern Coastal Plain endemic: se. VA south to FL and west to TX. Qu. fusiformis Small of TX has sometimes been treated as a variety of Qu. virginiana, but is best separated as a species. Flowering before Qu. geminata when growing together. [= C, FNA, GW, K, S; < Qu. virginiana – RAB, G (also see Qu. geminata); < Qu. virginiana var. virginiana – F; Qu. sempervirens Walter]

Quercus boyntonii Beadle, Boynton Oak. Dry forests. Ne. AL and (possibly) TX. [= FNA, K, S; Q. stellata Wangenheim var. boyntonii (Beadle) Sargent] * Quercus robur Linnaeus, English Oak, is cultivated in our area and sometimes persists or escapes in ne. United States, south at least to s. PA (Rhoads & Klein 1993). [= FNA, K] Quercus texana Buckley, Nuttall Oak, Texas Red Oak. Floodplain swamps and bottomlands. AL, TN, w. KY (Clark et al. 2005), west to e. TX. [= FNA, K; = Qu. nuttallii E.J. Palmer – F, GW; = Qu. shumardii Buckley var. texana (Buckley) W.W. Ashe] {not keyed at this time}

FUMARIACEAE Augustin de Candolle 1821 (Fumitory Family)

This family includes 15-20 genera and 500-600 species, herbs, mostly north temperate. The Fumariaceae should likely be merged into the Papaveraceae (Lidén 1981, 1986; Lidén et al. 1997; Judd, Sanders, & Donoghue 1994). References: Stern in FNA (1997); Hill (1992); Lidén (1986, 1981); Lidén et al. (1997); Lidén in Kubitzki, Rohwer, & Bittrich (1993).

1 Corolla with the 2 outer petals spurred or saccate at their bases; [tribe Corydaleae]. 2 Caulescent herbaceous vine (acaulescent in its first year, and appearing to be an herb); ultimate leaf segments 5-10 mm wide ...... Adlumia 2 Acaulescent herb with basal leaves; ultimate leaf segments 1-4 mm wide. 3 Leaves basal only...... Dicentra 3 Leaves cauline and basal...... [Lamprocapnos] 1 Corolla with only 1 outer petal spurred or saccate at its base. 4 Ovary and fruit subglobose, with 1 seed; [tribe Fumarieae]...... Fumaria 4 Ovary and fruit elongate, with several to many seeds; [tribe Corydaleae]. 5 Flowers pink, the petals tipped with yellow; biennial; stem erect, 3-8 (-10) dm tall; capsules erect, 25-35 mm long ...... Capnoides 5 Flowers yellow; annual; stem erect, decumbent, or prostrate, 1-3 (-4) dm tall; capsules erect, ascending, divergent, or pendent, 10-20 (-25) mm long ...... Corydalis

Adlumia Rafinesque ex Augustin de Candolle 1821 (Climbing Fumitory)

A genus of 2 species, herbs, of e. North America, Korea, and Manchuria. References: Boufford in FNA (1997); Lidén in Kubitzki, Rohwer, & Bittrich (1993).

Adlumia fungosa (Aiton) Greene ex Britton, Sterns, & Poggenburg, Alleghany-vine, Cliff-Harlequin, Climbing Fumitory. Mt (NC, VA): cliffs, talus, rocky slopes, rich stream-bottom forests, cool rocky forests; rare (NC Rare, VA Watch List). June- September. Québec west to WI and MN, south to DE, NC, TN, and IN. [= RAB, C, F, FNA, G, K, S, W]

Capnoides P. Miller 1754 (Rock Harlequin) FUMARIACEAE 409

The genus is monotypic, an herb, of n. North America. Recent studies have emphasized its distinction from Corydalis, and its closer relationship to Adlumia and Dicentra than to Corydalis (Lidén 1981, 1986; Lidén et al. 1997). References: Stern in FNA (1997); Ownbey (1947)=Z; Lidén (1981, 1986); Lidén et al. (1997); Lidén in Kubitzki, Rohwer, & Bittrich (1993).

Capnoides sempervirens (Linnaeus) Borkhausen, Rock Harlequin, Tall Corydalis, Pink Corydalis, Pale Corydalis. Mt (GA, NC, SC, VA), Pd (NC, VA): rock outcrops, especially granitic exfoliation domes, but also quartzite, greenstone, and sandstone; uncommon (rare in NC and VA Piedmont) (GA Special Concern). April-June; May-July. Newfoundland west to AK, south to NJ, PA, in and near the mountains to ne. GA, n. OH, n. IN, MN, MT, and British Columbia. [= S; = Corydalis sempervirens (Linnaeus) Persoon – RAB, C, F, FNA, G, K, W, Z]

Corydalis Augustin de Candolle 1805 (Corydalis) (also see Capnoides)

A genus of about 400 species, herbs, of temperate regions of the Northern Hemisphere (especially China and the Himalayas). References: Stern in FNA (1997); Ownbey (1947)=Z; Lidén in Kubitzki, Rohwer, & Bittrich (1993).

1 Flowers pink, the petals tipped with yellow; biennial; stem erect, 3-8 (-10) dm tall; capsules erect, 25-35 mm long ...... [Capnoides] 1 Flowers yellow; annual; stem erect, decumbent, or prostrate, 1-3 (-4) dm tall; capsules erect, ascending, divergent, or pendent, 10-20 (-25) mm long. 2 Fruits pendent or divergent; spurred petal 7-9 mm long; pedicels 6-15 mm long; seeds 2-2.5 mm wide, with a narrow, acute ring-margin...... C. flavula 2 Fruits erect or ascending; spurred petal 10-15 mm long; pedicels 1-6 mm long (5-10 mm long in C. aurea); seeds 1.0- 2.0 mm wide, without a narrow, acute ring-margin. 3 Capsules mostly 15-20 mm long, ca. 1.0 mm in diameter, strongly constricted between the seeds at maturity; inflorescence long, usually far exceeding the poorly-developed upper leaves; ultimate leaf segments 0.5-1.5 (-3.0) mm wide; seeds < 1.5 mm wide; plant slightly to strongly glaucous; [of sandy soils of the outer Coastal Plain]...... C. halei 3 Capsules mostly 10-15 mm long, 1.5-2.0 mm in diameter, slightly or not at all constricted between the seeds at maturity; inflorescence relatively short, barely (if at all) overtopping the upper leaves; ultimate leaf segments 1.0- 2.0 (-4.0) mm wide; seeds > 1.5 mm wide; plant green to slightly glaucous; [of circumneutral rock outcrops of the upper Piedmont and Mountains] ...... C. micrantha

Corydalis flavula (Rafinesque) Augustin de Candolle, Short-spurred Corydalis. Pd, Mt, Cp (GA, NC, SC, VA): rich moist forests, especially alluvial forests, glades and outcrops over mafic rocks (such as greenstone); common (rare in NC and SC Coastal Plain) (GA Special Concern). March-April; May-June. S. CT, NY, and s. Ontario west to SD, south to NC, AL, LA, and OK. [= RAB, C, F, FNA, G, K, W, Z; = Capnoides flavulum (Rafinesque) Kuntze – S] Corydalis halei (Small) Fernald & Schubert, Southern Corydalis. Cp (GA, NC, SC): sandy roadsides and disturbed areas; uncommon. March-April; May-June. E. NC south to FL, west to TX, and inland north to MO and OK. F and S recognized it as a species distinct from C. micrantha; Ownbey reduced it to a subspecies, citing inadequate morphological differences and some alleged intermediates in OK and MO. The two taxa appear readily separable on morphological, ecological, and geographical grounds; species status seems warranted. [= F; = Corydalis micrantha (Engelmann ex A. Gray) A. Gray ssp. australis (Chapman) G.B. Ownbey – RAB, FNA, K, Z; = Corydalis micrantha (Engelmann ex A. Gray) A. Gray var. australis (Chapman) Shinners – C; < Corydalis micrantha – G; = Capnoides halei Small – S] Corydalis micrantha (Engelmann ex A. Gray) A. Gray, Slender Corydalis. Mt, Pd (NC): circumneutral rock outcrops and adjacent glades and woodlands; rare (NC Rare). April; June. Ssp. micrantha is primarily midwestern, ranging from IL, WI, MN, and SD south to AR, TX, and OK, with disjunct outliers in e. TN and w. NC. Ownbey (1947) had no records of Southern Appalachian populations of C. micrantha, and considered ssp. micrantha to range no further east than IL and MO; RAB included montane populations in ssp. australis, stating "this is the only [subspecies] in our range." Morphologically, however, these populations closely resemble ssp. micrantha; their association in the Brushy Mountains with other species disjunct from western or prairie ranges (Anemone berlandieri, Arabis hirsuta, Pellaea wrightiana) provides phytogeographic corroboration. [= F; = Corydalis micrantha (Engelmann ex A. Gray) A. Gray ssp. micrantha – FNA, K, Z; = Corydalis micrantha (Engelmann ex A. Gray) A. Gray var. micrantha – C; < Corydalis micrantha – G; = Capnoides micranthum (Engelmann ex A. Gray) Britton – S]

Corydalis aurea Willdenow, south and east to MD, WV (?), and PA (Kartesz 1999). [= G, K; = Corydalis aurea var. aurea – C, F; = Corydalis aurea ssp. aurea – FNA; = Capnoides aureum (Willdenow) Kuntze – S] {not keyed at this time} Corydalis crystallina Engelmann, a species of the sc. United States, was collected in 1930 from an oat field at the Georgia Experiment Station in Laurens County. Presumably it was a one-time contaminant in seed and is a waif. Not considered a component of the flora of our area. [= FNA, F, G, K]

Dicentra Bernhardi 1833 FUMARIACEAE 410

A genus of about 12 species, perennial herbs, with a relictual north temperate distribution: e. North America, w. North America, and e. Asia. References: Stern in FNA (1997); Stern (1961)=Z; Lidén in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves cauline and basal...... [see Lamprocapnos spectabilis] 1 Leaves basal only. 2 Flowers pink, in panicles; rootstock lacking bulblets; ultimate leaf segments generally 3-parted, each part 2-5 mm wide at base, gradually tapering to the tip. 3 Reflexed portions of the outer sepals 4-8 mm long; [native and cultivated] ...... D. eximia 3 Reflexed portions of the outer sepals 2-5 mm long; [cultivated]...... [D. formosa ssp. formosa] 2 Flowers white or yellowish (very rarely pinkish), in racemes; rootstock with bulblets; ultimate leaf segments not generally 3-parted, about 1 (-3) mm wide, with parallel sides for most of their length, then tapering suddenly to the tip. 4 Spurs of the corolla rounded, incurved, 2-5 mm long; bulblets yellow, spherical...... D. canadensis 4 Spurs of the corolla elongate, divergent, 7-9 mm longl bulblets white to pink, tear-shaped (narrowed upward)...... D. cucullaria

Dicentra canadensis (Goldie) Walpers, Squirrel Corn. Mt (GA, NC, VA), Pd, Cp (NC, VA): rich, moist forests, especially rich cove forests in the mountains; common (uncommon in VA Piedmont, rare in Coastal Plain, rare or extirpated in NC Piedmont) (GA Special Concern). April-May; June. S. ME west to s. MN, south to w. NC, n. GA, TN, and MO. [= RAB, C, F, FNA, G, K, W; = Bicuculla canadensis (Goldie) Millspaugh – S] Dicentra cucullaria (Linnaeus) Bernhardi, Dutchman's Britches. Mt (GA, NC, SC, VA), Pd (NC, SC, VA), Cp (NC): rich, moist forests, especially rich cove forests in the mountains; common (rare in NC and SC Piedmont and NC Coastal Plain) (SC Rare). March-April; May-June. Nova Scotia west to n. MN, south to GA, AR, and KS; disjunct in WA, OR, and ID. [= RAB, C, F, FNA, G, K, W; = Bicuculla cucullaria (Linnaeus) Millspaugh – S] Dicentra eximia (Ker-Gawler) Torrey, Wild Bleeding Heart. Mt (GA, NC, SC, VA), Pd (NC, VA): cliffs, talus slopes, rocky slopes, rock outcrops, shale slopes; common in VA mountains (rare elsewhere) (GA Special Concern, NC Rare, SC Rare). April-June; July-August. An Appalachian endemic: NY and NJ south to NC and TN. [= RAB, C, F, FNA, G, K, W; = Bicuculla eximia (Ker-Gawler) Millspaugh – S]

* Dicentra formosa (Haworth) Walpers ssp. formosa, native from s. British Columbia south to c. CA, is frequently cultivated and resembles our native D. eximia. It has the reflexed portion of the outer petals 2-4 mm long. A variety of cultivars, some apparently derived from hybrids between the 2 species, make identification uncertain in some cases. [= FNA]

Fumaria Linnae 1753 (Fumitory)

A genus of about 50 species, annual herbs, primarily Eurasian. References: Boufford in FNA (1997); Lidén in Kubitzki, Rohwer, & Bittrich (1993).

* Fumaria officinalis Linnaeus, Fumitory, Earthsmoke. Cp (GA, NC, SC, VA), Pd (NC, SC, VA), Mt (VA): sandy fields, disturbed places, escaped from gardens; rare, introduced from Europe. March-May. [= RAB, C, F, FNA, G, K, S; > F. officinalis ssp. officinalis – K; > F. officinalis ssp. wirtgenii (W.D.J. Koch) Arcangeli – K]

Lamprocapnos Endlicher (Asian Bleeding Heart)

A monotypic genus, a perennial herb of e. Asia. References: Lidén et al. (1997).

Identification notes: Lamprocapnos differs from other "bleeding hearts" (the native Dicentra eximia and the western American Dicentra formosa ssp. formosa) in its leafy stem, the inflorescence borne terminally or opposite a leaf, the leaves much less finely divided, and the flowers about as broad as long (vs. much longer than broad in Dicentra eximia and Dicentra formosa).

* Lamprocapnos spectabilis (Linnaeus) Fukuhara, Bleeding Heart, native to e. Siberia, Korea, and n. China, is frequently cultivated and may persist or weakly naturalize. [= K; Dicentra spectabilis (Linnaeus) Lem.]

GARRYACEAE Lindley 1834 (Garrya Family)

Garryaceae is here circumscribed to include Aucuba (Bremer et al. 2002). References: Bremer et al. (2002)

Aucuba Thunberg (Aucuba, Japanese-laurel)

FUMARIACEAE 411 * Aucuba japonica Thunberg, Aucuba, Japanese-laurel, Spotted-laurel. Pd (NC): commonly planted throughout our area, rarely escaping and naturalizing in suburban woodlands; rare, introduced from Japan and se. Asia. The most frequently planted cultivars have the dark green leaves prominently speckled with yellow. [= K]

GELSEMIACEAE (G. Don) Struwe & V. Albert 1995 (Jessamine Family)

A family of 2 genera and about 10 species, shrubs and vines, of tropical and warm temperate America, Africa, and Asia. There is persuasive evidence that Gelsemium and Mostuea Didr., traditionally treated as part of a heterogeneous Loganiaceae, should be accorded family status as Gelsemiaceae (Backlund, Oxelman, & Bremer 2000; Struwe, Albert, & Bremer 1994; Sennblad & Bremer 1996). The Gelsemiaceae form a clade most closely related to the Apocynaceae (Backlund, Oxelman, & Bremer 2000). References: Backlund, Oxelman, & Bremer (2000); Struwe, Albert, & Bremer (1994); Sennblad & Bremer (1996); Rogers (1986).

Gelsemium Antoine Laurent de Jussieu (Yellow Jessamine)

A genus of 3 species, vines, our 2 species in se. North America (and also Central America) and 1 species in e. Asia. References: Wyatt et al. (1993); Duncan & Dejong (1964); Godfrey (1988); Rogers (1986)=Z; GW.

1 Sepals acuminate apically, persistent on the fruit; capsule elliptical, 1.0-1.6 cm long, 6-8 mm broad, the tapering tip bearing a definite beak about 3 mm long; seeds wingless; flowers odorless (rarely fragrant), usually golden-yellow...... G. rankinii 1 Sepals obtuse to broadly acute, not persistent on the fruit; capsule oblong, 1.5-2.5 cm long, 8-12 mm broad, very abruptly narrowed to a beak 1.5-2 mm long; seeds with a prominent membranous wing sharply differentiated from the body of the seed; flowers fragrant, usually lemon-yellow ...... G. sempervirens

Gelsemium rankinii Small, Swamp Jessamine. Cp (GA, NC, SC): swamps of blackwater rivers, restricted in NC to the se. corner of the state, most notably the swamps of the Waccamaw and Black rivers; rare north of GA (NC Rare). March-April; September-October. Se. NC south through SC and GA to the FL panhandle, and west to e. LA. See Wyatt et al. (1993) and Duncan & Dejong (1964) for extensive discussions of morphology, habitat, pollination, genetics, distribution, and evolutionary relationships of our 2 species of Gelsemium. [= RAB, GW, K, S, Z] Gelsemium sempervirens St. Hilaire, Carolina Jessamine. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (GA): in a wide range of habitats, from swamp forests to dry uplands and thickets, also commonly planted as an ornamental; common. March- early May; September-November. VA, se. TN, and AR south to c. peninsular FL and e. TX; disjunct in Guatemala and Mexico (Chiapas, Oaxaca, Puebla, and Veracruz). Jessamine climbs to the tops of trees. [= RAB, F, G, GW, K, S, W, Z]

GENTIANACEAE A.L. de Jussieu 1789 (Gentian Family) (also see MENYANTHACEAE)

A family of about 87 genera and over 1600 species, herbs, shrubs, and trees, cosmopolitan (Struwe & Albert 2002). References: Wood & Weaver (1982); Struwe & Albert (2002).

1 Leaves all scale-like, 1-3 (-5) mm long, appressed to the stem; [tribe Gentianeae, subtribe Swertiinae] ...... Bartonia 1 Leaves larger, spreading or ascending. 2 Stem leaves whorled; plants robust, 1-3 m tall; [tribe Gentianeae, subtribe Swertiinae]...... Frasera 2 Stem leaves opposite; plants generally < 1 m tall. 3 Calyx lobes 2; stem leaves obovate, widest near the rounded tip), 0.5-1.5 cm long, crowded anear the tip of the stem;, basal rosette never present; [of nutrient-rich, mesic forests]; [tribe Gentianeae, subtribe Swertiinae]...... Obolaria 3 Calyx lobes 4-5; stem leaves lanceolate, ovate, elliptic or narrowly elliptic (widest near the middle or toward the base, the tip acute or acuminate), mostly > 1.5 cm long, distributed fairly evenly along the stem, basal rosettes sometimes present; [of various more-or-less open habitats (except some species of Gentiana, which can occur in nutrient-rich, mesic forests)]. 4 Corolla lobes 5-14, much longer than the corolla tube, pink or white; [tribe Chironieae, subtribe Chironiinae] ...... Sabatia 4 Corolla lobes 4-5, shorter than the corolla tube, blue, lavender, pink or white. 5 Corolla tube < 2 mm wide; [tribe Chironieae, subtribe Chironiinae] ...... Centaurium 5 Corolla tube > 3 mm wide. 6 Corolla lobes alternating with corolla appendages (appearing as plaits or lobes, these often toothed, notched, or lacerate, sometimes as long as or longer than the true corolla lobes); main stem leaves cuneate at the base; perennial; [tribe Gentianeae, subtribe Gentianinae]...... Gentiana 6 Corolla lobes not alternating with corolla appendages; main stem leaves rounded to cordate at the base; biennial or annual; [tribe Gentianeae, subtribe Swertiinae]. GENTIANACEAE 412 7 Corolla lobes 4, finely fringed; main stem leaves rounded at base, with lateral veins obscure; biennial ...... Gentianopsis 7 Corolla lobes 5, entire, not fringed; main stem leaves cordate (the cordate bases often overlapping the opposite leaf), with 2-3 well-developed lateral veins (prominently visible on the lower surface); annual...... Gentianella

Bartonia Muhlenberg ex Willdenow (Bartonia)

A genus of 3 species, herbs, of e. North America. The genus has coralloid mycorrhizae and lacks root hairs, and is thus presumably partially mycotrophic. References: Gillett (1959)=Z.

1 Corolla lobes white, 5-10 mm long, spreading, spatulate to obovate, rounded at the apex; flowering in early spring (rarely to early summer) ...... B. verna 1 Corolla lobes green to creamy white, 2-3 (-5) mm long, ascending or erect, oblong to ovate or lance-ovate, acuminate or rounded-mucronate at the apex; flowering in summer or fall. 2 Mid-cauline scale leaves alternate; corolla lobes acuminate at the apex, their margins entire; anthers 0.3-0.5 mm long ...... B. paniculata ssp. paniculata 2 Mid-cauline scale leaves opposite; corolla lobes rounded at the apex, abruptly narrowed to a mucro, their margins erose (uncommonly entire); anthers 0.5-1.1 mm long...... B. virginica

Bartonia paniculata (Michaux) Muhlenberg ssp. paniculata, Screwstem Bartonia. Cp (GA, NC, SC), Mt (GA, VA), Pd (SC, VA): swamps, bogs, pocosins, pocosin ecotones, sphagnous seepages, sinkhole ponds; rare (NC Watch List, VA Watch List). August-October. Ssp. paniculata ranges from MA south to FL and west to e. TX, chiefly on the Coastal Plain, but with scattered occurrences inland (to c. VA, w. NC, KY, and AR). Ssp. iodandra (B.L. Robinson) J. Gillett is more northern, ranging from Newfoundland south to MA. These taxa have been variously treated (and ignored). Intermediates are alleged to occur in NC, MS, AL, and northward to MA (Gillett 1959, Wood & Weaver 1982). [= K, Z; < B. paniculata – RAB, GW; = B. paniculata – G; = B. paniculata var. paniculata – C, F; = B. lanceolata Small – S] Bartonia verna (Michaux) Rafinesque ex Barton, Spring Bartonia, White Bartonia. Cp (GA, NC, SC, VA): wet pine savannas, shores of Coastal Plain depression ponds, other moist sands; uncommon (NC Watch List, VA Rare). February-April (- June). VA (1 site known from City of Virginia Beach) (Belden et al. 2004) and se. NC (Carteret County) south to s. FL, west to se. TX. Perhaps not truly congeneric with the other 2 species; Wood & Weaver (1982) commented that "the species [of Bartonia] can be divided into two remarkably distinct groups." [= RAB, GW, K, S, Z] Bartonia virginica (Linnaeus) Britton, Sterns, & Poggenburg, Virginia Bartonia. Cp, Mt (GA, NC, SC, VA), Pd (NC, SC, VA): bogs, swamps, savannas, pocosin ecotones, pocosins; uncommon (rare in Mountains). July-October. Nova Scotia and Québec west to WI, south to n. FL and LA. [= RAB, C, F, G, GW, K, S, Z]

Centaurium Hill (Centaury)

A genus of about 20 species, herbs, mainly north temperate.

1 Flowers pedicellate, the pedicels 3-5 mm long ...... C. pulchellum 1 Flowers sessile or nearly so (sometimes appearing stalked but with bracteal leaves immediately below the calyx). 2 Inflorescence a corymbiform cyme (about as broad as long, the central axis poorly developed)...... C. erythraea 2 Inflorescence a spikelike cyme (distinctly elongate, the central axis straight)...... C. spicatum

* Centaurium erythraea Rafn, Common Centaury, Forking Centaury. Pd (NC, VA), Cp (VA), {GA}: lawns, disturbed areas; rare, introduced from Europe and w. Asia. July-September. [= C, K; = C. minus – RAB, later homonym; = C. umbellatum – F, G, later homonym] * Centaurium pulchellum (Swartz) Druce, Lesser Centaury, Branching Centaury. Cp (VA): disturbed areas; rare, introduced from Europe. June-September. [= C, F, G, K, S] * Centaurium spicatum (Linnaeus) Fritsch, Spiked Centaury. Cp (VA): disturbed areas; rare, introduced from s. Europe. July-August. [= C, F, G, K]

Eustoma Salisbury ex G. Don (Prairie-gentian)

A genus of 3 species, annual to perennial herbs, of se., c., and sw. North America south to Mexico and Belize and in the West Indies. References: Shinners (1957)=Z; Wood & Weaver (1982)=Y.

Eustoma exaltatum (Linnaeus) Salisbury ex G. Don, Prairie-gentian. Alkaline prairies, saline coastal areas. AL and FL west to TX, south to Mexico and Belize; West Indies. June-November. [= GW, S, Y, Z; = Eu. exaltatum ssp. exaltatum – K] {not yet keyed in generic key} GENTIANACEAE 413

Frasera Walter (Columbo)

A genus of 15 species, herbs, primarily of w. North America. References: Threadgill & Baskin (1978)=Z; Horn (1997).

Frasera caroliniensis Walter, American Columbo. Mt (GA, NC), Pd (GA, SC): rich forests over mafic rocks, upper slopes of cove forests, floodplain forests; rare (NC Rare, SC Rare). Late May-June; September-October. W. NY, nw. PA, and s. Ontario west to IL, MI, MO, and e. OK, south to w. SC, n. GA, and LA, primarily west of the Blue Ridge. Horn (1997) studied the ecology of this species in the Piedmont of SC. [= C, K, S, W, Z; = Swertia caroliniensis (Walter) Kuntze – RAB, F, G]

Gentiana Linnaeus 1753 (Gentian)

A genus of about 350-400 species, herbs, primarily temperate and arctic. Even following the removal of Gentianopsis and Gentianella, Gentiana is a large and apparently heterogeneous group, perhaps not monophyletic. No satisfactory comprehensive treatment is available, however. All of the species treated here as Gentiana are in the distinctive group often treated as section, subgenus, or genus Pneumonanthe. References: Pringle (1967)=Z; Halda (1996)=Y; Ho & Liu (2001)=X; Ho & Liu (1990); Yuan, Küpfer, & Doyle (1996); Pringle (1977). Key adapted from Z.

Identification Notes: In some species it may be somewhat difficult to interpret the corolla lobes and the corolla appendages. The filaments are alternate to the corolla lobes, and are therefore attached to the lower portion of the corolla appendages.

1 Flower solitary (rarely 2 or 3); corolla spotted within; leaves twisted, oblanceolate to oblinear; [subgenus Pneumonanthe, series Angustifoliae]...... G. autumnalis 1 Flowers clustered; corolla not spotted within; leaves planar, mostly lanceolate, elliptic, or ovate (rarely linear); subgenus Pneumonanthe, series Pneumonanthe]. 2 Calyx lobes keeled, the keel decurrent on the calyx tube; corolla greenish-white or yellowish-white; leaves ovate to lanceolate, acuminate...... G. alba 2 Calyx lobes not keeled; corolla white, greenish-white, or variously blue. 3 Corolla greenish-white (sometimes somewhat purplish); seeds wingless; lower leaves spatulate to obovate...... G. villosa 3 Corolla blue, purplish, pale blue, or nearly white; seeds winged; lower leaves linear, lanceolate, elliptic, or ovate. 4 Margins of leaves and calyx lobes entire to minutely denticulate; corolla appendages obliquely triangular, broader than high (sometimes with a minute deflexed segment)...... G. linearis 4 Margins of leaves and calyx lobes conspicuously ciliate (as seen at 10×); corolla appendages with 2 teeth, as long as broad or longer (sometimes with a deflexed segment, if so, the deflexed segment about as long as the erect one). 5 Anthers separate at anthesis; outer surfaces of petals suffused with green; calyx lobes linear-subulate, about as long as the tube; corolla lobes 6-14 mm long, about 2× as long as the the free portions of the corolla appendages ...... [G. puberulenta] 5 Anthers connate at anthesis; outer surfaces of petals not suffused with green; calyx lobes various; corolla lobes usually shorter. 6 Calyx lobes linear-subulate, broadest at the base, 4× or more as long as broad, shorter than the densely puberulent calyx tube; stems densely puberulent; corolla appendages very unevenly bifid, the narrower segment often deflexed into the corolla tube...... G. decora 6 Calyx lobes lanceolate, oblanceolate, ovate, or orbicular, 1-5× as long as broad, longer or shorter than the glabrous or puberulent calyx tube; stems glabrous or puberulent; corolla appendages subequally bifid, both segments erect. 7 Corollas open to loosely closed; involucral and upper leaves obtuse to acute (rarely acuminate); calyx lobes lanceolate. 8 Leaves ovate, widest near the base, bright green; calyx lobes longer than the calyx tube; corolla lobes spreading, usually 2-4 mm longer than the appendages ...... G. catesbaei 8 Leaves linear to elliptic, widest near the middle, dark green; calyx lobes shorter than or about equal to the calyx tube; corolla lobes usually incurved, rarely exceeding the appendages by > 2 mm ...... G. saponaria 7 Corollas tightly closed; involucral and upper leaves acuminate; calyx lobes ovate-orbicular. 9 Corolla lobes reduced to a minute mucro or triangular tooth, much exceeded by the corolla appendages ...... [G. andrewsii var. andrewsii] 9 Corolla lobes aboiut as long as the corolla appendages. 10 Calyx tubes densely puberulent; stems puberulent; filaments 7-12 mm long; corolla lobes often triangular, about 1/2 as wide as the corolla appendages...... G. austromontana GENTIANACEAE 414 10 Calyx tubes glabrous; stems glabrous; filaments 10-15 mm long; corolla lobes usually rounded, about as wide as the corolla appendages ...... G. clausa

Gentiana alba Muhlenberg ex Nuttall, Pale Gentian. Mt (NC): habitat unknown (not collected in NC in this century); rare (NC Rare). August-October. Mainly distributed in midwestern United States, from MI west to MN, south to n. AR, G. flavida occurs as scattered disjunctions eastward to PA, OH, WV, KY, and w. NC. G. alba is the older name; there is controversy, however, over whether it was validly published and applies clearly to the species at hand (see Wilbur 1988c for discussion). [= RAB, K, W, X, Y, Z; = Gentiana flavida A. Gray – C, F, G; = Dasystephana flavida (A. Gray) Britton – S; = Pneumonanthe flavida (A. Gray) Greene] Gentiana austromontana Pringle & Sharp, Blue Ridge Gentian. Mt (NC, VA): high elevation forests and grassy balds; rare (NC Watch List, VA Watch List). September-October. A Southern Appalachian endemic: s. WV and sw. VA south to w. NC and ne. TN. The flowers of G. austromontana are usually a deeper and more intense blue-violet than the similar G. clausa and G. decora. See Pringle & Sharp (1964) for additional discussion. [= C, K, W, X, Y, Z; < G. clausa Rafinesque – RAB, F, G, GW; < Dasystephana decora (Pollard) Small – S] Gentiana autumnalis Linnaeus, Pinebarren Gentian. Cp (NC, SC, VA): savannas, pine flatwoods, sandhills, in a variety of sites varying from moist to very xeric, nearly always associated with Pinus palustris and Aristida stricta; uncommon, rare in VA (SC Rare, VA Rare). Late September-early January (rarely at other times of the year, such as spring, in response to fire). This species is a "bimodal endemic," occurring in s. NJ and adjacent DE, and from se. VA south through e. NC to nc. SC. The related G. pennelliana Fernald (sometimes reduced to a subspecies of G. autumnalis) is endemic to the FL panhandle; other siblings occur in Mexico: G. bicuspidata (G. Don) Briq., G. hooperi Pringle, and G. longicollis Nesom. G. autumnalis is often overlooked, since it is very inconspicuous except when in flower, it usually flowers at a season when few botanists are about, and sterile plants greatly outnumber fertile ones. Vegetatively it is extremely distinctive once learned; the leaves are glossy, dark- green, opposite, oblanceolate to "oblinear," and twisted and curved in a manner reminiscent of an airplane propellor. [= RAB, C, F, GW, K, X, Z; = Gentiana porphyrio J.F. Gmelin – G; = Dasystephana porphyrio (J.F. Gmelin) Small – S; = Gentiana autumnalis ssp. autumnalis – Y; = Pneumonanthe porphyrio (Linnaeus) Greene] Gentiana catesbaei Walter, Coastal Plain Gentian. Cp (GA, NC, SC, VA), Pd (VA): pocosins, moist savanna edges, edges of moist hardwood forests; common (rare in Piedmont). Late September-November. S. NJ south to n. FL, on the Coastal Plain. [= RAB, C, G, GW, K, X, Y, Z; > G. catesbaei var. catesbaei – F; > G. catesbaei var. nummulariifolia Fernald – F; > Dasystephana latifolia (Chapman) Small – S; > D. parvifolia (Chapman) Small – S; = Pneumonanthe catesbaei (Walter) F.W. Schmidt] Gentiana clausa Rafinesque, Meadow Closed Gentian, Meadow Bottle Gentian. Mt, Pd (NC, VA): forests; uncommon. September-October. Mostly Appalachian: ME south to w. NC and ne. TN, extending east and west to adjacent physiographic provinces. [= C, K, W, X, Y, Z; < G. clausa – RAB, F, G, GW (also see G. austromontana); < Dasystephana decora (Pollard) Small – S; = Pneumonanthe clausa (Rafinesque) Greene] Gentiana decora Pollard, Appalachian Gentian. Mt, Pd (GA, NC, SC, VA): forests; common (rare in Piedmont). September-October. A Southern Appalachian endemic: c. WV south through w. VA to w. NC, e. TN, nw. SC, ne. GA. [= RAB, C, F, G, K, W, X, Y, Z; < Dasystephana decora (Pollard) Small – S; = Pneumonanthe decora (Pollard) Greene] Gentiana linearis Frölich, Narrowleaf Gentian. Mt (VA): openings in spruce-fir forest near the summit of Whitetop Mountain, VA; rare (VA Rare). September-October. Mainly occurring in ne. United States and e. Canada, west to Lake Superior, and south (scattered) in the Appalachians to sw. VA and e. TN (Chester, Wofford, & Kral 1997). On Mount LeConte (Sevier County, TN), G. linearis occurs in thin soils around high elevation outcrops of Anakeesta Slate. See Pringle (1977) for extensive discussion of actual and putative southern occurrences of this species. [= C, F, G, K, W, X, Y, Z; = Pneumonanthe linearis (Frölich) Greene] Gentiana saponaria Linnaeus, Soapwort Gentian. Cp, Pd, Mt (GA, NC, SC, VA): bogs, marshes, wet hardwood forests, other moist to wet habitats; uncommon. Late September-November. NY west to n. IL, south to w. FL and e. TX. A peculiar form with very narrow leaves has been found at several localities in Ashe and Watauga counties, NC and in the South Mountains, NC; it may warrant taxonomic recognition after further study. [= RAB, C, GW, K, W, X, Y, Z; > G. saponaria – F, G; > G. cherokeensis (W.P. Lemmon) Fernald – F, G; > G. saponaria var. saponaria – K; > G. saponaria var. latidens House – K; = Dasystephana saponaria (Linnaeus) Small – S; = Pneumonanthe saponaria (Linnaeus) F.W. Schmidt] {investigate varieties} Gentiana villosa Linnaeus, Striped Gentian. Pd, Mt, Cp (GA, NC, SC, VA): upland forests, sandhill/pocosin ecotones; uncommon. Late August-November. Se. PA west to n. KY and w. TN, south to w. FL and e. LA. [= RAB, C, F, G, K, W, X, Y, Z; = Dasystephana villosa (Linnaeus) Small – S; = Pneumonanthe villosa (Linnaeus) F.W. Schmidt]

Gentiana andrewsii Grisebach var. andrewsii, a northern species, ranges south to s. MD and WV; earlier reports of it as far south as GA or NC (as by F and G) are apparently based on misidentifications. An additional variety, var. dakotica A. Nelson, occurs from Manitoba and Saskatchewan south in the Great Plains to MO and IL. [= C, K, X, Y, Z; < G. andrewsii – F, G; < Dasystephana andrewsii (Grisebach) Small – S; = Pneumonanthe andrewsii (Grisebach) W.A. Weber var. andrewsii] Gentiana puberulenta J. Pringle, Prairie Gentian, ranges from w. NY west to ND, south to WV, KY, sc. TN (Coffee County) (Chester, Wofford, & Kral 1997), LA, n. AR, and KS. [= C, K, X, Y, Z; = G. puberula – F, G, misapplied; = Dasystephana puberula (Michaux) Small – S, misapplied]

Gentianella Moench (Agueweed)

GENTIANACEAE 415 A genus of about 125 species, herbs, temperate. The separation of Gentianella from Gentiana appears to be well warranted; some characters suggest that Gentianella is more closely allied to Swertia, Halenia, and Lomatogonium than to Gentiana (Wood & Weaver 1982). A molecular analysis has confirmed this (Yuan & Küpfer 1995). References: Gillett (1957)=Z. Key based on Gillett (1957).

1 Calyx 8-10 mm long; calyx tube 3.0-3.5 mm long; calyx lobes 5-6 mm long, elliptic-lanceolate to oblanceolate with thickened margins, and with broadly flattened, frequently nerve-like keels; corolla ca. 20 mm long...... G. quinquefolia var. occidentalis 1 Calyx 4-5 mm long; calyx tube 1.5-2.0 mm long; calyx lobes 2.0-2.5 mm long, narrowly triangular with hyaline margins and very prominent keels; corolla 16-18 mm long ...... G. quinquefolia var. quinquefolia

Gentianella quinquefolia (Linnaeus) Small var. occidentalis A. Gray, Western Agueweed. Mt (VA): calcareous barrens, dry and dry-mesic limestone woodlands; rare (VA Rare). Late August-October. Var. occidentalis A. Gray is more western, from OH and s. Ontario west to MN, east and south to w. VA, sc. KY, AR, and se. KS. [= C, G; < Gentiana quinquefolia Linnaeus – RAB, GW, W; = Gentiana quinquefolia var. occidentalis (A. Gray) Hitchcock – F; = Gentianella quinquefolia ssp. occidentalis (A. Gray) J. Gillett – K, Z; = Gentianella occidentalis (A. Gray) Small – S] Gentianella quinquefolia (Linnaeus) Small var. quinquefolia, Eastern Agueweed. Mt (GA, NC, SC, VA): forests, grassy balds; common. Late August-October. Var. quinquefolia is primarily Appalachian, from ME west to w. NY and s. Ontario, south to n. GA and sc. TN. [= C, G; < Gentiana quinquefolia Linnaeus – RAB, GW, W; = Gentiana quinquefolia var. quinquefolia – F; = Gentianella quinquefolia ssp. quinquefolia – K, Z; = Gentianella quinquefolia – S]

Gentianopsis Ma 1951 (Fringed-gentian)

A genus of about 20 species, herbs, of north temperate Asia and North America. The reasons for the recognition of Gentianopsis are enumerated by Ma (1951), Iltis (1965), and Wood & Weaver (1982). References: Gillett (1957)=Z; Iltis (1965); Ma (1951).

Gentianopsis crinita (Frölich) Ma, Eastern Fringed-gentian. Mt (GA, NC, VA): sunny or semi-shaded seepage areas over calcareous, mafic, or ultramafic rocks (such as limestone, amphibolite, or serpentinized olivine); rare (GA Threatened, NC Endangered, VA Rare). September-October. ME, s. Ontario, and ND south to NJ, IN, and IA (mostly north of the glacial maximum) and from PA south to nw. NC and ne. GA in the unglaciated Appalachians. Certainly one of the most beautiful of our native plants. [= C, K; = Gentiana crinita Frölich – RAB, F, G, GW, W; = Anthopogon crinitum (Frölich) Rafinesque – S; = Gentianella crinita (Frölich) G. Don ssp. crinita – Z]

Obolaria Linnaeus (Pennywort)

A monotypic genus, herb, of e. North America. References: Gillett (1959)=Z.

Obolaria virginica Linnaeus, Pennywort. Mt, Pd (GA, NC, SC, VA), Cp (NC, VA): nutrient-rich, moist forests; common (rare in Coastal Plain south of VA). March-May; May-June. NJ west to OH, s. IN, and s. IL, south to panhandle FL and se. LA (reported from TX). The small purplish-green plants are inconspicuous, often nearly hidden under fallen leaves. Obolaria has well-developed mycorrhizae and may be substantially mycotrophic. [= RAB, C, F, G, K, S, W, Z]

Sabatia Adanson 1763 (Sabatia, Rose-gentian, Marsh-pink, Sea-pink) (contributed by B.A. Sorrie and A.S. Weakley)

A genus of about 20 species, of North America and the West Indies. References: Wilbur (1955)=Z.

1 Flowers with (7-) 8-12 (-14) corolla lobes. 2 Pedicels < 5 mm long; calyx subtended by linear bracts that usually exceed the corolla lobes; terminal flowers in capitate clusters (less commonly single). 3 Basal leaves similar in shape and size to the stem leaves; cauline leaves (25-) 35-50 (-65) mm long, (7-) 10-20 (- 25) mm wide, 2-4× as long as wide; corolla lobes pale rose or white; [of mountain slopes, restricted in our area to sw. NC and nw. GA]...... S. capitata 3 Basal leaves much broader and shorter than the stem leaves; cauline leaves (15-) 40-80 (-100) mm long, 1-3 mm wide, 20-60× as long as wide; corolla lobes medium rose to deep rose; [of bogs and savannas, of the Coastal Plain] ...... S. gentianoides 2 Pedicels > 10 mm long; calyx not subtended by long bracts; terminal flower single. 4 Upper stem leaves about as wide as the diameter of the stem, or narrower; calyx lobes terete or semi-terete; stems 6-12 dm tall; [of Taxodium ascendens-Nyssa biflora depressions and wet pine flatwoods in se. SC] .....S. bartramii GENTIANACEAE 416 4 Upper stem leaves much wider than the diameter of the stem; calyx lobes flat, linear to narrowly oblanceolate; stems 3-12 dm tall; [of various habitats, primarily along the shores of blackwater rivers or ponds, or in tidal marshes]. 5 Primary branches opposite; terminal flower short-stalked (much shorter than the first internode of the adjacent branch); stems 5-12 dm tall; [of drawdown blackwater riverbanks and similar situations] ...... S. kennedyana 5 Primary branches usually alternate; terminal flower long-stalked (usually longer than the first internode of the adjacent branch); stems 3-7 dm tall; [of brackish marshes or openings along blackwater streams]. 6 Surficial stolons usually absent or poorly developed; internodes commonly much longer than leaves; [of tidal brackish or freshwater marshes]...... S. dodecandra 6 Surficial stolons usually present and well-developed, elongate; internodes shorter than to equaling the leaves; [of openings of blackwater streams]...... S. foliosa 1 Flowers with 5-6 (-7) corolla lobes. 7 Upper branches of main stem alternate. 8 Calyx tube strongly winged; corolla lobes pink; [very rare introduction from c. United States]...... S. campestris 8 Calyx tube not winged; corolla lobes pink or white; [native, primarily of the Coastal Plain, except S. campanulata which has disjunct occurrences in the Mountains]. 9 Calyx lobes foliaceous, 5-8 mm wide, oblong to oblanceolate, mostly exceeding the corolla lobes...... S. calycina 9 Calyx lobes linear-setaceous, 0.5-2 mm wide, if equaling the corolla lobes then very narrow and not foliaceous. 10 Calyx lobes (3-) 4-7 (-8) mm long; corolla lobes white; [of the Coastal Plain from se. SC southward]...... S. brevifolia 10 Calyx lobes (4-) 6-17 (-23) mm long; corolla lobes pink (rarely white in individual plants); [of the Coastal Plain of GA, NC, SC, and VA, and rarely the Mountains of NC and VA]. 11 Plants perennial, often with several stems from a caudex; calyx lobes > 3/4× as long as the corolla lobes, and sometimes exceeding them; [of saturated soils from Coastal Plain savannas to Mountain bogs]...... S. campanulata 11 Plants annual, solitary; calyx lobes up to 3/4× as long as the corolla lobes; [of brackish marshes and interdune swales]...... S. stellaris 7 Upper branches of main stem opposite. 12 Corolla lobes pink (rarely white); pedicels at least in part > 5 mm long. 13 Lower half of stem winged; leaves ovate, clasping, < 2× as long as wide; [widespread in our area]...... S. angularis 13 Lower half of stem not winged; leaves elliptic to lanceolate, more or less tapered to the base, mostly > 3× as long as wide; [of the Coastal Plain or very rarely Piedmont] ...... S. brachiata 12 Corolla lobes white or creamy white; pedicels (above the uppermost bracts or branches) ca. 1-2 (-5) mm long. 14 Lower portion of stem quadrangular, narrowly winged; plants annual or biennial, with 1 (-several) stems arising from a taproot ...... S. quadrangula 14 Lower portion of stem terete, not winged (though the upper stem is quadrangular or angled in S. difformis); plants perennial, with several stems arising from a short rhizome; [section Eusabatia, subsection Difformes]. 15 Leaves and upper stem not glaucous; stem terete below, becoming quadrangular or quadrangular-angled above; corolla lobes (5-) 7-15 (-21) mm long; [widespread in our area]...... S. difformis 15 Leaves and upper stem glaucous; stem terete throughout; corolla lobes (4-) 5-7 (-8) mm long; [of GA southward and westward]. 16 Calyx-lobes erect, (0.1-) 0.2-1.5 (-2.0) mm long, as long as or shorter than the calyx-tube; [of sw. GA and n. FL westward to e. LA] ...... S. macrophylla var. macrophylla 16 Calyx lobes strongly recurved, (1.0-) 1.5-3 mm long, longer than the calyx-tube; [of e. and sc. GA south to ne. FL] ...... S. macrophylla var. recurvans

Sabatia angularis (Linnaeus) Pursh, Bitter-bloom, Common Marsh-pink. Cp, Pd, Mt (GA, NC, SC, VA): forests, woodlands, marshes, fields; common. July-August; September-October. NY west to s. MI, IL, and e. KS, south to Panhandle FL and e. TX. [= RAB, C, F, GW, K, S, W, Z] Sabatia bartramii Wilbur, Bartram's Rose-gentian. Cp (GA, SC): margins of Taxodium ascendens-Nyssa depressions, wet pine flatwoods; rare north of GA (SC Rare). June-August; August-October. Ne. SC south to s. FL, west to s. AL and se. MS. [= GW, K, Z; = S. dodecandra var. coriacea (Elliott) Ahles – RAB; = S. decandra (Walter) Harper – S] Sabatia brachiata Elliott, Narrowleaf Rose-pink. Cp (GA, NC, SC, VA), Pd (GA, NC), Mt (GA): sandhills, pine savannas, pine flatwoods; uncommon (VA Watch List). Late May-July; August-September. Se. VA south to s. GA, west to LA, north in the interior to c. TN and se. MO. [= RAB, C, F, GW, K, S, W, Z] Sabatia brevifolia Rafinesque. Cp (GA, SC): pine savannas; rare north of GA. September-October; October-November. E. SC south to peninsular FL, west to s. AL. [= RAB, GW, K, Z; = S. elliottii Steudel – S] GENTIANACEAE 417 Sabatia calycina (Lamarck) Heller, Coastal Rose-pink. Cp (GA, NC, SC, VA): swamp forests, river banks; common (VA Watch List). June-October; July-October. Se. VA south to peninsular FL, west to se. TX; e. Cuba and Hispaniola. [= RAB, C, F, GW, K, S, Z] Sabatia campanulata (Linnaeus) Torrey, Slender Marsh-pink. Cp (GA, NC, SC, VA), Mt (GA, NC, VA): pine savannas, bogs; common (rare in Mountains) (VA Rare). June-August; September-October. MA south to n. FL, west to LA and AR; scattered inland as in w. VA, w. NC, c. TN, and KY. [= RAB, C, GW, K, S, W, Z; > S. campanulata var. campanulata – F; > S. campanulata var. gracilis (Michaux) Fernald – F] * Sabatia campestris Nuttall, Western Marsh-pink, Prairie Rose-gentian, Prairie Sabatia. Mt (NC): roadsides and woodland edges; rare (NC Watch List), introduced from c. United States. July-August; September-October. [= RAB, C, F, GW, K, Z] Sabatia capitata (Rafinesque) Blake, Cumberland Rose-gentian. Mt (GA, NC): sloping woodlands and meadows, over sandstone or shale; rare (GA Rare, NC Watch List). July-August; September-October. Sw. NC and se. TN south to nw. GA and c. AL. Apparently present in NC, at least formerly, based on a specimen collected "from Cherokee", probably Cherokee County, NC, a remarkably poorly botanized area. [= K, Z; = Lapithea capitata (Rafinesque) Small – S] Sabatia difformis (Linnaeus) Druce. Cp (GA, NC, SC, VA), Pd (NC, SC): pine savannas, bogs, pocosins; common (rare in Piedmont) (VA Rare). May-September; September-December. S. NJ south to peninsular FL, west to s. AL. [= RAB, C, F, GW, K, S, Z] Sabatia dodecandra (Linnaeus) Britton, Sterns, & Poggenburg, Perennial Sea-pink, Large Marsh Rose-pink. Cp (GA, NC, SC, VA): tidal brackish and freshwater marshes; common (VA Watch List). June-August; August-October. CT south to e. SC and e. GA (Sorrie 1998b). [= F, S; < S. dodecandra var. dodecandra – RAB (also see S. foliosa); = S. dodecandra var. dodecandra – C, GW, K, Z] Sabatia foliosa Fernald. Cp (GA, SC): openings along blackwater rivers, rare north of GA. June-August; August-October. E. SC south to n. FL, west to se. TX. [< S. dodecandra var. dodecandra – RAB; = S. dodecandra (Linnaeus) Britton, Sterns, & Poggenburg var. foliosa (Fernald) Wilbur – GW, K, Z; > S. foliosa – S; > S. harperi Small – S] Sabatia gentianoides Elliott. Cp (GA, NC, SC): pine savannas, bogs; common. July-August; September-October. NC south to n. FL, west to se. TX. [= RAB, GW, K, Z; = Lapithea gentianoides (Elliott) Grisebach – S] Sabatia kennedyana Fernald, Plymouth Gentian. Cp (NC, SC, VA*): seasonally exposed drawdown banks of the Waccamaw River, in adjacent ditches and disturbed flats (in se. NC and ne. SC), and very rarely on shores of beaver ponds (in e. VA, by introduction); rare (NC Threatened, SC Rare). June-August; August-October. This species has a strange, disjunct range, likely related to Pleistocene refugia on the (now) Continental shelf, present in s. Nova Scotia; e. MA and RI; se. NC and ne. SC. The record of the species in e. VA (Caroline County) reported by Fleming & Ludwig (1996) has now been determined to be a deliberate introduction. Studies underway suggest that the Carolina plants may differ varietally from those in New England (Sorrie, pers. comm.). [= C, F, GW, K, Z; = S. dodecandra var. kennedyana (Fernald) Ahles – RAB] Sabatia macrophylla Hooker var. macrophylla, Large-leaf Rose-gentian. Cp (GA): wet savannas; uncommon. Sw. GA west to e. LA. [= K, Z; < S. macrophylla – GW; = S. macrophylla Hooker – S] Sabatia macrophylla Hooker var. recurvans (Small) Wilbur, Small's Rose-gentian. Cp (GA): wet savannas; common. E. and c. GA south to ne. FL; it may occur in se. SC. [= K, Z; < S. macrophylla – GW; = S. recurvans Small – S] Sabatia quadrangula Wilbur, Four-angle Sabatia. Cp, Pd (GA, NC, SC, VA): sandhills, moist forests, pocosin ecotones; uncommon (NC Watch List). June-September; August-November. E. VA south to FL, west to s. AL. [= RAB, C, GW, K, Z; = S. paniculata Michaux – F, S, misapplied] Sabatia stellaris Pursh, Annual Sea-pink. Cp (GA, NC, SC, VA): brackish marshes; common. July-October; August- November. S. MA south to s. FL, west to LA; Bahama Islands, Cuba, c. Mexico. [= RAB, C, F, GW, K, Z]

GERANIACEAE A.L. de Jussieu 1789 (Geranium Family)

A family of about 11 genera and 700 species, herbs and shrubs, mostly temperate.

1 Leaves pinnately cleft or compound; fertile stamens 5, staminodia 5...... Erodium 1 Leaves palmately cleft or compound; fertile stamens 10 (except in G. pusillum, and note that anthers are readily deciduous in all species) ...... Geranium

Erodium Linnaeus (Stork's-bill, Filaree)

A genus of about 60 species, herbs, mainly Old World.

1 Leaves simple, deeply lobed but not divided ...... E. texanum 1 Leaves compound, with 3 or more leaflets. 2 Primary leaflets sessile or nearly so, sometimes connected by blade tissue; blades of the primary leaflets divided nearly or quite to the base; apical pits of mericarp lacking sessile glands ...... E. cicutarium 2 Primary leaflets petiolulate; blades of the primary leaflets divided <0.75× to the base; apical pits of mericarp with sessile glands...... E. moschatum var. moschatum

GERANIACEAE 418 * Erodium cicutarium (Linnaeus) L'Héritier, Heron's-bill, Common Stork's-bill, Redstem Filaree, Alfileria, Pin-clover. Mt, Pd, Cp (GA, NC, SC, VA): disturbed areas, fields, lawns; common, introduced from Europe. March-June; April-July. [= RAB, C, F, G, S, W; > E. cicutarium ssp. cicutarium – K] * Erodium moschatum (Linnaeus) L'Héritier var. moschatum, Whitestem Filaree. Cp (SC): waste area near wool-combing mill; rare, perhaps merely a waif, native of Mediterranean Europe. April-September. Naturalized south to DE and PA; also in SC Coastal Plain. [= F, K; < E. moschatum – C, G, S] * Erodium texanum A. Gray, Texas Stork's-bill. Cp (SC): waste areas near wool-combing mill; rare, perhaps merely a waif, native of sc. and sw. United States. [= K]

Geranium Linnaeus 1753 (Geranium, Crane's-bill)

A genus of about 300-430 species, herbs, mainly temperate. House plants called 'geranium' are members of the genus Pelargonium. References: Aedo, Aldasoro, & Navarro (1998); Yeo (1984).

1 Petals 12-18 mm long; perennial, from a stout rhizome; anthers >2 mm long; [subgenus Geranium, section Geranium] ...... G. maculatum 1 Petals 2-13 mm long; annual or biennial, from a taproot; anthers <1 mm long. 2 Leaves compound, at least the terminal segment (and often also the two lateral segments) petiolulate, not connected to the lateral segments by leaf tissue; petals 9-13 (-15) mm long; [rare in our area, restricted to Mountains of VA]; [subgenus Robertium, section Robertium] ...... G. robertianum 2 Leaves dissected, but not compound, all segments interconnected by leaf tissue; petals 2-10 mm long; [collectively common and widespread in our area]. 3 Sepals blunt or acute, or terminating in a minute callus tip; [subgenus Robertium, section Batrachioidea]. 4 Mericarps glabrous, reticulately ridged; stem pubescence an admixture of long eglandular hairs (1.0-1.7 mm long) and short (< 0.5 mm long) gland-tipped and eglandular hairs; stamens (all 10) fertile (note that anthers may fall readily)...... G. molle 4 Mericarps appressed pubescent, not ridged; stem pubescence of short (< 0.3 mm long), gland-tipped and eglandular hairs; stamens partly sterile (the inner 5 fertile, the outer 5 lacking anthers)...... G. pusillum 3 Sepals awned or subulate, the subulate awn 0.7-3 mm long. 5 Mature pedicels < 1.5× as long as the calyx. 6 Mericarps with spreading hairs about 0.5 mm long, these often gland-tipped; [subgenus Geranium, section Dissecta] ...... G. dissectum 6 Mericarps with long appressed hairs about 1 mm long, these not gland-tipped; [subgenus Geranium, section Geranium]. 7 Inflorescence diffusely corymbiform (because of long upper internodes), mostly 4-12-flowered; pubescence of stem mostly < 0.5 mm long...... G. carolinianum var. carolinianum 7 Inflorescence a compact corymb (because of notably short upper internodes), mostly 5-25- flowered; pubescence of stem mostly >0.75 mm long...... G. carolinianum var. confertiflorum 5 Mature pedicels >2× as long as the calyx; [subgenus Geranium, section Geranium]. 8 Pedicels spreading pubescent, the hairs not glandular; mature stylar beak ca. 1 mm long....[G. sibiricum] 8 Pedicels either retrorse-strigose or glandular-villous; mature stylar beak 3-5 mm long. 9 Pedicels glandular-villous ...... [G. bicknellii] 9 Pedicels retrorsely strigose...... G. columbinum

Geranium carolinianum Linnaeus var. carolinianum, Southern Carolina Crane's-bill. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, lawns, pastures, gardens, disturbed areas; common. March-June (and sometimes later). [= C, F, G; < G. carolinianum – RAB, S, W; < G. carolinianum var. carolinianum – K] Geranium carolinianum Linnaeus var. confertiflorum Fernald, Northern Carolina Crane's-bill. Mt (NC, VA), Pd (VA): fields, roadsides, lawns, gardens, disturbed areas; common. March-June (and sometimes later). [= C, F, G; < G. carolinianum – RAB, S, W; < G. carolinianum var. carolinianum – K] * Geranium columbinum Linnaeus, Long-stalk Crane's-bill. Mt (GA, NC, VA), Pd (VA): roadsides, pastures, disturbed areas; common, introduced from Europe. May-July. [= RAB, C, F, G, K, S, W] * Geranium dissectum Linnaeus, Cutleaf Crane's-bill. Mt (GA, NC, SC, VA), Pd (NC, SC, VA), Cp (GA, NC, VA): roadsides, pastures, disturbed areas; common, introduced from Europe. April-June. [= RAB, C, F, G, K, S, W] Geranium maculatum Linnaeus, Wild Geranium. Mt, Pd, Cp (GA, NC, SC, VA): cove forests, bottomland forests, other mesic, base-rich forests; common (rare in Coastal Plain). April-June. ME west to Manitoba, south to SC, GA, and ne. OK. Sometimes cultivated. [= RAB, C, F, G, K, S, W] * Geranium molle Linnaeus, Dove's-foot Crane's-bill. Pd (GA, NC, SC, VA), Mt (NC, SC, VA), Cp (VA): roadsides, pastures, disturbed areas; common, introduced from Europe and w. Asia. April-July. [= RAB, C, F, G, K, S, W] * Geranium pusillum Linnaeus, Small-flowered Crane's-bill. Mt (NC, VA), Pd (SC, VA), Cp (VA): roadsides, pastures, disturbed areas; common, introduced from Europe. May-June. [= RAB, C, G, K, S, W] GERANIACEAE 419 Geranium robertianum Linnaeus, Herb Robert. Mt (VA): rocky woodlands, especially over calcareous rocks; rare (VA Rare). June-October. Circumpolar, ranging south in North America to w. VA, nc. TN (Chester, Wofford, & Kral 1997), OH, IN, and IL. Considered by C and G to be introduced in North America, but apparently native. [= C, F, G, K, W]

Geranium bicknellii Britton. South to PA, IN, IL, c. TN. No documentation is known for the reports for VA and WV by Kartesz (1999). [= C, G, K, W; > G. bicknellii var. bicknellii – F] * Geranium ibericum Cavanilles, Iberian Crane's-bill. Mt (NC): rare, spread from horticultural use, native of Europe. Recently been found in Great Smoky Mountains National Park, in both NC and TN (K. Langdon, pers. comm.). [= F, K; G. nepalense – C] {not keyed at this time} * Geranium sanguineum Linnaeus, Blood-red Crane's-bill. Mt (NC): roadbank, rare, probably persistent or spread from cultivation. [= C, F, G, K] {not keyed at this time; add to synonymy} * Geranium sibiricum Linnaeus, native to Asia, is naturalized south to s. PA (Rhoads & Klein 1993) and is likely to occur in at least the northern part of our area. [= C, F, K] * Geranium thunbergii Siebold & Zuccarini ex Lindley & Paxton. Mt (NC), Pd (VA): lawn along Blue Ridge Parkway; rare, native of e. Asia. Reported for NC by Nesom (2000) on the basis of a 1936 specimen. [= K; G. ibericum Cavanilles – C, apparently misapplied; = G. nepalense Sweet var. thunbergii (Siebold & Zuccarini ex Lindley & Paxton) Kudo – F, G] {not keyed at this time, and 'promote' to main text}

GROSSULARIACEAE DC. 1805 (Currant Family) (also see ITEACEAE)

A family of one genus, of the northern hemisphere and montane South America (Andes). The familial distinction from the Saxifragaceae is supported by recent molecular data, though the affinities of Grossulariaceae and Saxifragaceae (sensu stricto) are closer than those of many other groups traditionally included in the Saxifragaceae, such as Parnassia, Lepuropetalon, and Penthorum (Morgan & Soltis 1993).

Ribes Linnaeus 1753 (Currant, Gooseberry)

A genus of about 150 species, temperate, of the Northern Hemisphere and montane South America. The genus is separated into distinctive subgenera, these sometimes maintained as full genera (as by S). Of the species treated or mentioned here, the currants (subgenus Ribes) include R. americanum, R. aureum var. villosum, R. glandulosum, R. lacustre, R. nigrum, R. rubrum, and R. triste. The gooseberries (subgenus Grossularia) include R. curvatum, R. cynosbati, R. echinellum, R. hirtellum, R. missouriense, R. rotundifolium, and R. uva-crispa var. sativum. The dried "currants" commonly available in stores are actually raisins made from a small variety of grape, and have nothing to do with Ribes. A molecular study suggests that recognition of Grossularia as a genus distinct from Ribes is not warranted, though it does form a monophyletic group nested within Ribes s.l. (Senters & Soltis 2003; Weigend, Mohr, & Motley 2002). References: Sinnott (1985)=Z; Weigend, Mohr, & Motley (2002); Spongberg (1972); Schultheis & Donoghue (2004); Senters & Soltis (2003). Key adapted from C, F, and Z.

1 Flowers solitary or in corymbs of 2-4; pedicels not jointed just beneath the ovary or fruit, the fruit not disarticulating at maturirty; stems with (0-) 1-3 nodal spines and sometimes also with internodal bristles (especially on young, vigorous growth); [subgenus Grossularia]. 2 Ovary and fruit bristly or spiny; stamens (at full anthesis) either 9-15 mm long, exserted well beyond the calyx lobes (R. echinellum) or 1-3 mm long, shorter than the calyx lobes (R. cynosbati). 3 Bristles of the ovary and fruit not gland-tipped; stamens (at full anthesis) 1-3 mm long, shorter than the calyx lobes; calyx lobes 2.5-4 mm long; petals 1-2 mm long; [of the Mountains] ...... R. cynosbati 3 Bristles of the ovary and fruit gland-tipped; stamens (at full anthesis) either 9-15 mm long, exserted well beyond the calyx lobes; calyx lobes 4-7 mm long; petals 2-3 mm long; [of the Piedmont or, potentially, Coastal Plain]...... R. echinellum 2 Ovary and fruit glabrous; stamens (at full anthesis) 6-12 mm long, exserted well beyond the calyx lobes. 4 Calyx tube white; stamens 9-12 mm long; nodal spines 1-3 per node, 7-18 mm long, stout ...... R. missouriense 4 Calyx tube purplish or purplish-green; stamens 6-8 mm long; nodal spines 0-1 (-2) per node, 3-11 mm long, slender...... R. rotundifolium 1 Flowers in racemes of 4-many; pedicels jointed just beneath the ovary or fruit, the fruit disarticulating at maturity; stems lacking nodal spines and internodal bristles (except R. lacustre); [subgenus Ribes]. 5 Stems (especially young, vigorous growth) with internodal bristles and sometimes internodal spines; fruit bristly, purple or black when mature; racemes drooping...... R. lacustre 5 Stems lacking internodal bristles and nodal spines; fruit glabrous or glandular-hispid, red or black when mature (but not both dark and bristly); racemes ascending, spreading, or drooping. 6 Ovary and fruit bristly or spiny with glandular hairs; fruit dark red when mature; racemes ascending; leaves mostly 5-10 cm across, the 5-7 lobes more-or-less acute...... R. glandulosum 6 Ovary and fruit glabrous; fruit red or black when mature; racemes spreading to drooping; leaves mostly 3-8 cm across, the 3-5 lobes more-or-less obtuse. GROSSULARIACEAE 420 7 Leaves with resinous glands beneath; fruits black when mature; [native]...... R. americanum 7 Leaves not glandular beneath; fruits red when mature; [introduced, rarely naturalized]...... R. rubrum

Auxiliary Key to widely distributed Ribes of the Mountains

1 Leaves usually 5-10 cm long and wide, serrate or doubly serrate with sharp teeth; plants usually without nodal spines; inflorescence a raceme of 4-15 flowers; pedicel jointed below the fruit...... R. glandulosum 1 Leaves 1-5 cm long and wide, serrate with rounded teeth; plants usually with nodal spines; inflorescence a raceme of 1-4 flowers; pedicel not jointed below the fruit. 2 Ovary with glandular hairs which become stiff spines on the mature fruit; leaf bases cordate to deeply cordate (rarely truncate or cuneate), the angle of leaf tissue mostly 190-230 degrees, moderately to sparsely silvery-pilose beneath, usually on the surface as well as on the veins and in the vein axils; stamens at full anthesis equalling the petals...... R. cynosbati 2 Ovary and fruit glabrous; leaf bases rounded or cuneate (rarely truncate or cordate), the angle of leaf tissue mostly 130- 170 degrees, glabrescent to sparsely pubescent beneath (mostly on the veins and in the vein axils); stamens at full anthesis exceeding the petals ...... R. rotundifolium

Ribes americanum P. Miller, American Black Currant. Mt (VA): moist forests, marl marshes; rare (VA Rare). April-June. Nova Scotia west to Alberta, south to w. VA, e. and nc. KY (Clark et al. 2005), IN, NE, and CO. [= C, F, G, K, W] Ribes cynosbati Linnaeus, Prickly Gooseberry, Dogberry. Mt (GA, NC, VA): moist slopes, periglacial boulderfields, grassy balds, mostly at high elevations; common. May-June; July-September. Ne. United States and s. Canada south to w. NC, e. TN, n. GA, n. AL, AR, and OK. [= RAB, C, G, K, W, Z; > R. cynosbati var. cynosbati – F; > R. cynosbati var. glabratum Fernald – F; = Grossularia cynosbati (Linnaeus) P. Miller – S] Ribes echinellum (Coville) Rehder, Miccosukee Gooseberry. Pd (SC): mesic, nutrient-rich forests; rare (US Endangered, SC Rare). March-April; June-September. This species has a remarkable range, known only from a small area of McCormick County, SC and the vicinity of Lake Miccosukee, Jefferson County, FL. Godfrey (1988) has a detailed description of R. echinellum. Catling, Dumouchel, & Brownell (1998) discuss its pollination biology. [= K, Z; = Grossularia echinella Coville – S] Ribes glandulosum Grauer, Skunk Currant, Mountain Currant. Mt (NC, VA): periglacial boulderfields, high elevation seeps, spruce-fir forests; uncommon, but locally abundant (VA Watch List). May-June; June-September. Newfoundland and British Columbia, south to VT, MI, and MN, and in the mountains to w. NC and e. TN. [= RAB, C, F, G, K, S, W] Ribes lacustre (Persoon) Poiret, Bristly Black Currant, Spiny Swamp Currant. Mt (VA): forests, acid swamps; rare (VA Rare). May-June. Labrador to AK, south to MA, PA, w. VA, TN (allegedly), n. OH, MI, MN, CO, UT, and CA. The documentation for the inclusion of R. lacustre in the flora of VA is a sterile specimen not definitely identifiable (Wieboldt, pers. comm.). [= C, F, G, K, W] Ribes missouriense Nuttall, Missouri Gooseberry. Mt (VA): forests, rock outcrops; rare, in VA probably introduced from further west (VA Watch List), but native in KY and TN. April-June; May-July. W. WV, sw. OH, IN, WI, MN, and e. ND south to KY, e. TN (Roane and Grainger counties), s. IL, MO, n. AR, and KS, with scattered occurrences (perhaps escapes from cultivation) in CT, NJ, PA, MD, and VA. [= C, F, G, K, Z; = Grossularia missouriensis (Nuttall) Coville & Britton – S] Ribes rotundifolium Michaux, Appalachian Gooseberry. Mt (NC, SC, VA), Pd (VA): moist slopes, balds, boulderfields, rocky forests, mostly at high elevations south of VA; common (rare in Piedmont). April-May; June-September. An Appalachian endemic: MA, CO, and NY south to w. NC and e. TN. [= RAB, C, F, G, K, W, Z; = Grossularia rotundifolia (Michaux) Coville & Britton – S] * Ribes rubrum Linnaeus, Garden Red Currant. Mt (NC, VA), Pd (VA): persistent from cultivation and rarely escaped to adjacent fence-rows and disturbed areas; rare, native of Europe. [= K; > R. sativum Syme – C, F, G]

Ribes aureum Pursh var. villosum Augustin de Candolle, Buffalo Currant, of midwestern United States is cultivated in ne. United States, rarely as far south as our area; it may escape. It also is reported as occurring as a native species as far east as Montgomery County in nc. TN (Chester, Wofford, & Kral 1997). It will key to couplet 6, where trouble will be encountered. It has the hypanthium long-tubular, the flowers golden yellow, and fruits black (rarely yellow). [= K; = R. odoratum H. Wendland – C, F, G] {not keyed at this time} Ribes curvatum Small, Granite Gooseberry. Pd, Mt (GA): rocky upland forests; rare (GA Special Concern). Native from c. and nw. GA (Jones & Coile 1988) and e. TN, in the Cumberland Plateau (Chester, Wofford, & Kral 1997) westward. Also reported for NC by Sinnott (1985), and his report is backed up by specimens. The specimens, however, indicate that the species was cultivated in a botanist's garden; there is no evidence that R. curvatum is a native or naturalized component of NC's flora. It will key best to R. missouriense; it can be distinguished from all our species of gooseberries by its glandular-punctate leaves, and additionally from R. missouriense by its calyx lobes 7.5-9 mm long, sparsely hairy to villous (vs. 5-7.5 mm long, glabrous to sparsely pubescent). [= K; = Grossularia curvata (Small) Coville & Britton – S] {not keyed at this time} Ribes hirtellum Michaux, Northern Gooseberry, ranges south to WV (Tucker County), n. NJ, s. PA, and OH. It will key to couplet 2, where trouble will be encountered. It has glabrous ovaries and fruits, but the stamens are short and included. [= C, K; > R. hirtellum var. hirtellum – F, G] {not keyed at this time} * Ribes nigrum Linnaeus, Garden Black Currant or Cassis, native to Europe, is cultivated in ne. United States, rarely as far south as our area; it may escape. It will key to R. americanum, from which it can be distinguished by its pedicels 2-8 mm long, GROSSULARIACEAE 421 much longer than subtending ovate bracts (vs. pedicels 0-2 mm long, shorter than the subtending lanceolate bracts). [= C, F, G, K] {not keyed at this time} Ribes triste Pallas, Swamp Red Currant, ranges south to WV (Mineral, Pocahontas, and Randolph counties); it may occur in our area. It is very similar to R. rubrum, and will key there. It differs in the following ways: decumbent or straggling shrub (vs. erect), axis and pedicels of inflorescence often with stipitate glands (vs. not glandular). If found in our area, it should be in an obviously native habitat, likely in boggy forests or seepage wetlands at high elevations, and probably in VA. [= C, F, G, K] {not keyed at this time} * Ribes uva-crispa Linnaeus var. sativum Augustin de Candolle, Garden Gooseberry, native of Europe, is cultivated in ne. United States, rarely as far south as our area; it may escape. It will key best to R. cynosbati, but differs in the fruits being glandular-pubescent (vs. hispid), and the peduncles and pedicels being short (vs. peduncles 7-25 mm long, pedicels 5-16 mm long). [= K; > R. uva-crispi – C; = R. grossularia Linnaeus – F, G] {not keyed at this time}

HALORAGACEAE R. Brown 1814 (Water-milfoil Family)

A family of about 9 genera and 145 species, aquatic and wetland herbs, but also shrubs and trees, cosmopolitan but centered in the Southern Hemisphere. The family is sometimes spelled "Haloragidaceae."

1 Leaves whorled or alternate; stamens 4 or 8; carpels 4; emersed leaves bract-like and much-reduced (except in M. aquaticum) ...... Myriophyllum 1 Leaves alternate; stamens 3; carpels 3; emersed leaves foliaceous, little if at all reduced ...... Proserpinaca

Myriophyllum Linnaeus 1753 (Water-milfoil) (contributed by B.A. Sorrie and A.S. Weakley)

A genus of about 60 species, aquatic and wetland herbs, cosmopolitan, but centered in Australia. References: Crow & Hellquist (2000)=Z; Aiken (1981)=Y.

Identification notes: Stranded plants of M. heterophyllum and M. humile (and perhaps others) produce leaves that are reduced in size. Leaves and bracts become pectinate or pinnate, so that plants resemble M. pinnatum. Such plants are the source of nearly all inland records of M. pinnatum in the VA-NC-SC-GA area. M. heterophyllum usually flowers and fruits when stranded and may be distinguished from M. pinnatum by its much denser disposition of leaves and bracts, and by its dull red fruits obscurely tuberculate (vs. tan or pale brown fruits strongly tuberculate). From stranded M. humile, M. heterophyllum may be distinguished by leaves and bracts which are clearly whorled and much more densely disposed. M. humile differs from M. pinnatum by its wholly alternate leaves and bracts, and by its smooth fruits.

1 Leaves reduced to small scales or absent; stems short, erect from substrate...... M. tenellum 1 Leaves well-developed, pinnately divided with filiform segments; stems elongate, suspended in the water column and/or floating. 2 Flowers/fruits absent and emersed shoots with leaves closely similar in size and shape to submersed ones; widespread alien ...... M. aquaticum 2 Flowers/fruits present; emersed shoots present or not. 3 Flowers/fruits in axils of leaves. 4 Leaves whorled; emersed stems present and with feathery leaves ...... M. aquaticum 4 Leaves strictly alternate; flowers/fruits on submersed stems (forma capillaceum) or on emersed stems with pinnatifid or pectinate leaves (forma natans) ...... M. humile 3 Flowers/fruits in erect spikes emersed from water, f/f subtended by bracts much smaller than the normally submersed leaves. 5 Uppermost flowers/fruits alternate; leaves alternate or whorled or both. 6 Bracts much shorter than floral internodes, varying from pectinate to entire; fruit surface smooth or papillose ...... M. laxum 6 Bracts usually longer than floral internode, pinnatifid to pectinate; fruit surface strongly tuberculate ...... M. pinnatum 5 Uppermost flowers/fruits opposite; leaves whorled (technically pseudo-whorled in many M. heterophyllum) (note that early season plants of M. pinnatum may have flowers opposite, but at least some leaves will be alternate). 7 Bracts usually > 2× as long as pistillate flowers; stems drying brown, pale brown, or reddish. 8 Bracts throughout inflorescence pectinate to pinnatifid; winter buds scattered along stem, clavate, falling by early winter; [of DE and northward] ...... M. verticillatum 8 Distal bracts subentire to serrate, proximal bracts pectinate to serrate; winter buds developed only at base of stem or on rhizomes, usually persisting; widespread...... M. heterophyllum 7 Bracts usually < 2× as long as pistillate flowers; stems drying pale tan or whitish. HALORAGACEAE 422 9 Midstem leaves with 11 or fewer segments on each side of rachis; leaves rounded at apex; stem diameter more-or-less uniform; stem tips usually green; winter buds produced; [native, of DE and northward]...... M. sibiricum 9 Midstem leaves with 12 or more segments on each side of rachis; many leaves appear truncate or clipped at apex; stem diameter below inflorescence is up to 2× diameter of lower stem; stem tips usually reddish; no winter buds; [widespread alien]...... M. spicatum

Alternate key

1 Leaves reduced to small scales or absent; stems short, erect from substrate...... M. tenellum 1 Leaves well-developed, pinnately divided with filiform segments; stems elongate, suspended in the water column and/or floating. 2 Flowers/fruits produced in axils of submersed leaves...... M. humile 2 Flowers/fruits produced in axils of emersed leaves or on emersed shoots with bracts (reduced bracteal leaves). 3 Emersed shoots with feathery leaves about same size and shape as submersed leaves; flowers/fruits rarely produced; [widespread alien] ...... M. aquaticum 3 Emersed shoots with bracts subtending flowers/fruits; these bracts much different in shape than submersed leaves. [stranded plants may produce bracts and leaves of similar size and shape, but these not feathery]. 4 All flowers/fruits alternate; fruits smooth ...... M. humile 4 All flowers/fruits opposite or whorled (or the lower opposite and the upper alternate in M. pinnatum). 5 Bracts usually longer than the internodes. 6 Leaves whorled or pseudo-whorled; fruits with low bumps...... M. heterophyllum 6 Leaves strictly alternate; fruits strongly tuberculate...... M. pinnatum 5 Bracts usually shorter than the internodes. 7 All bracts pectinate to pinnatifid ...... M. verticillatum 7 Bracts vary from entire to pectinate. 8 Leaves alternate, pseudo-whorled, or both; plain green; [of se. VA and southward] ...M. laxum 8 All leaves whorled, grayish green; [collectively widespread] 9 Midstem leaves with 11 or fewer segments on each side of rachis; leaves rounded at apex; stem diameter more-or-less uniform; stem tips usually green; winter buds produced; [native, of DE and northward] ...... M. sibiricum 9 Midstem leaves with 12 or more segments on each side of rachis; many leaves appear truncate or clipped at apex; stem diameter below inflorescence is up to 2× the diameter of the lower stem; stem tips usually reddish; no winter buds; [widespread alien] ...... M. spicatum

* Myriophyllum aquaticum (Vell. Conc.) Verdc., Parrot-feather. Cp, Pd, Mt (GA, NC, SC, VA): ditches, slow-moving rivers, pools, ponds; common (rare in Piedmont and Mountains), native of South America. April-June. An introduced species now widespread in se. United States, north to NY, WV, and MO. [= C, GW, K, W, Y, Z; = M. brasiliense Cambessedes – RAB, F, G; = M. proserpinacoides Gillies ex Hooker & Arnott – S] Myriophyllum heterophyllum Michaux, Southern Water-milfoil. Cp (GA, NC, SC, VA), Pd, Mt (GA): ditches, slow- moving waters of rivers and streams, pools, ponds; common. April-July. NY west to Ontario and MN, south to FL and TX. [= RAB, C, F, G, GW, K, S, Y, Z] Myriophyllum humile (Rafinesque) Morong. Cp (VA): floating in an artificial pond; rare (VA Rare). [= C, F, G, K, Y, Z] Myriophyllum laxum Shuttleworth ex Chapman, Loose Water-milfoil. Cp (GA, NC, SC, VA): limesink depression ponds (dolines), spring-runs, rarely also in lakes; rare (GA Threatened, NC Threatened, SC Rare). June-October. Se. VA south to n. FL, s. AL, and s. MS (Sorrie & Leonard 1999). M. laxum and M. heterophyllum both have reddish submersed stems and present difficulties in identification when in sterile condition. M. laxum has a total of 7-15 (-17) segments per leaf, vs. (15-) 17-31 (-37) segments in M. heterophyllum. Documented for VA by a 1922 specimen from Princess Anne County at GH (Sorrie, pers. comm.). [= RAB, GW, K, S, Y] Myriophyllum pinnatum (Walter) Britton, Sterns, & Poggenburg, Alternate-leaved Water-milfoil. Cp (GA, NC, SC, VA), Mt? (GA, VA): pools, ditches; uncommon (VA Watch List). June-October. MA west to IA and SD, south to GA and TX. [= RAB, C, F, G, GW, K, S, W, Y, Z] * Myriophyllum spicatum Linnaeus, Eurasian Water-milfoil. Cp (GA, NC, SC, VA): ponds and impoundments; uncommon? {habitat and range in our area uncertain}, native of Eurasia, confused with M. sibiricum, (see below). An introduced species, now widespread in e. United States. Reported for South Carolina by Hill & Horn (1997). [= C, GW, K, W, Y, Z] Myriophyllum tenellum Bigelow, Leafless Water-milfoil. Cp (NC, VA): natural lakes (Carolina bay lakes), typically growing on the sandy bottoms in water 1-2 meters deep; rare (NC Rare, VA Rare). Newfoundland west to MN, south to PA and NJ, and disjunct south to a few occurrences in VA and NC. [= C, F, G, K, Y, Z]

Myriophyllum sibiricum Komarov. South to DE, and reported for VA. [= C, G, K, Z; > M. exalbescens Fernald – F, Y] Myriophyllum verticillatum Linnaeus. A circumboreal species, south in North America to DE, MD, IN, NE, TX, and CA. [= C, G, K, Y, Z; > M. verticillatum var. pectinatum Wallroth – F] HALORAGACEAE 423

Proserpinaca Linnaeus (Mermaid-weed)

A genus of 2-3 species, aquatic and wetland herbs, of e. North America and the West Indies. References: Catling (1998)=Z.

1 Bracteal (emersed) leaves serrate; submersed pectinate leaves with 8-14 pairs of divisions 5-30 mm long; fruits 2.3-6.0 mm wide. 2 Fruit 2.3-4.0 mm wide, acutely angled, not winged, the sides of the capsule more-or-less planar...... P. palustris var. crebra 2 Fruit (3.5-) 4.0-6.0 mm wide, sharply angled (to somewhat winged), the sides of the capsule concave...... P. palustris var. palustris 1 Bracteal (emersed) leaves pinnatifid to pectinate; submersed pectinate leaves with 4-12 pairs of divisions 2-7.5 mm long; fruits 2.0-3.6 mm wide. 3 Leaves with a flattened rachis 1-4 mm wide, the 7-12 pairs of divisions 2.0-3.5 mm long; fruits 2.3-3.6 mm wide ...... P. intermedia 3 Leaves with a filiform rachis (midrib) 0.2-1.0 mm wide, the 4-9 pairs of divisions 2.0-7.5 mm long; fruits 2.0-2.8 mm wide ...... P. pectinata

Proserpinaca intermedia Mackenzie, Intermediate Mermaid-weed. Cp (NC, SC, VA), Mt (VA); {GA}: wet places; rare (NC Watch List). July-September. Nova Scotia to SC on the Coastal Plain; disjunct in sc. TN. This taxon is intermediate in morphology between P. palustris and P. pectinata; whether it warrants species status is unclear. If merely a rarely produced first-generation hybrid, it should be treated as a hybrid binomial (P. ×intermedia); if it forms independent, self-reproducing populations, it should probably be treated as a species. [= RAB, C, F, G, K, Z; < P. palustris – GW] Proserpinaca palustris Linnaeus var. crebra Fernald & Griscom, Common Mermaid-weed. Cp (GA, NC, SC, VA): wet places, swamp forests; uncommon. June-October. Throughout e. North America and south to the Caribbean and Central America. [= C, F, G, K, Z; < P. palustris – RAB, S, W; < P. palustris – GW (also including P. intermedia)] Proserpinaca palustris Linnaeus var. palustris, Coastal Mermaid-weed. Cp (GA, NC, SC, VA), Pd (NC, SC), Mt? (GA?): wet places, swamp forests; common. June-October. MA (?) to FL and west to LA, on the Coastal Plain. [= C, F, G, K, Z; < P. palustris – RAB, S, W; P. palustris – GW (also including P. intermedia)] Proserpinaca pectinata Lamarck, Feathery Mermaid-weed. Cp (GA, NC, SC, VA), Pd (GA): bogs, savannas, ditches, other wet places; common. June-October. Nova Scotia south to s. FL and west to w. LA, mostly on the Coastal Plain, but scattered inland as well, as in c. TN. [= RAB, C, F, G, GW, K, S, Z]

Proserpinaca palustris Linnaeus var. amblyogona Fernald occurs east to KY, TN, and GA. [= C, F, G, K; < P. palustris – GW, S] {not keyed at this time}

HAMAMELIDACEAE R. Brown 1818 (Witch Hazel Family) (also see ALTINGIACEAE)

A family of ca. 27 genera and ca. 87 species, trees and shrubs, tropical to temperate, and especially e. Asian. References: Meyer in FNA (1997); Endress in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves 5-7-palmately lobed and palmately veined, glabrous...... [see Liquidambar in ALTINGIACEAE] 1 Leaves unlobed, pinnately veined, stellate-pubescent beneath (at least when young). 2 Petals 0; stamens 12-32; flowers numerous in dense globose or elongate spikes; leaves with a symmetric or asymmetric (oblique) base, the lateral veins marginal for a distance of at least 2-3 mm; [tribe Fothergilleae] ...... Fothergilla 2 Petals 4; stamens 4; flowers few in small clusters; leaves with a weakly to strongly asymmetric (oblique) base, the lateral veins included in the blade tissue or barely exposed for a distance of <1 mm; [tribe Hamamelideae] ...... Hamamelis

Fothergilla Murray in Linnaeus 1774 (Witch-alder)

A genus of 2 species, shrubs, of temperate e. North America. References: Meyer in FNA (1997); Weaver (1969)=Z; Endress in Kubitzki, Rohwer, & Bittrich (1993).

Identification notes: Fothergilla major often occurs with Hamamelis virginiana, with which it is easily confused in vegetative condition; a reliable character is the base of the lateral veins (marginal in Fothergilla, included in leaf tissue in Hamamelis).

1 Leaves stellate-pubescent above, up to 6 cm long and 5 cm wide (the largest < 5.2 cm wide); stamens 12-24; capsules 6.5- 10-5 (-13) mm long, the persistent hypanthium 3.4.5 mm long; seeds 4.8-6.3 mm long; [of wet savannas, pocosins, and pocosin margins of the Coastal Plain]...... F. gardenii HAMAMELIDACEAE 424 1 Leaves glabrous or sparsely stellate-pubescent above, up to 12 cm long and 10 cm wide (the largest > 5.2 cm wide); stamens (18-) 22-32; capsules 8-15.2 mm long, the persistent hypanthium 4-9.2 mm long; seeds 6.2-7.8 mm long; [of rocky habitats of the Mountains and Piedmont]...... F. major

Fothergilla gardenii Linnaeus, Coastal Witch-alder. Cp (GA, NC, SC): wet savannas, pocosins, and pocosin margins; uncommon (GA Threatened). March-May; September-October. Se. NC (and allegedly se. VA) south to panhandle FL and s. AL. An ornamental prized for its small size and attractive fall color. [= RAB, F, FNA, GW, K, Z; > F. gardeni – S, orthographic variant; > F. parvifolia Kearney – S] Fothergilla major (Sims) Loddiges, Large Witch-alder. Mt, Pd (GA, NC, SC): dry ridgetop forests of middle elevation ridges in the mountains, especially along the Blue Ridge Escarpment, summits and upper slopes of Piedmont monadnocks, north- facing bluffs in the lower Piedmont; rare (GA Special Concern, NC Rare, SC Rare). April-May; July-October. C. NC west to ne. TN, south to nc. GA and nc. AL; disjunct in AR. [= RAB, FNA, K, S, W, Z; > F. monticola Ashe]

Hamamelis Linnaeus 1753 (Witch-hazel)

A genus of 5-6 species, shrubs and small trees, of e. North America and e. Asia (China and Japan). The other North American species, H. vernalis Sargent, is endemic to the Ozark/Ouachita region of AR, OK, and MO. References: Leonard (2006)=X; Meyer in FNA (1997); Lane (2005)=Z; Jenne (1966)=Y; Wen & Shi (1999); Endress in Kubitzki, Rohwer, & Bittrich (1993).

1 Outer surface of calyx scarlet; petals 7-14 mm long, red or reddish (often yellow-tipped), flowering late December to early February; leaves 12-24 cm long, 5-17 cm wide, densely stellate-pubescent below, usually with 11 lateral veins (6 on one side of the leaf, 5 on the other); [plants of pineland ravines in s. MS]...... [H. ovalis] 1 Outer surface of calyx yellow; petals 6-8 mm long, yellow, flowering October-January; leaves 3.7-16.7 cm long, 2.5-13 cm wide, glabrous to moderately stellate-pubescent beneath, usually with 9 or 10 lateral veins; [plants widespread in our area] 2 Stellate trichomes of the leaves moderately dense to dense, averaging 0.09 mm across, with 7-11 rays; leaves (3.6-) avg. 6.4 (-10.3) cm long, (1.8-) avg. 4.1 (-6.2) cm wide; petals 7-15 mm long, 0.5-0.8 mm wide; [e. SC south to Panhandle FL, west to se. LA in the Coastal Plain] ...... H. virginiana var. henryae 2 Stellate trichomes of the leaves sparse to moderately dense, averaging 0.16-0.40 mm across, with 3-6 (-8) rays; leaves (4.7-) avg. 9.9 (-14.0) cm long, (3.9-) avg. 6.6 (-9.2) cm wide; petals 15-20 mm long, 1 mm wide; [widespread in our area] ...... H. virginiana var. virginiana

Hamamelis virginiana Linnaeus var. henryae Jenne ex C. Lane, Small-leaved Witch-hazel. Cp (GA, SC): sandhill margins, xeric hammocks, streamheads; uncommon. November-January. E. SC (Horry and Hampton counties), s. GA, and panhandle FL west to se. LA. Though cited in Lane (2005) as var. henryi, the honoree is collector Mary G. Henry; thus the honorific epithet should be corrected to the feminine form. Additional study is needed of these small-leaved Coastal Plain populations. [< H. virginiana – FNA, GW, K, S; = H. virginiana var. henryi Jenne ex C. Lane – Y, Z, orthographic error; = H. virginiana var. henryae] Hamamelis virginiana Linnaeus var. virginiana, Witch-hazel. Mt, Pd, Cp (GA, NC, SC, VA): forests; common. October- December; October-November (of the following year). Québec and Nova Scotia west to n. MI and MN, south to FL and TX. The bark is still gathered in large quantities in the Southern Appalachians, as the source for witch hazel liniment. The name "witch-hazel" alludes to its superficial resemblance to Corylus, the true hazel, and to its "perverse" habit of flowering in the fall, as it drops its leaves. [= Y, Z; < H. virginiana – RAB, C, FNA, G, GW, K, S, W; > H. virginiana var. parvifolia Nuttall – F; > H. virginiana var. virginiana – F]

Hamamelis ovalis S.W. Leonard. Apparently endemic to dry-mesic pineland ravines in sc. MS (Perry County) (Leonard 2006). Late December-early February. [= X]

HELIOTROPACEAE Schrader 1819 (Heliotrope Family)

A family of 5 genera and ca. 400 species, trees, shrubs, and herbs, semicosmopolitan.

Heliotropium Linnaeus (Heliotrope, Turnsole)

A genus of ca. 250 species, widespread in tropical and temperate regions. Probably better placed in the family Heliotropiaceae, as it is apparently more closely related to Hydrophyllaceae than to Boraginaceae. Currently under study and additional taxonomic changes may be forthcoming (Hilger & Diane 2003). References: Al-Shehbaz (1991)=Z; Hilger & Diane 2003).

1 Flowers solitary at the ends of short branches; [of limestone habitats from nw. GA westward]; [section Orthostachys, subsection Bracteata]...... H. tenellum 1 Flowers in secund, helicoid cymes. 2 Leaves glabrous, succulent, < 7 mm wide; [of saline coastal situations]; [section Halmyrophila] ...... HELIOTROPACEAE 425 ...... H. curassavicum var. curassavicum 2 Leaves pubescent, not succulent, > 10 mm wide; [of a variety of mostly disturbed, inland situations]. 3 Mericarps separating at maturity; fruit 4-lobed prior to maturation; leaves petiolate, ca. 2× as long as wide; [section Heliotropium] ...... H. europaeum 3 Mericarps cohering in pairs at maturity; fruit 2-lobed prior to maturation; leaves petiolate or sessile to subsessile, ca. 2-5× as long as wide. 4 Fruit tuberculate, 4-seeded; leaves sessile to subsessile, ca. 4-5× as long as wide; corolla throat and tube densely villous within; [section Heliophytum] ...... H. amplexicaule 4 Fruit longitudinally ribbed, 2-seeded; leaves petiolate, ca. 2× as long as wide; corolla throat and tube glabrous within; [section Tiaridium]...... H. indicum

* Heliotropium amplexicaule M. Vahl, Wild Heliotrope. Cp, Pd (GA, NC, SC, VA): disturbed areas, roadsides, fields; uncommon (rare north of SC), introduced from South America. April-September. [= RAB, C, F, G, K, Z] Heliotropium curassavicum Linnaeus var. curassavicum, Seaside Heliotrope. Cp (GA, NC, SC, VA): edges of brackish and salt marshes, estuarine shores; rare (NC Rare). June-September. Var. curassavicum ranges from DE (and farther north as an introduction) south to the New World tropics. Considered by some authors to be introduced and naturalized in our area. Other varieties occur inland in the mw. and w. United States. [= C, K, Z; < H. curassavicum – RAB, GW; = H. curassavicum – F, G; = Heliotropium curassavicum ssp. curassavicum] * Heliotropium europaeum Linnaeus, European Heliotrope. Cp (GA, NC, SC, VA), Pd (NC, VA): roadsides, disturbed areas; rare, introduced from s. Europe. June-September. [= RAB, C, F, G, K, Z] * Heliotropium indicum Linnaeus, Turnsole. Cp, Pd (GA, NC, SC, VA): roadsides, woodland borders, swamps, ditches; uncommon, introduced from South America. July-November. [= RAB, C, F, G, GW, K, Z; = Tiaridium indicum (Linnaeus) Lehm. – S] Heliotropium tenellum (Nuttall) Torrey, Delicate Heliotrope. Mt (GA): limestone glades and barrens; rare (GA Special Concern). WV, KY, IN, IL, IA, and KS south to nw. GA, AL, MS, LA, and TX. [= C, F, G, K, Z; = Lithococca tenella (Nuttall) Small – S]

HIPPOCASTANACEAE (Buckeye Family) (see SAPINDACEAE)

HYDRANGEACEAE Dumortier 1829 (Hydrangea Family)

A family of about 17 genera and 190-220 species, trees, shrubs, vines, and herbs, primarily north temperate. As here interpreted, the family Hydrangeaceae includes two well-marked groups, the Hydrangeae (including Decumaria and Hydrangea) and the Philadelpheae (including Deutzia and Philadelphus). This group has been shown by molecular research to be unrelated to the Saxifragaceae, and to have its closest affinities to the Loasaceae, Cornaceae, and Nyssaceae (Xiang et al. 2002; Soltis, Xiang, & Hufford 1995; Morgan & Soltis 1993). References: Spongberg (1972); Soltis, Xiang, & Hufford (1995); Morgan & Soltis (1994); Xiang et al. (2002); Hufford in Kubitzki (2004).

1 Woody vine, climbing by aerial rootlets; petals 7-10; [tribe Hydrangeae] ...... Decumaria 1 Shrub; petals 5 (rarely 10 in the cultivars of the alien Deutzia). 2 Pubescence of leaves and twigs stellate; stamens 10; [a cultivated alien, rarely escaped]; [tribe Philadelpheae] ...... Deutzia 2 Pubescence of leaves and twigs simple; stamens 8-10 (Hydrangea) or 25-90 (Philadelphus); [natives and aliens]. 3 Leaf blades 10-30 cm long; inflorescences of 25-many flowers; stamens 8-10; [tribe Hydrangeae]...... Hydrangea 3 Leaf blades 3-8 cm long; inflorescences of 1-7 flowers; stamens 25-90; [tribe Philadelpheae] ...... Philadelphus

Decumaria Linnaeus (Climbing Hydrangea, Woodvamp)

A genus of 2 species, vines, of e. North America and e. Asia (China). References: Hufford in Kubitzki (2004).

Identification notes: Decumaria is readily distinguished from the other opposite-leaved, woody vines in our flora (Gelsemium, Trachelospermum, Lonicera, Bignonia, Campsis, and Clematis) by its leaves (simple, ovate, and usually serrate) and climbing structures (adventitious roots).

Decumaria barbara Linnaeus, Climbing Hydrangea, Woodvamp. Cp (GA, NC, SC, VA), Pd (GA, SC), Mt (GA, NC): swamp forests and bottomlands in the Coastal Plain (and Piedmont of SC), moist forests in the mountains of n. GA, nw. SC, sw. NC; common (rare in Mountains). May-June; July-October. Se. VA south to FL and west to LA and e. TX (Singhurst, Keith, & Holmes 2005), inland to nw. SC, se. TN, and w. TN. This handsome vine climbs to the tops of trees via adventitious roots. The opposite leaves are somewhat fleshy in texture. [= RAB, C, F, G, GW, K, S, W]

HYDRANGEACEAE 426

Deutzia Thunberg (Deutzia)

A genus of about 60 species, shrubs, mainly Asian. References: Hufford in Kubitzki (2004).

* Deutzia scabra Thunberg, Deutzia, Pride-of-Rochester. Mt, Pd (NC, VA), Cp (VA): fairly commonly cultivated, persistent around old homesites and escaping to adjacent woodlands; rare, native of Japan and China. First reported for NC (Jackson Co., NC) by Pittillo & Brown (1988). D. crenata Siebold & Zuccarini, Chinese Deutzia, is reported as introduced in GA by Kartesz (1999); this may not be taxonomically distinct from D. scabra. [= C, F; > D. scabra – K; > D. crenata Siebold & Zuccarini – K; > D. scabra var. candisissima (Froebel) Rehder]

Hydrangea Linnaeus (Hydrangea, Sevenbark)

A genus of about 25 species, shrubs, of e. North America and e. Asia. Recent molecular analyses suggest that Hydrangea as usually interpreted is polyphyletic (Soltis, Xiang, & Hufford 1995); future taxonomic changes are to be expected. See Dirr (2004) and van Gelderen & van Gelderen (2004) for information on cultivated hydrangeas. References: Pilatowski (1982)=Z; McClintock (1957)=Y.

1 Leaves pinnately lobed, the lobes toothed; inflorescence a panicle; large sterile flowers many (> 20 per inflorescence), borne throughout the inflorescence...... H. quercifolia 1 Leaves unlobed, merely toothed; inflorescence a corymb (except H. paniculata); large sterile flowers absent to relatively few (0-15 per inflorescence), borne around the periphery of the corymb (except H. paniculata). 2 Inflorescence a panicle; large sterile flowers many (> 20 per inflorescence), borne throughout the inflorescence; [large shrub to small tree (to 8 m tall)]; [alien, cultivated and sometimes persistent] ...... H. paniculata 2 Inflorescence a corymb; large sterile flowers absent to relatively few (0-15 per inflorescence), borne around the periphery of the corymb; [small to medium shrub, to 3 m tall]; [native]. 3 Lower leaf surface glabrous or inconspicuously puberulent, appearing green; trichomes of the lower leaf surface restricted to the midrib and major veins; sterile flowers absent, or, if present, usually < 1 cm in diameter ...... H. arborescens 3 Lower leaf surface variously pubescent, appearing white or gray; trichomes of the lower leaf surface on veins and interveinal areas; sterile flowers usually present, large and showy, usually greater than 1 cm in diameter. 4 Lower leaf surface velutinous, pilose, or tomentose, appearing gray; trichomes usually not dense enough to entirely mask the green leaf surface; trichomes with prominent tubercles (as seen at 40× magnification); sterile flowers generally very few (0-3 per inflorescence) ...... H. cinerea 4 Lower leaf surface densely floccose-velutinous, felt-like, appearing bright white or silver; trichomes dense enough to entirely mask the green leaf surface; trichomes without tubercles, or with small and inconspicuous tubercles (as seen at 40× magnification); sterile flowers generally fairly many (2-15 per inflorescence)...... H. radiata

Hydrangea arborescens Linnaeus, Smooth Hydrangea. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): forests, especially around rock outcrops and along streambanks; common (rare in Coastal Plain). May-July. NJ, s. NY, OH, IN, IL, MO, and se. KS south to e. NC, c. SC, c. GA, panhandle FL, s. AL, LA, and OK. [= K, S, W, Z; = H. arborescens ssp. arborescens – RAB, Y; = H. arborescens var. arborescens – C, G; > H. arborescens var. arborescens – F; > H. arborescens var. oblonga Torrey & A. Gray – F] Hydrangea cinerea Small, Ashy Hydrangea. Mt, Pd (GA, NC, SC): rocky forests and rock outcrops, roadbanks, perhaps strictly or mostly associated with mafic or calcareous rocks; uncommon. May-July. Sw. NC, c. IN, c. IL, and c. MO south to n. SC, sc. AL, and c. AR. [= K, S, W, Z; = H. arborescens ssp. discolor (Seringe) McClintock – RAB, Y; = H. arborescens var. discolor Seringe – C, G; = H. arborescens var. deamii E. St. John – F] * Hydrangea paniculata Siebold, Panicle Hydrangea. Mt (NC, VA): persistent after cultivation at old home-sites, sometimes appearing naturalized; rare, native of e. Asia. July-August. [= C, F, G, K] Hydrangea quercifolia Bartram, Oakleaf Hydrangea. Pd, Mt, Cp (GA, *NC, SC): native in rich foothill forests (very rare), also in disturbed areas, thickets, or forests adjacent to urban or suburban areas; rare, introduced from further south. May-July. C. and sw. TN and nw. SC, south through w. GA, AL, and MS to panhandle FL and e. LA; scattered elsewhere as a remnant or escpae from cultivation. Boufford & Wood (1977) describe an apparently native occurrence in nw. SC. This southeastern native is a spectacular garden plant, frequently planted, rarely escaping or persisting. [= C, F, G, K, S] Hydrangea radiata Walter, Snowy Hydrangea, Silverleaf. Mt (GA, NC, SC), Pd (NC, SC): rocky forests and rock outcrops, often common and conspicuous on roadbanks; common. May-July. A Southern Appalachian endemic: sw. NC (in the valley of the French Broad River and to its southwest), nw. SC, ne. GA, and se. TN, with outliers (perhaps escaped from cultivation?) in Stokes County, NC and Calhoun County, SC. This attractive species is especially typical of the escarpment gorge region near the tricorner of NC, SC, and GA, in the vicinity of the towns of Highlands, Cashiers, and Rosman, NC, where it is conspicuous along roadbanks. [= K, S, W, Z; = H. arborescens ssp. radiata (Walter) McClintock – RAB, Y]

HYDRANGEACEAE 427 Philadelphus Linnaeus (Mock-orange)

A genus of 65 (or fewer) species, shrubs, of north temperate areas. The most recent monographer of the genus, Hu (1954-1955) recognizes many species and varieties on the basis of minor differences in pubescence. Many of the recognized taxa are based only on cultivated material. The native distributions of the varieties have little phytogeographic coherence, and several varieties are often reported from the same site, suggesting that they reflect merely variation within a population (if genetically based at all). For instance, Hu recognizes three varieties in Ph. hirsutus and five in Ph. inodorus, but these seem to be no more than forms. As Hu writes, "the formerly recognized species, Ph. grandiflorus Willd., and Ph. laxus Schrad., are merely different forms of a species with heterogeneous leaf shape, size, and margins. Fostered by growers, propagated and distributed through cuttings, these forms have maintained their distinction in gardens since their discoveries. But when they are projected on the spectrum of variations exhibited by a large number of specimens collected from the homeland of Ph. inodorus Linn. they appear to be nothing but a few transitional forms. In this paper, these forms are treated as varieties." Hu's "varieties" should be treated as forms or cultivars, if recognized at all. I have taken a conservative approach, though variation in several of our native species could use additional study. References: Hu (1954-1955)=Z; A.E. Weakley (2002).

1 Axillary buds exposed; twigs of the current year villous-hirsute; seeds not caudate; [subgenus Deutzioides, section Deutzioides] ...... Ph. hirsutus 1 Axillary buds enclosed; twigs of the current year glabrous; seeds with caudate tails about as long as the embryo; [subgenus Philadelphus]. 2 Flowers 1-3 (-9) in a cymule; stamens 60-90; [subgenus Philadelphus, section Pauciflorus] ...... Ph. inodorus 2 Flowers 5-9 in a determinate raceme; stamens 20-50; [subgenus Philadelphus, section Philadelphus]. 3 Bark of the current year brown, exfoliating in its second year; flowers fragrant...... Ph. coronarius 3 Bark of the current year gray, not exfoliating later; flowers not fragrant or only slightly so ...... Ph. pubescens

* Philadelphus coronarius Linnaeus, European Mock-orange. Pd (NC), {GA, SC, VA}: cultivated (though moreso in the past than now), and sometimes escaped or persisting around old homesites; rare, native of Europe. May-July. Ph. coronarius is the most commonly cultivated Philadelphus in our area, though it is currently considered rather old-fashioned. [= C] Philadelphus hirsutus Nuttall, Hairy Mock-orange, Cumberland Mock-orange. Mt (GA, NC, SC, VA), Pd (SC): bluffs, rock outcrops, rocky woodlands, often with seepage, over mafic or calcareous rocks; rare (NC Watch List, SC Rare, VA Watch List). April-May; June-August. A Southern Appalachian species: sw. VA and KY south and west to w. NC, TN, n. GA, and n. AL. Ph. sharpianus Hu, known from e. TN and nc. AR, is similar to Ph. hirsutus, allegedly differing in the hypanthium glabrous (vs. more or less pubescent), the leaves strigose-pilose above, glabrous or sparsely strigose or with the nerves only villous beneath (vs. scabrous-hirsute above, uniformly villous beneath); it is probably best considered only a form of Ph. hirsutus. Ph. hirsutus is cultivated and it may escape outside of the range stated. [= RAB, C, F, G, S, W; > Ph. hirsutus – K, Z; > Ph. sharpianus Hu – K, Z; > Ph. hirsutus var. intermedius Hu – Z; > Ph. hirsutus var. nanus Hu – Z; > Ph. sharpianus Hu var. parviflorus Hu – Z] Philadelphus inodorus Linnaeus, Appalachian Mock-orange. Mt, Pd, Cp (GA, NC, SC, VA): rich forests and woodlands, rocky bluffs over mafic or calcareous rocks, and also cultivated and persistent; uncommon (rare in Coastal Plain). April-May; June-August. VA and TN south to panhandle FL, GA, and s. AL (and according to C, also in e. PA). Ph. floridus Beadle, known from nw. GA, is similar to Ph. inodorus, allegedly differing in the pedicels and hypanthium pubescent (vs. glabrous); it is probably only a form of Ph. inodorus. [= RAB, C, G, W; > Ph. inodorus var. inodorus – F, S, Z; > Ph. inodorus var. carolinus Hu – Z; > Ph. inodorus var. grandiflorus (Willdenow) A. Gray – F, Z; > Ph. inodorus var. laxus (Schrader) Hu – Z; > Ph. inodorus var. strigosus Beadle – S, Z; > Ph. grandiflorus Willdenow – S; > Ph. gloriosus Beadle – S; > Ph. inodorus – K; > Ph. floridus Beadle – K, S, Z] Philadelphus pubescens Loiseleur, Ozark Mock-orange, Hairy Mock-orange. Mt (GA): limestone bluffs; rare (GA Special Concern). E. TN, KY, nw. GA (Jones & Coile 1988), AL, s. IL, MO, OK, and AR, west of the Blue Ridge. It has been documented from TN counties adjacent to both VA and NC, and is likely to be found in VA, at least. [> Ph. intectus Beadle – S; > Ph. latifolius Schrader ex Augustin de Candolle – S; > Ph. intectus var. intectus – Z; > Ph. intectus var. pubigerus Hu – Z; > Ph. pubescens var. verrucosus (Schrader) Hu – Z; > Ph. pubescens var. pubescens – K, Z; > Ph. pubescens var. intectus (Beadle) A.H. Moore – K]

HYDRASTIDACEAE Martinov 1820 (Golden-seal Family)

A family of 2 genera and 2 species, perennial herbs, of temperate e. North America and Japan. In chemistry, morphology, and anatomy, Hydrastis shows some relationship to Podophyllum and Diphylleia of the Podophyllaceae (often included in the Berberidaceae). Though usually placed in the Ranunculaceae, Tobe & Keating (1985) present evidence from morphology, anatomy, embryology, palynology, chemistry, and cytology that suggests that Hydrastis is best recognized as a monotypic family. They contend that "Hydrastis represents a relictual primitive group which very early diverged from a common ancestral stock of the Ranunculaceae, Berberidaceae and probably of Circaeasteraceae, and that Hydrastis has evolved in its own evolutionary line parallel with other lines leading to the modern representatives of these families." In recent papers on classification of the flowering plants, Thorne (1992) and Reveal (1993) have also accepted Hydrastidaceae as a distinct family. Tobe in Kubitzki & Bayer places Hydrastis with Glaucidium Siebold & Zuccarini in a bigeneric Hydrastidaceae. References: Tamura in Kubitzki, Rohwer, & Bittrich (1993); Tobe in Kubitzki & Bayer (2002). HYDRASTIDACEAE 428

Hydrastis Linnaeus 1759 (Golden-seal)

A monotypic genus, an herb, endemic to e. North America. References: Ford in FNA (1997); Tamura in Kubitzki, Rohwer, & Bittrich (1993).

Hydrastis canadensis Linnaeus, Golden-seal. Mt (GA, NC, SC, VA), Pd (GA, NC, VA): mesic, very nutrient-rich forests, with circumneutral soils, over calcareous or mafic rocks such as limestone, amphibolite, and dolostone, sometimes forming large colonies after canopy disturbance such as logging; rare (GA Endangered, NC Endangered, VA Watch List). April; May-June. VT and MN south to w. and c. NC, n. GA, TN, and AR. Exploited for the herbal trade (and still often used as a home remedy in more remote parts of the mountains, though too rare in our area to support any economically significant wild collection. The root is bitter in taste and contains several alkaloids. Reported for SC (P. McMillan, pers.comm. 2002). [= RAB, C, F, FNA, G, K, S, W)

HYDROLEACEAE Berchtold & J. Presl 1820 (Hydrolea Family)

The Hydroleaceae is not closely related to Hydrophyllaceae; recent molecular data confirm the prevailing view through most of the 19th century that Hydrolea was in its own family. References: Ferguson (1998); Hilger & Diane (2003); Angiosperm Phylogeny Group (1998, 2003).

Hydrolea Linnaeus

A genus of about 11 species, aquatic and wetland herbs, of tropical and subtropical regions.

1 Flowers in axillary cymes; leaves 3-14 cm long, 1.5-4 cm wide; axillary spines present in the axils of some leaves; corolla 7- 8 mm long. 2 Calyx and stem sparsely pubescent with spreading hairs 2-3 mm long...... H. quadrivalvis 2 Calyx and stem glabrous, or minutely puberulent or with sessile glands...... [H. uniflora] 1 Flowers in terminal cymes or corymbs; leaves 2-6 cm long, 0.6-2.5 cm wide; axillary spines present or absent; corolla 10-15 mm long. 3 Leaves elliptic to lanceolate, 2-3 cm long, 0.6-1.0 cm wide; axillary spines absent or rudimentary...... H. corymbosa 3 Leaves ovate to ovate-lanceolate, 3-6 cm long, 1.5-2.5 cm wide; axillary spines preset, well-developed, to 1.5 cm long ...... H. ovata

Hydrolea corymbosa J. Macbride ex Elliott, Skyflower. Cp (GA, SC): pond cypress savannas, depression meadows; rare. Ne. SC south to sw. GA and FL. See Nelson (1993). The author is sometimes stated as J.F. Macbride, but this is an error. [= RAB, GW, K; = Nama corymbosum (Macbride ex Elliott) Kuntze – S] Hydrolea ovata Nuttall ex Choisy, Ovate False-fiddleleaf. Cp (GA): swamps, ponds, ditches; rare. June-August. C. GA west to TX, north in the interior to sc. TN and MO. [= C, F, G, GW, K; = Nama ovatum (Nuttall ex Choisy) Britton – S] Hydrolea quadrivalvis Walter, Waterpod. Cp (GA, NC, SC, VA), Pd (NC, VA): swamp forests, backwater sloughs, marshes, ditches; common (VA Watch List). June-September. Se. VA south to FL, west to LA. [= RAB, C, F, G, GW, K; = Nama quadrivalve (Walter) Kuntze – S]

Hydrolea uniflora Rafinesque. Swamp forests, sloughs, marshes. June-September. Mainly in the Mississippi River Alluvial Plain, west to e. TX and east to AL, TN, and KY. [= C, F, G, GW, K; = Nama affine (A. Gray) Kuntze – S; = Hydrolea affinis A. Gray]

HYDROPHYLLACEAE R. Brown 1817 (Waterleaf Family) (also see HYDROLEACEAE)

A family of about 18 genera and 270 species, herbs and shrubs, nearly cosmopolitan, concentrated in w. North America. References: Wilson (1960a); Constance (1963).

1 Leaves entire, simple; styles 2, distinct to the summit of the ovary. 2 Flowers in axillary cymes; capsule subglobose; leaves 2-12 cm long...... [see Hydrolea in HYDROLEACEAE] 2 Flowers solitary or paired in the leaf axils; capsule cylindrical leaves 0.8-1.5 (-3.5) cm long...... Nama 1 Leaves dissected, lobed, or toothed (sometimes the basalmost leaves simple); style fused for a portion of its length, 2-cleft toward the tip; ovary with 1 locule. 3 Flowers solitary opposite the leaves on the upper portion of the stem (sometimes also terminal in a lax, (1-) 2-6- flowered cyme). HYDROPHYLLACEAE 429 4 Leaves opposite below, alternate above; petals 5-8 mm long; leaves elliptical in outline, pinnatifid into 7-13 lanceolate segments; calyx lobes to 10 mm long in fruit; capsule 4-seeded...... Ellisia 4 Leaves all alternate; petals 2-4 mm long; leaves broadly triangular in outline, divided into 3-5 obovate segments; calyx lobes 1-3 mm long; capsule 1-2 (-3)-seeded...... Nemophila 3 Flowers all terminal in 3-many-flowered cymes. 5 Inflorescence repeatedly branched subdichotomously; larger leaf blades > 8 cm wide; stamens well exserted from the corolla (3 mm or more beyond the corolla); plants perennial from fibrous roots ...... Hydrophyllum 5 Inflorescence with a strong central axis (some secondary branching in Ph. bipinnatifida, but not as above); larger leaf blades < 5 cm wide (except Ph. bipinnatifida); stamens slightly exserted from the corolla (< 3 mm beyond the corolla) (except well-exserted in Ph. bipinnatifida, included in Ph. covillei); plants annual (biennial in Ph. bipinnatifida) from a taproot...... Phacelia

Ellisia Linnaeus (Waterpod)

Ellisia is considered to be a monotypic genus, an herb of c. and e. North America, but generic limits in the Hydrophyllaceae are badly in need of critical reassessment. References: Constance (1940)=Z.

Ellisia nyctelea (Linnaeus) Linnaeus, Waterpod, Aunt Lucy. Pd, Mt (VA): moist shaded forests, especially bottomlands; uncommon. May-June. IN and MI west to Alberta, south to AR and OK; disjunct in e. North America from s. NY and NJ south to sc. VA. Likely to occur in nc. NC. [= C, F, G, GW, K, W, Z; = Nyctelea nyctelea (Linnaeus) Britton – S]

Hydrolea (see HYDROLEACEAE)

Hydrophyllum Linnaeus (Waterleaf)

A genus of 8 species, herbs, of e. and w. North America. References: Constance (1942)=Z; Beckmann (1979)=Y; Alexander (1941)=X.

1 Principal cauline leaves palmately lobed, maple-like, differing from the pinnately divided basal leaves. 2 Sepals with small reflexed appendages exserted at each sinus of the calyx; plant biennial from a taproot; stamens exserted 1-3 mm beyond the corolla; leaf lobing relatively shallow and irregular...... [H. appendiculatum] 2 Sepals with appendages absent or rudimentary; plant perennial from fibrous roots; stamens exserted 3-6 mm beyond the corolla; leaf lobing relatively deep and regular...... H. canadense 1 Principal cauline leaves pinnately divided, similar to the basal leaves. 3 Inflorescence and upper stem densely hirsute with spreading hairs 1-2 mm long; leaves pinnatifid with (7-) 9-13 segments, these toothed but not lobed...... H. macrophyllum 3 Inflorescence and upper stem glabrate to strigose with appressed to ascending hairs < 0.5 mm long; leaves with 5-7 (-9) segments, some of them sometimes deeply 2-lobed. 4 Corolla deep purple to maroon; lower stem glabrous to slightly (rarely moderately) pubescent with retrorse hairs; [of low to high elevations in the Mountains]...... H. virginianum var. atranthum 4 Corolla white to lavender or pale purple; lower stem slightly to densely pubescent with retrorse to spreading hairs; [of low elevations of the Piedmont, Mountains, and Coastal Plain]...... H. virginianum var. virginianum

Hydrophyllum canadense Linnaeus, Mapleleaf Waterleaf, Canada Waterleaf, Broadleaf Waterleaf. Mt (GA, NC, SC, VA), Pd (NC, VA), Cp (VA): cove forests, rocky streambanks, other moist and nutrient-rich forests; common (rare in Coastal Plain and in Piedmont south of VA) (SC Rare). May-June; August. VT and s. Ontario west to MI and WI, south to n. GA, AL, AR, and MO. [= RAB, C, F, G, K, S, W, Y, Z] Hydrophyllum macrophyllum Nuttall, Hairy Waterleaf. Mt (GA, NC, VA): cove forests and other moist rocky forests, especially over calcareous or mafic rocks; rare (GA Special Concern, NC Rare, VA Watch List). May; July. WV west to OH, and IL, south to sw. VA, w. NC, n. GA, and n. AL; reports from AR are erroneous, and are based on material of Hydrophyllum brownei Kral & Bates (Peck 2003). The w. North American H. occidentale (S. Watson) A. Gray is rather closely related. [= RAB, C, F, G, K, S, W, Y, Z] Hydrophyllum virginianum Linnaeus var. atranthum (E.J. Alexander) Constance, Appalachian Waterleaf. Mt (NC, VA): cove forests and other moist rocky forests; common. May-June; July-August. N. WV south through w. and sw. VA and e. KY to w. NC and e. TN. Since its naming as a species (Alexander 1941) and subsequent reduction to a variety (Constance 1942) there has been little consensus about this taxon, some regarding it as merely a color form. Alexander lists numerous characters additional to that of flower color; they need further investigation. "H. atranthum differs from H. virginianum in the dark-violet flowers, the brown hairs on the appendages, brown filaments, corolla-lobes longer than the tube, stamens shorter [11.5 mm] and more slender, and the more numerous leaf-lobes. H. virginianum has flowers white to pale lavender or pinkish, white hairs on the appendages, white filaments, corolla-lobes and tube equal in length, filaments longer (13.5 mm) and stouter, and 5-7 leaf- HYDROPHYLLACEAE 430 segments." Beckmann (1979) did not accept the variety, stating that "this pigment combination appears sporadically in other sectors of the species range." Based on herbarium material I have seen darker than usual flowers are found outside of the Southern Appalachians; they do not, however, approach in darkness the flowers of Southern Appalachian material and the somewhat darker-flowered plants outside the Southern Appalachians do not share the stem pubescence character stated in the key above. The general correlation of flower color and lower stem pubescence and the tight geographical range of var. atranthum incline me to accept it provisionally as a variety. It is not, however, limited to high elevations, as stated or implied by some authors. The two varieties provisionally accepted here need more careful study, including either statistical studies of morphology, or electrophoretic or molecular studies. [= C, F, G, Z; < H. virginianum – RAB, K, W, Y; < H. virginicum – S, orthographic error; = H. atranthum E.J. Alexander – X] Hydrophyllum virginianum Linnaeus var. virginianum, Eastern Waterleaf, Virginia Waterleaf. Mt, Pd (NC, VA), Cp (VA): cove forests, moist rocky forests, alluvial forests; common. April-June; July-August. NH and Québec west to ND, south to e. VA, c. NC, KY, s. IN, s. IL, nc. AR, and ne. OK. As discussed by Beckmann (1979) and Constance (1941), H. virginianum is a closely related vicariant of H. tenuipes Heller of British Columbia south to CA. See Phacelia bipinnatifida for additional suggestions on distinguishing it from this species. [= C, F, G, Z; < H. virginianum – RAB, K, W, Y; < H. virginicum – S, orthographic error; = H. virginianum – X]

Hydrophyllum appendiculatum Michaux, Biennial Waterleaf. S. Ontario and MN, south to sw. PA, WV, e. TN, n. AL (Jackson Co.), MO, and e. KS. It was attributed to NC by Small (1933) on unknown grounds. [= C, F, G, K, Y, Z; = Decemium appendiculatum (Michaux) Small – S]

Nama Linnaeus (Fiddleleaf) (also see Hydrolea in HYDROLEACEAE)

A genus of about 45 species, herbs, of sw. North America, tropical America, and Hawaii.

* Nama jamaicense Linnaeus, Jamaica Weed. Cp (SC): lawns; rare, introduced from tropical America (including FL and TX). May. [= K; = N. jamaicensis – RAB, orthographic variant; = Marilaunidium jamaicense (Linnaeus) Kuntze – S]

Nemophila Nuttall (Baby Blue-eyes)

A genus of 11 species, herbs, of North America (mostly w. North America). References: Constance (1941).

Identification notes: Nemophila is superficially similar to Phacelia covillei and Ph. ranunculacea, with which it often co- occurs. They can be distinguished with the following key.

1 Flowers solitary, on pedicels opposite the leaves, the pedicels mostly > 12 mm long; corolla white (sometimes aging to pale lavender), 2.5-3.5 mm long; fruits ovoid, longer than thick, turning purple at maturity, exceeding the 2-4 mm long calyx lobes...... Nemophila aphylla 1 Flowers borne in 2-6-flowered terminal cymes, the pedicels mostly < 12 mm long; corolla pale blue or lavender, 4-5 mm long; fruits depressed globular and weakly 4-lobed, the apex depressed, remaining green at maturity, shorter than the calyx, the lobes of which expand to 5-8 mm long. 2 Pubescence of the lower stem consisting of appressed to ascending, stiff, pointed hairs; [of rivers in the Atlantic drainage] ...... Phacelia covillei 2 Pubescence of the lower stem spreading or even retrorse, most of the hairs weak and twisted, and many of them glandular-tipped; [of rivers in the Mississippi drainage]...... [Phacelia ranunculacea]

Nemophila aphylla (Linnaeus) Brummitt. Cp, Pd (GA, NC, SC, VA): moist, nutrient-rich floodplain forests; uncommon, though often locally abundant. March-April. MD south to panhandle FL and west to TX, north in the interior to e. TN, w. KY, and se. MO. [= GW, K; = N. microcalyx (Nuttall) Fischer & Meyer – RAB, F, G, S; = N. triloba (Rafinesque) Thieret – C]

Phacelia A.L. de Jussieu 1789 (Phacelia)

A genus of about 150 species, of North America and South America, concentrated in w. North America. References: Constance (1949)=Z; Levy (1991)=Y; Murdy (1966); Gillett (1968, 1964).

Identification notes: 1. Phacelia bipinnatifida and Hydrophyllum virginianum are sometimes confused. Ph. bipinnatifida has the larger and more basal leaves distinctly bipinnatifid, the lower pinnae often stalked (vs. pinnatifid, the basal or terminal pinnae sometimes 2-lobed, all the pinnae more-or-less sessile), pubescence of the upper stem and inflorescence in part glandular (pubescence nonglandular), and seeds 4 per capsule, black (vs. 2 per capsule, light brown). 2. Phacelia covillei and Ph. ranunculacea are superficially similar to and sometimes confused with Nemophylla aphylla, which see for discussion.

HYDROPHYLLACEAE 431 1 Corolla lobes fimbriate; seeds 4 per capsule. 2 Corolla white (rarely slightly lavender); pubescence of the stem spreading; lobes of cauline leaves mostly obtuse; seeds 3.0-3.5 mm long...... Ph. fimbriata 2 Corolla lavender to blue; pubescence of the stem appressed; lobes of cauline leaves mostly acute; seeds 1.5-3.0 mm long ...... Ph. purshii 1 Corolla lobes entire; seeds 4-15 per capsule. 3 Stamens 1.5-2 mm long; style 1.5-2 mm long; corolla tubular; seeds globose-ovoid, nearly spherical, 4 per capsule. 4 Pubescence of the lower stem consisting of appressed to ascending, stiff, pointed hairs; [of rivers in the Atlantic drainage] ...... Ph. covillei 4 Pubescence of the stem spreading or even retrorse, most of the hairs weak and twisted, and many of them glandular-tipped; [of rivers in the Mississippi drainage]...... [Ph. ranunculacea] 3 Stamens 3-10 mm long; style 3-15 mm long; corolla rotate to broadly campanulate; seeds ovoid-angled, 4-15 per capsule. 5 Corolla 10-15 mm across, blue; plant 10-60 cm tall; seeds 2.5-4 mm long, black; ultimate segments of the leaf 15- 45 mm long, 10-25 mm wide; pedicels recurved in fruit; [of moist forests of the Mountains and (very rarely) Piedmont]...... Ph. bipinnatifida 5 Corolla 5-11 mm across, white to blue; plant 5-40 cm tall; seeds 1.5-2.2 mm long, brown; ultimate segments of the leaf 5-15 mm long, 5-9 mm wide; pedicels ascending to spreading in fruit; [of alluvial forests, granitic flatrocks, and other habitats, of the Piedmont, Coastal Plain, and Mountains]. 6 Sepals 4-8 mm long, linear or oblanceolate; marginal bristles of sepals spreading, 1.0-1.5 mm long; plants mostly erect...... Ph. maculata 6 Sepals 2-4 mm long, narrowly ovate; marginal bristles of sepals appressed, 0.3-1.0 mm long; plants mostly decumbent, branched from the base. 7 Sepals 2.6-4.0 mm long; petals 4-6 mm long; marginal bristles of sepals 0.6-0.9 mm long; basal leaves with 1-3 pairs of lateral leaflets. the terminal leaflet larger and usually 3-lobed; cauline leaves with 1-3 pairs of rather broad lobes; [of various habitats (including granitic flatrocks and domes) in SC, NC, and VA] ...... Ph. dubia var. dubia 7 Sepals 2.0-3.0 mm long; petals 3.5-5 mm long; marginal bristles of sepals 0.4-0.7 mm long; basal leaves with 4-5 pairs of lateral leaflets, the terminal leaflet about the same size and unlobed; cauline leaves with 2-4 pairs of narrow lobes; [of granitic flatrocks and domes of the Piedmont of SC and southwestward] ...... Ph. dubia var. georgiana

Phacelia bipinnatifida Michaux, Fernleaf Phacelia, Forest Phacelia. Mt (GA, NC, SC, VA): cove forests, especially where rocky; common (SC Rare, VA Watch List). April-May; June. W. VA west to s. OH, n. IN, n. IL, and c. MO, south to w. NC, nw. SC, n. GA, c. AL, and n. AR. Ph. bipinnatifida var. plummeri (= Ph. brevistyla) is "based on a variation with sparser pubescence, larger and less divided leaf segments, smaller flowers, and sub-included stamens and style. These variations are not concomitant, and the distribution of forms showing a complete or partial combination of them is sporadic" (Constance 1949). The matter deserves additional study. [= RAB, C, G, K, W, Z; > Ph. bipinnatifida var. bipinnatifida – F; > Ph. bipinnatifida var. plummeri Wood – F; > Ph. brevistyla Buckley – S; > Ph. bipinnatifida – S] Phacelia covillei S. Watson ex A. Gray, Eastern Buttercup Phacelia. Pd (NC, VA): rich soils of floodplains, and contiguous terraces and slopes; rare (NC Rare, VA Rare). April; May. Ranging in two disjunct areas – c. NC and sc. VA (in the drainages of the Cape Fear, Tar, and Roanoke rivers) and DC, n. VA, and sc. MD (in the drainage of the Potomac River). Most recent authors have included this taxon within the closely similar Ph. ranunculacea; as thus broadly defined, Ph. ranunculacea was considered to occur in 3 peculiarly disjunct areas; one centered around St. Louis, MO (w. KY, w. TN, e. MO, ne. AR, se. MO, s. IL, and s. IN), one near Washington, DC (DC, n. VA, and sc. MD), and a third in c. NC and sc. VA. Chuang & Constance (1977) reported that the western population center has a chromosome number of n=6, and that the two eastern population centers have n=14. The disparate cytotypes of eastern and western Ph. ranunculacea sensu lato led Chuang & Constance to seek morphological differences that would warrant the recognition of separate taxa, but they reported that "no consistent morphological distinction has been found between the two cytotypes." Stem pubescence does, however, show consistent, though subtle, differences between eastern and western populations of this complex. Eastern material has the stem pubescence relatively sparse, consisting of appressed to ascending, stiff, pointed hairs. Western populations have stem pubescence relatively dense, much of it spreading or even retrorse, most of the hairs weak and twisted, and many of them glandular-tipped. Given the disparate cytotypes, correlated with allopatric distribution and slight but consistent morphological differences, it seems best to provisionally recognize two taxa; further study, using chemical and molecular techniques would be valuable. Recognition at the species level is nomenclaturally the more conservative (and here followed) because of the pre- existence of Watson's binomial; varietal status might be the more appropriate. Ph. covillei and Ph. ranunculacea (sensu stricto) have numerous characteristics that render their inclusion in Phacelia uncomfortable (see discussion in Chuang & Constance 1977, Constance 1949, Gillett 1968). See Nemophila aphylla for suggestions on distinguishing these two superficially similar species. [= K; < Ph. ranunculacea (Nuttall) Constance – RAB, C, F, G, Z] Phacelia dubia (Linnaeus) Trelease var. dubia, Appalachian Phacelia. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): floodplain forests, rocky forests, fields, roadsides, granitic flatrocks; common. April-May; June. Var. dubia ranges from NY and PA west to WV, south to nc. SC, sw. NC, and se. TN. The Phacelia dubia complex has been under detailed biosystematic study by Foster Levy and associates (Levy 1991a, 199b, 1997; Levy et al. 1996; Levy & Malone 2001; Levy & Neal 2001; Taylor & Levy 2002; del Castillo 1994, 1998). Male sterile cytotypic variants are common in some populations but formal taxonomic HYDROPHYLLACEAE 432 recognition ius not warranted (Levy 1991a, 1991b; del Castillo 1994, 1998). Additionally, an incipient variety, informally termed "imitator", occurs in c. SC (Levy 1991a; Levy & Malone 2001). These populations are morphologically variable, some more similar to var. georgiana, others more similar to var. dubia; see Levy (1991a) for further discussion. They may warrant taxonomic recognition, as they are allopatric from each of the 3 named varieties, and show degrees of sterility when bred with each of the three, but morphologic differences have not evolved (Levy & Malone 2001). [= K, Y; < Ph. dubia – RAB, C, F, S, W; > Ph. dubia var. dubia – G; > Ph. dubia var. fallax (Fernald) Gleason – G; > Ph. dubia var. dubia – Z (also including var. interior)] Phacelia dubia (Linnaeus) Trelease var. georgiana McVaugh, Georgia Phacelia. Pd (GA): granitic flatrocks; rare. April- May; June. Var. georgiana ranges from GA west to ec. AL, in the Piedmont. It has sometimes been attributed to SC, and Levy found plants in SC which morphologically resemble var. georgiana, but he concluded that this "imitator" genotype was largely sterile when bred with var. georgiana. See var. dubia for additional discussion. [= K, Y, Z; < Ph. dubia – RAB, C, F, S, W; ? Ph. dubia var. dubia – G] Phacelia fimbriata Michaux, Fringed Phacelia, Blue Ridge Phacelia. Mt (GA, NC, VA): moist forests on slopes and floodplains, at low to high elevations, perhaps mainly over circumneutral soils; uncommon, but locally abundant (GA Special Concern, VA Rare). April-May. Sw. VA south to w. NC, e. TN, and n. GA (Jones & Coile 1988), a Southern Appalachian endemic. [= RAB, C, F, G, K, S, W, Z] Phacelia maculata Wood, Flatrock Phacelia. Pd (GA, NC, SC): bottomlands, granitic flatrocks; uncommon (NC Rare List). April; May. Sc. NC south to GA and west to ec. AL. [= RAB, K, W, Y, Z; ? Ph. hirsuta – S, misapplied] Phacelia purshii Buckley, Miami-mist. Mt (GA, NC, SC, VA): moist forests on floodplains and slopes; rare (GA Special Concern, VA Watch List). May-June. S. PA west to s. Ontario, OH and MO, south to nw. SC, nw. GA, and c. AL. Plants "with smaller flowers, shorter pedicels, and smaller capsules and seeds" are the basis of Ph. boykinii and Ph. bicknellii (Constance 1949). A study of the matter was initiated and specimens annotated as "Ph. purshii ssp. boykinii," but the research was not completed and the name was never published; further study is warranted. [= RAB, C, F, G, K, W, Z; > Ph. purshii – S; > Ph. boykinii (A. Gray) Small – S; > Ph. bicknellii Small – S]

Phacelia dubia (Linnaeus) Trelease var. interior Fernald, endemic in c. TN. [= K; < Ph. dubia – C, F, G, S; < Ph. dubia var. dubia – Z] {not keyed at this time} Phacelia ranunculacea (Nuttall) Constance, Western Buttercup Phacelia. In the Mississippi and Ohio river drainages, centered around St. Louis, MO (w. KY, w. TN, e. MO, ne. AR, se. MO, s. IL, and s. IN). [= K; < Ph. ranunculacea – RAB, C, F, G, Z] Phacelia strictiflora (Engelmann & Gray) Gray var. robbinsii Constance. East to AL. [= K, Z] {not keyed at this time}

HYPERICACEAE A.L. de Jussieu 1789 (St. John's-wort Family)

It appears from molecular analysis that recognition of the Hypericaceae may (after all) be warranted. Hypericum is in a clade with Podostemum and Bonnetia, sister to a clade including Clusiaceae s.s. (Savolainen et al. 2000). References: Adams (1973)=Z; Godfrey (1988)=Y; Wood & Adams (1976).

1 Petals yellow; stamens fascicled or not, if fascicled then not into 3 fascicles of 3 stamens each; staminodia (hypogynous glands) lacking; perianth 4-5-merous; [tribe Hypericeae] ...... Hypericum 1 Petals pale pink; stamens fascicled, in 3 fascicles of 3 stamens each; staminodia (hypogynous glands) present, alternating with the fascicles of stamens; perianth 5-merous; [tribe Cratoxyleae]...... Triadenum

Hypericum Linnaeus 1753 (St. John's-wort)

A genus of about 370 species, trees, shrubs, and herbs, primarily temperate. Hypericum in our area is a large, complex, and interesting genus, with a number of unresolved questions remaining. The species treated in Key B have often been treated in the segregate genus Ascyrum. Evidence from a variety of disciplines now suggests that they should be included in Hypericum (Adams & Robson 1961; Calie, Schilling, & Webb 1983; Robson 1996). {The curious bimodal distribution of H. prolificum (more than one taxon?). Basis of Small's H. lobocarpum in Blue Ridge of NC. And more....} References: Adams (1973)=Z; Godfrey (1988)=Y; Robson (1977, 1981, 1990, 1996, 2001, 2002, 2006)=X; Adams (1962)=V; Adams (1957); Webb (1980); Robson & Adams (1968); Adams & Robson (1961); Calie, Schilling, & Webb (1983); Culwell (1970). Key based in part on Adams (1973), Godfrey (1988), C, and GW.

1 Leaves with an articulation at the very base, this appearing as a narrow line, groove, or abrupt change of color and texture which extends across the petiole; shrub; [section Myriandra]. 2 Leaves needle-like, 0.5-1.5 (-2) mm wide, the margins essentially parallel (H. galioides keyed here and below); [subsection Centrosperma] ...... Key A1 or A2 2 Leaves, at least the largest on the plant, not needle-like, wider than 2 mm, the margins not parallel, the widest point often beyond the middle. 3 Petals 4; sepals 4 (rarely 2); plant 5-100 cm tall; leaves 2-40 mm long; [subsection Ascyrum]...... Key B 3 Petals 5; sepals 5; plant 50-250 cm tall; leaves (10-) 20-70 mm long; [subsections Centrosperma and Brathydium] HYPERICACEAE 433

...... Key C 1 Leaves without an articulation at the very base, the petiole merging gradually into the stem with no break, groove, or abrupt change in color or texture; decumbent shrub (H. buckleyi) or an annual or perennial herb (suffruticose at the base in H. cistifolium, H. dolabriforme, H. nudiflorum, H. apocynifolium, H. sphaerocarpum). 5 Leaves ascending or appressed, 1-nerved, < 1 mm wide; inflorescence a compound raceme; [section Brathys] .....Key D 5 Leaves spreading or ascending, generally multi-nerved, > 1 mm wide; inflorescence a dichasial cyme. 6 Capsule 3 (-4) locular; stamens connate at the base into 3 or 5 fascicles; leaves with black glandular dots as well as translucent glandular dots when backlit (except in H. perforatum); sepals and/or petals marked with black glandular dots or lines; [section Hypericum]...... Key E 6 Capsule 1-locular; stamens separate or connate at the base, but not grouped into fascicles; leaves with translucent glandular dots, without black glandular dots (when backlit); sepals and petals with translucent glandular lines or dots only, not marked with black glandular dots or lines. 7 Decumbent shrub or suffruticose herb; [section Myriandra, subsections Pseudobrathydium and Suturosperma]...... Key F 7 Herb; [section Trigynobrathys, and section Myriandra subsection Suturosperma]...... Key G

Key A1 – shrubby St. John's-worts with needle-like leaves and flowers with 5 petals and 5 sepals [section Myriandra, subsection Centrosperma]

1 Longest leaves 5-16 mm. 2 Capsules 6-9 mm long; longest leaves 5-10 (-11) mm; flowers 13-15 mm diameter; [of spodosol pine flatwoods]...... H. tenuifolium 2 Capsules 3-4.5 mm long; longest leaves 7-16 mm; flowers 13-17 diameter; [of alfisols and ultisols of wet pine savannas, seepage bogs]. 3 Flowers 16-17 mm diameter; longest leaves 11-16 mm; plants 2.5-6.5 dm tall; [of wet pine savannas in se. NC to ne. SC] ...... H. species 2 3 Flowers 13-14 (-15?) mm diameter; longest leaves 7-13 mm; plants 0.5-1.0 dm (rarely to 1.5); [of seepage bogs and wet flatwoods of s. GA to s. FL to w. LA] ...... H. brachyphyllum 1 Longest leaves 13-30 mm. 4 Plant a low shrub <5 dm tall, more-or-less decumbent, forming dense clumps; inflorescence elongate (flowers at up to 5 nodes); flowers 10-12 mm diameter; [of dry to mesic soils of lower piedmont and inner coastal plain of sc. VA-NC-SC- GA-AL; disjunct to rock outcrops of s GA]...... H. lloydii 4 Plant an erect shrub 0.5-4 m tall, with single main stem branched above; inflorescence elongate (3-7 nodes) or short (1- 3 nodes in H. fasciculatum and H. chapmanii); flowers 13-26 mm diameter; [of wet soils of the Coastal Plain]. 5 Undersurface of most leaves easily seen on both sides of midrib, veins obvious on undersurface, leaves narrowly oblanceolate to oblinear, 1.5-5 (-7) mm wide; inflorescence elongate (3-7 nodes)...... H. galioides 5 Undersurface usually not seen except for midrib (leaf margins nearly touch midrib its whole length), if undersurface visible then no veins visible, leaves linear, needle-like, 0.5-1.5 mm wide; inflorescence elongate or short. 6 Plant short, <1 m tall; stem <1 cm wide at base; plant unbranched or few-branched, wand-like with narrow crown; [endemic to FL panhandle]...... [H. exile] 6 Plant tall, normally >0.8 m; stem 1-several cm wide at base; crown broader with many ascending to spreading branches. 7 Young branches, leaves, and sepals strongly glaucous; bark of upper stem and branches silvery gray and smooth; mature plant 2-4 m tall with ascending branches imparting tree-like or vase-like aspect; [restricted to shores of sinkhole ponds in Bay and Washington Counties, FL] ...... [H. lissophloeus] 7 Young branches, leaves, and sepals not glaucous; bark of upper stem and branches not silvery gray and smooth (except some chapmanii); mature plants variously shaped. 8 Inflorescence elongate (3-7 nodes); stem bark tight, thin, not exfoliating or exfoliating in narrow strips, not revealing buff or pale cinnamon color; if leaf undersurface is exposed it is distinctly paler than upper surface; [usually associated with flowing water (blackwater streams and impoundments)] ...... H. nitidum 8 Inflorescence short (1-3 nodes); stem bark corky-thickened to spongy, exfoliating in broad strips or sheets revealing buff or pale cinnamon color; if leaf undersurface is exposed it is about the same color as upper surface; [usually associated with static water (Carolina bays, impoundments, beaver ponds, borrow pits, flatwoods depressions, cypress-gum ponds)]. 9 Mature plant 2-3 (-4) m tall; branches ascending and imparting a tree-like or vase-like aspect (younger plants may be bushy); youngest internodes terete; [of flatwoods depressions and cypress-gum ponds and stringers of FL panhandle only] ...... [H. chapmanii] 9 Mature plant 0.8-1.5 (-2) m tall; branches spreading and imparting a bushy or gumdrop aspect; youngest internodes with distinct winged ridge on either side; [of Carolina bays, impoundments, beaver ponds, borrow pits, widespread]...... H. fasciculatum

HYPERICACEAE 434

Key A2 – shrubby St. John's-worts with needle-like leaves and flowers with 5 petals and 5 sepals [section Myriandra, subsection Centrosperma]

1 Plants < 0.6 m tall, erect, decumbent, or matted and with ascending/erect branches. 2 Longest leaves 5-13 mm; flowers 13-15 mm diameter. 3 Mature capsule 6-9 mm long; longest leaves 5-10 (-14) mm; [plants of spodosol flatwoods] ...... H. tenuifolium 3 Mature capsule 3-6 mm long; longest leaves 6-13 mm; [of seepage bogs and wet flatwoods of s GA-s FL-w LA]...... H. brachyphyllum 2 Longest leaves 11-25 mm; flowers 9-17 mm diameter. 4 Plant unbranched or few-branched, wand-like with narrow crown; restricted to FL panhandle flatwoods...... [H. exile] 4 Plant densely branched, bushy with broad crown; occurrring only north of FL. 5 Flowers 10-12 mm diameter; longest leaves 13-25 mm; [plants of dry to mesic soils of lower piedmont and inner coastal plain of se VA-NC-SC-GA-AL, disjunct to rock outcrops of s GA]...... H. lloydii 5 Flowers 16-17 mm diameter; longest leaves 11-16 mm; [plants of wet pine savannas in se NC-ne SC] ...... H. species 2 1 Plants > 0.6 m tall, plants erect. 6 Longest leaves 6-13 mm, linear, needle-like, permanently tightly revolute with usually only the midrib showing on underside; flowers 13-15 mm diameter; [s GA-s FL-w LA]...... H. brachyphyllum 6 Longest leaves (12-)15-30 mm, linear and needle-like or narrowly oblanceolate to oblinear (H. galioides), permanently tightly revolute with usually only the midrib showing on underside OR margins revolute during drying but leaving considerable exposed undersurface (H. galioides); flowers 13-26 mm diameter. 7 Leaves narrowly oblanceolate to oblinear, 1.5-5 (-7) mm wide, margins revolute during drying but leaving considerable exposed undersurface, veins obvious on undersurface; flowers 13-15 mm diameter in elongate inflorescences of 3-7 nodes...... H. galioides 7 Leaves linear, needle-like, 0.5-1.5 mm wide, permanently tightly revolute with usually only the midrib showing on underside, if undersurface visible there are no veins; flowers 14-26 mm diameter, inflorescences elongate or short. 8 Plant short, < 1 m tall; stem < 1 cm wide at base; plant unbranched or few-branched, wand-like with narrow crown; [restricted to FL panhandle] ...... [H. exile] 8 Plant tall, normally > 0.8 m; stem 1-several cm wide at base; crown broader with many ascending to spreading branches. 9 Plant to 1.5 m tall, with broad bushy or gumdrop aspect; plants widespread. 10 Inflorescence short (1-3 nodes); bark corky-thickened, exfoliating in sheets and wide strips exposing buff or pale cinnamon color; if leaf undersurface is exposed it is about the same color as upper surface; [usually associated with static water of Carolina bays, impoundments, beaver ponds, borrow pits] ...... H. fasciculatum 10 Inflorescence elongate (3-7 nodes); bark tight, thin (not corky thickened), not exfoliating in sheets or wide strips; if leaf undersurface is exposed it is distinctly paler than upper surface; [associated with moving water of blackwater streams and impoundments]...... H. nitidum 9 Plant 2-4 m tall, with tree-like or vase-like aspect; restricted to FL panhandle. 11 Young branches, leaves, and sepals not glaucous; bark of upper stem and branches normally not silvery gray, variously roughened; inflorescence short (1-3 nodes); [flatwoods depressions and cypress-gum ponds of FL panhandle]...... [H. chapmanii] 11 Young branches, leaves, and sepals strongly glaucous; bark of upper stem and branches silvery gray and smooth; inflorescence elongate (3-7 nodes); [restricted to shores of sinkhole ponds in Bay and Washington Counties, FL]...... [H. lissophloeus]

Key B – shrubby St. John's-worts with 4 petals and 4 (rarely 2) sepals [section Myriandra, subsection Ascyrum]

1 Styles and carpels 3 (rarely 4); leaves (5-) 7-20 mm wide, rounded, subcordate, or cordate-clasping at the base; plant an erect shrub. 2 Leaves rounded or subcordate at the base; [widespread in our area]...... H. crux-andreae 2 Leaves cordate-clasping at the base; [of e. GA southward] ...... H. tetrapetalum 1 Styles and carpels 2 (3 in H. microsepalum); leaves 1-7 mm wide, mostly cuneate (or if rounded the leaves < 8 mm long and 3 mm wide); erect or decumbent shrub. 3 Sepals nearly equal in size; styles 3; [s. GA south to n. FL] ...... H. microsepalum 3 Sepals markedly unequal, one opposite pair large and enclosing the capsule; styles 2; [collectively widespread]. 4 Pedicels 6-13 mm long, soon reflexed; subtending bractlets located near the last pair of leaves; decumbent shrub, to 2 dm tall ...... H. suffruticosum 4 Pedicels 1-5 mm long, erect; subtending bractlets located midway between the base of the flower and the last pair of leaves; erect or decumbent shrub, mostly 1-15 dm tall. HYPERICACEAE 435 5 Erect shrub, usually with a single stem, freely branched well above ground level (or from ground level if injured, as by fire, but then the multiple branches still erect rather than decumbent), to 1 m or more tall; leaves usually variable in size and shape, widest near the middle...... H. hypericoides 5 Decumbent, matted shrub, with several prostrate stems arising from a primary rootstock near ground level, each with numerous erect branchlets, rarely over 3 dm tall; leaves usually relatively uniform in size and shape, widest above the middle...... H. stragulum

Key C – shrubby St. John's-worts with broader leaves (mostly lanceolate or oblanceolate) and flowers with 5 petals and 5 sepals

1 Leaves cordate-clasping at the base, ovate; [of s. SC southward]; [section Myriandra, subsection Brathydium]...... H. myrtifolium 1 Leaves cuneate at the base, oblanceolate, oblong, elliptic, or narrowly elliptic; [collectively widespread]; [section Myriandra, subsection Centrosperma]. 2 Leaves mostly narrowly oblanceolate, the larger 2-3 cm long, 2-5 (-7) mm wide, mostly 5-10× as long as wide; seeds 0.4-0.8 mm long, dark brown...... H. galioides 2 Leaves mostly oblong, elliptic, narrowly elliptic, or broadly oblanceolate, the larger (2-) 3-7 cm long, 5-15 mm wide, mostly 2.5-5× as long as wide; seeds 0.8-1.3 mm long, amber to medium brown. 3 Flowers solitary, terminal (or in 3-flowered terminal cymes); petals 10-20 mm long; sepals 7-15 mm long; shrubs to 1 m tall ...... H. frondosum 3 Flowers (1-) 3-many in terminal cymes; petals 5-10 mm long; sepals 1.5-8 mm long; shrubs to 3 m tall. 4 Flowers (1-) 3-7 per inflorescence; capsules (6-) 7-14 mm long; larger leaves (4-) 7-14 mm wide...... H. prolificum 4 Flowers 7-many per inflorescence; capsules (3-) 4.5-6 mm long; larger leaves 1-7 (-11) mm wide. 5 Leaves (1.8-) 2.8-8.3 (-11) mm wide, the widest on a plant always over 4 mm wide; [plants widespread] ...... H. densiflorum 5 Leaves 1.0-3.7 (-4.1) mm wide; [plants of the Ridge and Valley of nw. GA, c. and nw. AL, and e. TN]...... H. interior

Key D – herbaceous St. John's-worts with leaves ascending or appressed, 1-nerved, < 1 mm wide and with a diffuse, racemose inflorescence

1 Leaves linear-subulate, (5-) 8-20 mm long; capsules 1-1.75× as long as the sepals; seeds coarsely rugose-areolate ...... H. drummondii 1 Leaves scale-like, 1-5 mm long; capsules ca. 2-3× as long as the sepals; seeds minutely and inconspicuously reticulate ...... H. gentianoides

Key E – herbaceous St. John's-worts with broad leaves, 3 (-4) locular capsules, stamens connate at base into 3 or 5 fascicles, leaves with black dots as well as translucent glands (except in H. perforatum), and sepals and/or petals marked with black dots or lines

1 Smaller stems strongly wing-angled; seeds 1.0-1.3 mm long; leaves of the main stem (8-) 11-20 (-26) mm long, those of the lateral branches typically much smaller; leaves punctate primarily with translucent glands; [alien, usually in disturbed habitats]; [section Hypericum]...... H. perforatum 1 Smaller stems not wing-angled; seeds 0.6-1.1 mm long; leaves of the main stem (11-) 21-48 (-64) mm long, those of the lateral branches nearly to quite as large; leaves punctate with black glands; [native, in a variety of habitats]; [section Graveolentia]. 2 Petals (3.0-) 4.3-12.2 (-14.0) mm long; sepals 1.5-6 mm long, conspicuously punctate with black glands (sometimes also black-lined); capsules (2.5-) 3.0-5.4 (-6.0) mm long; [collectively widespread, occurring in the Coastal Plain, Piedmont, and Mountains of NC, SC, and VA]. 3 Sepals 3-6 mm long; styles (2.5-) 5.4-7.4 (-9.0) mm long; petals (6.0-) 9.2-12.2 (-14.0) mm long; leaf apices acute ...... H. pseudomaculatum 3 Sepals 1.5-4.0 mm long; styles (1.0-) 1.4-2.4 (-3.0) mm long; petals (3.0-) 4.3-5.9 (-9.0) mm long; leaf apices obtuse to slightly retuse...... H. punctatum 2 Petals 6-18 mm long; sepals 4-10 mm long, with or without black lines (sometimes also black-punctate); capsules (3.0- ) 4.0-7.7 (-10.0) mm long; [endemic to moderate to high elevations of w. NC, sw. VA, and e. TN]. HYPERICACEAE 436 4 Styles (3.0-) 5.6-10.0 (-12.0) mm long; sepals without black lines; petals (5.0-) 11.5-16.1 (-18.0) mm long, without black lines and with round black glands only along the petal margin; longest stamens (8.0-) 10.7-16.3 (- 22.0) mm long; cymes relatively few-flowered, (2-) 5-14 (-22) flowers per plant...... H. graveolens 4 Styles (1.5-) 1.9-2.9 (-5.0) mm long; sepals with black lines; petals (6.0-) 7.0-9.2 (-11.0) mm long, with black lines and round black glands scattered over the surface of the petal; longest stamens (4.0-) 6.1-8.5 (-10.0) mm long; cymes relatively many-flowered, (5-) 13-61 (124) flowers per plant...... H. mitchellianum

Key F – shrubby and subshrubby St. John's-worts

1 Plant a matted, decumbent shrub, 0.5-3 (rarely to 5) dm tall; leaves 1.5-2.5× as long as wide, without axillary fascicles of leaves; flowers solitary or in small simple cymes; [endemic to rock outcrops at moderate to high elevations in the Mountains of sw. NC, nw. SC, and ne. GA]; [section Myriandra, subsection Pseudobrathydium]...... H. buckleyi 1 Plant an erect suffrutescent herb, 1.5-10 dm tall; leaves 1.5-5× as long as wide, with or without axillary fascicles of leaves; flowers in compound cymes; [mostly of the Coastal Plain and Piedmont, very rarely in the Mountains and then at low elevations]; [section Myriandra, subsection Suturosperma]. 2 Larger leaves 4-10 mm wide, 3-5× as long as wide; axillary leaf fascicles present in main leaf axils; seeds pale brown, faintly reticulate, 0.4-0.5 mm long...... H. cistifolium 2 Larger leaves 10-30 mm wide, 1.5-3× as long as wide; axillary leaf fascicles absent; seeds dark brown, strongly reticulate, 1.5-2 mm long. 3 Flowers in simple 3-flowered cymes or in compound cymes with up to 8 flowers; sepals 3 mm long, oblong, obtuse apically; capsules ovoid, 8-10 mm long (excluding the styles) and 5-7 mm broad; seeds 1.8-2.0 mm long, cylindric, sometimes slightly falcate, dull brown when mature ...... H. apocynifolium 3 Flowers usually in many-flowered cymes terminating branches; sepals 1.5-2.0 mm long, usually triangular-acute; capsules ovoid to subglobose, 4-5 mm long (excluding the styles) and 4-5 mm broad; seeds 1.5-1.8 mm long, usually falcate-cylindric, dark purplish-brown and lustrous when mature...... H. nudiflorum

Key G – herbaceous St. John's-worts with broad leaves, 1-locular capsules, stamens separate or connate at base, but not grouped into fascicles, leaves with translucent dots, without black dots, sepals and petals with translucent lines or dots only, not marked with black dots or lines

1 Stems and leaves pubescent; [section Trigynobrathys]...... H. setosum 1 Stems and leaves glabrous. 2 Styles united, persistent as a single straight beak on the capsule; [section Myriandra, subsection Suturosperma]. 3 Leaves 3-6 cm long, 4-6× as long as wide, the margins revolute; [plants (in our area) of low elevations in the Coastal Plain]...... H. adpressum 3 Leaves 1-3 (-4) cm long, 2-3× as long as wide, the margins not revolute; [plants (in our area) of high elevations in the Mountains] ...... H. ellipticum 2 Styles separate, more or less divergent, not persistent as a beak on the capsule; [section Trigynobrathys]. 4 Styles 2-4 mm long; stamens 50-80. 5 Punctate glands absent on the stem (rarely very few on the internodes of the inflorescence); punctate glands of the leaves small, round, distributed on the lower leaf surface, becoming sparse toward the base of the leaf and toward the midrib; midstem leaves mostly broadest at or beyond the middle ...... H. virgatum 5 Punctate glands frequent on the stem; punctate glands of the leaves and stem large, oval, distributed evenly and densely on the lower leaf surface, also dense on the upper leaf surface in H. denticulatum and H. harperi (absent on upper leaf surface in H. species 1); midstem leaves usually broadest at or below the middle. 6 Upper surface of the leaf with no punctate glands; inflorescence branches typically with 3-12 pairs of bracteal leaves about 1/2 as large as the foliage leaves; [of shallow soil mats on granitic domes in the Piedmont of NC] ...... H. species 1 6 Upper surface of the leaf with abundant punctate glands; inflorescence branches with at most a few pairs of very small bracts; [of Coastal Plain wetlands, very rarely disjunct inland and then in wetlands]. 7 Leaves 5-20 (-24) mm long, 5-15 mm wide, 1.5-3× as long as wide, ovate to obovate to narrowly elliptic, mostly appressed to the stem, mostly shorter than the internodes; sepals 4.0-8.0 mm long, 2.0-4.0 mm wide, acute; lower stem not spongy-thickened with aerenchymatous tissue; [of moist pinelands of the Coastal Plain, very rarely disjunct inland to bog habitats in the Piedmont and Mountains] ...... H. denticulatum 7 Leaves 10-35 (-40) mm long, 3-8 (-12) mm wide, 3-10× as long as wide, lanceolate to linear- lanceolate, mostly ascending to spreading, often equalling the internodes; sepals 3.0-5.0 mm long, 0.8-2.5 mm wide, acute to acuminate; lanceolate to linear-lanceolate; upper portion of stem with numerous axillary branches; lower stem usually spongy-thickened with aerenchymatous tissue; [of upland depression ponds of the Coastal Plain, growing where seasonally inundated] ...... H. harperi 4 Styles 0.5-1.5 mm long; stamens 5-22. HYPERICACEAE 437 8 Leaves lanceolate to linear, 6-30 mm long, 0.5-3 mm wide, the leaf base attenuate to cuneate...... H. canadense 8 Leaves ovate to elliptic, 3-35 mm long, 2-15 mm wide, the leaf base rounded to cordate-clasping. 9 Sepals broadest near the base; inflorescence with few or no normally sized leaves, these only low in the inflorescence, giving the inflorescence a naked appearance; [of Coastal Plain pinelands]...... H. gymnanthum 9 Sepals broadest near the middle; inflorescence with many normally sized leaves and leaflike bracts, giving the inflorescence a leafy appearance; [collectively widespread]. 10 Ultimate bracts of the inflorescence elliptic, much like the leaves; leaves not paler beneath; sepals obtuse, much shorter than the capsule; capsule 3-5 mm long...... H. boreale 10 Ultimate bracts of the inflorescence linear, differing conspicuously from the leaves; leaves paler beneath; sepals acute, about equalling the capsule; capsule 2-3.5 mm long. 11 Inflorescence branches from the upper 1-6 nodes of the stem, the further branching repeatedly monochasial; stem with apical internode well developed, usually longer than the internode below; sepals broader above the middle, more-or-less imbricate; [of the Coastal Plain]...... H. mutilum var. latisepalum 11 Inflorescence branches from the upper 2-10 nodes of the stem, the further branching mostly dichasial; stem with apical internode shorter than the the internode below or even essentially absent; sepals broader below the middle, not imbricate (rarely broader above the middle and imbricate); [widespread]...... H. mutilum var. mutilum

Hypericum adpressum Rafinesque ex Barton, Bog St.-John's-wort, Creeping St.-John's-wort. Cp (GA, NC, SC, VA): boggy depressions; rare (US Species of Concern, GA Special Concern, NC Rare, SC Rare, VA Rare). July-August. E. MA south to sw. GA in the Coastal Plain; disjunct inland in WV, IN, IL, and sc. TN. See discussion on its habitats and rarity in Sorrie (1998b). [= RAB, C, F, G, GW, K, S, V, X, Z] Hypericum apocynifolium Small. Cp (GA): mesic bluffs and ravines, ridges and natural levees in floodplains; rare. C. and s. GA west to se. AR and e. TX. [= S, V, X, Y; < H. nudiflorum – GW, K, Z] Hypericum boreale (Britton) Bicknell, Dwarf St.-John's-wort, Northern St.-John's-wort. Cp (VA), Mt (NC?, VA): sinkhole ponds in the Mountains, interdune ponds in the outer Coastal Plain, boggy places; rare (NC Watch List, VA Watch List). Newfoundland and Québec west to w. Ontario, south to VA, NC (?), OH, IN, and n. IL. Hybrids with H. canadense have been called H. ×dissimulatum Bicknell (pro sp.). [= C, F, G, K; = H. mutilum Linnaeus ssp. boreale (Britton) J.M. Gillett – X] Hypericum brachyphyllum (Spach) Steudel. Cp (GA, SC?): ponds and wet pinelands; common. E. GA (near SC) south to s. FL, west to s. MS. Also reported from SC (Kartesz 1999); needs confirmation. [= GW, K, V, X, Y, Z; < H. aspalathoides – S] Hypericum buckleyi M.A. Curtis, Granite Dome St.-John's-wort. Mt (GA, NC, SC): thin soil in seasonal seepage around rock outcrops, particularly granitic exfoliation domes; rare (GA Special Concern, NC Watch List, SC Rare). June-August. Sw. NC south to nw. SC and ne. GA, a Southern Appalachian endemic. Wilbur (1995) showed that Curtis's spelling of the epithet, "buckleii", should be maintained; however, changes in the International Code of Botanical Nomenclature have reversed this (Robson 1996). [= RAB, GW, S, W, V, X, Z; = H. buckleii – K, orthographic variant] Hypericum canadense Linnaeus, Canada St. John's-wort Cp (GA, NC, SC, VA), Mt (NC, SC, VA), Pd (NC, VA); bogs, pine savannas, ditches; common (rare in Piedmont and VA Mountains). July-September. Newfoundland and Québec west to MN, south to s. GA, n. FL, and MS; also in Holland and Ireland, where considered by some to be native. Hybrids with H. mutilum and/or H. boreale have been called H. ×dissimulatum Bicknell (pro sp.). [= RAB, C, G, GW, K, S, W, X, Z; > H. canadense var. canadense – F; > H. canadense var. galiiforme Fernald – F] Hypericum cistifolium Lamarck. Cp (GA, NC, SC): pine savannas, wet pine flatwoods; common. June-August. E. NC south to s. FL, west to e. TX. [= RAB, GW, K, V, X, Y, Z; > H. cistifolium – S, in a narrower sense; > H. opacum Torrey & A. Gray – S] Hypericum crux-andreae (Linnaeus) Crantz, St. Andrew's Cross, St. Peter's-wort. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): dry forests and woodlands, pine flatwoods; common (rare in Piedmont and Mountains). June-October. NY (Long Island) and NJ south to s. FL, west to e. TX, primarily on the Coastal Plain, but scattered inland to w. NC and n. GA, also north in the interior to c. TN, s. KY, c. AR, and se. OK. [= GW, K, W, X, Y; = H. stans (Michaux ex Willdenow) P. Adams & Robson – RAB, C, V, Z; = Ascyrum stans Michaux ex Willdenow – F, G; > Ascyrum stans – S; > Ascyrum cuneifolium Chapman – S] Hypericum densiflorum Pursh, Mountain Bushy St. John's-wort. Mt (GA, NC, VA), Pd (NC, VA), Cp (GA, NC, SC): bogs, streambanks, dry to moist forests, rock outcrops, moist forests, pine savannas; common (rare in the VA Piedmont). June- August. Sw. PA south to n. GA and c. AL in and near the Mountains; NJ south to SC in the Coastal Plain; s. GA west to TX in the Coastal Plain. The related H. lobocarpum Gattinger is more western, extending east to TN; the basis for attribution of H. lobocarpum to "Blue Ridge, N.C." by Small (1933) is unknown. [< H. densiflorum – RAB, C, GW, K, W, X, Z (also see H. interior); < H. densiflorum var. densiflorum – F, G (also see H. interior); > H. densiflorum Small – S; > H. glomeratum Small – S] Hypericum denticulatum Walter, Coppery St.-John's-wort. Cp (GA, NC, SC, VA), Pd, Mt (NC): savannas, wet pine flatwoods, adjacent ditches, borrow scrapes, blackwater stream shores; common (GA Special Concern, VA Watch List). July- September. S. NJ south to e. GA (McIntosh County) (Sorrie 1998b) on the Coastal Plain; disjunct inland in c. and w. NC, sc. TN, HYPERICACEAE 438 and in s. AL. See discussion under H. virgatum. [= K, S; = H. denticulatum var. denticulatum – RAB, C, F, G, Z; < H. denticulatum – GW (also see H. virgatum); = H. denticulatum ssp. denticulatum – X] Hypericum drummondii (Greville & Hooker) Torrey & A. Gray, Nits-and-lice, Drummond's St.-John's-wort. Pd, Cp (GA, NC, SC, VA), Mt (NC, SC, VA): dry woodlands, woodland borders, fields; uncommon. July-September. MD west to OH, IL, and se. KS, south to panhandle FL and c. TX. [= RAB, C, F, G, GW, K, W, X, Z; = Sarothra drummondii Greville & Hooker – S] Hypericum ellipticum Hooker, Pale St.-John's-wort. Mt (NC?): wet places; rare (NC Watch List). Newfoundland and Nova Scotia west to w. Ontario, south to CT, NY, MI, and MN, and in the mountains to WV, NC (?), and ne. TN (Johnson County) (Chester, Wofford, & Kral 1997), and NC (?). The documentation for C's attribution of H. ellipticum to NC is unknown. [= C, F, G, K, V, X] Hypericum fasciculatum Lamarck, Peelbark St.-John's-wort. Cp (GA, NC, SC): wet pine savannas, beaver ponds, upland depression ponds; uncommon (NC Watch List). May-September. E. NC south to s. FL, west to s. MS. [= RAB, GW, K, V, X, Y, Z; < H. fasciculatum – S (also see H. nitidum and H. chapmanii)] Hypericum frondosum Michaux. Mt (GA, NC*, VA*), Pd (GA), Cp (GA): rock outcrops and rocky woodlands; rare, native in GA, introduced in NC and VA from further south and west. Late May-July. This species is native and widespread as far east as e. TN (Chester, Wofford, & Kral 1997). [= C, F, G, K, V, W, Y, Z; > H. aureum Bartram – S; > H. splendens Small – S] Hypericum galioides Lamarck. Cp (GA, NC, SC): wet pine savannas, wet pine flatwoods, pools, edges of bottomlands; common. June-August. E. NC south to n. FL, west to se. TX. [= RAB, GW, K, V, X, Y, Z; > H. ambiguum Elliott – S; > H. galioides – S, in a narrower sense] Hypericum gentianoides (Linnaeus) Britton, Sterns, & Poggenburg, Pineweed, Orange-grass. Mt, Pd, Cp (GA, NC, SC, VA): fields, rock outcrops, woodland borders, eroding areas; common. July-October. ME and Ontario west to MN, south to FL and TX. [= RAB, C, F, G, K, W, X, Z; = Sarothra gentianoides Linnaeus – S] Hypericum graveolens Buckley, Mountain St.-John's-wort. Mt (NC): grassy balds, grassy openings, forests, at high elevations (1200 m or more); rare (NC Watch List). July-August. Nw. NC and ne. TN south to sw. NC, a Southern Appalachian endemic. This and the related H. mitchellianum (another narrow endemic to the Southern Appalachians) hybridize, forming local hybrid populations with intermediate characteristics (Culwell 1970). [= RAB, GW, K, S, W, X, Z] Hypericum gymnanthum Engelmann & A. Gray, Clasping-leaf St.-John's-wort. Cp (GA, NC, SC, VA), Pd (GA, VA), Mt (VA): pine savannas, wet pine flatwoods, sinkhole ponds (Augusta and Rockingham counties, VA), other wet to moist habitats; common (rare in Piedmont). June-September. S. NJ south to n. FL, west to c. TX, and scattered inland in PA, WV, sc. TN, OH, IN, IL, MO, and e. KS; also disjunct in Guatemala (introduced?). [= RAB, C, F, G, GW, K, S, X, Z] Hypericum harperi R. Keller, Harper's St.-John's-wort. Cp (GA, SC): clay-based Carolina bays, other upland depression ponds, with Taxodium ascendens; rare. July-September. E. and c. SC south to sw. GA and panhandle FL. H. harperi should be sought in sc. and se. NC, where it may well occur. This species has generally been considered a part of H. denticulatum or H. virgatum, but Webb (1980) makes a convincing argument for its recognition, including the ecological differentiation and absence of intermediates or hybrids when growing in proximity to H. denticulatum. See H. virgatum for additional discussion. [= X; < H. denticulatum var. acutifolium – RAB, Z; < H. denticulatum – GW; < H. harperi – K; < H. acutifolium – S] Hypericum hypericoides (Linnaeus) Crantz, St. Andrew's Cross. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): dry forests and woodlands; common (uncommon in Mountains). May-August. NJ, w. VA, c. KY, se. MO, and c. OK, south to s. FL and e. TX; also in the West Indies, Mexico, and Central America. [= RAB, C, GW, V, W, Y, Z; > Ascyrum hypericoides Linnaeus var. hypericoides – F, G; > Ascyrum hypericoides Linnaeus var. oblongifolium (Spach) Fernald – F, G; = H. hypericoides ssp. hypericoides – K, X; > Ascyrum hypericoides Linnaeus – S; > Ascyrum linifolium Spach – S] Hypericum interior Small, Interior Bushy St. John's-wort. Mt (GA): rocky forests, riverbanks; uncommon. E. and c. TN, nw. GA south to c. AL. Perhaps best treated as a variety of H. densiflorum. [= S; < H. densiflorum – K, V, X, Z; ? H. revolutum R. Keller] Hypericum lloydii (Svenson) P. Adams, Lloyd's St.-John's-wort. Pd (GA, NC, SC, VA), Cp (GA, NC, SC): dry woodlands, sandhills, edges of granitic flatrocks, edges of Altamaha Grit outcrops, roadbanks; uncommon. June-September. Sc. VA south to c. AL. [= RAB, K, V, X, Z; = Hypericum galioides Lamarck var. lloydii Svenson] Hypericum microsepalum (Torrey & A. Gray) A. Gray ex S. Watson. Cp (GA): moist to wet pine flatwoods; rare. S. GA south to n. FL. [= GW, K, V, X, Y, Z; = Crookea microsepala (Torrey & A. Gray) Small – S] Hypericum mitchellianum Rydberg, Blue Ridge St.-John's-wort. Mt (NC, VA): grassy balds, grassy openings, forests, seepages, at moderate to high elevations (1000-1900 m or more); rare (NC Watch List, VA Rare). July-August. W. VA, e. WV, and e. TN south to sw. NC, a Southern Appalachian endemic. Robson (2006) interprets this as a hybrid of H. graveolens and H. punctatum but offers no evidence other than its general morphological intermediacy. [= RAB, C, F, G, GW, K, S, W, Z; = H. ×mitchellianum Rydberg pro sp. – X] Hypericum mutilum Linnaeus var. mutilum, Common Dwarf St.-John's-wort Mt, Pd, Cp (GA, NC, SC, VA): bogs, marshes, other wet habitats; common. June-October. Newfoundland and Québec west to Manitoba, south to s. FL and c. TX; scattered (probably as an adventive) farther west in North America, in Central and South America, and Europe. Hybrids with H. canadense have been called H. ×dissimulatum Bicknell (pro sp.). [= F; < H. mutilum – RAB, C, G, GW, K, S, W, Z; = H. mutilum ssp. mutilum – X] Hypericum mutilum Linnaeus var. latisepalum Fernald, Southern Dwarf St.-John's-wort. Cp (GA, SC): marshes and other wet habitats; rare. June-October. Se. SC south to peninsular FL, west to TX (and, according to F, north to s. NJ). Hybrids with H. canadense have been called H. ×dissimulatum Bicknell (pro sp.). [= F; < H. mutilum – RAB, G, GW, K, S, W, Z; = H. mutilum ssp. latisepalum (Fernald) N. Robson – X] HYPERICACEAE 439 Hypericum myrtifolium Lamarck, Myrtle-leaf St.-John's-wort. Cp (GA, SC): ponds; common (rare north of GA). Small (1933) reports this species from SC; this distribution is now documented by a specimen from Jasper Co., SC (P. McMillan, pers. comm.). Se. SC south to sc. FL, west to se. MS, a Southeastern Coastal Plain endemic. [= GW, K, S, V, X, Y, Z] Hypericum nitidum Lamarck. Cp (GA, SC): pine savannas; rare (NC Rare, SC Rare). June-August. C. SC south to panhandle FL, west to sw. AL. [= RAB, GW, K, V, Y, Z; H. nitidum ssp. nitidum – X] Hypericum nudiflorum Michaux ex Willdenow. Pd, Cp (GA, NC, SC, VA), Mt (GA, NC): streambanks, moist forests; common (uncommon in VA Piedmont). June-July. Se. VA south to panhandle FL, west to e. TX, s. AR, and se. OK; disjunct in Cumberland Plateau of TN. [= RAB, C, F, G, S, W, V, X, Y; < H. nudiflorum – GW, K, Z (also see H. apocynifolium of the deep South)] * Hypericum perforatum Linnaeus, European St. John's-wort. Pd (GA, NC, SC, VA), Mt, Cp (NC, SC, VA): fields, pastures, roadsides, woodland borders; common, introduced from Europe. June-September. See Duncan (1985) for documentation for GA. [= RAB, C, F, G, K, S, W, Z; = H. perforatum ssp. perforatum – X] Hypericum prolificum Linnaeus, Shrubby St-John's-wort. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): bogs, seepages, rocky forests, rock outcrops; common (rare in Coastal Plain). June-October. NY west to s. MI and MN, south to GA and LA. [= RAB, C, G, K, W, S, V, X, Z; = H. spathulatum (Spach) Steudel – F] Hypericum pseudomaculatum Bush. Cp, Pd (GA, SC), Mt (GA): wet, moist, or dry forests; rare. June-September. SC south to panhandle FL, west to TX, north in the interior to e. TN, c. IL, s. MO, and c. OK. {records east of the Ozarks need to be studied more carefully} [= RAB, C, G, K, S, X, Z; = H. punctatum Lamarck var. pseudomaculatum (Bush) Fernald – F] Hypericum punctatum Lamarck, Spotted St.-John's-wort. Mt, Pd, Cp (GA, NC, SC, VA): fields, woodland borders; common. June-September. Québec west to MN, south to c. peninsular FL and TX. [= RAB, C, G, K, W, X, Z; = H. punctatum var. punctatum – F; > H. punctatum – S; > H. subpetiolatum Bicknell ex Small – S] Hypericum setosum Linnaeus. Cp (GA, NC, SC, VA): pine savannas, wet pine flatwoods, boggy areas, adjacent ditches, fireplow lines, and scrapes; common (VA Rare). May-September. Se. VA south to c. peninsular FL, west to se. TX. [= RAB, C, F, G, GW, K, S, X, Z] Hypericum species 1 Weakley, Radfords' St. John's-wort. Pd (NC): shallow circumneutral soil mats of granitic domes in the Brushy Mountains; rare (NC Watch List). Apparently endemic to the Brushy Mountains of Alexander and Wilkes counties, NC. This taxon, included in H. denticulatum var. acutifolium by Webb (1980), differs from typical H. virgatum in being profusely branched from the medial and upper nodes (rather than being little if at all branched, and then only from the uppermost nodes), in having leaves with acuminate (rather than acute to obtuse) apices, and electrophoretically (Webb 1980). Additionally, these plants have numerous bracteal leaves along the inflorescence branches (vs. few or none), the punctate glands of the foliage are large and oval, resembling those of H. denticulatum (vs. small and round), and the punctate glands are distributed on the lower leaf surface and stem (vs. lower leaf surface only). It may be notable that these same outcrops are phytogeographically interesting, with other disjunct and weakly differentiated races (see Allium cuthbertii). Further study is planned. [< H. virgatum included in concept of H. virgatum (= H. denticulatum var. acutifolium, H. denticulatum ssp. acutifolium) by most earlier authors] Hypericum species 2 LeBlond. Cp (GA, NC, SC?): wet pine savannas, over ultisols or alfisols; rare? June-September. [included in concept of H. tenuifolium (=reductum) by earlier authors] Hypericum stragulum P. Adams & Robson. Mt, Pd (GA, NC, SC, VA), Cp (NC, VA): dry rocky or sandy woodlands; common (uncommon in Coastal Plain). May-August. MA (Nantucket I.), NY (Long Island), west to s. PA, s. OH, s. IN, s. IL, c. MO, se. KS, and c. OK, south to ne. NC, c. SC, c. GA, n. AL, n. MS, n. LA, and c. TX. [= C, V, W, Z; = H. stragalum – RAB, misspelling; = Ascyrum hypericoides Linnaeus var. multicaule (Michaux ex Willdenow) Fernald – F, G; = H. hypericoides (Linnaeus) Crantz ssp. multicaule (Michaux ex Willdenow) Robson – K, X] Hypericum suffruticosum P. Adams & Robson, Pineland St.-John's-wort. Cp (GA, NC, SC): pine savannas and flatwoods; common (rare in NC and SC) (NC Rare). April-June. Se. NC south to c. peninsular FL, west to se. LA. [= RAB, K, V, X, Y, Z; = Ascyrum pumilum Michaux – S] Hypericum tenuifolium Pursh, Sandhill St.-John's-wort. Cp (GA, NC, SC): pine flatwoods, pine savannas, sandhills; common. June-September. Se. NC south to c. peninsular FL, west to panhandle FL and se. AL. Robson (1996) indicates that the older name H. tenuifolium Pursh has now been adequately shown to apply to this taxon. [= X; = H. reductum (Svenson) P. Adams – RAB, GW, K, V, Y, Z; < H. aspalathoides Willdenow – S (also including H. brachyphyllum)] Hypericum tetrapetalum Lamarck. Cp (GA): wet pinelands and in depressional wetlands (open or dominated by Taxodium ascendens); uncommon. E. GA (within a few counties of se. SC), south to s. FL, west to panhandle FL. [= GW, K, V, X, Y; = Ascyrum tetrapetalum (Lamarck) Vail – S] Hypericum virgatum Lamarck. Pd, Mt (GA, NC, SC, VA), Cp (GA, SC): woodlands, rock outcrops, woodland borders; common (VA Watch List). Late June-September. MD west to s. OH, s. IN, and s. IL, south to c. NC, c. SC, sw. GA, panhandle FL, s. MS, and se. LA. Though treated by most recent authors as a variety of H. denticulatum, H. virgatum is better considered as a distinct species. Webb (1980) recognized H. harperi as a separate species (it had previously been considered a part of H. virgatum), and continued to recognize this taxon as a variety of H. denticulatum. However, based on the nature of the punctate glands, size of seeds, inland distribution, etc., it appears that H. virgatum is more distantly related to H. denticulatum and H. harperi than they are to one another; recognition at the species level is warranted for H. virgatum. As pointed out by Webb, H. denticulatum is primarily tetraploid (n = 24), while H. virgatum and H. harperi are (as far as is known) strictly diploid. Additionally, the aberrant populations from granitic outcrops in the Brushy Mountains of Alexander and Wilkes counties, NC referred by Webb (1980) to this taxon are distinct, and more closely allied to H. denticulatum and H. harperi. See Hypericum species 1 for additional discussion. [= K; < H. denticulatum Walter var. acutifolium (Elliott) Blake – RAB, C, F, G, W, Z (also HYPERICACEAE 440 see H. harperi); > H. denticulatum var. recognitum Fernald & Schubert – RAB, F; < H. denticulatum – GW; < H. acutifolium Elliott – S (also see H. harperi); = H. denticulatum ssp. acutifolium (Elliott) N. Robson – X]

Hypericum ascyron Linnaeus ssp. pyramidatum (Aiton) N. Robson, American Great St.-John's-wort. The species is of e. North America and e. Asia; ssp. pyramidatum occurs from Québec west to MN, south to s. PA (Rhoads & Klein 1993), MD (Robson 2000), and WV (Kartesz 1999). [= X; < H. ascyron Linnaeus – K; = H. pyramidatum Aiton – C, F, G] {not keyed at this time} {section Roscyna} Hypericum chapmanii W.P. Adams, Apalachicola St. John's-wort, Tree St. John's-wort. Margins of pond-cypress ponds, pond-cypress stringers, often growing with Cyrilla parviflora and Nyssa ursina. Endemic to Panhandle FL (9 counties). [= GW, K, V, X, Y, Z; < H. fasciculatum – S; = H. arborescens Chapman] Hypericum dolabriforme Ventenat, Glade St. John's-wort. Mt (GA): limestone glades and barrens; rare (GA Special Concern). In nw. GA (Jones & Coile 1988) and e. TN (Chester, Wofford, & Kral 1997); this species should be sought in sw. VA. [= C, F, G, K, S, V, X, Z] {not keyed at this time} Hypericum exile W.P. Adams. Pine flatwoods. Endemic to Panhandle FL (Bay, Franklin, Gulf, Liberty, and Washington counties). There seems nothing in particular to recommend Robson's reduction of H. exile to a subspecies of H. nitidum. [= GW, K, V, Y, Z; = H. nitidum Lamarck ssp. exile (W.P. Adams) N. Robson – X] Hypericum lissophloeus W.P. Adams. Margins of sinkhole ponds. Endemic to Panhandle FL (Bay and Washington counties). [= GW, K, V, X, Y, Z] Hypericum lobocarpum Gattinger. Streambanks, river bottoms, pinelands. C. TN (Chester, Wofford, & Kral 1997) and MS west to s. IL, se. OK, e. TX; credited to SC by Robson (1996), based on specimens debated and dismissed by Adams (1973). Late May-September. [= C, K, S, V, X, Z; = H. densiflorum var. lobocarpum (Gattinger) Svenson – F, G; < H. densiflorum – GW] {not keyed at this time} Hypericum majus (A. Gray) Britton. South to nw. PA (Rhoads & Klein 1993), NJ, DE, and OH (Kartesz 1999). [= C, F, G, K, X] {not keyed at this time} Hypericum species 3 J. Allison, Georgia St, John's-wort. Cp (GA): seepage bogs, roadside ditches; rare (GA Special Concern). Apparently endemic to the Altamaha Grit region of the GA Coastal Plain. {not keyed at this time} Hypericum sphaerocarpum Michaux, Barrens St. John's-wort. Mt (GA): limestone barrens; rare (GA Special Concern). East to GA (GAHP 2003), e. and c. TN (Chester, Wofford, & Kral 1997), sw. PA (Rhoads & Klein 1993). [= C, F, G, K, V, X, Z; > H. turgidum Small – S; > H. sphaerocarpum var. turgidum (Small) Svenson] {not keyed at this time}

Triadenum Rafinesque (Marsh St-John's-wort)

A genus of 6-10 species, herbs, of e. North America and e. Asia. Although Triadenum has sometimes formerly been included in Hypericum, Robson (1977) and others consider it to be more closely related to the tropical Asian shrub Cratoxylum Blume. References: Adams (1973)=Z.

1 Leaves narrowed to the cuneate or broadly cuneate (rarely truncate) base. 2 Lower leaves sessile; sepals acute; leaves lacking translucent or dark glands or punctae...... T. tubulosum 2 Lower leaves petiolate; sepals obtuse; leaves with translucent glands and dark punctae...... T. walteri 1 Leaves clasping, cordate, or subcordate at the base. 3 Sepals 2.5-5 mm long at maturity, obtuse or acute; styles 0.5-1 (-1.5) mm long (best seen in fruit) ...... T. fraseri 3 Sepals 5-8 mm long at maturity, acute or acuminate; styles 1.8-3 mm long (best seen in fruit) ...... T. virginicum

Triadenum fraseri (Spach) Gleason, Marsh St.-John's-wort. Mt (NC, VA): bogs, peaty wetlands; rare (VA Rare). July- August. Newfoundland and Québec west to MN, south to NY, PA, w. VA, ne. TN, w. NC, OH, n. IN, and NE. Closely related to T. virginicum and reduced to a variety of (or included in) that species by some authors. [= C, G, K; = Hypericum virginicum Linnaeus var. fraseri (Spach) Fernald – F; < T. virginicum – W, Z, in part; = Hypericum fraseri Spach] Triadenum tubulosum (Walter) Gleason, Marsh St.-John's-wort. Cp (GA, NC, SC, VA), Pd (VA): bogs, peaty wetlands, drawdown sloughs along rivers; uncommon (GA Rare, VA Rare). August-September. Se. VA south to n. FL, west to LA, and north in the interior to se. and c. TN, s. IL and s. OH. [= C, G, GW, K, Z; = Hypericum tubulosum Walter – RAB; = Hypericum tubulosum Walter var. tubulosum – F; = T. longifolium Small – S] Triadenum virginicum (Linnaeus) Rafinesque, Marsh St.-John's-wort. Cp, Mt (GA, NC, SC, VA), Pd (GA, VA): bogs, peaty wetlands; common (rare in Mountains and Piedmont). July-September. Nova Scotia west to OH and s. Ontario, south to FL and MS, mostly on the Coastal Plain but scattered inland. [= C, G, GW, K, S; = Hypericum virginicum Linnaeus – RAB; = Hypericum virginicum var. virginicum – F; < T. virginicum – W, Z, in part only (also see T. fraseri)] Triadenum walteri (J.G. Gmelin) Gleason, Marsh St.-John's-wort. Cp, Pd, Mt (GA, NC, SC, VA): swamp forests and marshes; common (uncommon in VA Piedmont, rare in VA Mountains). August-September. MD south to n. FL, west to e. TX, north in the interior to s. MO, s. IL, and OH. [= C, G, GW, K, W, Z; = Hypericum walteri J.G. Gmelin – RAB; = Hypericum tubulosum Walter var. walteri (J.G. Gmelin) Lott – F; = T. petiolatum (Walter) Britton – S]

ILLICIACEAE A.C. Smith 1947 (Star-anise Family)

HYPERICACEAE 441 A family of 1 genus and about 42 species, shrubs and trees, of temperate and subtropical se. Asia and se. North America (se. United States, Cuba, Haiti, and e. Mexico). The family is most closely related to the Schisandraceae and Winteraceae. References: Keng in Kubitzki, Rohwer, & Bittrich (1993).

Illicium Linnaeus 1759 (Star-anise)

A genus of about 42 species, shrubs and trees, of temperate and subtropical se. Asia and se. North America (se. United States, Cuba, Haiti, and e. Mexico). References: Vincent in FNA (1997); Keng in Kubitzki, Rohwer, & Bittrich (1993); Stone & Freeman (1968).

1 Flowers 2-5-5 cm across; tepals 21-33, red-maroon (rarely white or pinkish); leaf tips acute to acuminate; [section Illicium]...... I. floridanum 1 Flowers 0.8-1.2 cm across; tepals 11-16, yellowish green; leaf tips obtuse or rounded; [section Cymbostemon]...... I. parviflorum

Illicium floridanum Ellis, Florida Star-anise. Cp (GA): acid ravines and small stream swamps; rare (GA Endangered). Sw. GA west to e. LA. [= FNA, GW, K, S] * Illicium parviflorum Michaux ex Ventenat, Swamp Star-anise, Yellow Anise-tree. Cp (GA): cultivated and persistent; rare (GA Special Concern). April-June. This species occurs in swampy forests, evergreen hammocks, and bayheads and is endemic to scattered localities in central FL; it is in the horticultural trade and has been introduced in various places, including sw. and se. GA. [= FNA, K, S]

ITEACEAE J. Agardh 1858 (Sweetspire Family)

A family of 1 genus and about 20 species, shrubs, of e. Asia, e. North America, and Mexico.

Itea Linnaeus (Virginia-willow, Sweetspire, Tassel-white)

Variously treated in a very broadly-conceived Saxifragaceae (RAB, F, G, GW, W), a less comprehensive Grossulariaceae (C, K), a narrow Escalloniaceae, or a very narrow (single genus) Iteaceae (S), the relationships of Itea remain problematic. Recent molecular data suggest that the relationship between Itea and other woody "saxifragaceous" genera (including Escallonia) is only distant (Morgan & Soltis 1993). Itea is here conservatively treated in a narrow Iteaceae. The remainder of the genus (about 20 species) is found in e. Asia and w. Malaysia; the only close relative of our species is I. japonica Oliver, of Japan. References: Spongberg (1972); Morgan & Soltis (1993); Bohm et al. (1999).

Identification notes: Sometimes confused needlessly with Clethra, whose much more coarsely serrate, obovate leaves contrast with the serrulate, elliptic leaves of Itea. Also often confused with Leucothoe racemosa in vegetative condition.

Itea virginica Linnaeus, Virginia-willow, Sweetspire, Tassel-white. Cp (GA, NC, SC, VA), Pd (GA, NC, SC, VA), Mt (GA, NC, SC): moist forests and thickets, especially along the banks of small streams; common (uncommon in Piedmont and Mountains). May-June. S. NJ south to FL and west to e. TX, north in the interior (especially in the Mississippi Embayment) to s. IL and se. MO. [= RAB, C, F, K, G, GW, S, W]

JUGLANDACEAE A. Richard ex Kunth 1824 (Walnut Family)

A family of about 8 genera and 60 species, trees and shrubs, mostly temperate. References: Stone in FNA (1997); Manos & Stone (2001); Elias (1972); Stone in Kubitzki, Rohwer, & Bittrich (1993).

1 Fruit with husk dehiscent into 4 valves; pith of twigs continuous; leaves with (3-) 5-17 (-19) leaflets, the largest usually the terminal or final 2 lateral; nut with shell smooth, ridged, or irregularly wrinkled (but not deeply furrowed); terminal buds with imbricate (overlapping) or valvate scales; [tribe Juglandeae, subtribe Caryinae] ...... Carya 1 Fruit with husk indehiscent; pith of twigs chambered (not always developing until autumn of the first year's growth); leaves with (7-) 11-19 (-23) leaflets, the largest usually about halfway up the leaf; nut with shell deeply furrowed in a complex corrugated pattern; terminal buds with valvate; [tribe Juglandeae, subtribe Juglandinae]...... Juglans

Carya Nuttall (Hickory) (by A.S. Weakley & R.K. Peet)

JUGLANDACEAE 442 A genus of about 18 species, trees, of e. North America (south into s. Mexico), and e. Asia. Carya in our area is separated into two sections, section Apocarya (C. aquatica, C. cordiformis, C. illinoinensis) and section Carya (C. alba, C. carolinae- septentrionalis, C. glabra, C. laciniosa, C. myristiciformis, C. ovalis, C. ovata, C. pallida). The southeastern United States is the center of diversity of Carya. Our four-state area includes 11 of 13 North American species (including the naturalized C. illinoinensis and lacking only a more southern species, C. floridana Sargent of peninsular FL, and a south-central species, C. texana Buckley, of sc. United States), and 11 of 18 species worldwide. Section Rhamphocarya includes a single Asian species. The remaining 4 species in the genus are all in section Apocarya: C. palmeri Manning of Mexico and 3 Asian species.

C. cordiformis, C. aquatica, C. illinoinensis, C. myristiciformis, C. laciniosa, C. ovata, and C. carolinae-septentrionalis are diploids, with n = 16. C. pallida, C. glabra, C. ovalis, and C. alba are tetraploids with n = 32 (Stone 1961). As suggested by Stone, Adrouny, & Flake (1969), it seems possible that reticulate evolution involving extant or extinct diploid species is responsible for some of the difficulties in the C. glabra-ovalis complex. Many hybrids have been described, but some are questionable. Additionally, Hardin & Stone (1984) state that "most of these hybrids are localized and have not led to introgressive populations, or at least none that have been recognized".

Ecologically, Carya is one of the more diverse and ubiquitous genera of trees in our area, surpassed in number of species, abundance, and ecological amplitude only by Quercus and Pinus. This has led to a long tradition of describing large parts of our area (in particular the Piedmont) as being characterized by "oak-hickory" or "oak-pine-hickory" forests (e.g. Küchler 1964; Greller 1988; Schafale & Weakley 1990; Skeen, Doerr, & Van Lear 1993). Ware (1992) and others have recently questioned this tradition, pointing out that Carya only rarely dominates or codominates, primarily in specialized circumstances (such as in soils with greater cation concentrations, derived from mafic rocks).

The association of many (but certainly not all) species of hickories with soils with high base status was noted in print as early as 1820 in an account of the landscape of North Carolina. "The sandy pine barrens, and all the lands on which pine is the exclusive growth, are unfriendly to agriculture; but where the pine is intermixed with oak and hickory, the soil is good. Some of our strongest lands have tall pine, mixed not only with hickory and oak, but also with walnut and cherry, and such trees that indicate the best soil. Where hickory prevails, the land is strong" (Guthrie 1820).

Note that the C. glabra-C. ovalis portion of this treatment is tentative. In our area, this group has been variously treated as consisting of between 1 and 10 (or more) taxa. For testing, we present here a plausible middle ground treatment in the key and species accounts (4 taxa). References: Stone in FNA (1997); Hardin (1992); Hardin & Stone (1984); Elias (1972); Stone, Adrouny, & Flake (1969); Stone (1961); Mohlenbrock (1986); Sargent (1918); Manning (1950); Hardin (1952); Little (1969); Harrar & Harrar (1962); Stone in Kubitzki, Rohwer, & Bittrich (1993). Key based in part on an unpublished manuscript prepared by Stone & Hardin for the Flora of the Southeastern United States.

Identification notes: Surface vestiture of leaves and bud scales is useful in distinguishing species of Carya. Some use of these characters can be made with a 10× or 20× hand lens; better still is a dissecting microscope. It is important to understand the different trichome types mentioned in the key (terminology follows Hardin 1990 and Hardin & Stone 1984). Short acicular trichomes are simple, unicellular trichomes tapered to a pointed tip, 0.10-0.35 mm long and with rough walls. Long acicular trichomes ("solitary" of Hardin & Stone 1984) are similar to short acicular, but are much larger, 0.45-1.6 mm long, and have smooth walls. Fasciculate trichomes are multicellular and have 2-8 straight or curled rays radiating from a clustered base. Multiradiate trichomes are similar to fasciculate, but have 8-17 rays, the inner (and usually more upright) rays attached basally above the outer (and usually more spreading) rays. Capitate glandular trichomes are unicellular or multicellular, and are distinguished by their bulbous or expanded tip; they are usually 0.02-0.1 mm long. Peltate scales are flat or dome-shaped shields or disks, slightly to strongly glandular, (sometimes regularly or irregularly lobed) and can be either sessile or stalked (they are often referred to as scales, resin dots, peltate glands, or lepidote scales). On the lower surfaces of leaflets, peltate scales are of two types: large peltate scales are 0.08-0.3 mm in diameter and are round, with smooth or slightly irregular margins, while small peltate scales are 0.025-0.12 mm in diameter and are either round, irregularly lobed or regularly 2- or 4-lobed.

1 Terminal buds with 4-6 valvate scales; leaves with 7-13 (-19) leaflets, these symmetrical to strongly falcate; fruit sutures narrowly winged. 2 Leaves with 7-9 (-11) leaflets, these symmetrical to slightly falcate; fasciculate trichomes with 2-4 rays; terminal bud elongate, flattened, 9-19 mm long, bright orangy yellow to dull orange-tan; [common and widely distributed tree in our area, typically in floodplain and slope forests]...... C. cordiformis 2 Leaves with (7-) 9-19 leaflets, these slightly to strongly falcate; fasciculate trichomes with 2-8 rays. 3 Leaves with (7-) 9-11 (-13) leaflets; bark shaggy; lateral petiolules 0-2 mm long; nut flattened and angled in cross- section; kernel bitter; [native, of swamp forests, primarily in the Coastal Plain]...... C. aquatica 3 Leaves with 11-19 leaflets; bark scaly, with small exfoliating plates; lateral petiolules 0-7 mm long; nut round in cross-section; kernel sweet; [introduced, frequently cultivated, long persistent, and occasionally naturalized] ...... C. illinoinensis 1 Terminal buds with 6-15 imbricate scales; leaves with (3-) 5-9 (-11) leaflets, these symmetrical to slightly falcate; fruit sutures not winged (except C. myristiciformis). JUGLANDACEAE 443 4 Bark shaggy (on large trees separating in segments to a meter in length); leaves with (3-) 5 (-7) leaflets; serrations of the leaflets densely (or only moderately) ciliate when young, most densely so just below the tooth apex, the hairs sloughing with age but leaving a subapical tuft of white trichomes on at least some teeth. 5 Twigs slender, hardened first-year growth or second-year growth 1-3 mm in diameter; terminal bud 6-15 mm long, glabrous to sparsely puberulent (except for ciliate fringe on the scales), reddish-brown (usually turning black on drying); lower surface of leaflets nearly glabrous, except for tufts of trichomes in the main vein axils, and only slightly lepidote with a few, scattered scales, the large peltate scales yellow and round, the small peltate scales brown, 2- and 4-lobed; terminal leaflet 2-5 (-6) cm wide ...... C. carolinae-septentrionalis 5 Twigs stout, hardened first-year growth or second-year growth (2.5-) 3-6 mm in diameter; terminal bud 9-18 mm long, tomentose, tan to brown (rarely turning black on drying); lower surface of leaflets moderately to densely hirsute with acicular and fasciculate hairs (sometimes the hairs more or less limited to the main veins), and also moderately lepidote, the large peltate scales yellow and round, the small peltate scales dark brown and mostly round; terminal leaflet (4-) 6-15 cm wide ...... C. ovata 4 Bark tight (the ridges typically forming an interlocking diamond pattern), scaly, or shaggy (when shaggy, the separated segments normally much < 1 meter long); leaves with (3-) 5-9 (-11) leaflets; serrations of the leaflets glabrous or ciliate, but lacking subapical tufts of trichomes. 6 Twigs stout; terminal buds 8-20 mm long; leaves with (5-) 7-9 (-11) leaflets; lower surface of leaflets densely hirsute with a mixture of acicular (single), fascicled (2-8 rays), and multiradiate (8-many rays) hairs; small peltate scales of the lower surface of leaflets all round; fruit husk 4-13 mm thick; nuts slightly to strongly 4-angled toward the apex. 7 Bark tight; petiole hirsute; leaflet apex acute; lower surface of leaflets densely hirsute with acicular (single) and abundant 2-8-rayed fascicled and multiradiate hairs; fruit husk glabrous, with pustulate bumps; fruit 3.5-5 cm long; nut 2.5-3.5 cm long; [common in our area]...... C. alba 7 Bark shaggy; petiole hirtellous; leaflet apex acuminate; lower surface of leaflets hirsute with acicular (single), 2-6-rayed fascicled, and occasional multiradiate hairs; fruit husk pubescent, lacking pustulate bumps; fruit 4- 7 cm long; nut 3-6 cm long; [rare in our area]...... C. laciniosa 6 Twigs slender; terminal buds 3-15 mm long; leaves with (3-) 5-7 (-9) leaflets; lower surface of leaflets mostly glabrous, except for along the midrib and primary veins, and sometimes hirsute on the surface with acicular (single) and infrequent fascicled (2-8 rays) hairs (lacking multiradiate trichomes); small peltate scales of the lower surface of leaflets of various types, 4-lobed and/or irregular scales often more frequent than round scales; fruit husk 2-5 mm thick; nuts not 4-angled toward the apex. 8 Terminal bud 4-10 mm long, predominantly lepidote (also pubescent); leaves with (5-) 7 (-9) leaflets; lower surface of spring leaflets densely lepidote with 4-lobed, irregular, and round peltate scales, giving the undersurface a reflective, silvery-tan, rusty-brown, or bronze sheen. 9 Lepidote scales initially silver, soon turning bronze, and giving the buds, young twigs, and undersurface of the leaves a metallic bronze sheen; fruit 2-3 cm long; [of calcareous swamps, bottomlands and slopes of the Coastal Plain of se. NC, e. SC, and GA] ...... C. myristiciformis 9 Lepidote scales silvery-tan or rusty-brown, giving the buds, young twigs, and undersurface of the leaves a dull or slightly shiny tan or rusty-brown color; fruit 3-5 cm long; [usually of upland and acidic forests and woodlands, collectively widespread in our area]. 10 Undersurface of the leaflets with dense, silvery-tan large peltate scales, and fewer and less conspicuous fewer small peltate scales (thus the leaves appearing overall silvery-tan); petiole and rachis hirsute with fasciculate trichomes, and also with concentrations of hairs near the leaflet insertions; [widespread in our area, of upland and acidic forests and woodlands] ...... C. pallida 10 Undersurface of the leaflets with dense, rusty-brown small peltate scales, and fewer and less conspicuous sivery-tan large peltate scales (thus the leaves appearing overall rusty-brown); petiole and rachis with few fasciculate hairs (but densely scaly), and lacking concentrations of hairs near the leaflet insertions; [of the sc. United States, east to MS, and perhaps AL, GA, and western KY and TN, of upland or lowland, acidic or calcareous forests and woodlands]...... [C. texana] 8 Terminal bud 5-15 mm long, predominantly pubescent (also sparsely lepidote); leaves with (3-) 5-7 (-9) leaflets; lower surface of spring leaflets slightly to densely lepidote with irregular and round peltate scales (4- lobed peltate scales uncommon). 11 Fruit husk spitting to base at maturity along 2-4 sutures; leaves with (5-) 7 leaflets, pubescent beneath; petiole reddish; fruits typically ellipsoidal; bark tight or often scaly or somewhat shaggy ...... C. ovalis 11 Fruit husk indehiscent at maturity or tardily splitting to base along 1 suture; leaves with (3-) 5 (-7) leaflets, glabrous to pubescent beneath; petiole usually green; fruits ellipsoidal, pyriform, or subglobose; bark tight. 12 Rachis (and also often the petiole and lower surfaces of leaflets) densely pubescent...... C. glabra var. hirsuta 12 Rachis, petiole, and lower surfaces of leaflets glabrous or glabrescent. 13 Husk of fruit 1.5-2.5 mm thick; fruit ellipsoidal, subglobose, to obovoid, 1.5-3.5 cm long; terminal leaflet usually 8-17 cm long; [primarily of the Piedmont and Mountains] ...... C. glabra var. glabra JUGLANDACEAE 444 13 Husk of fruit about 3.5 mm thick; fruit pyriform, 2.5-5 cm long; terminal leaflet usually 20-25 cm long; [primarily of the Coastal Plain] ...... C. glabra var. megacarpa

Carya alba (Linnaeus) Nuttall ex Elliott, Mockernut Hickory, White Hickory. Pd, Cp, Mt (GA, NC, SC, VA): forests and woodlands; common. April-May; October. MA west to IN and IA, south to FL and TX. One of the most common forest trees of much of our area. There has been confusion and controversy for several centuries over the specific epithet. The oldest basionym available is Juglans alba Linnaeus, which apparently included disparate elements, including this taxon and C. ovata. Following a more circumscribed typification by Crantz in 1766, the epithet "alba" should have been applied to this taxon, but continued to be applied in various ways. Rehder (1945) proposed that C. alba should be considered a nomen ambiguum, but agreed that it applied correctly to what has often been called C. tomentosa. He argued that the use of C. alba should be rejected "in order to avoid confusion and ambiguity." However, C. alba has not been officially rejected, its application appears to be nomenclaturally clear, and there is no alternative under the International Code of Botanical Nomenclature (Greuter 1988) to its use. For further discussion see Rehder (1945), Howard & Staples (1983), and Wunderlin, Hansen, & Hall (1985). [= K; = C. tomentosa (Lamarck ex Poiret) Nuttall – RAB, C, F, FNA, G, W; = Hicoria alba (Linnaeus) Britton – S] Carya aquatica (Michaux f.) Nuttall, Water Hickory, Bitter Pecan. Cp (GA, NC, SC, VA), Pd (GA, SC): swamp forests, where flooded during the winter months; uncommon. April-May; October. Se. VA south to c. peninsular FL, west to e. TX, north inland to se. MO, s. IL, and se. OK. [= RAB, C, F, FNA, G, GW, K; = Hicoria aquatica (Michaux f.) Britton – S] Carya carolinae-septentrionalis (Ashe) Engler & Graebner, Carolina Shagbark Hickory, Carolina Hickory. Pd (GA, NC, SC, VA), Mt (GA): upland flats, especially those weathered from mafic rocks and with shrink-swell soils dominated by montmorillonitic clays, less typically on slopes and bottomlands; uncommon (rare in VA). April-May; October. Sc. VA (Halifax County) south to GA, AL, and MS, and inland northward to c. TN and sc. KY. First reported for VA by Wieboldt et al. (1998). The taxonomic status of C. carolinae-septentrionalis has been controversial, with some workers reducing it to variety of C. ovata or not recognizing it at all. It seems to us morphologically and ecologically distinctive and to represent an independent evolutionary lineage. Hardin & Stone (1984) found differences in trichomes, and in a study of nut oils, Stone, Adrouny, & Flake (1969) found C. ovata "surprisingly distant" from C. carolinae-septentrionalis. There are reports that the two taxa are also phenologically separated, C. carolinae-septentrionalis leafing out about two weeks earlier than C. ovata, when growing together in the c. Piedmont of NC. Though usually ecologically and/or geographically segregated, the two species sometimes occur together or in close proximity to one another; they maintain their distinctness. [= RAB, C, G, K; = C. ovata (P. Miller) K. Koch var. australis (Ashe) Little – FNA; = Hicoria carolinae-septentrionalis Ashe – S; = C. ovata var. carolinae-septentrionalis (Ashe) Reveal; = C. australis Ashe] Carya cordiformis (Wangenheim) K. Koch, Bitternut Hickory. Mt, Pd, Cp (GA, NC, SC, VA): forests and woodlands, especially in rich, moist alluvial or slope forests; common. April; October. ME and s. Québec west to MN and NE, south to panhandle FL and e. TX. [= RAB, C, F, FNA, G, GW, K, W; = Hicoria cordiformis (Wangenheim) Britton – S] Carya glabra (P. Miller) Sweet var. glabra, Pignut Hickory. Mt, Pd, Cp (GA, NC, SC, VA): forests and woodlands; common. April-May; October. C. glabra ranges from s. NH west to s. MI, se. IA, and se. KS, south to c. peninsular FL and e. TX; the ranges of the varieties are poorly known. [= F, G; < C. glabra – RAB, C, FNA, GW, K; < C. glabra var. glabra – W; = Hicoria glabra (P. Miller) Britton var. glabra – S] Carya glabra (P. Miller) Sweet var. hirsuta (Ashe) Ashe, Hairy Pignut Hickory. Mt, Pd? (GA, NC, SC, VA): forests and woodlands; common? April-May; October. Var. hirsuta is apparently primarily Appalachian, variously described as being endemic to the Southern Appalachians or ranging north to s. NH. It needs additional taxonomic assessment. [< C. glabra – RAB, C, FNA, GW, K; = C. ovalis (Wangenheim) Sargent var. hirsuta (Ashe) Sargent – F; = Hicoria glabra (P. Miller) Britton var. hirsuta Ashe – S; < C. glabra var. glabra – W] Carya glabra (P. Miller) Sweet var. megacarpa (Sargent) Sargent, Coastal Pignut Hickory. Cp (GA, NC, SC, VA?): maritime forests and other forests of the outer Coastal Plain; uncommon? April-May; October. Var. megacarpa is apparently primarily a tree of the se. United States Coastal Plain, ranging from s. NY south to FL, west to TX, and north in the interior to s. IL. It needs additional taxonomic assessment. [= F, G; < C. glabra – RAB, C, FNA, GW, K; ? Hicoria austrina Small – S] * Carya illinoinensis (Wangenheim) K. Koch, Pecan. Cp, Pd (GA, NC, SC, VA): persistent around dwellings and in pecan orchards, escaped to suburban woodlands, rural forest edges and floodplains; commonly cultivated, rarely naturalized. April- May; October. Native to the sc. United States, now more widespread in the se. United States as a result of cultivation. The spelling of the specific epithet has been a source of controversy. [= C, FNA, K; = C. illinoensis – RAB, F, G, GW, orthographic variant; > Hicoria pecan (Marshall) Britton – S; > Hicoria texana LeConte – S] Carya laciniosa (Michaux f.) G. Don, Kingnut Hickory, Big Shellbark Hickory. Mt (GA), Cp, Pd (NC): moist, circumneutral, alluvial levee forests along brownwater rivers of the Coastal Plain (NC), streams of the Piedmont (NC) and Mountains (GA); rare (NC Rare). April-May; October. NY and s. Ontario west to IA, south to NC, nw. GA, MS, and OK. This species is sometimes planted, but occurs native in nw. GA, along the Roanoke River (Halifax and Northampton counties, NC) and New Hope Creek (Durham County, NC). [= RAB, C, F, FNA, G, GW, K, W; = Hicoria laciniosa (Michaux f.) Sargent – S] Carya myristiciformis (Michaux f.) Nuttall, Nutmeg Hickory. Cp (GA, NC, SC): nonriverine swamps over calcareous substrates, including calcareous clays and coquina limestone ("marl"), oak flatwoods; rare (GA Special Concern, NC Threatened, SC Rare). April; October. Se. NC south to GA, and from wc. AL west to e. TX and se. OK; disjunct in Mexico (Nuevo Léon and Tamaulipas). The bronze sheen of the leaflets of this species is diagnostic. First reported for NC by Leonard (1971b). [= FNA, K; = C. myristicaeformis – RAB, GW, orthographic variant; = Hicoria myristicaeformis (Michaux f.) Britton – S] Carya ovalis (Wangenheim) Sargent, Red Hickory. Mt, Pd, Cp (GA, NC, SC, VA): forests and woodlands; common. April-May; October. MA west to WI, south to GA, MS, and MO. [= RAB, C, K; > C. ovalis var. obcordata (Muhlenberg & JUGLANDACEAE 445 Willdenow) Sargent – F, G; > C. ovalis var. obovalis Sargent – F, G; > C. ovalis var. odorata (Marshall) Sargent – F, G; < C. glabra – FNA; = Hicoria microcarpa (Nuttall) Britton – S; = C. glabra (P. Miller) Sweet var. odorata (Marshall) Little – W] Carya ovata (P. Miller) K. Koch, Common Shagbark Hickory. Mt (GA, NC, VA), Pd, Cp (NC, VA): rich moist bottomlands, slopes, occasionally on dry upland flats; uncommon. May; October. S. ME and s. Québec west to MN and NE, south to GA and TX; also disjunct in Mexico. [= RAB, C, F, G, GW, K, W; > C. ovata var. ovata – F; > C. ovata var. pubescens Sargent – F; = C. ovata var. ovata – FNA; = Hicoria ovata (P. Miller) Britton – S] Carya pallida (Ashe) Engler & Graebner, Sand Hickory, Pale Hickory. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): dry sandy or rocky forests and woodlands; common. April-May; October. S. NJ south to FL, west to TX, inland in the interior to w. NC, KY, s. IL, and AR. [= RAB, C, F, FNA, G, K, W; = Hicoria pallida Ashe – S]

Carya texana Buckley, Black Hickory. Reported to occur as far east as KY, TN, and GA (Kartesz 1999), an eastern extent not reported by FNA, which accepts it as far east as MS. Jones (2005) states that w. KY material of C. pallida is transitional to C. texana. [= FNA, K; > C. texana var. arkansana (Sargent) Little – C; > C. texana var. texana – F; > C. buckleyi Durand var. arkansana (Sargent) Sargent – G]

Juglans Linnaeus (Walnut)

A genus of about 21 species, trees and shrubs, of Mediterranean Europe to e. Asia, and North America to Andean South America. Stanford, Harden, & Parks (2000) present a molecular phylogeny and a discussion of biogeography; our two species are distantly related within the genus, with J. nigra most closely related to sw. North American J. microcarpa and J. major, and J. cinerea most closely related to several e. Asian species. References: Whittemore & Stone in FNA (1997); Stanford, Harden, & Parks (2000); Stone & Hardin in SE (in prep.); Stone in Kubitzki, Rohwer, & Bittrich (1993); Stanford (1998).

1 Lower surface of the leaflets densely hirsute with 4-8-rayed fascicled hairs; fruit ellipsoid, densely pubescent with reddish- brown glandular hairs; leaf scars with a velvety ridge along the upper margin; leaves with (7-) 11-17 leaflets; pith dark brown; terminal buds 12-18 mm long; bark of mature trees pale; [section Trachycaryon]...... J. cinerea 1 Lower surface of the leaflets hirsute with single and 2-rayed fascicled hairs; fruit spherical or nearly so, lepidote with peltate scales and occasional glandular hairs; leaf scars without a velvety ridge along the upper margin; leaves with (9-) 15-19 (-23) leaflets; pith light brown; terminal buds 8-10 mm long; bark of mature trees dark; [section Rhysocaryon]...... J. nigra

Juglans cinerea Linnaeus, Butternut, White Walnut. Mt (GA, NC, SC, VA), Pd (NC, SC, VA), Cp (VA): moist, nutrient- rich forests; uncommon (US Species of Concern, GA Special Concern, NC Watch List, SC Rare, VA Watch List). April-May; October. New Brunswick west to MN, south to n. GA and AR. This tree, formerly common, is afflicted with butternut canker disease, which now threatens its continued existence. [= RAB, C, F, FNA, G, K, W; = Wallia cinerea (Linnaeus) Alefeld – S] Juglans nigra Linnaeus, Black Walnut. Mt, Pd, Cp (GA, NC, SC, VA); moist, nutrient-rich forests; common (uncommon in the Coastal Plain of NC and SC). April; October. MA west to MN, south to nw. FL and TX. The dark brown wood is famous for cabinetry and other uses; it is one of the most prized of North American hardwoods. The nuts, though difficult to crack, are prized for their intense flavor. The husk is used as a dye. Country people dehusk the nuts by putting them in dirt or gravel driveways where the passage of car tires removes the husk but does not crack the nut. [= RAB, C, F, FNA, G, K, W; = Wallia nigra (Linnaeus) Alefeld – S]

KRAMERIACEAE Dumortier 1829 (Krameria Family)

A family of a single genus and about 15-18 species, herbs, shrubs, and trees, of warm (and usually dry) parts of s. North America to South America. References: Robertson (1973); Simpson et al. (2004).

Krameria Linnaeus (Ratany)

A genus of about 15 species, herbs, shrubs, and trees, hemiparasitic by haustoria. References: Robertson (1973)=Z.

Krameria lanceolata Torrey, Trailing Ratany, Sandspur. Cp (GA): sandhills; uncommon. AR, TX, and s. KS west to se. CO, se. AZ, Chihuahua, and Coahuila; disjunct eastward in the Coastal Plain of FL and GA (east to Bulloch, Bryan, Evans, and Emanuel counties, GA). [= K, Z; > K. spathulata Small – S]

LAMIACEAE Lindley 1836 or LABIATAE A.L. de Jussieu 1789 (Mint Family)

A family of about 230-250 genera and 6700-7170 species, herbs, shrubs, vines, and trees, cosmopolitan. The placement in the Lamiaceae of several genera traditionally placed in Verbenaceae (e.g. Clerodendrum) is strongly supported by several lines of evidence. References: Harley et al. in Kadereit (2004). subfamily Viticoideae: Vitex. LAMIACEAE 446 subfamily Ajugoideae: Ajuga, Clerodendrum, Teucrium, Trichostema. subfamily Scutellarioideae: Scutellaria. subfamily Lamioideae: Galeopsis, Lamiastrum, Lamium, Leonurus, Macbridea, Marrubium, Physostegia, Sideritis, Stachys, Synandra. subfamily Nepetoideae: tribe Elsholtzieae: Collinsonia, Elsholtzia, Mosla, Perilla. tribe Mentheae: subtribe Salviinae: Rosmarinus, Salvia. subtribe Menthinae: Blephilia, Clinopodium, Conradina, Cunila, Dicerandra, Hedeoma, Hyssopus, Lycopus, Mentha, Monarda, Origanum, Piloblephis, Prunella, Pycnanthemum, Thymus. subtribe Nepetinae: Agastache, Dracocephalum, Glechoma, Meehania, Nepeta. incertae sedis: Melissa. tribe Ocimeae: subtribe Hyptidinae: Hyptis. subtribe Ociminae: Ocimum. incertae sedis: Callicarpa.

Acinos (see Clinopodium)

Agastache Clayton ex Gronovius 1762 (Giant-hyssop)

A genus of about 22 species, herbs, of c. and e. Asia, and North America to Mexico. References: Vogelmann (1985); Lint & Epling (1945); Harley et al. in Kadereit (2004).

1 Leaves densely white tomentose below; corolla blue; [cultivated as an ornamental and rarely naturalized].... [A. foeniculum] 1 Leaves glabrous to villous beneath, appearing green; corolla yellow, greenish-yellow, or pinkish; [native]. 2 Corolla yellow or greenish-yellow; calyx lobes obtuse or subacute, 1-1.5 mm long at anthesis; calyx lobes and bracts green; midstem internodes glabrous or minutely pubescent; lower surface of the leaf pubescent on the veins and surface ...... A. nepetoides 2 Corolla pinkish; calyx lobes acute or acuminate, 2-2.5 mm long at anthesis; calyx lobes and bracts with white or pink margins; midstem internodes at least sparsely long-pubescent; lower surface of the leaf pubescent mainly on the veins...... A. scrophulariifolia

Agastache nepetoides (Linnaeus) Kuntze, Yellow Giant-hyssop. Pd (NC, SC, VA), Cp (NC, VA), Mt (GA, NC, VA): woodlands and forests, generally over calcareous or mafic rocks; uncommon (GA Special Concern, NC Watch List). July- September; September-October. VT west to MN, south to nw. GA and OK. In our area, this species occurs mostly in the Piedmont. [= RAB, C, F, G, K, S, W] Agastache scrophulariifolia (Willdenow) Kuntze, Purple Giant-hyssop. Mt (GA, NC, VA), Pd (NC, VA): rich woodlands and forests, bottomlands; uncommon (GA Special Concern). July-September; September-October. VT west to MN, south to NC, e. TN, n. GA, and e. KS. [= K; = A. scrophulariaefolia – RAB, C, G, S, W, an orthographic variant; > A. scrophulariaefolia var. scrophulariaefolia – F; > A. scrophulariaefolia var. mollis (Fernald) Heller – F]

* Agastache foeniculum (Pursh) Kuntze, Lavender Giant-hyssop, native of w. North America, is cultivated as an ornamental and naturalized in scattered locations in PA (Rhoads & Klein 1993), KY (Kartesz 1999), and elsewhere. [= C, F, G, K]

Ajuga Linnaeus (Bugle, Bugleweed)

A genus of about 40-50 species, herbs, of the temperate Old World. References: Harley et al. in Kadereit (2004)

1 Leaves deeply cleft into narrow segments; corolla yellow; annual...... A. chamaepitys 1 Leaves entire to shallowly lobed; corolla blue (to white or pink); perennial. 2 Plants not stoloniferous; stems hairy all around...... [A. genevensis] 2 Plants stoloniferous; stems hairy in lines ...... A. reptans

* Ajuga chamaepitys (Linnaeus) Schreber, Yellow Bugle, Ground-pine Bugle. Cp (VA): disturbed areas; rare, introduced. May-September. [= C, F, G, K] * Ajuga reptans Linnaeus, Carpet Bugle. Pd, Mt (NC, VA), Cp (VA): lawns and roadsides; commonly cultivated, uncommonly established. March-June. [= RAB, C, F, G, K]

LAMIACEAE 447 * Ajuga genevensis Linnaeus, Standing Bugle, is cultivated and rarely escaped in ne. North America, at least as far south as PA (Rhoads & Klein 1993) and MD (Kartesz 1999). [= C, F, G, K]

Blephilia Rafinesque 1819 (Woodmint, Pagoda-plant)

A genus of 3 species, herbs, of e. North America. References: Simmers & Kral (1992)=Z; Harley et al. in Kadereit (2004).

1 Stem glabrate below the middle; leaf lower surface glabrous or with a few unicellular hairs on the midvein; [of moist forests over limestone in n. AL] ...... [B. subnuda] 1 Stem strongly pubescent below the middle; leaf lower surface distinctly pubescent, at least on the larger nerves; [of various moist to dry forests, woodlands, and meadows, collectively widespread in our area]. 2 Lobes of the lower lip of the calyx linear, approaching the sinuses of the upper lip; outer bracteoles acute; leaves with rounded to acutish tips (rather Prunella-like); petioles 1-7 (-12) mm long; stem canescent, rarely with intermixed long trichomes; [primarily in the Piedmont] ...... B. ciliata 2 Lobes of the lower lip of the calyx deltoid, not reaching the sinuses of the upper lip; outer bracteoles long-acuminate; leaves with acuminate to acute tips (rather Monarda-like); petioles 9-42 mm long; stem densely to sparsely pubescent with long, spreading trichomes; [primarily in the Mountains) ...... B. hirsuta

Blephilia ciliata (Linnaeus) Bentham. Pd, Mt (GA, NC, SC, VA), Cp (NC, SC, VA): woodlands, meadows, forests, usually in circumneutral soils (over diabase, limestone, etc.); uncommon. May-early July; August-October. MA and WI south to c. GA and AR. [= RAB, C, F, G, K, S, W, Z] Blephilia hirsuta (Pursh) Bentham. Mt, Pd (NC, VA), {GA}: rocky or alluvial forests, montane forests up to at least 5000 feet elevation; common (VA Watch List). Late June-October; August-November. Québec and MN south to NC, AL, AR, and e. TX. [= RAB, C, G, S, W, Z; > B. hirsuta var. hirsuta – F, K]

Blephilia subnuda R.W. Simmers & Kral is endemic (so far as is known) to the Cumberland Plateau of nc. AL (Jackson and Madison counties). [= K, Z]

Calamintha (see Clinopodium)

Callicarpa Linnaeus 1753 (Beautyberry)

A genus of about 140 species, small trees, shrubs, and lianas, mainly tropical and subtropical. References: Moldenke (1980)=Z; Harley et al. in Kadereit (2004).

1 Leaves 7-23 cm long, stellate-scurfy beneath; peduncles 1-5 mm long...... C. americana 1 Leaves 2-6.5 (-7) cm long, glabrous or nearly so beneath (except on the midrib); peduncles 10-20 mm long. 2 Inflorescence supra-axillary, diverging from the twig 1-4 mm above the leaf axil ...... C. dichotoma 2 Inflorescence axillary, borne directly in the axil of the leaf ...... C. japonica

Callicarpa americana Linnaeus, Beautyberry, American Beautyberry, French-mulberry. Cp (GA, NC, SC, VA), Pd, Mt (GA, NC, SC): maritime forests, other forests (especially with sandy or rocky soils), disturbed areas; common (rare in Mountains). June-July; August-October (persisting into the winter). MD and AR south to FL, TX, Mexico; West Indies. [= RAB, C, F, G, K, S, W] * Callicarpa dichotoma (Loureiro) K. Koch, Chinese Beautyberry. Pd (NC, VA), Cp, Mt (NC), {SC}: roadsides, powerline rights-of-way, woodland edges, suburban woodlands, bogs; uncommon, native of Asia. September-November. This species is beginning to spread more rapidly in the Southeast. [= RAB, C, K] * Callicarpa japonica Thunberg, Japanese Beautyberry. Pf (NC): suburban woodlands; rare, native of e. Asia. Reported for Durham Co., NC by Moldenke (1980); corroborated by specimens from Orange County, NC in 2005. [= K, Z]

Chaiturus Willdenow 1787

* Chaiturus marrubiastrum (Linnaeus) Reichenbach, Horehound Motherwort. Mt, Pd (VA): disturbed areas; rare, introduced from Europe and n. Asia. June-September. [= K; = Leonurus marrubiastrum Linnaeus – C, F, G, S]

Clerodendrum Linnaeus (Glory-bower)

LAMIACEAE 448 A genus of about 400-500 species, trees and shrubs, mostly tropical and warm temperate, e. and w. hemispheres. References: Steane et al. (1999); Hsiao & Lin (1995); Steane, de Kok, & Olmstead (2004); Harley et al. in Kadereit (2004).

* Clerodendrum bungei Steudel. Pd (GA, SC), Cp (GA): roadsides and suburban woodlands; rare, introduced from e. Asia. August-September. First reported from South Carolina by Hill & Horn (1997); also reported for our area by W. Duncan (pers. comm.). [= K] * Clerodendrum indicum (Linnaeus) Kuntze, Tubeflower, Turk's-turban. Cp (GA, SC): disturbed areas, roadsides; rare, introduced from the Malaysian Archipelago. August-October; November-December. [= K; = Clerodendron indicum – RAB, orthographic variant] * Clerodendrum trichotomum Thunberg var. ferrugineum Nakai, Harlequin Glory-bower. Pd, Mt (NC), {GA}: roadsides, streambanks; rare, cultivated and strongly naturalized, native of e. Asia. [= K]

* Clerodendrum chinense (Osbeck) Mabberley, Stickbush, is cultivated and naturalized in FL, including the Panhandle (Escambia County) (Wunderlkin & Hansen 2004). [= K; ? C. japonicum (Thunberg) Sweet var. pleniflorum (Schauer) Maheshwari] * Clerodendrum japonicum (Thunberg) Sweet is also cultivated and may be encountered in our area. It is naturalized in MD (Staff of the Bailey Hortorium 1976). [= K]

Clinopodium Linnaeus 1753 (Calamint)

A genus of about {??} 20 species (as here circumscribed), herbs and shrubs, of temperate and subtropical areas of the w. and e. hemispheres. References: Cantino & Wagstaff (1998)=Y; Shinners (1962a)=Z; Shinners (1962f)=X. Key adapted in part from Z.

1 Plant a shrubby perennial, not flowering the first year; [of sandy or rocky habitats of the Coastal Plain and Piedmont, from s. NC southward]. 2 Corolla bright scarlet, 27-50 mm long; calyx 8-18 mm long ...... C. coccineum 2 Corolla light lavender or pink with darker spots, 10-20 mm long; calyx 6.0-7.5 mm long. 3 Leaves ovate or elliptic, sharply serrate, not revolute; leaves distinctly petioled; leaf surfaces glabrous ...... C. georgianum 3 Leaves linear to linear-elliptic, entire, strongly revolute; leaves subsessile; leaf surfaces minutely and densely pubescent ...... C. ashei 1 Plant an herbaceous to suffrutescent perennial, often flowering the first year; [of various habitats, collectively widely distributed in our area]. 4 Stem glabrous or pubescent at the nodes only; leaves of flowering stems linear to oblanceolate; [native, of limestone glades, barrens, and bluffs]. 5 Plant stoloniferous, bearing leafy stolons with ovate leaves; leaves of the flowering stems 1-2 cm long, 1-5 mm wide, entire...... C. arkansanum 5 Plant not bearing leafy stolons; leaves of the flowering stems oblanceolate, 2.5-5 cm long, 5-17 mm wide, with several teeth on each side...... [C. glabellum] 4 Stem pubescent; leaves of flowering stems elliptic to ovate; [alien or native, generally of disturbed or weedy situations]. 6 Axillary flower clusters sessile, dense. 7 Whorls with 8 or fewer flowers; calyx 4.5-7 mm long; corolla 7-10 mm long ...... C. acinos 7 Whorls with > 8 flowers; calyx 7-10 mm long; corolla 12-22 mm long ...... C. vulgare 6 Axillary flower clusters peduncled, contracted cymes. 8 Calyx 6.0-10.2 mm long, the hairs inside the throat barely or not exserted; blades of larger stem leaves 2-5 cm long...... C. ascendens 8 Calyx 2.8-6.0 mm long, the hairs inside the throat exserted, prominent; blades of the larger stem leaves 0.8- 2.4 cm long...... C. calamintha

* Clinopodium acinos (Linnaeus) Kuntze, Mother-of-thyme, Basil-thyme. Mt (VA): cultivated, rarely escaped or persisting; rare, introduced from Europe. [= Satureja acinos (Linnaeus) Scheele – C, F, G; = Acinos arvensis (Lamarck) Dandy – K] Clinopodium arkansanum (Nuttall) House, Arkansas Calamint. Mt (VA): dry limestone glades; rare (VA Rare). Ontario west to MN, south to w. NY, nw. PA, w. VA, WV, IL, c. TN, and s. WI; also in MO, OK, AR, and TX. There appears to be confusion about the identities and distributions of this taxon and C. glabellum. [= K, Y; = Satureja glabella (Michaux) Briquet var. angustifolia (Torrey) Svenson – C, G; = Satureja arkansana (Nuttall) Briquet – F; < Calamintha arkansana (Nuttall) Shinners – GW (also see Clinopodium glabellum); = Calamintha arkansana (Nuttall) Shinners – Z; < Clinopodium glabellum (Michaux) Kuntze – S] * Clinopodium ascendens (Jord.) Samp., Common Calamint. Cp (VA): rich calcareous slope; rare, introduced from Europe. August. [= Calamintha sylvatica Bromf. ssp. ascendens (Jord.) P.W. Ball – K; ? Calamintha officinalis – Z] LAMIACEAE 449 Clinopodium ashei (Weatherby) Small, Ashe's Calamint, Ashe's Savory, Ohoopee Dunes Wild Basil. Cp (GA): xeric sandhills; rare (GA Threatened). Peninsular FL and disjunct in e. GA (Candler and Tatnall counties). [= K, S, Y; = Calamintha ashei (Weatherby) Shinners – Z; = Satureja ashei Weatherby] * Clinopodium calamintha (Linnaeus) Stace, Lesser Calamint, Basil-thyme. Mt, Pd, Cp (NC, VA), {GA}: disturbed areas; common (rarer in NC), introduced from Europe. July-October. [> Satureja calamintha (Linnaeus) Scheele var. nepeta (Linnaeus) Briquet – RAB, F, G, W; = Satureja calamintha (Linnaeus) Scheele – C; > Satureja calamintha var. calamintha – F; > Satureja calamintha var. nepetoides (Jord.) Briquet – F, G; > Satureja calamintha var. glandulosa (Riquien) Briquet – F; > Calamintha nepeta (Linnaeus) Savi ssp. nepeta – K; > Calamintha nepeta ssp. glandulosa (Riquien) P.W. Ball – K; = Clinopodium nepeta (Linnaeus) Kuntze – S; > Calamintha officinalis Moench – Z; > Calamintha nepeta (Linnaeus) Savi – Z] Clinopodium coccineum (Nuttall ex Hooker) Kuntze, Scarlet Calamint, Scarlet Wild Basil, Red Mint Shrub. Cp (GA): sandhills and flatwoods; uncommon. E. GA south to FL and west to s. MS. [= K, S, Y; = Calamintha coccinea (Nuttall ex Hooker) Bentham – Z; = Satureja coccinea (Nuttall ex Hooker) Bertolini] Clinopodium georgianum Harper, Georgia Calamint. Cp, Pd (GA, NC, SC): longleaf pine sandhills, dry rocky woodlands; rare (NC Rare). July-September. S. NC south to Panhandle FL and west to LA. [= K, S, Y; = Satureja georgiana (Harper) Ahles – RAB; = Calamintha georgiana (Harper) Shinners – Z] Clinopodium vulgare Linnaeus, Wild Basil. Mt, Pd, Cp (NC, VA): pastures, roadbanks, forests, thin soils around rock outcrops; common. July-September. Newfoundland to Manitoba, south to NC, sc. TN, and KS, scattered in the west, widespread in Europe. Plants in our area may reflect both native and introduced genotypes. [= K, S, Y, Z; = Satureja vulgaris (Linnaeus) Fritsch – RAB, C, F, G, W; > Satureja vulgaris var. vulgaris – F; > Satureja vulgaris var. diminuta (Simon) Fernald & Wiegand – F; > Satureja vulgaris var. neogaea Fernald – F; > Clinopodium vulgare var. neogaea (Fernald) C.F. Reed]

Clinopodium brownei (Swartz) Kuntze, Browne's Savory. Cp (GA, SC): floodplain forests, pondshores; rare (GA Special Concern). In sw. GA (Jones & Coile 1988). Reported for SC (Beaufort County, SC) (Daniel Payne, pers.comm. 2006, specimen at CLEMS). [= K; > Micromeria pilosiuscula (A. Gray) Small – S; > Micromeria brownei (Sw.) Bentham var. pilosiuscula A. Gray – GW, X] {not keyed at this time} Clinopodium dentatum (Chapman) Kuntze, of GA and Panhandle FL. [= K; = Satureja dentata (Chapman) Briquet; = Calamintha dentata Chapman] {not keyed at this time; add to synonymy} Clinopodium glabellum (Michaux) Kuntze. Dry-mesic to mesic shaley forests, limestone barrens of c. KY, c. TN, MO, AR, and c. AL. Reports of this for VA (Kartesz 1999) are apparently based on confusion with Clinopodium arkansanum. [= Y; = Satureja glabella (Michaux) Briquet var. glabella – C; = Clinopodium glabellum (Michaux) Kuntze – K; < Calamintha arkansana (Nuttall) Shinners – GW; < Clinopodium glabellum (Michaux) Kuntze – S; = Calamintha glabella (Michaux) Bentham] * Clinopodium gracile (Bentham) Kuntze, Slender Wild Basil, introduced in s. AL, FL, LA (Kartesz 1999; Woods, Diamond, & Searcy 2003) and MS (S.W. Leonard, pers. comm. 2005). [= K] {not keyed at this time; add to synonymy}

Collinsonia Linnaeus 1753 (Horsebalm, Richweed, Stoneroot)

A genus of about 4 species, perennial herbs, of e. North America. References: Peirson, Cantino, & Ballard (2006)=Y; Shinners (1962b)=Z; Harley et al. in Kadereit (2004). Key adapted from Y and Z.

1 Inflorescence an unbranched thyrse, the lower nodes with (3-) 6 flowers per node; floral bracts absent; pedicels flattened at base; leaves (2-) 4 (-6), the 4 upper (or only) leaves subverticillate; flowers light pink to lavender; flowering April-June; [subgenus Micheliella]...... C. verticillata 1 Inflorescence a panicle (rarely unbranched), the flowers 2 per node; floral bracts present, minute to large; pedicels not enlarged basally; leaves 6 or more, opposite; flowers cream to yellow; flowering July-September; [subgenus Collinsonia]. 2 Fertile stamens 4; fresh plants with anise scent; [of GA southward and westward]...... C. anisata 2 Fertile stamens 2; fresh plants with lemon scent; [collectively widespread in our area]. 3 Blades of the larger stem leaves 4.0-10.5 cm long, with 5-15 teeth on each margin, glabrous or hispidulous on the main veins beneath; plant from a small, rounded tuber-like crown, to 6 cm long and 5 cm in diameter...... C. tuberosa 3 Blades of the larger stem leaves 8-25 cm long, with 11-42 teeth on each margin, glabrous or variously pubescent beneath; plant from an elongate, woody, rhizome-like crown, to 15 cm long. 4 Calyx 2-5 mm long; calyx teeth lance-subulate to narrowly lanceolate; flowers 8-13 mm long...... C. canadensis 4 Calyx 4.5-7 mm long; calyx teeth broadly lanceolate; flowers 12-17 mm long...... C. punctata

Collinsonia anisata Sims, Southern Horsebalm, Anise Horsebalm. Pd, Cp (GA): rich forests; uncommon. Late July- September; September-October. C. GA south and west to Panhandle FL and west to s. MS, on the Piedmont and Coastal Plain. This species is apparently distinct, but Shinners's concept of it included hybrids with C. canadensis and aberrant C. canadensis (Peirson, Cantino, & Ballard 2006). [= Y; < Collinsonia serotina Walter – K, W, Z; < C. canadensis var. punctata (Elliott) A. Gray –F, misapplied; < C. punctata Elliott – S; ? Micheliella anisata (Sims) Briquet – S] LAMIACEAE 450 Collinsonia canadensis Linnaeus, Richweed, Northern Horsebalm. Mt, Pd, Cp (GA, NC, SC, VA): cove forests, rich forests, especially over calcareous or mafic substrates; common. Late July-September; September-October. Québec, MI, and WI south n. FL and LA. [= RAB, C, F, G, K, S, W, Z; < C. canadensis – Y (also see C. tuberosa)] Collinsonia punctata Elliott, Florida Horsebalm. Cp (GA, SC): rich woods; rare. Late August-mid October; September- October. S. SC (Barnwell County) to e. LA, on the Coastal Plain. [= Y; < Collinsonia serotina – K, Z] Collinsonia tuberosa Michaux, Stoneroot. Pd (GA, NC, SC), Mt (GA): rich forests, over calcareous or mafic substrates; rare (NC Watch List). Late July-September; September-October. C. NC west to c. TN, south to n. GA and MS (or LA?). Peirson, Cantino, & Ballard (2006) conclude that C. tuberosa should be merged into C. canadensis, a conclusion not followed here. [= RAB, K, S, W, Z; < C. canadensis – Y; = C. canadensis Linnaeus var. tuberosa (Michaux) A. Wood] Collinsonia verticillata Baldwin, Whorled Horsebalm. Pd (GA, NC, SC, VA), Mt (GA): rich forests, ranging from moist (cove) forests to rather dry oak forests over mafic or calcareous rocks; rare (NC Rare, SC Rare, VA Rare). Late April-early June; June-July. S. VA west to e. TN, south to w. NC, nw. SC, c. GA, and MS; disjunct in s. OH. The range is strangely scattered and fragmented. [= RAB, C, G, K, W, Y, Z; = Micheliella verticillata (Baldwin) Briquet – F, S]

Conradina A. Gray 1870

A genus of 6 species, shrubs and suffrutescent herbs, of temperate se. North America. References: Shinners (1962)=Z; Harley et al. in Kadereit (2004).

1 Leaves densely gray-pubescent above and below...... [C. canescens] 1 Leaves green above, glabrous or inconspicuously short-pubecsent. 2 Plants upright to 8 dm tall; calyx tube glabrous or minutely and inconspicuously puberulent; [of the Coastal Plain of Panhandle FL and s. AL] ...... [C. glabra] 2 Plants decumbent, rooting at the nodes; calyx tube densely short-pubescent and also pilose with longer glandular hairs; [of the Cumberland Plateau of KY and TN] ...... [C. verticillata]

Conradina canescens A. Gray, Gray Rosemary. Sandhills, scrub, flatwoods. January-May. Panhandle FL and s. AL west to s. MS. [= K, Z; > C. canescens – S; > C. puberula Small – S] Conradina glabra Shinners, Apalachicola Rosemary. Sandhills. Panhandle FL and s. AL. [= K, Z] Conradina verticillata Jennison, Cumberland Rosemary. Endemic to flood-scoured cobble bars of the Cumberland Plateau area of ne. TN and se. KY. It has an odor similar to rosemary, and showy purplish flowers. [= K, Z; = C. montana Small – S]

Cunila D. Royen ex Linnaeus 1759 (Stone-mint, American-dittany, Wild-oregano)

A genus of about 15 species, herbs, from e. North America to South America. References: Harley et al. in Kadereit (2004).

Cunila origanoides (Linnaeus) Britton, Stone-mint, American-dittany, Wild-oregano. Mt (GA, NC, SC, VA), Pd (NC, SC, VA), Cp (NC, VA): dry rocky slopes, other dry slopes; common (rare in NC Coastal Plain). S. NY and PA west to MO, south to c. SC, n. GA, OK, and ne. TX (Singhurst & Holmes 2004). [= RAB, C, F, G, K, W; = Mappia origanoides (Linnaeus) House – S]

Dicerandra Bentham 1830

A genus of 9 species, herbs, endemic to se. North America. References: Huck (1987)=Z; Huck (1984)=Y; Huck & Chambers (1997); Harley et al. in Kadereit (2004).

1 Corolla tubular, straight or slightly curved; superior lobe cucullate (hoodlike); stamens and style arching under the hooded upper lobe of the corolla, included or slightly exserted beyond its apex; filaments inserted at 2 levels within the corolla; odor of fresh plant cinnamon-like, spicy; [section Lecontea]. 2 Corolla tube ca. 18 mm long, the orifice ca. 2 mm wide; leaves (15-) avg. 25 (-45) mm long, linear, the margins entire; [of s. SC south through much of the Coastal Plain of GA] ...... D. odoratissima 2 Corolla tube >20 mm long, the orifice ca. 4 mm wide; leaves (19-) avg. 40 (-55) mm long, narrowly oblong, the margins often dentate; [endemic to McIntosh County, GA] ...... D. radfordiana 1 Corolla funnel-shaped, the tube geniculate; superior corolla lobe a lobed, flaring standard; stamens and style exserted, the stamens either widely flaring to the sides or declined along the lower lobe of the corolla; filaments inserted at the same level within the corolla; odor of fresh plant minty; [section Dicerandra]. 3 Cymes epedunculate; flowers nearly sessile in compact verticils; pollen white to pale yellow; anther spurs obtuse to barely acute, with domes of minute hairs ...... [D. densiflora] 3 Cymes on peduncles 3-6 mm long; flowers on pedicels (3-) avg. 5 (-9) mm long; pollen bright yellow; anther spurs acuminate, glabrous. LAMIACEAE 451 4 Leaves linear; cymes 3-flowered; corolla white to pale purple; anthers vivid yellow; [widespread in the Coastal Plain of GA south to ne. FL and s. AL]...... D. linearifolia var. linearifolia 4 Leaves narrowly rhombic; cymes 5-7-flowered; corolla purplish red to vivid purple; anthers strongly reddish brown; [of extreme s. GA south into e. Panhandle FL and ne. FL]...... D. linearifolia var. robustior

Dicerandra linearifolia (Elliott) Bentham var. linearifolia. Cp (GA): sandhills and flatwoods; uncommon. Mid September-late November. W. and ec. Coastal Plain of GA south to ne. FL and s. AL. This taxon is hexaploid. [= K, Y, Z; < D. linearifolia – S] Dicerandra linearifolia (Elliott) Bentham var. robustior R.B. Huck. Cp (GA): sandhills and flatwoods; uncommon. Late September-late November. Sc. Coastal Plain of GA (Brooks, Echols, Lowndes counties) (Huck 1987) south to e. Panhandle FL and ne. FL. This taxon is tetraploid. [= K, Y, Z; < D. linearifolia – S] Dicerandra odoratissima Harper. Cp (GA, SC): sandhills; uncommon (SC Rare). Late August-early October. S. SC south to se. GA. This taxon is tetraploid. [= RAB, K, S, Y, Z] Dicerandra radfordiana R.B. Huck, Radford's Dicerandra. Cp (GA): dry flatwoods and sandhills; rare (GA Special Concern). September-October. Endemic to e. GA (McIntosh County). This species was postulated to be a polyploid derivative of D. odoratissima by Huck (1984, 1987); later study has shown that this is not the case (Huck & Chambers 1997). Both taxa are tetraploid. [= K, Y, Z]

Dicerandra densiflora Bentham, reported for GA by Small (1933), but this report is apparently in error. Huck (1987) regards it as endemic to n. peninsular FL. October-early November. This taxon is tetraploid. [= K, S, Y, Z]

Dracocephalum Linnaeus 1753 (Dragon's-head) (also see Physostegia)

A genus of about 45-70 species, herbs, of Eurasia and North America. References: Harley et al. in Kadereit (2004).

* Dracocephalum parviflorum Nuttall, Dragon's-head. Pd (NC): cultivated ground; rare, native west of the Appalachians. May-July; July-September. [= C, F, G, K; = Moldavica parviflora (Nuttall) Britton – RAB]

Elsholtzia Willdenow 1790

A genus of about 35-40 species, herbs, of temperate e. hemisphere. References: Harley et al. in Kadereit (2004).

* Elsholtzia ciliata (Thunberg) Hylander. Mt (NC): disturbed areas; rare, introduced from Asia. First reported for NC by Leonard (1971b). [= C, F, G, K]

Galeopsis Linnaeus 1753 (Hemp-nettle)

A genus of about 10 species, herbs, of Eurasia. References: Stace (1997)=Z; Harley et al. in Kadereit (2004). Key adapted from Stace (1997).

1 Stem with soft, appressed hairs; stem not swollen at the nodes ...... [G. ladanum var. ladanum] 1 Stem with rigid, bristly hairs; stem swollen at the nodes. 2 Terminal lobe of lower lip of corolla clearly emarginate and also convex (the sides revolute); corolla 13-16 mm long...... G. bifida 2 Terminal lobe of the lower lip of the corolla entire to very slightly emarginate, essentially planar, not revolute; corolla 13-20 (-25) mm long...... [G. tetrahit]

* Galeopsis bifida Boenninghausen, Bifid Hemp-nettle. Mt (NC): streamsides, pastures, roadsides; rare, introduced from Eurasia. July-frost. [= K, Z; < G. tetrahit – RAB, S; = G. tetrahit Linnaeus var. bifida (Boenninghausen) Lejeune & Courtois – C, F, G]

* Galeopsis ladanum Linnaeus var. ladanum, Red Hemp-nettle is naturalized in ne. North America, south at least to se. PA (Rhoads & Klein 1993) and may occur in our area. [= F, K; > G. ladanum Linnaeus var. angustifolia (Ehrhart ex Hoffmann) Wallroth – C, G, misappied] * Galeopsis tetrahit Linnaeus, Common Hemp-nettle is naturalized in ne. North America and may occur in our area. Some of the material reported from our area may be this taxon. [= Z; = G. tetrahit var. tetrahit – C, F, G; > G. tetrahit var. tetrahit – K]

Glechoma Linnaeus 1753 (Gill-over-the-ground)

LAMIACEAE 452 A genus of about 4-10 species, herbs, of temperate Eurasia. References: Harley et al. in Kadereit (2004).

* Glechoma hederacea Linnaeus, Gill-over-the-ground, Ground-ivy. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): lawns, gardens, disturbed areas; common, introduced from Eurasia. Late March-June; May-July. [= C, K; = Glecoma hederacea – RAB, S, W, misspelled; > G. hederacea var. hederacea – F; > G. hederacea var. micrantha Moricand – F; > Glecoma hederacea var. parviflora (Bentham) House – G]

Hedeoma Persoon 1807 (American Pennyroyal)

A genus of about 38-42 species, herbs, of North America, Central America, and South America. References: Irving (1980)=Z; Harley et al. in Kadereit (2004).

1 Leaves elliptic, 4-11 mm wide, slightly to strongly crenate; nutlets subspherical, 0.7-1.0 mm long, 0.6-0.9 mm wide, the surface smoothish, mottled, not glaucous; [subgenus Hedeoma]...... H. pulegioides 1 Leaves linear to narrowly elliptic, 1-4 mm wide, entire; nutlets narrowly ovoid, 1.0-1.3 mm long, 0.4-0.6 mm wide, the surface areolate and strongly glaucous; [subgenus Saturejoides]. 2 Calyx teeth convergent, closing the orifice at maturity; bracteoles subtending the individual flower pedicels 1-2 mm long, about ½ as long as the pedicel; leaves (5.0-) avg. 7.7 (-11.0) mm long, (1.2-) avg. 2.2 (-4.0) mm wide, 3-5× as long as wide ...... [H. drummondii] 2 Calyx teeth spreading (the upper) to slightly convergent (th lower), not closing the orifice at maturity; bracteoles subtending the individual flower pedicels (1.5-) 2.5-6 mm long, generally as long as or longer than the pedicel; leaves (11.0-) avg. 16.4 (-21.0) mm long, (1.0-) avg. 2.2 (-3.0) mm wide, > 5× as long as wide ...... H. hispida

*? Hedeoma hispida Pursh, Rough Pennyroyal. Pd (GA, SC, VA), Mt, Cp (GA): disturbed areas, pastures, granitic flatrocks; rare, apparently adventive from further south and west. Irving (1980) shows no collections of H. hispida closer to our area than e. panhandle FL, c. AL, nc. TN, and s. OH; it may be recently arrived in our area or previously overlooked. First reported for SC by Hill & Horn (1997). [= C, F, G, K, Z] Hedeoma pulegioides (Linnaeus) Persoon, American Pennyroyal. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): dry soils of woodlands, roadbanks, woods-roads, especially common in shaly parts of the VA mountains; common (uncommon to rare in the Carolinas). Late July-October. Nova Scotia, s. Québec, s. Ontario, MI, WI, and IA south to c. SC, c. GA, and AR. The fragrant oil is apparently very similar to that of the European Pennyroyal, Mentha pulegium Linnaeus. The oil is a powerful insect repellant and insecticide, often used on pets to repel fleas. It is also poisonous to humans, however, at least in substantial quantities. It is sometimes used as a tea; native Americans are reputed to have used it as an abortion inducer. This plant should be used with great caution, if at all. [= RAB, C, F, G, K, S, W, Z]

Hedeoma drummondii Bentham. East to MS and AL, where it occurs in black belt prairies. [= K, Z]

Hyptis Jacquin 1786 (Cluster Bushmint)

A genus of about 280-300 species, herbs and shrubs, of warm temperate, subtropical, and tropical America. References: Harley et al. in Kadereit (2004).

1 Leaves lanceolate, narrowed to a narrowly cuneate, subpetiolar base; flowers borne in large, globose heads, 1.5-2.5 cm across, borne on peduncles 2-6 cm long ...... H. alata 1 Leaves ovate, narrowed to a broadly cuneate to truncate base and well-developed petiole (4-6 cm long on larger leaves); flowers borne in irregular verticillate spikes, sessile to pedunculate on peduncles 1-2 mm long ...... H. mutabilis

Hyptis alata (Rafinesque) Shinners, Musky Mint, Cluster Bushmint. Cp (GA, NC, SC): wet pine savannas, margins of swamp forests, wet powerline rights-of-way, ditches; common. Late June-September. Ne. NC south to FL, west to se. TX; also in the West Indies. [= RAB, GW, K; = H. radiata Willdenow – S] Hyptis mutabilis (A. Richard) Briquet, Tropical Bushmint. Cp (GA, SC, VA); rare, perhaps only a waif, introduced from South America. This species is naturalized in disturbed wetlands from e. GA south to s. FL, west to se. LA. [= GW, K, S]

Hyssopus Linnaeus 1753 (Hyssop)

A genus of 2-5 species, herbs, of s. Europe to c. Asia. References: Harley et al. in Kadereit (2004).

* Hyssopus officinalis Linnaeus, Hyssop. {NC} Reported for NC (see G and S); documentation not known. Introduced from Eurasia. July-October. [= RAB, C, F, G, K, S]

LAMIACEAE 453 Isanthus Rafinesque (see Trichostema)

Lamiastrum Heister ex Fabricius 1759 (Yellow Archangel)

A monotypic genus, an herb, of w. Europe to Iran, often included in Lamium. The generic name may be illegitimate. References: Mennema (1989)=Z; Harley et al. in Kadereit (2004).

* Lamiastrum galeobdolon (Linnaeus) Ehrendorfer & Polatschek, Yellow Archangel. Pd (VA): disturbed areas; rare, introduced from Europe and e. Asia. Several subspecies are recognized in Europe. [= K; = Lamium galeobdolon (Linnaeus) Linnaeus – Z; = Galeobdolon luteum Hudson]

Lamium Linnaeus 1753 (Dead-nettle, Henbit)

A genus of about 17-40 species, herbs, of n. Africa and Eurasia. References: Mennema (1989)=Z; Harley et al. in Kadereit (2004).

1 Corolla yellow; anthers glabrous; bracts present, reflexed...... [see Lamiastrum galeobdolon] 1 Corolla blue or white; anthers with tufts of hairs; bracts absent or present (if present not reflexed). 2 Perennial, with rhizomes or stolons; corolla 18-35 mm long, the tube curved; leaves all petioled; [section Lamiotypus]. 3 Corolla white; leaves not blotched with white; lower corolla lip with 2-3 teeth on each side; pollen light yellow...... L. album ssp. album 3 Corolla pinkish-purple (rarely white); leaves usually marked with white; lower corolla lip with 1 tooth on each side; pollen orange ...... L. maculatum 2 Annual, lacking rhizomes or stolons; corolla 10-18 (-20) mm long, the tube straight; leaves all petioled or upper leaves sessile and clasping. 4 Leaves subtending flower clusters sessile; [section Amplexicaule]...... L. amplexicaule var. amplexicaule 4 Leaves all petiolate; [section Lamium]. 5 Leaves subtending whorls deeply serrate, with many teeth > 2 mm long; nutlets (2.5-) 2.7-3.0 (-3.3) mm long ...... L. dissectum 5 Leaves subtending whorls crenate-serrate, with teeth < 2 mm long; nutlets (2.0-) 2.2-2.5 (-2.8) mm long ...... L. purpureum

* Lamium album Linnaeus ssp. album, White Dead-nettle, Snowflake. (VA): disturbed areas; rare, introduced from Eurasia. April-September. Reported from our area (VA) by many earlier manuals; not documented in Harvill et al. (1992). [= Z; < L. album – C, F, G, K] * Lamium amplexicaule Linnaeus var. amplexicaule, Henbit, Henbit Dead-nettle. Cp, Pd, Mt (GA, NC, SC, VA): lawns, fields, roadsides, disturbed areas, gardens, pastures; common, introduced from Eurasia and n. Africa. January-December. [= Z; < L. amplexicaule – RAB, C, F, G, K, W] * Lamium dissectum With., Cutleaf Dead-nettle. Mt (NC), Pd (VA): lawns, fields, roadsides, disturbed areas; rare, introduced from Eurasia. April-May. This taxon is apparently an allopolyploid derivative (2n=36), resulting from hybridization of L. purpureum and another species, perhaps L. amplexicaule. Because of its allopolyploid status, this taxon should not be treated as a variety of L. purpureum. It is, however, possible that some individuals identified here may be sterile hybrids (2n=18). [= L. hybridum Villars – RAB, C, F, G, misapplied; = L. purpureum Linnaeus var. incisum (Willdenow) Persoon – K, Z] * Lamium maculatum Linnaeus, Spotted Dead-nettle. Pd, Mt (NC?, VA): lawns, fields, roadsides, disturbed areas; rare, introduced from Eurasia. April-September. [= RAB, C, F, G, K, Z] * Lamium purpureum Linnaeus, Red Dead-nettle, Purple Dead-nettle. Mt, Pd (GA, NC, SC, VA), Cp (NC, VA): lawns, fields, roadsides, disturbed areas, pastures; common, introduced from Eurasia. March-October. Other varieties are found in the Old World. [= RAB, C, F, G, W; = L. purpureum var. purpureum – K, Z]

Leonotis (Persoon) R. Brown 1810 (Lion's-ears)

A genus of about 9 species, herbs, shrubs, and small trees, of sub-Saharan Africa. References: Iwarsson & Harvey (2003)=Z.

* Leonotis nepetifolia (Linnaeus) Aiton f. var. nepetifolia, Lion's-ears, Lightning-rod-plant. Cp, Pd (GA, NC, SC): pastures, disturbed areas; uncommon, introduced from s. Africa. Late August-October. [= Z; < L. nepetifolia – K; < L. nepetaefolia – RAB, S, orthographic variant]

Leonurus Linnaeus 1753 (Motherwort) LAMIACEAE 454 (also see Chaiturus)

A genus of 25 species, herbs, of temperate Eurasia. Though L. marrubiastrum and L. sibiricus are documented in our area only from VA, they are also documented from south of our area; they likely will be found to occur in all four states.

1 Calyx strongly 5-angled, the lower 2 lobes deflexed; upper corolla lip white-villous; leaves lacerately toothed and the larger shallowly lobed...... L. cardiaca 1 Calyx slightly 5-angled, no lobes notably deflexed; upper corolla lip with densely and finely puberulent; leaves either entire to few-toothed (but not lobed) or deeply 3-parted, the 3 divisions further lacerately toothed or lobed. 2 Leaves entire to few-toothed (but not lobed) ...... [see Chaiturus marrubiastrum] 2 Leaves deeply 3-parted, the 3 divisions further lacerately toothed or lobed ...... L. sibiricus

* Leonurus cardiaca Linnaeus, Motherwort, Lion's-tail. Pd (SC, VA), Mt (GA, VA), Cp (VA), {NC}: roadsides, pastures, disturbed areas; common, introduced from c. Asia. May-August; July-October. Nelson (1993) reports the occurrence of this species in SC. [= RAB, C, F, G, S, W; ? L. cardiaca ssp. cardiaca – K] * Leonurus sibiricus Linnaeus, Siberian Motherwort. Cp (VA): disturbed areas; rare, introduced from Asia. May-September. [= C, F, G, K, S]

Lycopus Linnaeus 1753 (Bugleweed, Water-horehound)

A genus of about 10-14 species, herbs, of temperate Eurasia, North America, and Australia. References: Sorrie (1997)=Z; Harley et al. in Kadereit (2004). Key adapted from Sorrie.

1 Calyx lobes acute at the apex, shorter than or equaling the nutlets. 2 Plant without tubers; leaf base tapered to a long, winged petiole; corolla lobes 4, erect; leaf teeth (6-) avg. 8.6 (-11) per side...... L. virginicus 2 Plant usually with tubers; leaf base subsessile or tapered to a short, winged petiole; corolla lobes 4 or 5, all or some spreading; leaf teeth (2-) avg. 5.0 (-7) per side. 3 Corolla lobes 4, one erect and three spreading; nutlet tubercles well-developed, deeply toothed; leaf teeth (2-) avg. 4.5 (-6) per side; [of the fall-line sandhills of NC and SC] ...... L. cokeri 3 Corolla lobes 5, spreading; nutlet tubercles weakly developed, undulate; leaf teeth (4-) avg. 5.4 (-7) per side; [of the Mountains, upper Piedmont, and VA Coastal Plain]...... L. uniflorus 1 Calyx lobes acuminate to subulate-tipped, much exceeding the nutlets. 4 Nutlet tubercles not developed or only weakly so. 5 Calyx 2.0-3.3 mm long; stems and branches glabrous to sparsely pubescent with hairs < 0.5 mm long; leaf teeth sharply acute to short-acuminate...... L. americanus 5 Calyx 3.0-4.5 mm long; stems and branches densely to sparsely pubescent with hairs 0.5-1.6 mm long; leaf teeth blunt to acute...... L. europaeus 4 Nutlet tubercles well developed. 6 Leaves evidently petiolate, the petioles narrowly winged...... L. rubellus 6 Leaves sessile or subsessile. 7 Leaves ovate to lanceolate, usually rounded at the base, scarcely reduced upward on the stem ...... L. amplectens 7 Leaves lanceolate to linear, cuneate at the base, upper leaves conspicuously narrower (and often also shorter) than the lower leaves...... L. angustifolius

Lycopus americanus Muhlenberg ex W. Barton, American Bugleweed. Cp (NC, SC, VA), Pd (NC, VA), Mt (GA, VA): marshes, bottomlands; common (GA Special Concern). June-November. Newfoundland west to British Columbia, south to FL and CA. [= RAB, C, GW, K, S, W, Z; > L. americanus var. americanus – F, G; > L. americanus var. longii Benner – F, G; > L. americanus var. scabrifolius Fernald – F] Lycopus amplectens Rafinesque, Clasping Water-horehound. Cp (GA, NC, SC), Mt? (NC?), {VA}: clay-based Carolina bays, other moist habitats; uncommon (NC Watch List). June-November. MA south to FL; disjunct inland around the Great Lakes and (allegedly) in w. NC. [= RAB, C, GW, K, W, Z; > L. amplectens var. amplectens – F, G; > L. amplectens var. pubens (Britton) Fernald – F, G; > L. pubens Britton – S; > L. sessilifolius A. Gray – S] Lycopus angustifolius Elliott, Narrowleaf Bugleweed, Southern Bog Water-horehound. Cp (GA, NC, SC, VA): bogs, marshes; uncommon (NC Watch List). June-November. Se. VA south to FL, west to e. TX, north in the interior to s. TN and s. MO. [= C, Z; = L. rubellus Moench var. angustifolius (Elliott) Ahles – RAB, GW; = L. rubellus Moench var. lanceolatus Benner – F; < L. rubellus – G, K, W] Lycopus cokeri Ahles ex Sorrie, Coker's Bugleweed, Carolina Bugleweed. Cp (NC, SC): sandhill pocosins, boggy streamheads, seepage bogs; uncommon (NC Watch List). July-November. Endemic to the fall-line sandhill region of sc. NC and SC. See Sorrie (1997) for a detailed discussion of this species. [= RAB, K, Z; < L. uniflorus Michaux – GW] LAMIACEAE 455 * Lycopus europaeus Linnaeus, Gypsywort, European Bugleweed. Cp (NC, VA): marshes, ditches; uncommon, introduced from Europe. June-November. [= RAB, C, G, K, S, Z; > L. europaeus var. europaeus – F; > L. europaeus var. mollis (Kern.) Briq. – F] Lycopus rubellus Moench, Stalked Bugleweed. Cp (GA, NC, SC, VA), Pd (NC, SC, VA), Mt (GA, VA): marshes, swamp forests, bottomlands; common. June-November. ME west to MI, south to FL and TX. [= C, S, Z; = L. rubellus var. rubellus – RAB, GW; < L. rubellus – G, K, W (also see L. angustifolius); > L. rubellus – S; > L. velutinus Rydberg – S] Lycopus uniflorus Michaux, Northern Bugleweed. Mt (NC, SC, VA), Pd (NC, VA): bogs, seeps, wet forests; common. July-October. Newfoundland west to AK, south to w. NC, AR, and CA. [= RAB, C, F, G, S, W, Z; < L. uniflorus – GW (also see L. cokeri); > L. uniflorus var. uniflorus – K] Lycopus virginicus Linnaeus, Virginia Bugleweed. Cp, Pd, Mt (GA, NC, SC, VA): swamps, bottomlands, other wet habitats; common. July-November. MA west to PA, s. IN, MO, and OK, south to n. FL and e. TX. [= RAB, C, F, G, GW, K, S, W, Z]

In the Great Lakes and St. Lawrence River regions, hybrid swarms involving L. americanus and L. europaeus are numerous (Webber & Ball 1980). However, to date there is no evidence that these species have hybridized within the Flora region.

Macbridea Elliott ex Nuttall 1818 (Birds-in-a-nest, Macbridea)

A genus of 2 species, herbs, of se. North America. References: Harley et al. in Kadereit (2004).

1 Corolla white (faintly marked with purple in the throat); leaf tips obtuse to rounded; [FL Panhandle]...... [M. alba] 1 Corolla lavender or pink; leaf tips acute; [se. NC south to s. GA] ...... M. caroliniana

Macbridea caroliniana (Walter) Blake, Carolina Birds-in-a-nest, Carolina Macbridea. Cp (GA, NC, SC): swamp forests, especially in sphagnous seepage areas away from direct flooding, savanna edges, ditches; rare (US Species of Concern, GA Special Concern, NC Proposed Threatened). July-November. Se. NC to s. GA; reported but undocumented from n. FL, AL, and MS. Apparently rare throughout its range. [= RAB, GW, K; = M. pulchra Elliott – S]

Macbridea alba Chapman, White Birds-in-a-nest, White Macbridea. Pine savannas. Panhandle FL. [= GW, K, S]

Marrubium Linnaeus 1753 (Horehound)

A genus of about 30-40 species, herbs, of Mediterranean Europe and Asia. References: Harley et al. in Kadereit (2004).

* Marrubium vulgare Linnaeus, Horehound. Mt (GA, NC, SC, VA), Pd (NC, SC, VA): fencerows, disturbed places; uncommon, introduced from Eurasia. Used for cough-syrups in folk medicine. [= RAB, C, F, G, K, S, W]

Meehania Britton 1894 (Meehania)

A genus of 2-6 species, herbs, ours in temperate e. North America, and the other species in e. Asia. References: Harley et al. in Kadereit (2004).

Meehania cordata (Nuttall) Britton, Meehania. Mt (NC, VA): moist, rocky, forested slopes; uncommon (NC Rare, VA Watch List). Late May-June; June-July. A Central and Southern Appalachian endemic: sw. PA and OH south to nw. NC and ne. TN. [= RAB, C, F, G, K, S, W]

Melissa Linnaeus 1753 (Balm)

A genus of 3-4 species, herbs, from Europe to Iran and c. Asia. References: Harley et al. in Kadereit (2004).

* Melissa officinalis Linnaeus, Lemon Balm, Common Balm. Mt, Pd (NC, SC, VA), Cp (VA): disturbed areas; rare, introduced from w. Asia. [= RAB, C, F, G, K, S, W]

Mentha Linnaeus 1753 (Mint)

A genus of about 20-25 species, herbs, of temperate Eurasia and n. North America. References: Stace (1997)=Z; Harley et al. in Kadereit (2004). Key largely adapted from C and Stace (1997).

1 Flowers in axillary verticils subtended by ordinary foliage leaves, and separated by internodes of ordinary length. LAMIACEAE 456 2 Calyx glabrous throughout, or pubescent toward the tips only; calyx 2-3.5 mm long; plants usually sterile; fresh plant usually with spearmint odor or flavor ...... M. ×gracilis 2 Calyx pubescent throughout its length; calyx 1.5-2.5 mm long; plants usually fertile; fresh plant usually with a rather unpleasant odor of flavor. 3 Leaves subtending the inflorescence mostly broadly rounded at the base; leaves of the inflorescence relatively narrow; [alien]...... M. arvensis var. arvensis 3 Leaves subtending the inflorescence mostly cuneate at the base; leaves of the inflorescence relatively broad; [native, though often in weedy situations] ...... M. arvensis var. canadensis 1 Flowers in terminal spikes or heads, the subtending leaves absent or distinctly smaller than the foliage leaves. 4 Inflorescence a terminal globose to ovoid head of 1-3 verticils. 5 Pedicels, calyx, and leaves pubescent; plants usually fertile...... M. aquatica 5 Pedicels and clayx glabrous, leaves glabrous or nearly so; plants usually sterile...... M. ×piperita var. citrata 4 Inflorescence a spike of several to many verticils. 6 Bracteal leaves much longer than the flowers, resembling the foliage leaves, but smaller or narrower...... M. ×gracilis 6 Bracteal leaves linear to laneolate, little surpassing the flowers. 7 Calyx tube glabrous; leaves glabrous, or with scattered hairs on the lower surface. 8 Petioles of the min leaves 4-15 mm long; spikes stout; plants sterile; fresh plant with peppermint odor or flavor...... M. ×piperita var. piperita 8 Petioles of the main leaves 0-3 mm long; spikes slender; plants fertile; fresh plant with spearmint odor or flavor...... M. spicata 7 Calyx tube pubescent; leaves moderately to densely hairy on the lower surface. 9 Leaves 2-3× as long as wide, broadly cuneate to rounded at the base; leaf serrations sharp; leaf surface nearly planar...... M. longifolia 9 Leaves 1-2× as long as wide, broadly roundec to subcordate at the base; leaf serrations rounded; leaf surface strongly rugose...... M. suaveolens

[Note: The distribution, habitats, phenology, and abundance of all Mentha species need herbarium investigation]

* Mentha aquatica Linnaeus, Water Mint, Lemon Mint. {NC, VA} introduced from Europe. [= C, F, G, S, Z; < M. aquatica – K (also see Mentha ×citrata)] Mentha arvensis Linnaeus var. canadensis (Linnaeus) Kuntze, Canada Mint. {NC, VA} [= C; ? M. arvensis – RAB, misapplied; ? M. gentilis Linnaeus – RAB; = M. arvensis var. villosa (Bentham) S.R. Stewart – F; > M. arvensis var. glabrata (Bentham) Fernald – G; > M. arvensis var. lanata Piper – G; = M. canadensis Linnaeus – S; = M. arvensis Linnaeus ssp. canadensis (Linnaeus) H. Hara; < M. arvensis – K] * Mentha ×gracilis Sole (pro sp.) [Mentha arvensis × spicata], Spearmint. {NC, SC, VA}: introduced from Europe. [= K, Z; ? M. cardiaca (S.F. Gray) Gerarde ex Baker – RAB; ? M. gentilis Linnaus (pro sp.) – C; > M. cardiaca – F, G; > M. gentilis Linnaeus – F] * Mentha longifolia (Linnaeus) Linnaeus, Horse Mint. {VA} Introduced from Europe. [= RAB, C, G; > M. longifolia (Linnaeus) Hudson var. longifolia – F; > M. longifolia var. undulata (Willdenow) Fiori & Paoletti – F] * Mentha ×piperita Linnaeus (pro sp.) var. citrata (Ehrhart) Briq. (pro sp.) [Mentha aquatica ×spicata], Lemon Mint. {VA} introduced from Europe. [= Z; = M. ×citrata Ehrhart – C; = M. citrata – F, G, S; < M. aquatica – K] * Mentha ×piperita Linnaeus (pro sp.) var. piperita [Mentha aquatica × spicata], Peppermint. {GA, NC, SC, VA} introduced from Europe. [= C, K, Z; = M. piperita – RAB, G, S; > M. piperita – F; > M. crispa Linnaeus – F] * Mentha ×rotundifolia (Linnaeus) Hudson [Mentha longifolia × suaveolens]. {GA, NC, SC, VA} [= C, K; = M. rotundifolia – G, S] * Mentha spicata Linnaeus, Spearmint. {GA, NC, SC, VA} introduced from Europe. [= RAB, C, F, G, K, S, Z] * Mentha suaveolens Ehrhart, Apple Mint, Pineapple Mint, Round-leaved Mint. {NC, SC} introduced from Europe. [= C, K, Z]

* Mentha arvensis Linnaeus var. arvensis, Field Mint. introduced from Europe. [= C, F, G; = M. arvensis Linnaeus – S, Z; = M. arvensis ssp. arvensis; < M. arvensis – K] * Mentha pulegium Linnaeus, European Pennyroyal, introduced in MD, PA, and NJ (Kartesz 1999). [= C, G, K] {not keyed at this time} * Mentha ×villosa Hudson (pro sp.) [Mentha spicata × suaveolens]. Introduced south to PA and KY. [= C, K; ? M. alopecuroides Hull – F]

Micromeria Bentham (see Clinopodium)

Moldavica (see Dracocephalum) LAMIACEAE 457

Monarda Linnaeus 1753 (Bergamot)

A genus of about 12-20 species, herbs, of North America. Many of our species are cultivated, especially M. didyma in various selected forms. Additional studies are needed on a number of taxonomic problems in Monarda. Most of the varieties recognized above have been considered valid by a succession of workers; they do seem to describe morphologically distinguishable (if not entirely discrete) entities which make phytogeographic sense. References: McClintock & Epling (1942)=Z; Scora (1967)=Y; Fosberg & Artz (1953)=X; Gill (1977); Prather & Keith (2003); Harley et al. in Kadereit (2004).

1 Flowers in 2-6 glomerules, terminal and at 2-5 successive nodes down the stem; stamens included; leaves lanceolate to narrowly elliptic, usually broadest near the middle and tapered to a cuneate base, (2.5-) 3-8× as long as wide. 2 Calyx lobes attenuated into a spinose awn 2-7 mm long; corolla white to pink; inner bracts subtending the flowers 4-9 mm wide, abruptly acuminate into a spinose bristle ...... M. citriodora var. citriodora 2 Calyx lobes narrowly to broadly triangular, acute or long-acuminate but not awned; corolla yellow, spotted with purple; inner bracts 8-14 mm wide, acuminate. 3 Lower leaf surface moderately to densely silvery-tomentose; stem densely villous with spreading or downwardly- curved coarse hairs, lacking coarse, horizontal bristles and short downwardly-curved hairs...... M. punctata var. villicaulis 3 Lower leaf surface pubescent mainly on the midvein and other main veins, appearing green; stem pubescent with short downwardly-curved hairs, also with coarse, horizontal bristles and/or upwardly-curved hairs. 4 Stem with many coarse horizontal bristles, also pubescent with short, downwardly-curved hairs; leaves (40-) 50-70 (-95) mm long, 10-28 mm wide (at least some over 15 mm wide), averaging ca. 3× as long as wide...... M. punctata var. arkansana 4 Stem with few or no coarse horizontal bristles, also pubescent with a mixture of upwardly-curved and downwardly-curved hairs; leaves (25-) 35-55 (80) mm long, 5-17 mm wide (the widest very rarely over 15 mm wide), averaging ca. 4× as long as wide...... M. punctata var. punctata 1 Flowers in 1 (-2) glomerule, terminal (rarely also 1 at the next node down the stem); stamens exserted; leaves ovate to ovate- lanceolate, broadest near the rounded, truncate, or subcordate base, 1.5-3 (-4)× as long as wide. 5 Corolla 30-45 mm long, scarlet-red, (3-) 4-8 mm broad at the expanded portion of the throat; [primarily of mountain seepages, streambanks, and boggy places]...... M. didyma 5 Corolla 14-33 (-36) mm long, white, lavendar, or purple, 1-3 (-4) mm broad at the expanded portion of the throat; [of various habitats, usually dryish to mesic]. 6 Leaves deltoid-ovate to ovate, 2-6 cm wide, usually ca. 2× as long as wide; orifice of the calyx glabrous to slightly hirsute with a few long hairs; upper lip of the corolla 5-8 mm long and not bearded (M. clinopodia) or 13-16 mm long and slightly bearded (M. media) near its apex; outer surface of the corolla glabrous to evenly pubescent with short curled hairs. 7 Corolla white, greenish, or pale pink, the lower lip purple-spotted; outer bracts subtending the inflorescence green or pale (rarely with a purplish midvein); upper lip of the corolla 5-8 mm long, not bearded ...... M. clinopodia 7 Corolla deep purple, the lower lip usually not spotted; outer bracts subtending the inflorescence purple to red; upper lip of the corolla 13-16 mm long and slightly bearded near its apex...... M. media 6 Leaves narrowly-deltoid, ovate-lanceolate to lanceolate, 1-4 cm wide, usually ca. 3× as long as wide; orifice of the calyx densely hirsute with numerous erect, stiff, white hairs; upper lip of the corolla prominently bearded near its apex; outer surface of the corolla evenly pubescent with short curled hairs. 8 Corolla deep purple; middle lobe of the lower corolla lip 4-6 mm long; outer bracts subtending the inflorescence reddish...... M. fistulosa var. rubra 8 Corolla lavendar, rose, or nearly white; middle lobe of the lower corolla lip 2-4 mm long; outer bracts subtending the inflorescence green (rarely the midvein only reddish). 9 Plants 10-30 cm tall; leaves subcoriaceous, glabrous, dark green, shiny; calyx 5-8 mm long, the lobes conspicuously pustulate-glandular; [of limestone glades and barrens]...... M. fistulosa var. brevis 9 Plants 30-130 cm tall; leaves herbaceous, pubescent, light to medium green, not shiny; calyx 7-11 mm long, the lobes not pustulate-glandular; [of various habitats]. 10 Pubescence of the petioles and lower leaf surface hirsute or villous, the trichomes spreading, 1-3 mm long ...... M. fistulosa var. fistulosa 10 Pubescence of the petioles and lower leaf surface canescent, the trichomes appressed (sometimes also with an admixture of longer, spreading trichomes)...... M. fistulosa var. mollis

* Monarda citriodora Cervantes ex Lagasca y Segura var. citriodora, Lemon Bergamot. Cp (GA, SC): disturbed places: rare, introduced from a native range centered in TX. June-July; July-August. [= Y; < M. citriodora – RAB, F, G; = M. citriodora ssp. citriodora var. citriodora – K; ? M. dispersa – S; = M. citriodora ssp. citriodora – Z] Monarda clinopodia Linnaeus, Basil Bergamot. Mt, Pd (NC, SC, VA): mesic, forested slopes; common. Late May- September; July-October. NJ, w. NY, and IL, south to n. GA and c. AL (some of the range perhaps accountable to cultivation). LAMIACEAE 458 There appear to be a number of chemical races in M. clinopodia which may warrant taxonomic status. [= RAB, C, F, G, K, S, W, Y, Z; = M. fistulosa Linnaeus var. clinopodia (Linnaeus) Cooperrider] Monarda didyma Linnaeus, Bee-balm, Oswego Tea. Mt (NC, SC, VA), Pd (NC, VA), Cp (NC): seepage slopes, periglacial boulderfields with abundant seepage, streambanks, boggy places, usually in strong to moderately filtered sunlight; common, rare in Piedmont, rare in Coastal Plain (where introduced only) (SC Rare). July-September; September-October. ME west to MI, south to PA and OH, and in the Appalachians south to sw. NC, se. TN, and ne. GA (part of the northern range is likely only by introduction). McClintock & Epling (1942) describe 2 forms of M. didyma: the "broad-leaved form," with leaves averaging 9.2 cm long and 5.2 cm wide and corollas averaging 35 mm long, ranging south to sc. PA and ne. WV, and the "narrow-leaved form," with leaves averaging 11.8 cm long and 4.4 cm wide and corollas averaging 39 mm long, occurring throughout the range of the species. Further study seems warranted. [= C, F, G, K, S, W, Y, Z; < M. didyma – RAB (also see M. media)] Monarda fistulosa Linnaeus var. brevis Fosberg & Artz, Smoke Hole Bergamot, Cedar Glade Bergamot. Mt (VA): limestone outcrops, cliffs, barrens, and glades, and on limestone talus; rare (US Species of Concern). June-August; July-October. Apparently endemic to w. VA and e. WV. This variety is seemingly very distinct (Kimball et al. 2002). It had been collected only a very few times prior to the work of Bartgis (1993), who found it to be a characteristic plant of limestone barrens and woodlands in localized areas in the Ridge and Valley Province of WV. It flowers about a month earlier than M. fistulosa in the vicinity (Bartgis, pers. comm.). [= X, Y; = M. fistulosa ssp. brevis (Fosberg & Artz) Scora, comb. nov. ined. – K; < M. fistulosa – W] Monarda fistulosa Linnaeus var. fistulosa, Appalachian Bergamot. Mt, Pd (NC, VA): moist wooded slopes, roadsides, woodland edges, old fields; common. June-September; August-October. CT south to sw. NC, nearly or entirely limited to the Appalachians. I have interpreted var. fistulosa and var. mollis somewhat differently than some previous workers. A more coherent geographic pattern is achieved by limiting var. fistulosa to plants with spreading hairs only. [= F, X, Y; < M. fistulosa – RAB, W; = M. fistulosa ssp. fistulosa var. fistulosa – K; < M. fistulosa var. fistulosa – C, G, Z (also see var. mollis); = M. fistulosa – S] Monarda fistulosa Linnaeus var. mollis (Linnaeus) Bentham, Eastern Bergamot. Mt, Pd (NC, SC), {GA, VA}: moist wooded slopes; common. June-September; August-October. See comments under var. fistulosa. ME west to MN, south to GA, AL, and se. TX. [= F, X, Y; < M. fistulosa – RAB, W; < M. fistulosa var. fistulosa – C, G, Z; = M. fistulosa ssp. fistulosa var. mollis (Linnaeus) Bentham – K; > M. mollis Linnaeus – S; > M. scabra Beck – S] Monarda fistulosa Linnaeus var. rubra A. Gray, Purple Bergamot. Mt (NC, VA), {GA}: moist slope forests; rare (NC Watch List). ME to NJ, and from nw. NC to n. GA, in the Appalachians. Perhaps native only in the Southern Appalachians. A problematic taxon; see M. media for comments. [= X, Y, Z; < M. fistulosa – RAB, W; = M. fistulosa ssp. fistulosa var. rubra A. Gray - K; < M. media – C, F, S] Monarda media Willdenow, Purple Bee-balm. Mt (NC), {GA, VA}: grassy balds, moist slopes, mostly at high elevations; rare (NC Watch List). July-September; September-October. VT west to IN, south to w. MD; disjunct in w. NC and sw. TN, part of the range perhaps the result of cultivation. M. media is a problematic taxon, especially in combination with M. fistulosa var. rubra. Many have suggested that M. media is the result of hybridization or introgression of M. didyma with either M. fistulosa or M. clinopodia, or both (see Scora 1967). Scora (1967) implies that M. media consists of hybrids, backcrosses, and "introgressive elements" involving all three pairwise combinations, and the three-way combination, but that M. fistulosa var. rubra is not of hybrid origin. Needed are studies of M. media, M. fistulosa var. rubra, and their possible parents which go beyond the herbarium and determine the genetics, origin, and population structure of these taxa. It seems best for the moment to recognize (or to attempt to!) M. media and M. fistulosa var. rubra in order to foster additional observation and study, hopefully leading to a more definite understanding of their taxonomic status(es). [= G, K, Z; < M. didyma – RAB; < M. media – C, F, S (also see M. fistulosa var. rubra); = M. ×media Willdenow (pro sp.) – W, Y] Monarda punctata Linnaeus var. arkansana (McClintock & Epling) Shinners, Arkansas Horse-mint. Mt (NC), Pd (GA): dryish forests over mafic rock; rare (NC Watch List). McClintock & Epling (1942) map and discuss this taxon as endemic to AR and immediately adjacent TX, but mention that "a specimen collected near Columbus, Polk County, North Carolina (Townsend, 1897) is scarcely different from subsp. arkansana." Scora (1967) treats var. arkansana as similarly endemic, though he cites (but does not map) a specimen from Cherokee County, GA and annotated (following the publication of his paper) a later collection from Polk County, NC as var. arkansana. The Polk County, NC material is manifestly var. arkansana and might be considered merely aberrant or a chance introduction, were it not for its repeated collection and the phytogeographic interest of the Blue Ridge Escarpment of Polk County, which harbors numerous Ozarkian and other Midwestern disjuncts, such as Veratrum woodii. [= Y; < M. punctata – RAB, S, W; = M. punctata ssp. punctata var. arkansana (McClintock & Epling) Shinners – K; = M. punctata ssp. arkansana McClintock & Epling – Z] Monarda punctata Linnaeus var. punctata, Eastern Horse-mint. Cp, Pd, Mt (GA, NC, SC, VA): maritime forests, dunes, roadsides, rocky or sandy woodlands; common, uncommon in Piedmont and Mountains. Late July-September; September- October. [= C, F, Y; < M. punctata – RAB, S, W; = M. punctata ssp. punctata – G, Z; = M. punctata ssp. punctata var. punctata – K] Monarda punctata Linnaeus var. villicaulis (Pennell) Palmer & Steyermark, Hairy-stem Horse-mint. Cp (NC): disturbed areas, rare, perhaps only adventive in our area. August; October. NY west to MN, south to n. IN and s. MO. [= C, F, Y; < M. punctata – RAB, S, W; = M. punctata ssp. villicaulis Pennell – G, Z; = M. punctata ssp. punctata var. villicaulis (Pennell) Palmer & Steyermark – K]

Monarda bradburiana Beck. East to c. TN (Chester, Wofford, & Kral 1997), KY, and AL. [= G, K; < M. russeliana – C, F] {not keyed at this time; synonymy incomplete} LAMIACEAE 459 Monarda russeliana Nuttall ex Sims, White Beebalm. East to AL and KY. [= G, K; < M. russeliana – C, F (also see M. bradburiana)] {not keyed at this time; synonymy incomplete}

Mosla (Bentham) Buchanan-Hamilton ex Maximowicz 1875 (Mosla)

A genus of about 10-22 species, of e. Asia. References: Harley et al. in Kadereit (2004).

* Mosla dianthera (Buchanan-Hamilton ex Roxburgh) Maximowicz, Mosla. Mt (GA, NC): disturbed areas; rare, introduced from e. Asia. August-September. This species is becoming a noxious weed west of our area (in KY and TN); it should be expected to become more widespread in our area. [= RAB, F, G, K; = Orthodon dianthera (Buchanan-Hamilton) Handel- Mazzetti – C]

Nepeta Linnaeus 1753 (Catnip, Catmint)

A genus of about 250 species, herbs, of Eurasia and n. Africa. References: Harley et al. in Kadereit (2004).

* Nepeta cataria Linnaeus, Catnip, Catmint. Mt (GA, NC, VA), Pd (NC, VA), Cp (NC, SC, VA): fencerows, barnyards, disturbed areas; uncommon, introduced from Eurasia. [= RAB, C, F, G, K, S, W]

Ocimum Linnaeus 1753 (Basil)

A genus of about 65 species, herbs and shrubs, of warm temperate and tropical areas. References: Harley et al. in Kadereit (2004).

* Ocimum basilicum Linnaeus, Basil. Cp, Pd (GA, NC, SC): commonly cultivated in gardens, rarely persistent for short times around gardens or as a waif on trash-heaps, probably not persistent; commonly cultivated, rarely persistent, introduced from tropical Asia and tropical Africa. [= C, G, K, S]

Origanum Linnaeus 1753 (Oregano, Marjoram)

A genus of about 36-40 species, herbs and dwarf shrubs, of Eurasia. References: Harley et al. in Kadereit (2004).

* Origanum vulgare Linnaeus, Wild Marjoram. Mt (NC, VA): cultivated in gardens, persistent around gardens or as a waif; commonly cultivated, rarely persistent, introduced from Eurasia. July-September. [= RAB, C, G, K, S]

Perilla Linnaeus 1764 (Perilla, Beefsteak-plant)

A genus of about 1-6 species, herbs, of s. and e. Asia. References: Harley et al. in Kadereit (2004).

* Perilla frutescens (Linnaeus) Britton, Perilla, Beefsteak-plant. Pd, Mt, Cp (GA, NC, SC, VA): moist disturbed areas; common, introduced from India. August-October; October-December. Two varieties are sometimes recognized. Var. crispa (Bentham) Deane (leaves purple above and below; leaf margins laciniate-dentate and also crisped) and var. frutescens (leaves purple below; leaf margins dentate, not crisped). These probably represent cultivars more than taxonomically distinct entities. [= RAB, C, G, S, W; > P. frutescens var. frutescens – F, K; > P. frutescens (Linnaeus) Britton var. crispa (Bentham) Deane]

Physostegia Bentham 1829 (Obedient-plant)

A genus of about 12 species, perennial herbs, of North America. References: Cantino (1982)=Z; Harley et al. in Kadereit (2004). Key adapted from Z and GW.

1 Leaves, 1 or more of them, conspicuously or inconspicuously clasping the stem. 2 Perennating buds borne directly on the primary rhizome or at the ends of short, vertical secondary rhizomes (horizontal secondary rhizomes lacking), the plant thus forming clumps 3 Most or all of the larger leaves sharply serrate; larger leaves usually < 2.5 cm wide and > 5× as long as wide ...... Ph angustifolia 3 Most or all of the larger leaves bluntly serrate to entire; larger leaves usually > 3 cm wide or < 5× as long as wide...... Ph. purpurea 2 Perennating buds borne at the ends of elongate, horizontal, secondary rhizomes, the plant thus forming clonal patches. LAMIACEAE 460 4 Flowers 22-35 mm long; larger stem leaves acute to attenuate at the tip; axis of raceme with at least some of the hairs 0.13-0.25 mm long; larger stem leaves mostly sharple serrate ...... Ph. angustifolia 4 Flowers smaller, or most of the leaves obtuse at the tip, or hairs of the raceme axis < 0.13 mm long; larger stem leaves bluntly toothed to entire. 5 Flowering calyx tube (1-) 2-4 mm long; flowers < 20 mm long ...... [Ph. intermedia] 5 Flowering calyx tube 3-7 (-8) mm long; flowers usually > 20 mm long. 6 Uppermost pair of leaves below the terminal raceme usually considerably larger than the floral bracts, the next pair of leaves down the stem (1.5-) 2.0-12.8 cm long and 0.3-2× as long as the internode above; principal stem leaves usually widest at or below the middle of the blade ...... Ph. leptophylla 6 Uppermost pair of leaves below the terminal raceme often no larger than the floral bracts, the next pair of leaves down the stem 0.4-3.2 cm long, generally 0.1-0.3× as long as the internode above; principal stem leaves usually widest at or above the middle of the blade...... Ph. purpurea 1 Leaves petiolate or sessile, none of them clasping the stem. 7 All or most of the largest leaves sharply serrate; apex of the leaves acute to attenuate. 8 Axis of raceme with at least some of the hairs 0.13-0.25 mm long; nutlets 2-3 mm long; flowering April to early July (or later if burned) ...... Ph. angustifolia 8 Axis of raceme with hairs < 0.1 mm long; nutlets usually 3-4 mm long; flowering July-October. 9 Perennating buds usually borne directly on the primary rhizome or at the ends of short, vertical secondary rhizomes (horizontal secondary rhizomes usually lacking), the plant thus forming clumps; nonglandular trichomes of the raceme axis < 0.1 (-0.13) mm long; sterile floral bracts usually present below flowers; flowers (16-) 18-37 mm long ...... Ph. virginiana ssp. praemorsa 9 Perennating buds usually borne at the ends of elongate, horizontal, secondary rhizomes, the plant thus forming clonal patches; nonglandular trichomes of the raceme axis frequently 0.15 (0.20 ) mm long; sterile floral bracts usually not present below flowers; flowers (13-) 14-28 mm long...Ph. virginiana ssp. virginiana 4 Half or more of the larger leaves bluntly toothed to entire; apex of the leaves obtuse, or acute to attenuate. 10 Calyx and rachis of the inflorescence bearing stalked glands (visible at 10× magnification or greater); nutlets 1.7- 2.0 mm long, usually warty over the surface...... [Ph. godfreyi] 10 Calyx and rachis lacking stalked glands; nutlets 2.0-3.6 mm long, smooth. 11 Uppermost pair of leaves below the terminal raceme often no larger than the floral bracts, the next pair of leaves down the stem 0.4-3.2 cm long, generally 0.1-0.3× as long as the internode above...... Ph. purpurea 11 Uppermost pair of leaves below the terminal raceme usually considerably larger than the floral bracts, the next pair of leaves down the stem (1.5-) 2.0-12.8 cm long and 0.3-2× as long as the internode above. 12 Leaves (some of them) present at or after anthesis usually petiolate, the petioles often > 2 cm long, the petiolate leaves typically the lowest and among the largest leaves present ...... Ph. leptophylla 12 Leaves present at or after anthesis usually sessile (rarely a few petiolate, but these with petioles < 2 cm long and the petiolate leaves usually not among the largest leaves present)...... Ph. virginiana ssp. virginiana

Physostegia angustifolia Fernald, Narrowleaf Obedient-plant. Cp (GA): calcareous openings; rare (GA Special Concern). Sw. GA and AL west to KS and TX. [= GW, K, Z] Physostegia leptophylla Small, Tidal Marsh Obedient-plant, Swamp Obedient-plant. Cp (GA, NC, SC, VA): bottomland hardwood forests, swamps, tidal freshwater or slightly brackish (oligohaline) marshes, rarely wet savannas (GA); uncommon (GA Threatened, SC Rare, VA Watch List). Late May-early August; June-September. Se. VA south to sc. peninsular FL, west to sw. GA and panhandle FL. P. leptophylla is a tetraploid; Cantino (1982) suggests that this species may be an allotetraploid, perhaps originating from P. purpurea × virginiana. [= C, GW, K, Z; < Dracocephalum purpureum (Walter) McClintock ex Gleason – RAB, G; > P. denticulata (Aiton) Britton – F, misapplied; > P. aboriginorum Fernald – F; > Dracocephalum leptophyllum Small – S; > Dracocephalum veroniciformis Small – S] Physostegia purpurea (Walter) Blake, Savanna Obedient-plant. Cp (GA, NC, SC): wet savannas, savanna-swamp ecotones, ditches adjacent to former pinelands; uncommon. Late May-early August; June-September. Ec. NC south to s. FL, west to sw. GA and panhandle FL. Cantino (1982) discusses clinal variation within P. purpurea. [= GW, K, Z; < Dracocephalum purpureum (Walter) McClintock ex Gleason – RAB (also see P. leptophylla); = P. obovata (Elliott) Godfrey ex Weatherby – F; = Dracocephalum denticulatum Aiton – S] Physostegia virginiana (Linnaeus) Bentham ssp. praemorsa (Shinners) Cantino, Southern Obedient-plant. Mt, Pd, Cp (NC, SC, VA), {GA}: woodlands, glades, seepages, especially over calcareous or mafic rock; common. July-October. OH west to n. IL, south to c. NC, n. FL, TX, NM, and Mexico. [= K, W, Z; < Dracocephalum virginianum Linnaeus – RAB, G, S; = P. virginiana var. arenaria Shimek – C; >< P. virginiana var. virginiana – F; >< P. virginiana var. speciosa – F; < P. virginiana – GW] Physostegia virginiana (Linnaeus) Bentham ssp. virginiana, Northern Obedient-plant. Mt, Pd, Cp (NC, SC, VA), {GA}: streambanks, seepages, marshes, grassy balds (native occurrences usually over mafic or calcareous rocks), other open or semi- open moist to wet habitats, disturbed areas, ditches; rare as a native, more common as an escape from cultivation. July-October. Native from Québec west to Manitoba, south to e. VA, nc. TN, and ne. KS; escaped elsewhere (as in most of our area). Cantino (1982) discusses ambiguous plants from a zone of intergradation between the 2 subspecies in sw. NC, n. GA, ne. AL, e. TN, and sc. KY. Moreover, garden escapes show some intermediacy between the 2 subspecies, and Cantino (1982) suggests that cultivars are likely inter-subspecific hybrids, stating "because the genetic background of modern cultivars is unknown, they cannot be LAMIACEAE 461 reasonably placed in either subspecies and should not be identified below the species level." [= K, Z; < Dracocephalum virginianum Linnaeus – RAB, G, S; = P. virginiana var. virginiana – C; >< P. virginiana var. virginiana – F; >< P. virginiana var. speciosa (Sweet) A. Gray – F; > P. virginiana var. granulosa (Fassett) Fernald – F; < P. virginiana – GW]

Physostegia godfreyi Cantino, Apalachicola Dragonhead. Wet savannas and flatwoods, adjacent ditches. Endemic to Panhandle FL. [= GW, K, Z] Physostegia intermedia (Nuttall) Engelmann & A. Gray. Il, KY, AR, and LA west to OK and TX. Also mapped as widespread in Coastal Plain of GA (Jones & Coile 1988); {investigate}. [= GW, K, Z; = Dracocephalum intermedium Nuttall]

Piloblephis Rafinesque 1838

A monotypic genus, a shrub, of se. North America. References: Harley et al. in Kadereit (2004).

Piloblephis rigida (Bartram ex Bentham) Rafinesque, Florida Pennyroyal. Cp (GA): xeric oak scrub, with Quercus myrtifolia; rare (GA Special Concern). S. GA to s. peninsular FL. [= K; = Pycnothymus rigidus (Bartram ex Bentham) Small – S; = Satureja rigida Bartram ex Bentham]

Prunella Linnaeus 1753 (Self-heal, Heal-all)

A genus of about 4-7 species, herbs, of n. temperate areas. References: Harley et al. in Kadereit (2004).

1 Upper leaves pinnatifid; flowers creamy yellow or white (rarely pale blue)...... P. laciniata 1 Upper leaves entire to obscurely toothed; flowers blue-violet (rarely pink or whitish). 2 Principal or median cauline leaves lanceolate to oblong, (2-) avg. 3 (-5)× as long as wide; leaf cuneate at the base ...... P. vulgaris var. lanceolata 2 Principal or median cauline leaves ovate to ovate-oblong, (1.5-) avg. 2 (-2.5)× as long as wide; leaf broadly rounded at the base ...... P. vulgaris var. vulgaris

* Prunella laciniata (Linnaeus) Linnaeus, Cutleaf Self-heal. Mt (NC?, VA), {GA}: disturbed areas; rare, introduced from Eurasia. [= RAB, C, G, K, S] Prunella vulgaris Linnaeus var. lanceolata (W. Barton) Fernald, American Self-heal. {Mt, Pd, Cp (NC, SC, VA): disturbed areas, pastures, roadsides, bottomland forests; other forests and woodlands; common. April-December. Additional herbarium work is needed to determine the relative ranges, distributions, habitats, and abundances of the two varieties.} Newfoundland west to AK, south to NC, SC?, TN, MO, KS, NM, AZ, and CA. [= C, F, G; < P. vulgaris – RAB, S, W; = P. vulgaris ssp. lanceolata (W. Barton) Hultén – K] * Prunella vulgaris Linnaeus var. vulgaris, Eurasian Self-heal. {Mt, Pd, Cp (NC, SC, VA): disturbed areas, pastures, roadsides, bottomland forests; other forests and woodlands; common. April-December. Additional herbarium work is needed to determine the relative ranges, distributions, habitats, and abundances of the two varieties. The possible recognition of var. hispida also needs assessment} Introduced from Eurasia. Var. hispida Bentham, considered to have been originally e. Asian, is alleged to be widespread in se. United States. It differs from P. vulgaris var. vulgaris in having the "stems, petioles, and often the lower surfaces of leaves densely villous-hispid" (vs. "only sparingly and not conspicuously pilose" – F). [< P. vulgaris – RAB, S, W; > P. vulgaris var. vulgaris – C, F, G; > P. vulgaris var. hispida Bentham – C, F, G; = P. vulgaris ssp. vulgaris – K]

Pycnanthemum Michaux 1803 (Mountain-mint, Wild-basil)

A genus of 20-25 species, herbs, of temperate North America. Pycnanthemum remains a complicated and difficult group, with speciation apparently having proceeded by allopolyploidy, autoploidy, and aneuploidy. Numerous aberrant forms and (probably) sterile hybrids complicate identification and understanding. References: Chambers (1993); Grant & Epling (1943)=Z; Chambers & Hamer (1992)=Y; Harley et al. in Kadereit (2004).

1 Leaves 1-15 mm wide (to 30 mm wide in P. setosum), mostly > 3× as long as wide (except in P. nudum); calyx lobes not tipped with a tuft of long, jointed bristles (except P. clinopodioides). 2 Longer calyx lobes 1.5-5 mm long, attenuate-aristate, stiff, whitened; [of Coastal Plain pinelands, rarely in Mountain bogs with Coastal Plain affinities]. 3 Principal stem leaves 5-15 mm wide...... P. flexuosum 3 Principal stem leaves 10-30 mm wide...... P. setosum 2 Longer calyx lobes 0.5-1.6 mm long, deltoid to narrowly triangular, not notably stiff (except in P. tenuifolium) or whitened; [widespread in our area, but mainly of the Piedmont and Mountains]. 4 Leaves 10-15 mm wide (or more often even wider, to 25 mm wide, in P. clinopodioides); longer calyx lobes 0.7- 1.6 mm long, tipped with a few long (1-3 mm) jointed bristles (P. clinopodioides) or not tipped (P. nudum). 5 Leaves 3-5× as long as wide, herbaceous; stems and leaves pubescent; [plants from NC northward]...... LAMIACEAE 462

...... P. clinopodioides 5 Leaves 1-2.5× as long as wide, coriaceous; stems and leaves glabrous; [plants from se. SC southward] ...... P. nudum 4 Leaves 1-12 (-15) mm wide; longer calyx lobes 0.5-1.5 mm long, variously pubescent but not tipped with a tuft of long jointed bristles. 6 Leaves glabrous on the lower and upper surface, with 2-3 pairs of lateral veins; stems glabrous on the faces and angles (rarely with a few small upwardly-curled hairs on the angles). 7 Leaves 5-15 mm wide, 1-2.5× as long as wide; calyx lobes and inner bracts of the inflorescence herbaceous...... P. nudum 7 Leaves 1-4 (-5.5) mm wide, 8-15× as long as wide; calyx lobes and inner bracts of the inflorescence semi-spinose, their tips subulate, thickened, and stiff ...... P. tenuifolium 6 Leaves pubescent at least on the lower surface along the midrib and main veins; leaves with 4-5 pairs of lateral veins; stems glabrous or pubescent on the faces, pubescent on the angles. 8 Stems pubescent on the angles only (or distinctly less pubescent on the faces); leaves 3-10 mm wide ...... P. virginianum 8 Stems pubescent on the faces and angles, the hairs distributed more-or-less evenly; leaves 8-12 (-15) mm wide. 9 Longer calyx teeth 1.0-1.5 mm long; bracts of the inflorescence and leaves glabrous or very sparsely pubescent on the upper surface...... P. torreyi 9 Longer calyx teeth 0.5-1.0 mm long; bracts of the inflorescence (and usually also the leaves) canescent on the upper surface ...... P. verticillatum var. verticillatum 1 Leaves broad, 15-40 mm wide, mostly 1.5-3× as long as wide; calyx lobes usually tipped with a tuft of long, jointed bristles (except P. curvipes, P. muticum, P. setosum). 10 Bracts of the inflorescence glabrous (or very sparsely pubescent) on the upper surface, the margins long-ciliate; calyx lobes and upper part (at least) of the tube with long spreading hairs (independent of the apical tufts) ...... P. montanum 10 Bracts of the inflorescence puberulent on the upper surface, the margins not ciliate; calyx lobes and tube variously glabrous or puberulent (independent of the apical tufts). 11 Calyx lobes not tipped with a tuft of long, jointed bristles. 12 Calyx lobes 1.5-3 mm long, attenuated into a subulate tip; [of the Coastal Plain] ...... P. setosum 12 Calyx lobes 0.5-1.2 mm long, triangular to narrowly triangular, acute to acuminate, but not subulate; [collectively widespread in our area]. 13 Petioles 5-15 mm long; inflorescence corymbose, loose, the branches apparent; [of dry rocky woodlands, in sw. NC, w. SC, and southward] ...... P. curvipes 13 Petioles 0-3 mm long; inflorescence capitate, tight, the branches within the clusters not apparent; [of moist habitats, widespread in our area] ...... P. muticum 11 Calyx lobes usually tipped with a tuft of long, jointed bristles. 14 Calyx not distinctly bilabiate, all of the calyx lobes about the same length, the sinuses about the same depth. 15 Longer calyx lobes 1-2 mm long; [of the Mountains] ...... P. beadlei 15 Longer calyx lobes 2.5-3 mm long; [of the Coastal Plain] ...... P. monotrichum 14 Calyx distinctly bilabiate, the lower 2 lobes 1.5-2.5× longer than the upper 3 lobes, and separated from each other and the upper 3 lobes by deeper sinuses. 16 Leaves lanceolate, (10-) 15-25 mm wide, > 3× as long as wide...... P. clinopodioides 16 Leaves ovate, 13-50 mm wide, < 3× as long as wide. 17 Leaves of the lower and middle stem with lower surfaces glabrate, glandular-punctate, similar in color to the dark green upper surface; calyx 5-7 mm long ...... P. beadlei 17 Leaves of the lower and middle stems with lower surface canescent, distinctly paler than the dark green upper surface; calyx 3-6.5 mm long. 18 Calyx lobes broadly triangular, their apices obtuse, acute, or somewhat acuminate; calyx tube > 2× as long as the longest (lower) calyx lobes. 19 Pubescence of the stem of dense, very small downwardly-curved hairs, usually mixed with scattered longer and spreading hairs...... P. incanum var. incanum 19 Pubescence of the stem of sparse, coarse, downwardly-curved hairs ...... P. incanum var. puberulum 18 Calyx lobes narrowly triangular, their apices acuminate to attenuate; calyx tube 1-2× as long as the longest (lower) calyx lobes. 20 Mericarps 0.5-1.3 mm long, with a smooth surface, glabrous or with a few short hairs at the tip...... P. loomisii 20 Mericarps 1.2-1.5 mm long, with a rugose or pitted surface, densely pubescent at the tip. 21 Stems and lower leaf surfaces canescent, the short hairs often intermixed with longer, spreading ones ...... P. pycnanthemoides var. pycnanthemoides 21 Stems and lower leaf surfaces with coarse, spreading hairs only...... P. pycnanthemoides var. viridifolium

LAMIACEAE 463 Pycnanthemum beadlei (Small) Fernald, Beadle's Mountain-mint. Mt (NC, SC, VA), Pd (NC): forests, woodland borders; uncommon (GA Special Concern, VA Watch List). August-September. A Southern Appalachian endemic: sw. VA and ne. TN south to sw. NC, nw. SC, and n. GA. A tetraploid species (n = 38), probably an allotetraploid derived from P. montanum × muticum. [= C, K, W, Y, Z; < P. incanum – RAB; = Koellia beadlei Small – S] Pycnanthemum clinopodioides Torrey & A. Gray. Cp, Pd (NC, VA): forests, woodland borders; rare (NC Watch List, VA Rare). July-September. MA south to NC, mostly on the Coastal Plain. A tetraploid species (n = 38). Probably an allotetraploid hybrid. [= C, F, K, Y, Z; < P. verticillatum – RAB; = Koellia clinopodioides (Torrey & Gray) Kuntze – S] Pycnanthemum curvipes (Greene) E. Grant & Epling, Tennessee Mountain-mint, Stone Mountain Mountain-mint. Mt (GA, NC), Pd (GA): dry rocky woodlands and rock outcrops (granite or mafic); rare (GA Special Concern). June-August. Sw. NC and se. TN south nc. GA and n. AL; disjunct in nc. TN (Chester, Wofford, & Kral 1997). A diploid species (n= 20). [= K, Y, Z; = Koellia curvipes Greene – S] Pycnanthemum flexuosum (Walter) Britton, Sterns, & Poggenburg, Savanna Mountain-mint. Cp (NC, SC, VA), Mt (NC): moist to wet pine savannas, pocosin margins, mountain bogs, seepage areas on low elevation granite domes; common (rare in Mountains). June-September; September-October. Se. VA south to n. FL, west to s. AL on the Coastal Plain; disjunct inland in bogs and rock outcrops of sw. NC with Coastal Plain affinities and in sc. TN. A diploid species (n = 18). Sometimes mistaken in vegetative condition for Eupatorium leucolepis, P. flexuosum can be distinguished by its square stem and aromatic odor. Koellia hugeri Small, alleged to differ details of the calyx, was established for the plants of bogs of the Blue Ridge; it apparently is not morphologically segregated from other variation within the species (Grant & Epling 1943). [= RAB, C, F, K, W, Y; = P. hyssopifolium Bentham – G, GW, Z; > Koellia hyssopifolia (Bentham) Britton – S; > Koellia hugeri Small – S] Pycnanthemum incanum (Linnaeus) Michaux var. incanum. Mt, Pd (NC, VA): forests and woodland borders; common (uncommon in NC). Late June-August; September-October. VT west to s. OH and s. IL, south to nc. NC, w. NC, and nc. TN. A tetraploid species (n = 38). [= F, K; < P. incanum – RAB (also see P. beadlei, P. loomisii, P. pycnanthemoides); < P. incanum – C, G, W, Y; > Koellia incana (Linnaeus) Kuntze – S; > Koellia dubia (Gray) Small – S; = P. incanum – Z; = P. incanum (Linnaeus) Michaux ssp. incanum] Pycnanthemum incanum (Linnaeus) Michaux var. puberulum (E. Grant & Epling) Fernald. Mt (NC, SC), Pd (NC): forests and woodland borders; rare. Late June-August; September-October. WV and NC south to FL and AL. A tetraploid species (n = 38). [= F, K; < P. incanum – RAB (also see P. beadlei, P. loomisii, P. pycnanthemoides); < P. incanum – C, G, W, Y; < Koellia incana (Linnaeus) Kuntze – S; = P. puberulum E. Grant & Epling – Z] Pycnanthemum loomisii Nuttall, Loomis's Mountain-mint. Pd, Mt (NC, SC, VA), Cp (NC, VA): forests and woodland borders; rare (VA Watch List). Late June-August; September-October. VA west to IL, south to n. FL. A diploid species (n = 19). [= C, K, Y, Z; < P. incanum – RAB; = P. incanum var. loomisii (Nuttall) Fernald – F; < P. pycnanthemoides var. pycnanthemoides – G; = P. incanum (Linnaeus) Michaux ssp. loomisii (Nuttall) Hamer] Pycnanthemum monotrichum Fernald. Cp (VA): sandy woodlands; rare (VA Rare). Allegedly endemic to se. VA. Perhaps only a hybrid or else likely more widespread and merely overlooked. [= F, G, K] Pycnanthemum montanum Michaux, Appalachian Mountain-mint. Mt (NC, SC, VA): balds, woodlands, forests, and forest edges; uncommon (Va Watch List). June-August; September-October. W. VA and WV south through w. NC and e. TN to nw. SC and n. GA, a Southern Appalachian endemic. A diploid species (n = 20). [= RAB, C, F, G, K, Y, Z; = Koellia montana (Michaux) Kuntze – S] Pycnanthemum muticum (Michaux) Persoon. Mt, Cp, Pd (NC, SC, VA): bogs, wet meadows, moist to wet forests; common (uncommon in Piedmont). June-August; September-October. MA west to MI and MO, south to FL and LA. A diploid, tetraploid, and hexaploid (?) species (n = 20, 40, ca. 54). [= RAB, C, F, G, GW, K, Y; = Koellia mutica (Michaux) Kuntze – S] Pycnanthemum nudum Nuttall, Smooth Mountain-mint. Cp (SC): wet pine flatwoods; rare. Se. SC south to n. FL and se. AL. Small (1933) attributes this species to NC; the documentation is unknown (and doubtful). This is a diploid species (n = 20). [= GW, K, Z; = Koellia nuda (Nuttall) Kuntze – S] Pycnanthemum pycnanthemoides (Leavenworth) Fernald var. pycnanthemoides. Mt, Pd (NC, SC, VA): forests and woodland borders; common. July-August. VA and IL south to w. SC and n. GA. A tetraploid species (n = 36). [= F, K; < P. incanum – RAB; < P. pycnanthemoides – C, Y; < P. pycnanthemoides var. pycnanthemoides – G (also see P. loomisii); < Koellia pycnanthemoides (Leavenworth) Kuntze – S; > P. tullia Bentham – Z; = P. incanum (Linnaeus) Michaux ssp. pycnanthemoides (Leavenworth) Hamer] Pycnanthemum pycnanthemoides (Leavenworth) Fernald var. viridifolium Fernald. Mt, Pd (NC, SC, VA), Cp (NC, VA): forests and woodland borders; uncommon. July-August. VA and WV south to ec. GA and AL. A tetraploid species (n = 36). [= F, G, K; < P. incanum – RAB; < P. pycnanthemoides – C, Y; > Koellia pycnanthemoides (Leavenworth) Kuntze – S; > Koellia dubia (A. Gray) Small – S; = P. viridifolium (Fernald) E. Grant & Epling – Z; P. incanum (Linnaeus) Michaux ssp. pycnanthemoides (Leavenworth) Hamer] Pycnanthemum setosum Nuttall. Cp (GA, NC, SC, VA): dry pinelands; uncommon (rare in VA) (NC Watch List, VA Rare). Mid June-August; August-October. NJ south to GA (FL?), on the Coastal Plain. See Wieboldt et al. (1998) for discussion of the taxonomy and rarity of this species. A tetraploid species (n = 38), probably an allotetraploid derived from P. flexuosum × muticum. [= RAB, C, GW, K, Y; > P. setosum – F, G; > P. umbratile Fernald – F, G; = Koellia aristata (Michaux) Kuntze – S; = P. aristatum Michaux – Z] Pycnanthemum tenuifolium Schrader. Mt, Pd, Cp (GA, NC, SC, VA): bogs, wet meadows, moist to wet forests; common. June-August; September-October. ME west to MN, KS, and OK, south to FL and TX. A diploid and tetraploid species (n = 20 and 40). [= RAB, C, F, GW, K, W, Y; = P. flexuosum – G, Z, misapplied; = Koellia flexuosa – S, misapplied] Pycnanthemum torreyi Bentham, Torrey's Mountain-mint. Mt (NC, SC, VA), Pd, Cp (VA), {GA?}: dry rocky woodlands, over mafic, ultramafic, or calcareous rocks, dry powerline rights-of-way; rare (NC Rare, VA Rare). NH west to IL, south to NC. LAMIACEAE 464 A tetraploid and hexaploid species (n = 40 and ca. 60). [= C, G, Y, Z; < P. verticillatum – RAB; = P. torrei – K, orthographic variant; > P. torrei var. torrei – F; > P. torrei var. leptodon (Gray) Boomhour – F; = Koellia leptodon (Gray) Small – S] Pycnanthemum verticillatum (Michaux) Persoon var. verticillatum. Mt (NC, SC, VA), Pd (NC, VA), Cp (VA): upland rocky woodlands; common. July-September. Var. verticillatum ranges from VT west to MI, south to NC and KY. Var. pilosum (Nuttall) Cooperrider ranges from s. Ontario west to MI and IA, south to TN, AR, and OK. It differs in having the stems thickly (vs. thinly pubescent), the lower surface of the leaves evenly pubescent (vs. pubescence chiefly restricted to the midrib). It should be sought in our area. A tetraploid species (n = 38-39). [= C, K; < P. verticillatum – RAB (also see P. clinopodioides, P. torrei); = P. verticillatum – F, G, Y, Z; > Koellia verticillata (Michaux) Kuntze – S; > Koellia leptodon (A. Gray) Small – S; < P. verticillatum – W] Pycnanthemum virginianum (Linnaeus) T. Durand & B.D. Jackson ex B.L. Robinson & Fernald, Virginia Mountain-mint. Mt (GA, NC, VA), Pd (NC, VA), Cp (VA?, NC?): wet meadows and marshes over calcareous or mafic rocks; uncommon (rare in VA Piedmont and VA Coastal Plain) (GA Special Concern, NC Watch List). June-September; September-October. ME west to ND, south to NC, nw. GA, n. AL, and OK. A tetraploid species (n = 40). [= RAB, C, F, G, GW, K, W, Y, Z; = Koellia virginiana (Linnaeus) MacMillan – S]

Pycnanthemum albescens Torrey & A. Gray, White-leaved Mountain-mint. Pd (GA): open, mesic forests; rare (GA Special Concern). Reported for NC by Small, as Koellia albescens. It is known from nc. GA (Jones & Coile 1988). [= C, F, G, K; = Koellia albescens (Torrey & A. Gray) Kuntze – S] {not keyed at this time; synonymy incomplete} Pycnanthemum floridanum E. Grant & Epling, north to e. GA. [= K] {not keyed at this time; synonymy incomplete} Pycnanthemum verticillatum (Michaux) Persoon var. pilosum (Nuttall) Cooperrider. {GA}. In c. TN, and reported from a single county in e. TN (Chester, Wofford, & Kral 1997), in se. PA (Rhoads & Klein 1993), and WV (K99). [= C, K; = P. pilosum Nuttall – F, G; = Koellia pilosa (Nuttall) Britton – S] {not keyed at this time; synonymy incomplete}

Rosmarinus Linnaeus (Rosemary)

A genus of 2-3 species, herb/shrubs, of Mediterranean Europe. Closely related to Salvia (Walker et al. 2004), and probably to be combined there. References: Harley et al. in Kadereit (2004).

* Rosmarinus officinalis Linnaeus, Rosemary. Cp, Pd (NC, SC): gardens; commonly cultivated, rarely persistent or established, introduced from Mediterranean Europe. October-December. [= K]

Salvia Linnaeus 1753 (Sage, Clary)

A genus of about 900 species, shrubs and herbs, almost cosmopolitan. Walker et al. (2004) have determined that Salvia as traditionally circumscribed is polyphyletic. References: Epling (1938)=Z; Walker et al. (2004).

1 Leaves predominantly basal. 2 Veins of the 3 upper calyx lobes parallel, the lobes themselves minute and widely-spaced (> 1 mm between the 2 lateral teeth), separated by flattish sinuses; basal leaves lobed; [native, though weedy, common throughout our area]...... S. lyrata 2 Veins of the 3 upper calyx lobes converging, the lobes themselves minute and spaced within a distance of 1 mm; basal leaves lobed or toothed; cauline leaves toothed (rarely lobed); [alien weeds, rarely naturalized in our area]. 3 Upper corolla-lip strongly arched; leaves serrate ...... S. pratensis 3 Upper corolla-lip straight; leaves lobed ...... S. verbenacea 1 Leaves predominantly cauline, not lobed. 4 Leaves lanceolate, linear, or narrowly elliptic, the base cuneate to attenuate. 5 Leaves canescent, gray; [introduced, rarely persistent from cultivation in gardens] ...... S. officinalis 5 Leaves puberulent, green; [native, of dry woodlands from sc. NC southward and westward]. 6 Stem usually with sparse, antrorse or somewhat spreading pubescence; calyx with antrorse hairs limited to major veins; flowers of mature inflorescences spaced out, most internodes elongate and ranging up to 25 (- 34) mm; [plants of Atlantic and Gulf Coastal Plain and adjacent piedmont, from south-central NC to central FL to southeast LA] ...... S. azurea var. azurea 6 Stem usually with dense, retrorse pubescence; calyx with dense antrorse pubescence; flowers of mature inflorescences densely arranged, internodes between flowers very short, only the lowermost 1-3 internodes elongate and ranging up to 12 (-17) mm; [plants of inland and prairie sites, ranging from IL, IA, NE, and eastern CO south to nw. AL, ne. MS, LA, southeastern and central TX]...... S. azurea var. grandiflora 4 Leaves rhombic-ovate, the base cordate, truncate, or broadly cuneate. 6 Petiole not clearly differentiated from the leaf blade (leaf tissue decurrent on the petiole for most or all its length); corolla blue; [native, of woodlands]...... S. urticifolia 6 Petiole clearly differentiated from the leaf blade; corolla blue, white, or scarlet; [introduced, weedy, rare]. 7 Corolla scarlet; larger leaves 3-6.5 cm long ...... S. coccinea 7 Corolla blue or whitish; leaves (8-) 12-20 cm long...... S. sclarea LAMIACEAE 465

Salvia azurea Michaux ex Lamarck var. azurea, Azure Sage. Cp, Pd (GA, NC, SC), Mt (GA): sandy or rocky woodlands; common (NC Rare). Late August-October; October-November. S. NC south to panhandle FL, west to TX. [= K; < S. azurea – RAB, S] *? Salvia coccinea P.J. Buchoz ex Etlinger, Scarlet Sage. Cp (GA, SC), Pd (GA): disturbed areas; rare, perhaps only introduced (at least in SC) from farther south and west. May-November. [= RAB, G, K, S] Salvia lyrata Linnaeus, Lyreleaf Sage. Cp, Pd, Mt (GA, NC, SC, VA): lawns, roadsides, woodlands; common. April-May; May-July. CT west to MO, south to FL and TX. A common and familiar weed. [= RAB, C, F, G, K, S, W] * Salvia officinalis Linnaeus, Garden Sage. Cp (VA): cultivated as a garden herb, rarely persistent; rare, introduced from Europe. [= C, F, G, K] * Salvia pratensis Linnaeus, Meadow Sage, Meadow Clary. Cp (VA): fields and disturbed areas; rare, introduced from Europe. [= C, F, G, K] * Salvia sclarea Linnaeus, Clary. Mt (NC?, VA): cultivated as a garden herb, rarely persistent; rare, introduced from Europe. [= C, G, K, S] Salvia urticifolia Linnaeus, Nettle-leaf Sage. Pd, Mt (GA, NC, SC, VA), Cp (GA, VA): woodlands and glades, usually over mafic or calcareous rocks; uncommon (rare in Coastal Plain). April-June; May-July. PA west to w. KY, south to SC, c. GA, and AL. Quite showy when in flower. [= RAB, C, F, G, K, S, W, Z] * Salvia verbenacea Linnaeus, Wild Clary. Mt (VA), {GA, NC?}: fields and disturbed areas; rare, introduced from Europe. [= C, G, K, S]

Salvia azurea Michaux ex Lamarck var. grandiflora Bentham. {GA}: IL, IA, NE, and eastern CO south to nw AL, ne MS, LA, southeastern and central TX. [= F, K; = S. pitcheri Torrey ex Bentham – C, G; < S. azurea – S; = S. azurea ssp. pitcheri (Torrey ex Bentham) Epling] {not keyed at this time; synonymy incomplete} Salvia chapmanii A. Gray. AL and FL. Uncertain status. [= K, S] {not keyed at this time; synonymy incomplete} * Salvia reflexa Hornemann, Lanceleaf Sage, Mintweed. In c. TN (Chester, Wofford, & Kral 1997). The apparent ascription by C of S. reflexa Hornemann to "N.C." is a typographic error for "N.D." This species is, however, sometimes adventive as far east as WV. [= C, F, G, K, Z] {not keyed at this time} * Salvia verticillata Linnaeus, Whorled Clary, is introduced as far south as scattered locations in PA (Rhoads & Klein 1993), MD, and WV (Kartesz 1999). [= C, F, G, K] {not keyed at this time}

Satureja (also see Clinopodium)

Scutellaria Linnaeus 1753 (Skullcap) (contributed by Bruce A. Sorrie and Alan S. Weakley)

A genus of about 350-360 species, herbs and shrubs, almost cosmopolitan. References: Pittman (1988)=Z; Collins (1976)=Y; Epling (1942)=X; Leonard (1892); Harley et al. in Kadereit (2004).

Identification notes: Recognizable by the "tractor seat"-shaped protuberance on the upper calyx.

1 Flowers axillary, bracts resembling stem leaves; stem leaves sessile or petioles < 4 mm. 2 Corollas 12-32 mm long ...... Sc. galericulata 2 Corollas 5-10 mm long. 3 Lower leaves hastate; plants glabrous ...... Sc. racemosa 3 Lower leaves ovate or deltoid-ovate; plants puberulent or pubescent. 4 Stems glabrate, the pubescence ascending, curled or appressed, eglandular. 5 Median leaves 10-15 mm long; corolla 6.5-9 mm long...... Sc. leonardii 5 Median leaves 20-40 mm long; corolla 8-10 mm long...... Sc. nervosa 4 Stems obviously hairy, pubescence spreading, glandular or not (or both). 6 Lower leaf surface with glandular hairs only; leaf veins tending to anastomose along leaf margins ...... Sc. australis 6 Lower leaf surface with glandular hairs or eglandular; leaf veins usually unbranched along margins...... Sc. parvula 1 Flowers in racemes, bracts much reduced (not leaf-like); stem leaf petioles > 4 mm. 7 Corolla tube glabrous within or sparsely hairy, lacking a sharply defined ring of hairs at bend of tube (non-annulate). 8 Racemes secund. 9 Corollas ca. 6 mm long; racemes terminal and axillary...... Sc. lateriflora 9 Corollas ca. 10 mm long; racemes terminal or terminating axillary branches...... Sc. saxatilis 8 Racemes not secund, flowers on more than one side of axis. LAMIACEAE 466 10 Stems and petioles with ascending hairs; at least some racemes from axillary branches; mid to upper leaves truncate basally ...... Sc. saxatilis 10 Stems and petioles with spreading or retrorse hairs; racemes terminal or in panicles; mid to upper leaves strongly cordate. 11 Margins of lower lip cleft and erose; lower lip with large lateral auricles (flabelliform) ...... [Sc. ovata ssp. bracteata] 11 Margins of lip entire; lip undulate or weakly auriculate. 12 Lower lip entirely white with a few blue spots; leaf surface smooth with sparse glandular hairs ...... Sc. ovata ssp. ovata 12 Lower lip blue with two longitudinal white bands; leaf surface rugose, usually densely glandular hairy (but may be eglandular). 13 [of the Ridge and Valley (especially shale barrens) of VA, WV, MD] ...... Sc. ovata ssp. rugosa var. rugosa 13 [of the Blue Ridge (moist talus slopes) of NC, TN] ...... Sc. ovata ssp. rugosa var. 1 7 Corolla tube with sharply defined ring of hairs (annulus) at bend of tube. 14 At least some upper leaves entire. 15 Leaves with stipitate glands ...... Sc. multiglandulosa 15 Leaves without stipitate glands. 16 Corolla glabrous, lower lip with immaculate white central band; leaf bases long-attenuate ...... Sc. glabriuscula 16 Corolla short pilose, lower lip with blue spots or lines on white central band; leaf bases cuneate to deltoid. 17 Lowest pedicels of main axis of inflorescence >4 mm, or if less, then subtending bracts < 13 mm...... Sc. arenicola 17 Lowest pedicels < 4 mm, or if more, then bracts >13 mm...... Sc. integrifolia 14 All leaves serrate or crenate. 18 Second internode below base of inflorescence stipitate glandular. 19 Corollas 24-33 mm long and upper surfaces of leaves punctate glandular ...... Sc. pseudoserrata 19 Corollas 14-23 mm long, or if longer, then upper surfaces of leaves eglandular. 20 Corollas 25-36 mm long; bracts elliptic to oblanceolate, apices acute ...... Sc. montana 20 Corollas 14-23 mm; bracts obovate to broadly oblanceolate, apices obtuse. 21 Bases of upper leaves cuneate to rounded; corollas 14-18 (-21) mm...... Sc. elliptica var. hirsuta 21 Bases of upper leaves cordate to rounded; corollas 18-23 mm...... Sc. ocmulgee 18 Second internode below base of inflorescence eglandular. 22 Corollas > 21 mm long*. 28 Stems glabrous or glabrate below inflorescence; calyces eglandular; [of the Mountains and Piedmont] ...... Sc. serrata 28 Stems canescent below inflorescence; calyces stipitate glandular or punctate glandular. 29 Lower lip with 20+ blue spots; calyces stipitate glandular; [of peninsular FL and s GA] ...... Sc. arenicola 29 Lower lip lacking blue spots; calyces punctate glandular; [of s SC-se GA; disjunct to c AL]...... Sc. mellichampii 22 Corollas < 21 mm long*. 23 Calyces densely to sparsely canescent, eglandular or with punctate glands (stipitate glands may also be present). 24 Leaves softly villous beneath; calyces and bracts eglandular ...... Sc. incana var. incana 24 Leaves glabrate, with appressed hairs on veins. 25 Stems canescent; calyces and bracts densely punctate glandular...... Sc. incana var. australis 25 Stems glabrate (rarely puberulent); calyces and bracts eglandular ...... Sc. incana var. punctata 23 Calyces pilose with spreading stipitate glandular hairs. 26 Bracts with stipitate glands; leaves eglandular ...... Sc. elliptica var. elliptica 26 Bracts without stipitate glands; leaves densely punctate glandular. 27 Corollas 19-22 mm long; [of the Mountains of AL]...... [Sc. alabamensis] 27 Corollas 11-16 mm long; [of the Coastal Plain of SC and GA]...... Sc. altamaha

*Note: in key break 22b, corollas of S. alabamensis may reach 22 mm long; its calyces are both stipitate glandular and punctate glandular, thus differing from S. arenicola and S. mellichampii. In key break 22a, corollas of S. mellichampii may be as short as 21 mm; its calyces are punctate glandular only, unlike S. incana var. australis which has both punctate glands and stipitate glands on calyces.

LAMIACEAE 467 Scutellaria altamaha Small, Altamaha Skullcap. Cp (GA), {NC, SC}: sandy deciduous forests; rare (GA Special Concern). [= K, S, Y; < S. mellichampii Small – RAB] Scutellaria arenicola Small, Sandhill Skullcap. Cp (GA): sandy scrub; rare (GA Special Concern). Endemic to GA and FL (Kartesz 1999). [= K, S, Y] Scutellaria australis (Fassett) Epling, Southern Skullcap. Pd (GA, NC, SC, VA), Cp (GA): bottomland forests; rare. VA, WV, KY, IN, IL, MO, and KS, south to Panhandle FL, LA, and e. TX. [= G, X; < S. parvula – RAB, S; = S. parvula Michaux var. australis Fassett – F, K] Scutellaria elliptica Muhlenberg ex Sprengel var. elliptica. Mt, Pd, Cp (GA, NC, SC, VA): mesic to dry forests; common. Late May-June; June-July. NY, KY and MO, south to s. GA, Panhandle FL, LA, and e. TX. [= C, F, G, K, W, Y; < S. elliptica – RAB; < S. ovalifolia – S; = S. ovalifolia ssp. mollis Epling – X] Scutellaria elliptica Muhlenberg ex Sprengel var. hirsuta (Short & Peter) Fernald. Mt (GA, NC, VA), Pd (VA): mesic to dry forests; uncommon. Late May-June; June-July. PA and MI south to n. VA, w. NC, nw. GA, s. AL, and e. TX. [= C, F, G, K, W, Y; < S. elliptica – RAB; < S. ovalifolia – S; = S. ovalifolia ssp. hirsuta (Short & Peter) Epling – X]] Scutellaria galericulata Linnaeus, Hooded Skullcap. Mt (NC, VA): spring-fed seepage; rare. The NC occurrence is based on a single specimen from the 19th century. Reported recently from MD (Steury, Tyndall, & Cooley 1996). [= C, G, K, X; > S. epilobiifolia A. Hamilton – F, S] Scutellaria glabriuscula Fernald, Georgia Skullcap. Cp (GA): {habitat}; rare. Sw. GA and FL Panhandle west through s. AL to s. MS. In GA, AL, MS, and FL. [= K, S, Y] Scutellaria incana Biehler var. incana. Pd, Cp (NC, VA): dry to mesic forests and woodlands; uncommon. NY, OH, IN, and IL, south to e. VA, c. NC, KY, w. TN, MS, and AR. [= C, F, G, K, Y; < S. incana – RAB, S; = S. incana – X] Scutellaria incana Biehler var. punctata (Chapman) C. Mohr. Mt (GA, NC, SC, VA): dry to mesic forests and woodlands; common. Southern Appalachian endemic: sw. VA south through w. NC, nw. SC, e. TN to n. GA and ne. AL. [= C, F, G, K, W, Y; < S. incana – RAB, S; S. punctata (Chapman) Leonard – X] Scutellaria integrifolia Linnaeus. Cp, Pd, Mt (GA, NC, SC, VA): wet pine savannas, seeps in forests, bottomlands, other moist sites; common. May-July; July-August. MA south to FL, west to TX, northward in the interior to OH, KY, and TN. [= C, G, GW, K, S, W, Y; > S. integrifolia var. hispida Bentham – RAB, F; > S. integrifolia var. integrifolia – RAB, F] Scutellaria lateriflora Linnaeus, Mad Dog Skullcap. Cp (NC, SC, VA), Pd, Mt (GA, NC, SC, VA): alluvial forests, bogs, seeps, marshes; common. July-November. Newfoundland west to British Columbia, south to GA and CA. [= RAB, C, F, G, GW, S, W; > S. lateriflora var. lateriflora – K] Scutellaria leonardii Epling, Shale-barren Skullcap, Glade Skullcap. Mt (GA, VA), Pd (NC, VA), Cp (VA): limestone glades, diabase barrens; rare (GA Special Concern). April-May; May-June. MA west to MI and ND, south to se. VA, nc. NC, AR, and OK. [= C, G, W, X; < S. parvula – RAB; > S. nervosa Pursh var. ambigua (Nuttall) Fernald – F; = S. parvula Michaux var. missouriensis (Torrey) Goodman & Lawson – K; = S. ambigua Nuttall – S; > S. parvula Michaux var. leonardii (Epling) Fernald – F] Scutellaria mellichampii Small, Mellichamp's Skullcap. Cp (GA, NC?, SC): sandy deciduous forests on river bluffs; rare (GA Special Concern). June; July. Se. SC (se. NC?) south to e. GA. [= S, X, Y; < S. mellichampii – RAB; = S. incana Biehler var. australis (Epling) Collins, comb. nov. ined. – K; = S. altamaha Small ssp. australis Epling] Scutellaria montana Chapman, Large-flowered Skullcap. Mt (GA): mesic hardwood (or hardwood-shortleaf pine) forests; rare (US Threatened, GA Endangered). Se. TN south to nw. GA. [= K, S, Y; = S. serrata Andrzedowski var. montana (Chapman) Penland – F] Scutellaria multiglandulosa (Kearney) Small ex Harper. Cp, Pd (SC, GA): sandhills, dry sandy bluff forests; rare. SC (Abbeville and Anderson counties) to e. GA, and in Panhandle FL. [= K, S, Y; = S. integrifolia Linnaeus var. multiglandulosa Kearney – F] Scutellaria nervosa Pursh, Bottomland Skullcap, Veined Skullcap. Pd (NC, SC, VA), Cp (NC, VA), Mt (VA), {GA}: alluvial forests, mesic forests; uncommon, rare south of VA (GA Special Concern). May-June; June-July. NY, MI, and IA, south to GA, AL, and LA. [= RAB, K, S, W; > S. nervosa var. nervosa – C, F, G; S. nervosa var. calvifolia Fernald – C, F, G] Scutellaria ocmulgee Small, Ocmulgee Skullcap. Cp (GA): bluff forests and other mesic hardwood forests; rare (GA Threatened). Endemic to e. GA. [= K, S, Y] Scutellaria ovata Hill ssp. bracteata (Bentham) Epling. Mt (GA): dry forests and woodlands; rare. MO south through AR and OK to c. TX; disjunct eastward in s. MS, c. and n. AL, and nw. GA. [= K, W, X; < S. ovata var. ovata – C, F, G; = S. ovata var. bracteata Bentham; > Scutellaria ovata Hill ssp. cuthbertii (Alexander) Epling – K, X; > S. cuthbertii Alexander – S; = S. ovata ssp. bracteata (Bentham) Epling var. bracteata – Z] {synonymy incomplete} Scutellaria ovata Hill ssp. ovata var. ovata. {GA, NC, SC, VA}. [= Z; > S. ovata ssp. ovata – K; < S. ovata – RAB, S; >< S. ovata var. ovata – C, F, G; > S. ovata var. calcarea (Epling) Gleason – C, G; > S. ovata var. versicolor (Nuttall) Fernald – C, G; = S. ovata ssp. ovata – W; > S. ovata ssp. calcarea Epling – X; > S. ovata ssp. versicolor (Nuttall) Epling – X; > Scutellaria ovata Hill ssp. venosa Epling – K, X] Scutellaria ovata Hill ssp. rugosa (Wood) Epling var. rugosa. Mt (VA): shale barrens, other dry woodlands; common. [= S. ovata var. rugosa – F; > S. ovata ssp. rugosa – K, W, X; > Scutellaria ovata Hill ssp. pseudoarguta Epling – K, X; < S. ovata – RAB, S; = S. ovata ssp. rugosa (Wood) Epling var. rugosa – Z; > Scutellaria ovata Hill ssp. virginiana Epling – K, X] Scutellaria ovata Hill ssp. rugosa (Wood) Epling var. 1, Appalachian Skullcap. Mt (GA, NC, VA): moist boulderfields at high elevations; rare. [Scutellaria arguta Buckley – C, G, K, S, W, X; = S. saxatilis Riddell var. pilosior Bentham – F; "S. ovata Hill ssp. rugosa (Wood) Epling var. arguta (Buckley) Pittman" – Z (not published)] Scutellaria parvula Michaux, Dwarf Skullcap. Pd (SC, VA). ME west to MN, south to GA and TX. In c. TN and scattered locations in e. TN (Chester, Wofford, & Kral 1997). [= G, W, X; = S. parvula var. parvula – C, F, K; < S. parvula – RAB, S] LAMIACEAE 468 Scutellaria pseudoserrata Epling. Mt, Pd (GA), {NC?, SC}: rich, rocky forests; rare. Also in e. TN (Chester, Wofford, & Kral 1997), nc. and c. GA (Jones & Coile 1988). Cultivated in Highlands, Macon Co., NC. [= K, W, X, Y] * Scutellaria racemosa Persoon, South American Skullcap. Cp (GA, SC), Pd (NC): disturbed areas, introduced from South America. Reported from FL, AL, GA, and SC by Kral (1981). Krings & Neal (2001a, 2001b) report it for Chatham Co., NC and discuss its occurrence in se. United States. [= GW, K] Scutellaria saxatilis Riddell, Rock Skullcap. Mt (GA, NC, SC, VA): June-August. [= RAB, C, G, K, S, W, X, Z; = S. saxatilis var. saxatilis – F] Scutellaria serrata Andrzedowski, Showy Skullcap, Serrate Skullcap. Mt, Pd (NC, VA), {GA, SC?}. Mid May-late June. [= RAB, C, G, K, S, W, X, Y; = S. serrata var. serrata – F]

Scutellaria alabamensis Alexander. AL (Epling 1942, Kartesz 1999). [= K, S, X, Y] Scutellaria drummondii Bentham var. drummondii, Drummond's Skullcap. Cp (GA): blackland prairies; rare. First reported for GA by Lee Echols in 2005 (pers. comm.). [= K]

Sideritis Linnaeus

A genus of about 140-150 species, herbs and shrubs, of temperate Eurasia. References: Harley et al. in Kadereit (2004).

* Sideritis romana Linnaeus, Ironwort, is introduced and naturalized as far south as PA (Rhoads & Klein 1991, Cronquist 1991) and WV (Cronquist 1991). [= C, K] {synonymy incomplete}

Stachys Linnaeus 1753 (Hedge-nettle) (contributed by John B. Nelson)

A genus of about 300 species, herbs and shrubs, mainly temperate, nearly cosmopolitan (except Australia and New Zealand). References: Nelson (1981)=Z; Nelson & Fairey (1979); Mulligan & Munro (1989); Pringle (2002); Harley et al. in Kadereit (2004).

Keying Note: This treatment will be revised substantially.

Key 1

1 Fruiting calyx lobes deltoid to broadly triangular. 2 Stem sides pubescent. 3 Petioles short to absent, blades mostly rounded to truncate; top of blooming stem stiff, erect ...... S. eplingii 3 Petioles well developed, blades commonly cordate at the base; top of blooming stem frequently flexuous, somewhat lax ...... S. cordata 2 Stem sides glabrous. 4 Leaf margins serrate or serrulate, but always with sharp teeth; nodes not bearded; stem angles with scattered, swollen-based, short, retrorse trichomes (not spreading pubescent); [plants of the Mountains or upper Piedmont]...... S. latidens 4 Leaf margins finely crenate; nodes bearded; stem angles abundantly pubescent with 3-celled, spreading hairs; [of the lower Piedmont of NC] ...... S. species 1 1 Fruiting calyx lobes lanceolate, narrowly triangular, or nearly subulate. 5 Leaf blades linear to lanceolate. 6 Stem sides with at least moderate pubescence on the highest sterile internode, moderately to densely pubescent on higher internodes; lower leaf surface abundantly pubescent to velvety-pilose...... S. pilosa 6 Stem sides without pubescence, except for internodes within the inflorescence, which may bear light villous or glandular hairs; lower leaf surface glabrous or pubescent, but not velvety. 7 Leaf blade margins entire to crenate; plants generally glabrous...... S. hyssopifolia 7 Leaf blade margins serrulate with at least a few teeth; plants glabrous or pubescent. 8 Stems strict or sparingly branched; leaves sessile or barely petioled, the blades crenate to serrate with shallow teeth...... S. aspera 8 Stems frequently branched from the upper nodes; leaves obviously petioled, the blades sharply toothed...... S. tenuifolia 5 Leaf blades wider, oblong to elliptic. 9 Petioles poorly developed, essentialy absent...... S. hispida 9 Petioles obvious, frequently 1/5 the length of the blade. 10 Leaf blades sharply serrate, nearly dentate; stem angles abundantly pubescent with spreading or somewhat retrorse, long (to 3 mm) hairs...... S. clingmanii LAMIACEAE 469 10 Leaf margins crenate to serrulate, but never dentate; stem angles glabrous or pubescent (if the latter, then with scattered, stiffish, retrorse hairs). 11 Plant producing thick, segmented, tuber-like rhizomes; stem sides and calyx usually strongly glandular; [weedy, mostly on the Coastal Plain] ...... S. floridana 11 Plant producing slender rhizomes; stem sides and calyx usually scarcely glandular, or glabrous; [not weedy, mostly of the Piedmont and Mountains]. 12 Calyx usually abundantly pubescent with stiff, eglandular hairs; fruiting calyx lobes straight ...... S. hispida 12 Calyx usually glabrous to sparingly pubescent; fruiting calyx lobes frequently curved or curling...... S. tenuifolia

Key 2 (Alternate Key)

1 Petioles obvious (at least some of those in the middle portion of the stem at least 1/5 as long as the leaf blade). 2 Calyx tubes glandular. 3 Leaf blade margins strongly serrate (dentate or nearly so); stem angles copiously pubescent with long (to 3 mm), spreading hairs ...... S. clingmanii 3 Leaf blade margins mostly crenate; stem angles glabrate or pubescent with mostly short, retrorse hairs. 4 Calyx lobes about as long as the tube, lanceolate; leaf blades never greater than 3 cm wide...... S. floridana 4 Calyx lobes shorter than the tube, deltoid; leaf blades frequently > 3 cm wide ...... S. cordata 2 Calyx not glandular, or very slightly so. 5 Calyx glabrous to sparsely pubescent; petioles well-developed, especially in shade forms...... S. tenuifolia 5 Calyx variously hairy, but at least hispidulous, frequently strongly hispid; petioles short or long. 6 Petioles usually well developed; blade margins commonly dentate...... S. clingmanii 6 Petioles frequently short to nearly absent; blade margins toothed, but never dentate ...... S. hispida 1 Petioles short or absent. 7 Leaves linear-lanceolate to narrowly lanceolate, usually widest at or near the base; leaf margins entire to crenulate, rarely serrulate, and then mostly toward the apex. 8 Corolla very pale pink to white; leaf blades abundantly pubescent below with appressed hairs, the surface soft- velvety...... S. pilosa 8 Corolla dark pink; leaf blades variously pubescent or glabrate, but never velvety. 9 Leaf blades narrow, 3-6 mm wide; plants generally glabrous to moderately pubescent; blade margins entire to obscurely crenulate...... S. hyssopifolia 9 Leaf blades broader, 5-8 mm wide; plants frequently hispidulous or at least moderately pubescent, or abundantly hairy; blade margins crenulate to serrulate ...... S. aspera 7 Leaves ovate to elliptic, widest near the center or toward the apex, oblong; leaf margins crenate to sharply serrate for nearly the entire length. 10 Stem sides commonly pubescent above ...... S. eplingii 10 Stem sides glabrous, the angles pubescent. 11 Fruiting calyx lobes lanceolate, about half as long as the calyx tube ...... S. hispida 11 Fruiting calyx lobes deltoid or triangular, shorter than half the length of the calyx tube. 12 Leaf margins serrate or serrulate, but always with sharp teeth; nodes not bearded; stem angles with scattered, swollen-based, short, retrorse trichomes (not spreading pubescent); [of the Mountains or upper Piedmont]...... S. latidens 12 Leaf margins finely crenate; nodes bearded; stem angles abundantly pubescent with 3-celled, spreading hairs; [of the lower Piedmont of NC] ...... S. species 1

Stachys aspera Michaux, Rough-leaved Hedge-nettle. Cp, Pd (GA, NC, SC, VA), Mt (VA): moist or wet sandy soil of savannas, marshes, or swamp forests; uncommon (VA Rare). June-August; August-September. [= C, G, K; = S. hyssopifolia Michaux var. ambigua A. Gray – RAB, F, GW, Z; = S. ambigua (A. Gray) Britton – S; ? S. grayana House] Stachys clingmanii Small, Clingman's Hedge-nettle. Mt (NC), Cp? (VA?), {SC?}: cove forests, especially periglacial boulderfields, mostly at high elevations (and see comments below); rare (NC Rare). June-August; September-October. A narrow Southern Appalachian endemic, known only from sw. NC and se. TN. Some plants similar to and perhaps referable to this species occur in Surry County VA (calcareous bushy thickets and ravines) and in IN. [= C, F, G, K, S, W, Z; < S. clingmanii – RAB] Stachys cordata Riddell, Heart-leaved Hedge-nettle, Nuttall's Hedge-nettle. Mt (GA, NC, VA), Pd (NC, VA): moist forests, especially alluvial bottomlands or over calcareous rocks; uncommon (rare in NC) (GA Rare, NC Watch List). June- August; September-October. Primarily montane, but extending east to Stokes County, North Carolina. See Pringle (2002) for a discussion of nomenclature. [= S; = S. nuttallii Shuttleworth ex Bentham – K, W, Z; < S. clingmanii – RAB; > S. cordata – C; > S. subcordata Rydberg – C, G; = S. riddellii House – F, G; > S. salvioides Small – S] Stachys eplingii J. Nelson, Epling's Hedge-nettle. Mt (GA, NC, SC, VA): mesic forests, bogs, wet meadows over calcareous or mafic substrates; rare (GA Rare, NC Rare, VA Rare). June-August; August-September. This species has a LAMIACEAE 470 scattered and sporadic range in the southern and central Appalachians, occurring also in the Ozarks. See Nelson & Fairey (1979) for a discussion of the nomenclatural change. [= C, GW, K, W, Z; = S. nuttallii – RAB, F, G, S, misapplied] * Stachys floridana Shuttleworth ex Bentham, Florida Betony, Rattlesnake-weed. Cp, Pd (GA, NC, SC, VA): disturbed sites, roadsides; uncommon, probably not native, introduced from Florida. April-July; May-August. The common name "Rattlesnake-weed" refers to the moniliform rhizomes. [= RAB, GW, K, S, Z] Stachys hispida Pursh, Hispid Hedge-nettle. Mt (NC, VA), Pd, Cp (VA), ?? (GA): wet meadows and mesic forests; uncommon (GA Rare). A highly variable taxon. [= C, G; = S. tenuifolia Willdenow var. hispida (Pursh) Fernald – F; < S. tenuifolia var. tenuifolia – K, Z] Stachys hyssopifolia Michaux var. hyssopifolia, Hyssop-leaved Hedge-nettle. Cp, Pd (GA, NC, SC, VA), Mt (VA): moist soils of savannas, marshes, seasonally flooded sinkhole ponds, roadside ditches; uncommon (VA Watch List). June-August; August-September. [= RAB, F, GW, Z; = S. hyssopifolia – C, G, K, S, W; ? S. atlantica Britton] Stachys latidens Small ex Britton, Broad-toothed Hedge-nettle. Mt, Pd (GA, NC, SC, VA): mesic forests in coves and on mountain slopes, mountain pastures and forest edges; common (GA Rare, SC Rare). June-August; September-October. [= RAB, C, F, G, S; = S. tenuifolia Willdenow var. latidens (Small ex Britton) J. Nelson – K, W, Z; < S. tenuifolia – GW] Stachys pilosa Nuttall, Woundwort. Mt, Pd (VA), Cp (SC): marl fens, roadsides, banks of waterfowl impoundments; rare, possibly adventive in part from further west, but some populations at least native. [= S. palustris Linnaeus var. pilosa (Nuttall) Fernald – C, F, G; = S. palustris Linnaeus ssp. pilosa (Nuttall) Epling; > S. pilosa var. arenicola (Britton) Mulligan & Monroe – K] Stachys species 1, Yadkin Hedge-nettle. Pd (NC, VA): in sandy alluvium along forest edges in river floodplain; rare. Stachys tenuifolia Willdenow, Smooth Hedge-nettle. Mt (NC), Pd (VA), Cp (NC, SC, VA): wooded alluvial river bottoms, swamp forests, and roadsides; rare (NC Watch List). June-August; September-October. [= RAB, C, G, K, S; > S. tenuifolia var. tenuifolia – F, Z; > S. tenuifolia var. perlonga Fernald – F, Z; > S. tenuifolia var. platyphylla Fernald – F; < S. tenuifolia – GW; = S. tenuifolia var. tenuifollia – W]

* Stachys annua (Linnaeus) Linnaeus. Cp (VA): disturbed area; rare, probably only a waif. Reported for VA (Kartesz 1999). [= C, F, G, K] {not keyed at this time} * Stachys arvensis (Linnaeus) Linnaeus. Reported for VA by C, G, and K; documentation uncertain (Virginia Botanical Associates 2006). [= C, F, K] {not keyed} * Stachys byzantina K. Koch ex Scheele. Pd (VA): roadside; rare, doubtfully established. Reported for VA (Virginia Botanical Associates 2006). [= C, K; = S. olympica Poiret – F, G] {not keyed at this time} Stachys crenata Rafinesque. Reported for AL, KY, FL (Kartesz 1999). [= K] * Stachys germanica Linnaeus. Mt (VA): roadsides; rare, doubtfully established. Reported for VA, TN, FL (Kartesz 1999). [= C, F, G, K] Stachys hyssopifolia Michaux var. lythroides (Small) J. Nelson. (GA Rare). E. Panhandle of FL. [= Z] * Stachys palustris Linnaeus. South to MD, PA, NJ. [= K; > S. palustris var. palustris – C, F, G]

Synandra Nuttall 1818 (Synandra)

A monotypic genus, an herb, of e. North America. References: Cantino (1985); Harley et al. in Kadereit (2004).

Synandra hispidula (Michaux) Baillon, Synandra, Gyandotte Beauty. Mt (NC, VA): moist, rich forests; rare (NC Rare, VA Rare). Late April-May; May-June. A broad Appalachian endemic: s. OH west to s. IL, south to sw. VA, w. NC, and n. AL. [= RAB, C, F, G, K, S, W]

Teucrium Linnaeus 1753 (Germander)

A genus of about 100-250 species, herbs and shrubs, nearly cosmopolitan in distribution. References: Harley et al. in Kadereit (2004).

Teucrium canadense Linnaeus var. canadense. {GA, NC, SC, VA}. [= C, F, G, K; < T. canadense – RAB, GW, W; = T. littorale Bicknell – S] Teucrium canadense Linnaeus var. hypoleucum Grisebach. {GA, NC, SC}. [= K; < T. canadense – RAB, GW, W; = T. nashii Kearney – S] Teucrium canadense Linnaeus var. occidentale (A. Gray) McClintock & Epling. (VA). Reported for VA (Kartesz 1999). {investigate} Occurs at least as far south and east as PA (Rhoads & Klein 1993). [= C, G, K; > T. occidentale A. Gray var. occidentale – F; > T. occidentale A. Gray var. boreale (Bicknell) Fernald – F] Teucrium canadense Linnaeus var. virginicum (Linnaeus) Eaton. [= C, F, G, K; < T. canadense – RAB, GW, W; = T. canadense – S]

Teucrium cubense Jacquin var. cubense. AL. [= K] {not keyed at this time; synonymy incomplete}

LAMIACEAE 471 Thymus Linnaeus (Thyme)

A genus of about 220-350 species, herbs and shrubs, of temperate Eurasia. References: Harley et al. in Kadereit (2004).

* Thymus praecox Opiz ssp. arcticus (Dur.) Jalas, Mother-of-Thyme. Pd (NC), Mt (GA): commonly cultivated and sometimes escaped or persisting; rare, introduced from Eurasia. July-September. [= K; ? Th. serpyllum Linnaeus – RAB, C, F, G, misapplied] * Thymus pulegioides Linnaeus, Lemon Thyme. Pd (VA). [= K]

Trichostema Linnaeus 1753 (Blue Curls)

A genus of about 18 species, shrubs, annual and perennial herbs, of temperate North America (especially diverse in w. North America, with a second center of diversity in se. North America). References: Weakley (in prep.)=Z; Harley et al. in Kadereit (2004).

1 Calyx lobes essentially equal; stamens straight, < 10 mm long; leaves acute to slightly acuminate, the two main lateral veins reconnecting to the midvein; [section Orthopodium]...... T. brachiatum 1 Calyx strongly bilabiate; stamens strongly arched, 12-20 mm long; leaves obtuse, the two main lateral veins not reconnecting to the midvein; [section Trichostema]. 2 Plants annual; larger leaves 3-7 cm long (including the petiole); plants with long internodes near the base, near-basal branches absent, the best-developed branches from the mid or upper stem; hairs on the upper stem long (0.5-2.0 mm long) or short (0.1-0.4 mm long); [collectively widespread, in a wide variety of habitats, primarily inland, though occasionally occurring as a weed in coastal areas]. 3 Leaves 2.5-4× as long as wide; longer hairs of the upper stem (0.3-) 0.5-2.0 mm long...... T. dichotomum 3 Leaves 5-15× as long as wide; longer hairs of the upper stem 0.1-0.3 (-0.4) mm long...... T. setaceum 2 Plants perennial; larger leaves 1-4 cm long (including the petiole); plants with short internodes near the base, near-basal branches well-developed, these often branching again; hairs on the upper stem short (0.1-0.4 mm long); [restricted to barrier islands, coastal peninsulas, and other maritime situations within 10 km of the ocean]. 4 Anthers lemon yellow; leaves ovate-rhombic, 1-1.5× as long as wide; corolla lavender (definitely with a pinkish tint); bark on older stems dark, tight; plants 1-4 dm tall, forming a tight, compact, hemispheric bush; [of maritime dunes, grasslands, and forest openings from 10 km north of Cape Hatteras (Dare County, NC) south to near Cape Romain (Georgetown County, SC] ...... T. species 1 4 Anthers blue; leaves spatulate, the petiole relatively well-developed, 1.5-3× as long as wide; corolla bluish (lacking a well-developed pinkish tint); bark on older stems yellow to tan, somewhat papery; plants 3-7 dm tall, often gangly and irregularly shaped; [of maritime dunes, grasslands, and coastal scrub from e. GA around the FL peninsula west to s. MS] ...... T. species 2

Trichostema brachiatum Linnaeus, Glade Blue Curls, False Pennyroyal. Mt (GA, NC, VA), Pd (NC, SC, VA): shale barrens, outcrops of calcareous or mafic rock, diabase barrens, calcareous dry prairies, disturbed rocky areas; uncommon (NC Rare). August-September. VT and s. Ontario west to MN and NE, south to c. NC, nw. GA, AL, TX, and AZ. Morphology, pollen, and phytogeography suggest the plausible recognition of Trichostema section Orthopodium (which includes this species andseveral from w. North America) as Isanthus, a genus distinct from section Trichostema (which includes all other eastern North American species). [= W; = Isanthus brachiatus (Linnaeus) Britton, Sterns, & Poggenburg – C, F, K, S; > Isanthus brachiatus var. brachiatus – G] Trichostema dichotomum Linnaeus, Common Blue Curls. Cp, Pd, Mt (GA, NC, SC, VA): dry woodlands, disturbed areas, thin soils around rock outcrops; common. August-November. Widespread in e. North America. [= RAB, C, K, S, W, Z; > T. dichotomum var. dichotomum – F; >< T. dichotomum var. puberulum Fernald & Griscom – F; = T. dichotomum var. dichotomum – G] Trichostema setaceum Houttuyn, Narrowleaf Blue Curls. Pd, Mt, Cp (GA, NC, SC, VA): thin soils around rock outcrops, especially granite flatrocks, dry sandy soils of the Coastal Plain; uncommon (NC Watch List, VA Rare). August-November. CT west to OH, south to FL and TX, primarily on the Coastal Plain. [= RAB, C, F, K, W, Z; = T. dichotomum var. lineare (Walter) Pursh – G; = T. lineare Walter – S] Trichostema species 1, Dune Blue Curls, Carolina Blue Curls. Cp (NC, SC): dunes on barrier islands, vegetated with perennial grasses (especially Uniola paniculata), openings in maritime scrub; rare (US Species of Concern, NC Rare). August- November. Endemic to barrier islands from slightly north of Cape Hatteras, NC south to North Island, Georgetown County, SC, north of Cape Romain. When growing together, the flowering period of T. species 1 is about 2-3 weeks later than that of T. dichotomum. Despite a considerable overlap of blooming period, only one hybrid has been seen, and that in common-garden cultivation in the Piedmont. [= Z] Trichostema species 2, Florida Blue Curls. Cp (GA): maritime dunes, grasslands, and coastal scrub; uncommon. E. GA around the FL peninsula west to s. MS; Bahamas. [= Z; >< T. dichotomum var. puberulum Fernald & Griscom – F]

Vitex Linnaeus 1753 (Chaste Tree) LAMIACEAE 472

A genus of about 250 species, trees and shrubs, tropical to temperate. References: Harley et al. in Kadereit (2004).

1 Leaves palmately compound; plant an upright small tree ...... V. agnus-castus 1 Leaves simple; plant a sprawling and spreading shrub/vine ...... V. rotundifolia

* Vitex agnus-castus Linnaeus, Chaste Tree. Cp (GA, NC, VA), Pd (GA, NC, VA), Mt (VA): pastures, woodland edges, suburban woodlands; rare, introduced from Mediterranean Europe. June-July. [= RAB, C, G, S; > V. agnus-castus var. agnus- castus – K; > V. agnus-castus var. caerulea Rehder – K] * Vitex rotundifolia Linnaeus f., Beach Vitex, Roundleaf Chaste-tree. Cp (GA?, NC, SC): coastal dunes; uncommon, planted for ornament and stabilization and now spreading aggressively as an invasive species. See Roecker & Socha (2004) for additional information. The runners are reported to reach 10 m in length. [= K]

LARDIZABALACEAE Decaisne 1839 (Lardizabala Family)

A family of about 8 genera and 35 species, shrubs and vines, primarily Asian, but also in s. South America. References: Thieret & Kartesz in FNA (1997); Cheng-Yih & Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

Akebia Decaisne 1837 (Akebia)

A genus of 5 species, vines, of temperate e. Asia. References: Cheng-Yih & Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

* Akebia quinata (Houttuyn) Decaisne, Five-leaf, Five-leaf Akebia, Chocolate-vine. Mt, Pd (NC, SC, VA), Cp (GA): escaped from cultivation to roadbanks, suburban woodlands, and floodplains; rare, native of Japan, China, and Korea, potentially invasive and difficult to eradicate. April-June; June-July. [= RAB, C, F, FNA, K]

LAURACEAE A.L. de Jussieu 1789 (Laurel Family)

A family of about 50 genera and 2500-3500 species, trees and shrubs, of tropical, subtropical, and (rarely) warm temperate regions. References: van der Werff in FNA (1997); van der Werff & Richter (1996); Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves evergreen; flowers perfect; [tribe Perseeae]. 2 Leaves glabrous, bright green, with yellow callosities in the principal vein axils; crushed leaves with the odor of camphor ...... Cinnamomum 2 Leaves pubescent to glabrate, dark green, without yellow callosities in the principal vein axils; crushed leaves with the odor of bay...... Persea 1 Leaves deciduous; flowers imperfect; [tribe Laureae]. 3 Some of the leaves with 1-2 (-5) rounded lobes; small to medium trees...... Sassafras 3 None of the leaves lobed; medium to large shrubs. 4 Leaves 4-16 cm long, 2-6 cm wide, obovate, ovate, or broadly elliptic ...... Lindera 4 Leaves 1.2-4 cm long, 0.5-1.5 (-1.9) cm wide, narrowly elliptic ...... Litsea

Cinnamomum Schaeff 1760 (Cinnamon)

A genus of about 350 species, trees and shrubs, of e. and se. Asia, Oceania, and tropical America. References: Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

* Cinnamomum camphora (Linnaeus) J. Presl, Camphortree. Cp (GA, NC, SC): disturbed areas; rare, native of eastern Asia, planted as an ornamental and rarely escaped to adjoining areas, such as in Southern Pines, Moore County, NC. April-May. Reported as escaped and apparently naturalized in South Carolina by Hill & Horn (1997). [= FNA, K; = Camphora camphora (Linnaeus) Karsten – S]

Laurus Linnaeus 1753 (Laurel, Bay)

A genus of 1-2 species, trees, of Mediterranean Europe, the Canary Islands, Madeira, and the Azores. References: Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

LAURACEAE 473 * Laurus nobilis Linnaeus, Laurel, Bay, native to the Mediterranean region of Europe and the bay leaf of commerce, is planted as an ornamental and spice, but is not known to escape in our area.

Lindera Thunberg 1783 (Spicebush, Benzoin)

A genus of about 100 species, trees and shrubs, of tropical and temperate Asia, Australia, and e. North America. References: Wofford (1983)=Z; Steyermark (1949); McCartney, Wurdack, & Moore (1989); Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves typically with a thick, subcoriaceous texture (though sometimes thinner in texture if growing in shade), 4-8 cm long, 2-3.5 cm wide, narrowly obovate to oblanceolate, pubescent and strongly whitened below; leaves and bark aromatic, the odor lemony...... L. subcoriacea 1 Leaves with a thin, membranous texture, 6-16 cm long, 2-6 cm wide, obovate, elliptic, or ovate, glabrous to pubescent below, but not strongly whitened; leaves and bark strongly aromatic, the odor spicy or like sassafras. 2 Leaf base widely cuneate to rounded; leaves narrowly ovate, reticulate-rugose, with an acute apex, pubescent above, drooping, fragrant when crushed with an odor like sassafras; shrubs colonial, short (to 2 m tall)...... L. melissifolia 2 Leaf base cuneate; leaves widely obovate, plane (not rugose), with a short-acuminate apex, glabrous above, borne horizontally, spicy-fragrant when crushed; shrubs not colonial, often multi-stemmed from base, short to tall (to 5 m tall). 3 Leaves glabrous beneath; young twigs glabrous ...... L. benzoin var. benzoin 3 Leaves pubescent beneath, at least on the midrib; young twigs pubescent ...... L. benzoin var. pubescens

Lindera benzoin (Linnaeus) Blume var. benzoin, Smooth Northern Spicebush. Mt (GA, NC, VA), Pd (VA): rich alluvial forests, mesic forests on slopes with circumneutral soils, bottomlands, swamps; common. March-April; August-September. The species is widespread in e. North America; var. benzoin is northern, ranging south to VA and MO, and in the mountains to GA. Where occurring on upland slopes, L. benzoin is an excellent indicator of base-rich soils, generally derived from calcareous sedimentary rocks or mafic metamorphic or igneous rocks. [= C, F, G, K; < L. benzoin – RAB, FNA, GW, W, Z; < Benzoin aestivale (Linnaeus) Nees – S] Lindera benzoin (Linnaeus) Blume var. pubescens (Palmer & Steyermark) Rehder, Hairy Northern Spicebush. Mt, Pd, Cp (GA, NC, SC, VA): rich alluvial forests, mesic forests on slopes with circumneutral soils, bottomlands, swamps; common. March-April; August-September. Var. pubescens is the more southern of the two varieties, ranging through much of se. North America, north to se. VA, sw. VA, s. OH, MI, and MO. Where occurring on upland slopes, L. benzoin is an excellent indicator of base-rich soils. [= C, F, G, K; < L. benzoin – RAB, FNA, GW, W, Z; < Benzoin aestivale (Linnaeus) Nees – S] Lindera melissifolia (Walter) Blume, Southern Spicebush, Pondberry. Cp (GA, NC, SC), Pd (NC): wet flats and depressions, generally with pocosin shrubs; rare (US Endangered, GA Endangered, NC Endangered, SC Rare). March-April; August-September. This species is southern in range, with a very scattered distribution in se. and c. NC, e. SC, e. & sw. GA, nw. FL, sw. AL (?), nw. MS, se. MO-AR, and se. AR-LA (recent collections unknown from FL and LA). It is nearly extirpated in NC, currently known only from three populations, in Sampson, Bladen, and Cumberland counties. A historic record from Orange County, NC (in the lower Piedmont), collected by Elisha Mitchell in 1820 and 1822, appears to be bonafide (McVaugh, McVaugh, & Ayers 1996). [= RAB, F, FNA, GW, K, Z; = Benzoin melissaefolium (Walter) Nees – S] Lindera subcoriacea B.E. Wofford, Bog Spicebush. Cp (GA, NC, SC, VA), Pd (NC): peaty seepage bogs in headwaters of blackwater streams, in the sandhills and immediately adjacent Piedmont, with other pocosin shrubs; rare (US Species of Concern, GA Special Concern, NC Endangered, SC Rare, VA Rare). March-April; July-August. The overall range of this newly described species is still poorly known; it appears to be a Southeastern Coastal Plain endemic, ranging from se. VA (perhaps s. NJ) south to FL and west to LA. Occurring in our area primarily in a scattering of small populations in the fall line Sandhills of NC and SC, with an outlier or two in "Piedmont pocosins" just west of the Sandhills. Distinctive characteristics of sun-grown plants include the rounded apex of the leaf, the leaf strongly whitened beneath and borne in an ascending to even appressed position in relation to the twigs, and a typically fastigiate or virgate branching pattern, with multiple stems or branches ascending vertically and nearly parallel to one another. Shade plants have a different form. [= FNA, K, Z]

Litsea Lamarck 1792 (Pondspice)

A genus of about 400 species, trees and shrubs, of warm temperate and tropical areas, especially se. Asia and Australia. The genus is very heterogeneous and probably needs division into more natural groups. References: Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

Litsea aestivalis (Linnaeus) Fernald, Pondspice. Cp (GA, NC, SC, VA): margins of limesink ponds and Carolina bays, less commonly in wet depressions dominated by shrubs; rare (US Species of Concern, GA Threatened, NC Rare, SC Rare, VA Rare). March-April; May-June. A Southeastern Coastal Plain endemic: e. MD (Dorchester County) and se. VA (York and Isle of Wight counties) south to n. FL (and allegedly also in LA, based on an old and poorly labeled specimen). The fine, zigzag twigs are distinctive. It grows to 6 m tall, characteristically forming a rounded bush. [= RAB, F, FNA, GW, K]

LAURACEAE 474 Persea P. Miller 1754 (Bay)

A genus of about 200 species, trees and shrubs, of Asia and America. The avocado is a member of this genus, Persea americana P. Miller. References: Godfrey (1988); Clewell (1985); Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

1 Twigs glabrous or glabrate; lower surfaces of leaves with minute, silvery to shining-golden hairs (the color depending on age), appressed to the surface; peduncles 1-3 cm long; leaves gtending to be smaller and blunter...... P. borbonia 1 Twigs densely rusty-pubescent; lower surfaces of leaves with longer, rusty, often crooked hairs, not appressed, especially evident along the midrib and principal veins; peduncles 4-7 cm long; leaves tending to be larger and more acute...... P. palustris

Persea borbonia (Linnaeus) Sprengel, Red Bay. Cp (GA, NC, SC): dunes, maritime forests, in dry sandy soils on barrier islands, known only north to Carteret County, NC; rare. May-June; September-October. E. NC (Carteret County) south to FL and west to se. TX; reports of the species north of NC are based on the inclusion of P. palustris in a broadly defined P. borbonia, or are simply in error, based on less hairy plants of P. palustris. This species is rare in our area and becoming rarer with the destruction of most maritime forests for the construction of vacation homes and tourist accomodations. [= FNA, G, GW, K; < P. borbonia – RAB, F (also see P. palustris); = Tamala borbonia (Linnaeus) Rafinesque – S; = P. borbonia var. borbonia] Persea palustris (Rafinesque) Sargent, Swamp Bay. Cp (GA, NC, SC, VA): swamps, pocosins, bay forests, maritime forests, generally in wet peaty soils, but also in fairly dry, sandy soils in maritime forests; common. May-June; September- October. A Southeastern Coastal Plain endemic: DE, e. MD, and se. VA south to FL and west to se. TX. Though variable in amount of hairs on the leaves, the hairs of P. palustris are always of a distinctly different character than those of P. borbonia. [= C, FNA, G, GW, K; < P. borbonia – RAB, F; = Tamala pubescens (Pursh) Small – S; = P. borbonia var. pubescens (Pursh) Little]

Sassafras Presl 1825 (Sassafras)

A genus of 3 species, trees, of temperate e. Asia (2 species) and e. North America (1 species). References: Rohwer in Kubitzki, Rohwer, & Bittrich (1993).

Sassafras albidum (Nuttall) Nees, Sassafras. Pd, Mt, Cp (GA, NC, SC, VA): a wide variety of forests, old fields, disturbed areas, fencerows; common. March-April; June-July. Widespread in e. United States. The original source of "root beer." [= RAB, C, FNA, G, K, W; > S. albidum var. molle (Rafinesque) Fernald – F; > S. albidum var. albidum – F]

LEITNERIACEAE (Corkwood Family) (see SIMAROUBACEAE)

LENTIBULARIACEAE Richard 1808 (Bladderwort Family)

A family of 3 genera and about 270-320 species, insectivorous herbs, cosmopolitan. References: Fischer et al. in Kadereit (2004).

1 Leaves ovate or elliptic, in a basal rosette; carnivory via the viscid-slimy upper leave surfaces; flowers solitary on bractless peduncles ...... Pinguicula 1 Leaves or leaf segments linear, borne along a subterranean or submersed stem; carnivory via specialized bladder-like traps; flowers in (1-) many-flowered racemes, each flower pedicel subtended by a bract...... Utricularia

Pinguicula Linnaeus 1753 (Butterwort)

A genus of about 46-80 species, herbs, of America, Mediterranean Europe, and circumboreal America and Eurasia. References: Schnell (2002b)=Z; Godfrey & Stripling (1961); Wood & Godfrey (1957); Schnell (1980a); Fischer et al. in Kadereit (2004). Key based in part on GW.

1 Expanded corolla < 1.5 cm across; palate not exserted from the throat of the corolla; rosettes usually 2-4 cm in diameter; flowers usually white to pale violet (rarely medium violet); seeds 0.4 mm long ...... P. pumila 1 Expanded corolla > 1.8 cm across; palate markedly exserted from the throat of the corolla; rosettes usually 5-10 (-15) cm across; corolla yellow, violet, or white; seeds (0.4-) 0.5-0.8 mm long. 2 Corolla yellow...... P. lutea 2 Corolla lavender-blue or white. LENTIBULARIACEAE 475 3 Hairs on the lower portion of the scape elongated, pointed, multicellular, nonglandular, transitioning upward to 1- celled glandular hairs; expanded portion of corolla markedly "veiny" (darker along the veins); [of se. NC southward to s. peninsular FL and e. Panhandle FL] ...... P. caerulea 3 Hairs throughout scape glandular; expanded portion of corolla not "veiny;" [collectively of sw. GA and FL Panhandle westward to s. MS]. 4 Fresh leaves dull red or reddish green; corolla lobes ca. 2× as long than broad, the lobes notched almost ½ their length ...... [P. planifolia] 4 Fresh leaves bright yellow-green; corolla lobes ca. 1× as long than broad, the lobes notched about 1/4 their length 5 Corolla tube violet, with darker violet veins; hairs of the inner corolla tube white ...... [P. ionantha] 5 Corolla tube yellow, with reddish-brown veins; hairs of the inner corolla tube yellow ...... P. primuliflora

Pinguicula caerulea Walter, Blue Butterwort. Cp (GA, NC, SC): pine savannas and wet pine flatwoods, mostly in the outer Coastal Plain, rarely extending inland to seepages and sandhill-pocosin ecotones in the fall-line Sandhills of NC and SC; uncommon. April-May. Se. NC (Carteret and Johnston counties) south to s. peninsular FL, west to e. panhandle FL. Schnell (1980a) discusses populations with white corollas. [= RAB, GW, K, S, Z; Pinguicula elatior Michaux] Pinguicula lutea Walter, Yellow Butterwort. Cp (GA, NC, SC): pine savannas and wet pine flatwoods, mostly in the outer Coastal Plain, rarely extending inland to seepages and sandhill-pocosin ecotones in the fall-line Sandhills of SC; uncommon (NC Watch List). Late March-May. Se. NC (Pender and New Hanover counties) south to s. FL, west to e. LA. [= RAB, GW, K, S, Z] Pinguicula primuliflora Wood & Godfrey, Clearwater Butterwort. Cp (GA): clearwater streams and seeps; rare (GA Threatened). Sw. GA, s. AL, Panhandle FL, and s. MS. [= GW, K, Z] Pinguicula pumila Michaux, Small Butterwort. Cp (GA, NC, SC): pine savannas and wet pine flatwoods; uncommon (rare in NC) (NC Rare). April-May. Se. NC (Carteret and Pender counties) south to s. FL, west to se. TX; and in the Bahamas. [= RAB, GW, K, S, Z]

Pinguicula ionantha Godfrey, Panhandle Butterwort. Pond margins, bogs, flatwoods. Panhandle of FL. [= GW, K, Z] Pinguicula planifolia Chapman, Chapman's Butterwort. Pond margins, bogs, flatwoods. S. AL, Panhandle of FL, and s. MS. [= GW, K, S, Z]

Utricularia Linnaeus 1753 (Bladderwort)

Utricularia, as monographed by Taylor (1989), consists of 214 species in 35 sections, with a nearly cosmopolitan distribution. In our area, 14 or 15 species in 5 sections are known to occur. References: Taylor (1989)=Z; Schnell (2002b)=Y; Müller & Borsch (2005); Fischer et al. in Kadereit (2004). Key based in part on Z and GW.

1 Flowers white or cream-white, 1-3 mm long; inflorescence peduncles very reduced, the pedicels appearing to arise direstly from the stolons; traps 0.3-0.8 mm long; plants floating unattached in water (sometimes deposited land by dropping water, but then the principal branch systems stranded on the soil surface); capsules ca. 1 mm long, fusiform, indehiscent, with 1 seed; seeds essentially smooth, unornamented; leaves absent; [section Utricularia]...... U. olivacea 1 Flowers yellow, pink, or purple (sometimes fading whitish), (2-) 5-20 mm long; inflorescence peduncles well-developed, the inflorescence clearly a raceme; traps 0.2-5.0 mm long (< 0.7 mm long only in the terrestrial species (see key lead 2); plants attached (with principal branch systems within the soil), or floating unattached in water (sometimes deposited on land by dropping water, but then the principal branch systems stranded on the soil surface); capsules 1-8 mm long, globose, subglobose, or ovoid, with many seeds; seeds reticulate, papillose, echinate, multi-angled, or winged (rarely more-or-less smooth); leaves present (sometimes absent in the terrestrial species). 2 Plants attached (with principal branch systems within the soil); leaves absent or simple, linear, grass-like aerial leaves; bladders 0.2-1.1 mm long, most or all on a plant usually < 1.0 mm long; seeds reticulate-alveolate (also angled in U. resupinata), 0.20-0.25 mm long. 3 Flowers pink; inflorescence 1 (-2)-flowered; bract at base of the pedicel tubular, attached circumferentially around the stem; aerial leaves (when present) terete, septate; [very rare in our area]; [section Lecticula]...... U. resupinata 3 Flowers yellow (sometimes fading whitish); inflorescence (1-) 2-15-flowered; bract at base of the pedicel peltate or ovate, attached on one side of the stem; aerial leaves (when present) flattened, not septate; [collectively common in our area]. 4 Bracts subtending the pedicels peltate (attached near their middles), unattached at either end; pair of bracteoles absent; spur of the corolla oriented forward, more-or-less appressed to the lower lip; aerial leaves (when present) with subacute to obtuse apex; [section Setiscapella] ...... U. subulata 4 Bracts subtending the pedicels ovate (attached at their bases), free only at their upper end; pair of bracteoles associated with each bract present, linear to lanceolate; spur of the corolla oriented downward or backward, at approximately a right angle to the lower lip; aerial leaves (when present) with acute apex; [section Stomoisia]. 5 Corolla 1.5-2.0 cm long; spur 8-12 mm long; raceme usually short, the (1-) 2-6 flowers crowded together, all of them chasmogamous ...... U. cornuta LENTIBULARIACEAE 476 5 Corolla 0.25-1.5 cm long; spur 5-7 (-9) mm long; raceme usually elongate, the (1-) 2-15 flowers well- spaced, often the lower (sometimes all) cleistogamous and much smaller than the chasmogamous flowers...... U. juncea 2 Plants floating unattached in water (sometimes deposited on land by dropping water, but then the principal branch systems stranded on the soil surface); leaves present and dissected into linear segments; bladders 0.7-5.0 mm long, most or all on a plant > 1.0 mm long; seeds papillose, reticulate, ridged, angled, or winged, 0.5-2.0 mm long. 6 Flowers purple; leaves divided into verticillate segments with terminal traps; [section Vesiculina]...... U. purpurea 6 Flowers yellow; leaves divided into alternate segments with lateral traps; [section Utricularia]. 7 Peduncle with whorl of inflated leaf-like organs (floats). 8 Floats 4-7, not fused basally to one another, fusiform, tapering gradually to base and apex from a widest point near the middle; leaves with the 2 primary divisions unequal; bracts of the scape longer than broad, entire; flowers (6-) 9-14 (-17) per scape; apex of corolla spur bifid...... U. inflata 8 Floats (5-) 6-8 (-10), fused basally to one another, cylindrical, more-or-less parallel-sided through most of their length, tapering abruptly to base and apex; leaves with the 2 primary divisions equal; bracts of the scape broader than long, the apex slightly to strongly 3-lobed; flowers (1-) 3-4 (-7) per scape; apex of corolla spur usually entire (rarely bifid)...... U. radiata 7 Peduncle without whorl of inflated leaf-like organs (floats). 9 Main axes distinctly flattened in cross-section, up to 10 mm wide ...... U. foliosa 9 Main axes round in cross-section. 10 Lower lip of corolla 3-lobed; seeds disk-shaped, not angular or winged; inflorescences of 2 types, the chasmogamous on erect peduncles 5-25 cm long bearing 2-8 flowers, the cleistogamous without a peduncle, the solitary pedicels borne directly on the stolons, 0.5-2 cm long, deflexed...... U. geminiscapa 10 Lower lip of corolla entire or slightly irregular, not 3-lobed; seeds angular or winged; inflorescences of 1 type (erect, chasmogamous). 11 Upper corolla lip smaller than the lower, entire; capsule circumscissilely dehiscent; seeds 0.7- 1.0 mm long, 4-6-angled; corolla without stipitate glands on its external surface. 12 Leaves of one kind only, divided into numerous capillary segments bearing lateral traps; bracts scarcely auriculate; plant distinctly aquatic, floating in water and only rarely stranded; [of the Coastal Plain]...... U. macrorhiza 12 Leaves of 2 or 3 kinds, some divided into capillary or narrowly linear segments and bearing few or no traps, others divided into fewer capillary segments and bearing more- or-less numerous traps; bracts distinctly auriculate; plants typically in boggy situations, in shallow water or frequently stranded; [either of the Mountains at high elevations or of various physiographic provinces northward]. 13 Broadest leaf segments with 9-20 lateral setae (use 10× magnification); spur of corolla cylindrical, distinctly longer than wide, the apex distinctly curved forward...... [U. intermedia] 13 Broadest leaf segments lacking lateral setae; spur of corolla shortly saccate to broadly conical, wider than long, the apex not curved forward ...... U. minor 11 Upper corolla lip larger than the lower, obscurely 3-lobed; capsule laterally 2-valved or indehiscent; seeds 0.8-2.5 mm long, lenticular, with an irregular, lobed, or continuous wing; corolla (or at least the spur) with a few to many short stipitate glands (sometimes patchily distributed). 14 Vegetative shoots of 2 kinds, some bearing leafy segments and few or no traps, others bearing reduced segments and more-or-less numerous traps; seeds 1.0-2.5 mm long, with an irregularly deeply lobed or partial wing. 15 Plant anchored in mud up to 100 cm below water surface; green leafy shoots up to 40 cm long and 5 cm wide; peduncle flexuous, to 100 cm long, only the uppermost ca. 10 cm emergent...... U. floridana 15 Plant in shallow water or stranded; green leafy shoots usually not > 10 cm long and 2 cm wide; peduncle erect, straight, to 30 cm long, the uppermost 10-25 cm emergent ...... U. striata 14 Vegetative shoots uniform, all bearing rather sparsely divided leaf segments bearing traps, seeds 0.8-1.1 mm long, with a continuous, circumferential wing, slightly to irregularly lobed. 16 Lower corolla lip 8-10 mm long, about equalling or slightly shorter than the conical 5-9 mm long spur; leaves usually forked twice...... U. biflora 16 Lower corolla lip 5-6 mm long, exceeding the blunt 3.5-4.5 mm long spur; leaves usually forked once...... U. gibba

Utricularia biflora Lamarck, Longspur Creeping Bladderwort. Cp (GA, NC, SC, VA), Pd, Mt (GA): ponds, lakes, and diches; common. June-October. This species may not be distinct from U. gibba (which see for discussion). E. MA south FL, west to TX and OK. [= RAB, C, F, G, GW, W; = U. pumila Walter – S, apparently misapplied; < U. gibba – K, Y, Z] LENTIBULARIACEAE 477 Utricularia cornuta Michaux, Horned Bladderwort. Cp (GA, NC, SC), Mt (GA, NC), Pd (GA): shores of limesink ponds (dolines), mountain bogs; rare (NC Watch List). (May-September. Newfoundland and Québec west to n. Ontario, Alberta, and MN, south to s. FL and e. TX; also in the Bahamas and Cuba. Taylor (1989) states that where sympatric with U. juncea, U. cornuta flowers much earlier. [= RAB, C, F, G, GW, K, W, Y, Z; = Stomoisia cornuta (Michaux) Rafinesque – S] Utricularia floridana Nash, Florida Bladderwort. Cp (GA, NC, SC): in deep water of natural Carolina bay lakes, other natural lakes, and limesink ponds (dolines); rare (NC Rare, SC Rare). July-August. Se. NC south to c. peninsular FL, west to panhandle FL and sw. GA. [= GW, K, S, Y, Z] Utricularia foliosa Linnaeus, Flatstem Bladderwort. Cp (GA, NC): in deep water of natural lakes and ponds; rare. Se. NC south to s. FL, west to TX (Brown & Marcus 1998); West Indies, South America, Africa. This species is reported for NC by Taylor (1989). See GW for a detailed description of this species. [= GW, K, S, Y, Z] Utricularia geminiscapa Benjamin, Two-flowered Bladderwort, Hidden-fruited Bladderwort. Cp (NC, VA), Mt (VA): beaver ponds, mucky seepages; rare (NC Rare, VA Watch List). Newfoundland and Québec west to n. MI and n. WI, south to PA and sc. NC. [= C, F, G, K, W, Y, Z] Utricularia gibba Linnaeus, Shortspur Creeping Bladderwort. Cp (GA, NC, SC, VA), Mt (GA, NC, VA), Pd (SC, VA): ponds, lakes, and diches; uncommon. May-September. Québec west to WI, south to FL and LA; also in the West Indies and Central America and apparently in the Old World. Taylor (1989) includes U. biflora in U. gibba. Other authors have expressed doubts about the distinction, including RAB ("doubtfully distinct"). Taylor suggests that "further research is clearly indicated, but to be at all meaningful, it must be conducted on a worldwide basis." I have here, for the moment, retained the 2 traditionally recognized species, though intermediates will be encountered. [= RAB, C, F, G, S, W; < U. gibba – K, Y, Z (also see U. biflora)] Utricularia inflata Walter, Swollen Bladderwort, Inflated Bladderwort. Cp (GA, NC, SC, VA), Mt (NC): ponds, lakes, ditches; common. May-November. NJ south to s. FL, west to e. TX; disjunct in WA (probably introduced). Also disjunct in an artificial pond in Henderson County, NC (Carl Sandburg Home National Historic Site). [= C, G, GW, K, S, Y, Z; = U. inflata var. inflata – RAB, F] Utricularia juncea M. Vahl, Southern Bladderwort. Cp (GA, NC, SC, VA), Pd (NC): shores of limesink ponds (dolines), borrow pits, wet sands; uncommon (VA Rare). July-September. NY (Long Island) and NJ south to FL, west to e. TX and se. AR; also in the West Indies, Central America and South America. [= RAB, C, F, G, GW, K, Y, Z; > Stomoisia juncea (M. Vahl) Barnhart – S; > Stomoisia virgatula Barnhart – S] Utricularia macrorhiza Le Conte, Greater Bladderwort. Cp (NC, SC, VA): pools and ponds; rare (NC Watch List, VA Watch List). May-September. Newfoundland west to AK, south to NC, SC, FL, TX, CA, and Mexico; also in e. Asia. See Taylor (1989) for a discussion of the differences between this species and U. vulgaris of Europe and w. Asia, with which it has often been combined or associated as a variety. [= K, S, Y, Z; < U. vulgaris Linnaeus – RAB, C, F, G, misapplied to American plants] Utricularia minor Linnaeus, Lesser Bladderwort, Small Bladderwort. Mt (NC): mountain bog at about 1400 meters elevation; rare (NC Rare). Circumboreal, south in North America to NJ, PA, IN, IL, IA, NE, CO, UT, NV, and CA; disjunct in w. NC. [= C, F, G, K, W, Y, Z] Utricularia olivacea Wright ex Grisebach, Dwarf Bladderwort, Minute Bladderwort. Cp (GA, NC, SC, VA): in floating mats (often algal) in water of limesink ponds (dolines), artificial lakes or beaver ponds; rare (NC Threatened, SC Rare, VA Rare). September-October. NJ south to FL, west to s. AL and s. MS (Sorrie & Leonard 1999), in the Coastal Plain; also in the West Indies (Cuba), Central America, and South America. [= RAB, GW, K, Y, Z; = Biovularia olivacea (Wright ex Grisebach) Kam. – S] Utricularia purpurea Walter, Purple Bladderwort. Cp (GA, NC, SC, VA): in water of ponds, ditches, other slow-moving water; uncommon (VA Rare). May-September. Nova Scotia and Québec west to MN, south to NY, n. IN, s. MI, and WI, and on the Coastal Plain south to s. FL, west to se. TX; also in Mexico, the West Indies, and Central America. [= RAB, C, F, G, GW, K, Y, Z; = Vesiculina purpurea (Walter) Rafinesque – S] Utricularia radiata Small, Floating Bladderwort, Small Swollen Bladderwort. Cp (GA, NC, SC, VA), Mt (VA): ponds, depression ponds, lakes, and ditches; uncommon (VA Watch List). May-November. Nova Scotia south to s. FL, west to TX; disjunct in w. VA, w. TN, nw. IN; reports of this species in Cuba and South America are apparently in error. [= C, G, GW, K, S, W, Y, Z; = U. inflata var. minor Chapman – RAB, F] Utricularia resupinata B.D. Greene ex Bigelow, Northeastern Bladderwort, Resupinate Bladderwort. Cp (GA, NC): shores of natural lakes; rare (NC Rare). Nova Scotia west to nw. WI, south (irregular and scattered in part) to FL and sw. GA; also in the Bahamas (Sorrie & LeBlond 1997). Although "the curious gap in the North American range" [NC, SC, and VA] (Taylor 1989) is no longer strictly a gap, U. resupinata does appear to have a strangely bimodal range, with a center of distribution in ne. United States and se. Canada and a second extending from se. United States south into the West Indies and Central America. [= C, F, G, GW, K, Y, Z; = Lecticula resupinata (B.D. Greene) Barnhart – S] Utricularia striata Le Conte ex Torrey, Fibrous Bladderwort. Cp (GA, NC, SC, VA), Mt (VA): ponds, lakes, and ditches; uncommon (VA Rare). May-November. Se. MA south to FL, west to e. TX and e. OK. [= K, Y, Z; = U. fibrosa Walter – RAB, C, F, G, GW, S, of uncertain application and likely misapplied] Utricularia subulata Linnaeus, Slender Bladderwort, Zigzag Bladderwort. Cp (GA, NC, SC, VA), Mt (GA, NC, VA), Pd (GA, NC): moist sands or peats of various kinds of acidic wetlands, including wet pine savannas and flatwoods, shores of limesink ponds (dolines), borrow pits, ditches; common (VA Watch List). March-July (-later). In North America primarily in the Coastal Plain, from Nova Scotia and e. MA south to s. FL, west to TX, north in the interior to TN and AR; also in the West Indies, Central America, South America, Africa, and Asia. Taylor (1989) terms this "the most widespread of Utricularia LENTIBULARIACEAE 478 species." [= RAB, C, F, G, GW, K, W, Y, Z; > Setiscapella subulata (Linnaeus) Barnhart – S; > Setiscapella cleistogama (A. Gray) Barnhart – S]

Utricularia intermedia Hayne, Northern Bladderwort. South to se. PA (Rhoads & Klein 1993), DE, and MD (Kartesz 1999). The report from sc. GA (Jones & Coile 1988) is in error. [= C, F, G, K, Y, Z]

LIMNANTHACEAE R. Brown 1838 (False-mermaid Family, Meadow-foam Family)

A family of 2 genera and 8 species, herbs, of temperate North America. References: Bayer & Appel in Kubitzki & Bayer (2003).

Floerkea Willdenow 1801 (False-mermaid)

A peculiar and monotypic genus, an annual herb, endemic to North America.

Floerkea proserpinacoides Willdenow, False-mermaid. Pd, Mt (VA): moist, rich floodplain forests; rare (VA Watch List). April-May. Nova Scotia and Québec west to British Columbia, south to n. VA, TN, and CA. [= C, F, G, K, S, W]

LINACEAE DC. ex Gray 1821 (Flax Family)

A family of about 14 genera and 250-350 species, trees, vines, shrubs, and herbs, cosmopolitan. References: Robertson (1971)=Y.

Linum Linnaeus 1753 (Flax)

A genus of about 180 species, herbs, of temperate and subtropical areas. References: Rogers (1984)=Z; Rogers (1963)=Y.

1 Petals blue; capsule 6-10 mm long; [section Linum]. 2 Inner sepals with minutely ciliate margins; stigmas slender, elongate; capsule 6-10 mm long...... L. usitatissimum 2 Inner sepals entire; stigmas capitate; capsule 5-7 mm long. 3 Flowers homostylous (flowers with stigmas at about the level to slightly above the anthers) ...... [L. lewisii var. lewisii] 3 Flowers heterostylous (some flowers with stigmas below the anthers, others with stigmas well above the anthers)...... [L. perenne] 1 Petals yellow; capsules 1-4 mm long; [section Linopsis]. 4 Inner and outer sepals all very conspicuously glandular-toothed; annual; leaves with 2 brownish glands flanking the attachment to the stem; styles united basally for (0.2-) 0.5-1.2 (-1.8) mm; [section Linopsis, subsection Rigida]. 5 Sepals 2.3-3.5 mm long, acute; inflorescence consisting of 1 or more elongate and racemiform branches; dried plants dark, purple-dotted...... L. harperi 5 Sepals (3.1-) 3.6-5 (-7.3) mm long, acuminate; inflorescence an open panicle; dried plants pale green...... L. sulcatum 4 Outer sepals entire (very rarely sparsely glandular-toothed), inner sepals entire or sparsely to conspicuously glandular- toothed; perennial; leaves without brownish glands flanking the attachment to the stem; styles free; [section Linopsis, subsection Linopsis]. 6 Fruit as long as broad or longer, its apex acute, apiculate, or obtuse, (2-) 2.2-3.2 (-3.3) mm long; leaves mostly 1.3-4.3 mm wide. 7 Leaves (1.2-) 2.3-4.3 (-5.6) mm wide, mostly 25-50 below the inflorescence; septa of the fruit sparsely but conspicuously ciliate; false septa incomplete; fruit apex acute, the exposed portions purple...... L. intercursum 7 Leaves (1.0-) 1.3-2.0 (-3.2) mm wide, mostly 50-120 below the inflorescence; septa of the fruit glabrous; false septa virtually complete; fruit apex rounded to apiculate, the exposed portions purple or yellow. 8 Fruit pyriform, (2.0-) 2.3-2.8 (-3.0) mm long, 1.7-2.6 mm in diameter, the apex rounded, the exposed portions purple; seeds (1.6-) 1.7-2.0 (2.1) mm long; anthers averaging 0.8 mm long...... L. floridanum var. floridanum 8 Fruit ovate, (2.8-) 3.0-3.2 (-3.3) mm long, 2.5-3.1 mm in diameter, the apex minutely apiculate, the exposed portions yellow; seeds 2.1-2.4 mm long; anthers averaging 1.2 mm long...... L. floridanum var. chrysocarpum 6 Fruit broader than long, its apex depressed, flattened, or broadly rounded, (1.3-) 1.5-2.1 (-2.3) mm long; leaves mostly 1.9-9.3 mm wide. 9 Margins of the inner sepals with conspicuous stalked glands; mature fruits of dried specimens usually adhering to the plant...... L. medium var. texanum LINACEAE 479 9 Margins of the inner sepals glandless, or with a few inconspicuous, sessile glands; mature fruits of dried specimens usually shattering and falling freely. 10 Inflorescence paniculate, the lower inflorescence branches not elongate, their tips not nearly reaching the tips of the upper inflorescence branches; branchlets striate-ridged; leaves mostly opposite (usually to beyond the midpoint from the base of the plant to the first inflorescence branch) ...... L. striatum 10 Inflorescence corymbose, some (at least) of the lower branches of the inflorescence elongate, their tips nearly equalling the tips of the upper inflorescence branches; branchlets terete or nearly so; leaves mostly alternate (usually the opposite leaves of the lower stem not extending beyond the midpoint from the base of the plant to the first inflorescence branch) ...... L. virginianum

Linum floridanum (Planchon) Trelease var. chrysocarpum Rogers, Yellow-fruited Yellow Flax. Cp (GA, NC, SC): wet savannas; rare (NC Watch List). June-October. Se. NC south to s. FL and west to s. MS. [= K, Y, Z; < L. virginianum var. floridanum Planchon – RAB; < L. floridanum – GW; < Cathartolinum floridanum (Planchon) Small – S] Linum floridanum (Planchon) Trelease var. floridanum, Florida Yellow Flax. Cp (GA, NC, SC): savannas, sandhill seeps; common. June-October. E. NC south to s. FL and west to LA, also in the West Indies, essentially limited to the Coastal Plain. [= K, Y, Z; < L. virginianum var. floridanum Planchon – RAB (also see L. floridanum var. chrysocarpum and L. intercursum); < L. floridanum – C, F, G, GW; < Cathartolinum floridanum (Planchon) Small – S] Linum harperi Small, Harper's Grooved Flax. Cp (GA): dry pinelands; rare (GA Special Concern). This is a rare taxon of longleaf pine woodlands or savannas in w. FL, sw. GA, and c. AL. It is probably distinct from L. sulcatum at the species level, needing additional study. [= L. sulcatum Riddell var. harperi (Small) Rogers – K, Y, Z; = Cathartolinum harperi (Small) Small – S] Linum intercursum Bicknell, Bicknell's Yellow Flax. Pd, Mt (GA, NC, SC, VA), Cp (NC, VA): dry to moist places; common. June-October. MA south to c. TN, nw. GA, and c. AL; from MA to MD, nearly restricted to the Coastal Plain, in VA, NC, SC, GA, AL, and se. TN, however it is primarily on the Piedmont and Mountains. It also occurs disjunctively in n. IN near the Great Lakes. [= C, F, G, K, W, Y, Z; < L. virginianum var. floridanum (Planchon) – RAB; = Cathartolinum intercursum (Bicknell) Small – S] Linum medium (Planchon) Britton var. texanum (Planchon) Fernald, Texas Yellow Flax. Cp, Pd, Mt (GA, NC, SC, VA): dry to moist places; common (rare in Mountains). Var. texanum ranges from s. ME MI, and n. IL south to c. FL and TX, and in the West Indies. Var. medium is limited to area around the Great Lakes. [= C, F, GW, K, W, Y, Z; < L. virginianum var. medium Planchon – RAB; < L. medium – G; < Cathartolinum medium (Planchon) Small – S] Linum striatum Walter, Ridgestem Yellow Flax. Mt, Cp, Pd (GA, NC, SC, VA): bogs, seepages, other wet places, often growing in Sphagnum; common. June-October. MA, PA, MI, and IL south to w. FL, LA, and e. TX. [= RAB, C, G, GW, K, W, Y, Z; > L. striatum var. striatum – F; = Cathartolinum striatum (Walter) Small – S] Linum sulcatum Riddell, Grooved Yellow Flax. Mt (GA, VA), Pd (NC): dry calcareous places in the mountains of VA (where also somewhat weedy in adjacent disturbed areas), diabase barrens in the Piedmont of NC; rare (NC Rare, VA Watch List). May-August. Primarily a species of the Great Plains of s. Manitoba, ND, and MN south through SD, IA, WI, NE, MO, IL, KS, and MO to OK, L. sulcatum occurs farther east as a rare disjunct on glades or barrens over rocks such as limestone or diabase. [= RAB, C, F, G, W; = Linum sulcatum Riddell var. sulcatum – K, Y, Z; = Cathartolinum sulcatum (Riddell) Small – S; = Mesynium sulcatum (Riddell) A. & D. Löve] * Linum usitatissimum Linnaeus, Common Flax. Pd, Cp (NC, SC, VA): disturbed places; rare, native of Europe. This is the flax of commerce, used both for its fiber, the source of flax, and the oil expressed from its seeds (linseed oil). [= RAB, C, F, G, K, S, Z] Linum virginianum Linnaeus, Virginia Yellow Flax. Mt, Pd (GA, NC, SC, VA), Cp (VA): dry or moist places; uncommon. June-October. MA, NY, Ontario, MI, and IL south to SC, GA, AL, and MO. [= C, F, G, GW, K, W; = L. virginianum var. virginianum – RAB; = Cathartolinum virginianum (Linnaeus) Reichenbach – S]

Linum lewisii Pursh var. lewisii, Prairie Flax, a western blue-flowered species, occurs as a disjunct at Smoke Hole Caverns, WV, and several adjacent counties. [= K; < L. perenne – C, apparently misapplied to WV material; < L. lewisii – F; < L. perenne Linnaeus var. lewisii (Pursh) Eaton & J. Wright – G; < Adenolinum lewisii (Pursh) A. & D. Löve] * Linum perenne Linnaeus, Perennial Flax, is cultivated and "rarely naturalized along roadsides" in scattered locations in PA (Rhoads & Klein 1993) and reported tentatively for VA (Kartesz 1999). [= K; < L. perenne – C (also see L. lewisii)]

LINDERNIACEAE Borsch, K. Müller, & Eb. Fischer 2005 (False-pimpernel Family)

References: Tank et al. (2006).

Lindernia Allioni (False-pimpernel)

A genus of about 80 species, of warm temperate and subtropical regions of the Old and New Worlds. Probably not a member of Plantaginaceae (Albach, Meudt, & Oxelman 2005). References: Cooperrider & McCready (1975)=Z; Qualls (1984)=Y; Lewis (2000)=X.

LINDERNIACEAE 480 1 Fertile stamens 4; calyx lobes connate at anthesis for > ½ their length, later separating; [section Torenioides]..... L. crustacea 1 Fertile stamens 2 (with 2 staminodia without anthers, or with rudimentary anthers); calyx lobes distinct to the base at anthesis and after; [section Brachycarpae]. 2 Pedicels shorter than or about as long as the subtending leaves...... L. dubia var. dubia 2 Pedicels longer than the subtending leaves (or bracteal leaves in some species). 3 Leaves nearly orbicular; stems creeping ...... L. grandiflora 3 Leaves distinctly longer than wide; stems erect (sometimes decumbent at the base and rooting if knocked down by water). 4 Leaves not glandular punctate; seeds 2-3× as long as wide...... L. dubia var. anagallidea 4 Leaves glandular punctate; seeds ca. 1× as long as wide. 5 Leaves primarily in a basal rosette, leaves of the stem strongly reduced upward to bracts; capsule (1.8-) 3.4-7 mm long; [primarily of seepage of flatrocks]...... L. monticola ("monticola" form) 5 Leaves primarily on the stem, not conspicuously reduced upward; capsule 1.4-3.4 mm long; [of stream or river banks] ...... L. monticola ("saxicola" form)

* Lindernia crustacea (Linnaeus) F. Mueller. Cp (GA, SC), Pd (SC), {NC}: lawns; uncommon, introduced from Malaysia. September. [= RAB, GW, K, P, X, Y] Lindernia dubia (Linnaeus) Pennell var. anagallidea (Michaux) Cooperrider. Cp, Pd, Mt (GA, NC, SC, VA): wet sandy or muddy areas; common (rare in Mountains). June-September. Nearly throughout North America, Central America, and South America. The extensive and essentially coincident ranges of the two varieties of L. dubia suggests that they may be merely forms, as suggested by Voss (1996). [= C, K, X, Y, Z; = L. anagallidea (Michaux) Pennell – RAB, F, G, GW, P; = Ilysanthes inequalis (Walter) Pennell – S; < L. dubia – W] Lindernia dubia (Linnaeus) Pennell var. dubia. Cp, Pd, Mt (GA, NC, SC, VA): wet sandy or muddy areas; common. May-November. Nearly throughout North America, Central America, and South America. [= C, X, Y, Z; = L. dubia (Linnaeus) Pennell – RAB, GW; > L. dubia var. dubia – F, G, K; > L. dubia var. riparia (Rafinesque) Fernald – F, G; > L. dubia var. inundata Pennell – F, G, K; > L. dubia var. major (Pursh) Pennell – P; > L. dubia var. typica – P; = Ilysanthes dubia (Linnaeus) Barnhart – S; < L. dubia – W; = Gratiola dubia Linnaeus] Lindernia grandiflora Nuttall. Cp (GA): depressional wetlands; rare. S. GA south to s. FL. [= GW, K, P, X, Y; = Ilysanthes grandiflora (Nuttall) Bentham – S] Lindernia monticola Muhlenberg ex Nuttall, Flatrock Pimpernel, Riverbank Pimpernel. Pd, Mt (GA, NC, SC), Cp (GA): in seasonal seepage on granitic flatrocks, and on river-scoured siliceous rocks; rare (GA Endangered – "saxicola", NC Watch List). April-June (-September). Nc. and sw. NC south to ne. FL and ec. AL. L. saxicola appears to be merely a form of L. monticola, the leafy stems the result of the basal leaves being covered by silt deposited by floodwaters (Qualls 1984; Lewis 2000); this needs additional study. [= K, X; > L. monticola – RAB, GW, P, W, Y; > L. saxicola M.A. Curtis – RAB, P, W, Y; > Ilysanthes monticola (Muhlenberg ex Nuttall) Rafinesque – S; > Ilysanthes saxicola (M.A. Curtis) Chapman – S]

Lindernia diffusa (Linnaeus) Wettstein, is reported for SC by Kartesz (1999) on the basis of specimens at NCU, but the specimens so labelled are actually L. dubia. {not keyed}

Micranthemum Michaux

The appropriate generic treatment is unclear. If treated (as here) as including Hemianthus, a genus of about 14 species, of tropical to warm temperate America.

1 Calyx lobes uneven, 3 of the sinuses extending about halfway to the base of the calyx, the lowermost sinus extending to the base...... M. micranthemoides 1 Calyx lobes even, all of the sinuses reaching nearly to the base of the calyx...... M. umbrosum

Micranthemum micranthemoides (Nuttall) Wettstein, Nuttall's Micranthemum. Cp (VA): muddy, freshwater intertidal shores; rare, possibly extinct (US Species of Concern, VA Rare). NY (Hudson River) south to VA (Chesapeake Bay, Potomac River, James River). [= F, K; = Hemianthus micranthemoides Nuttall – C, G, P] Micranthemum umbrosum (J.F. Gmelin) Blake, Shade Mudflower. Cp (GA, NC, SC, VA): shallow pools, stagnant streams, wet depressions in swamp forests; uncommon (VA Rare). May-October. Se. VA south to FL, west to TX, and south into tropical America. [= RAB, C, F, G, GW, K, P; = Globifera umbrosa J.F. Gmelin – S]

LINNAEACEAE (Rafinesque) A. Backlund 1998 (Twinflower Family)

A family of 5 genera and about 35 species, shrubs and suffrutescent herbs. Various segregate families (or reassignments) of taxa traditionally placed in the Caprifoliaceae have been proposed, including the transfer of Sambucus and Viburnum to the Adoxaceae, placement of Diervilla and Weigela in the Diervillaceae (Backlund & Pyck 1998), placement of Abelia, Linnaea, and Kolkwitzia in the Linnaeaceae (Backlund & Pyck 1998), and retention of Lonicera, Symphoricarpos, and Triosteum in a much LINNAEACEAE 481 more narrowly circumscribed Caprifoliaceae. Alternatively, all these taxa could be included in the Caprifoliaceae, along with Dipsacaceae and Valerianaceae, as a very broadly circumscribed Caprifoliaceae. References: Backlund & Pyck (1998).

1 Trailing shrubby herb; [native] ...... [Linnaea] 1 Upright shrub; [planted and persistent or weakly naturalizing]. 2 Sepals oblanceolate, the larger > 1 mm wide; fruit and ovaries free, not hirsute ...... Abelia 2 Sepals lanceolate to linear, < 1 mm wide; fruit and ovaries fused in pairs, densely hirsute...... Kolkwitzia

Abelia R. Brown

A genus of about 30 species, shrubs, primarily of s. and e. Asia.

* Abelia ×grandiflora (André) Rehder [chinensis × uniflora]. Cp (NC): suburban thickets; commonly planted in our area; sometimes persistent or rarely weakly naturalizing, parent species natives of China. [= K]

Kolkwitzia Graebner (Beautybush)

A monotypic shrub, of c. China.

* Kolkwitzia amabilis, Beautybush. Mt (NC): disturbed areas; planted as an ornamental shrub, rarely naturalized from plantings, native of c. China. [= K]

Linnaea Linnaeus (Twinflower)

A monotypic genus, a trailing weak shrub, circumboreal.

Linnaea borealis Linnaeus ssp. americana (Forbes) Hultén ex Clausen, American Twinflower, is documented by an early specimen (1892) from Sevier County, TN, presumably from the Great Smoky Mountains. This population (not recently seen) is disjunct from more northern areas. L. borealis ssp. americana also occurs in several counties in WV along or near the VA border, and the species might be expected in high elevation or cold sites in nw. VA. [= K; = L. borealis var. longiflora Torrey – C, G; = L. borealis var. americana (Forbes) Rehder – F; < L. borealis – W; = L. americana Forbes; = L. borealis ssp. longiflora (Torrey) Hultén]

LOGANIACEAE R. Brown ex Mart. 1827 (Logania Family) (also see GELSEMIACEAE, SCROPHULARIACEAE, and TETRACHONDRACEAE)

As here rather narrowly interpreted, Loganiaceae consists of 12 genera (Mitreola, Logania, and Mitrasacme) and about 300 species, herbs and subshrubs, of tropical, subtropical, and warm temperate areas of the Old and New Worlds. Genera in our area other than Mitreola which have traditionally been considered components of the Loganiaceae now are clearly better placed in the small families Tetrachondraceae (Polypremum), Gelsemiaceae (Gelsemium), and Scrophulariaceae (Buddleja), more closely related to other families (such as Rubiaceae) than to Loganiaceae sensu stricto (Struwe, Albert, & Bremer 1994). The affinities of Spigelia appear to be with a small group of tropical and subtropical genera, the largest of which is Strychnos. Struwe, Albert, & Bremer (1994) treated this group as the family Strychnaceae, based on a cladistic analysis of data. A later, more thorough analysis suggested that Strychnaceae is best recombined with Loganiaceae (Backlund, Oxelman, & Bremer 2000). References: Rogers (1986).

1 Woody vine...... Gelsemium [see GELSEMIACEAE] 1 Herb. 2 Corolla funnelform, 0.1-0.2 cm long, white...... Mitreola 2 Corolla tubular, 3-6 cm long, red and yellow...... Spigelia

Mitreola Linnaeus (Miterwort)

A genus of about 6 species, herbs, tropical, subtropical, and warm temperate. References: Nelson (1980)=Y; Rogers (1986)=Z.

1 Leaves 2-8 cm long petiolate or sessile and tapering to a cuneate base ...... M. petiolata 1 Leaves 1-4 cm long, sessile, the base rounded. 2 Mature seed reticulate; mature capsule smooth to slightly and finely tuberculate; larger leaves ca. 4× as long as wide ...... M. angustifolia LOGANIACEAE 482 2 Mature seed smooth; mature capsule markedly papillose-warty; larger leaves 1.5-2× as long as wide ...... M. sessilifolia

Mitreola angustifolia (Torrey & A. Gray) J.B. Nelson, Narrow-leaved Miterwort. Cp (GA, SC): clay-based Carolina bays, other Coastal Plain depressional wetlands; rare (GA Special Concern). June-August. Se. SC south to n. FL, and west to s. AL and se. MS (Sorrie & Leonard 1999). [= GW, Y; < M. sessilifolia – K, Z; = Cynoctonum angustifolium (Torrey & A. Gray) Small – S] Mitreola petiolata (J.F. Gmelin) Torrey & A. Gray, Caribbean Miterwort. Cp, Pd (GA, NC, SC, VA), Mt (GA): swamps, marshes, ditches, other wet habitats; uncommon (rare in the Piedmont) (VA Rare). July-September; September-November. Se. VA south to FL and west to AR and c. TX, north in the interior to nw. GA and c. and se. TN; Mexico, the West Indies, and n. South America. [= GW, K, Y; = Cynoctonum mitreola (Linnaeus) Britton – RAB, C, F, G, S] Mitreola sessilifolia (J.F. Gmelin) G. Don, Small-leaved Miterwort. Cp (GA, NC, SC), Pd (GA, VA): wet savannas, pocosins, ditches, margins of limesink depressions (dolines); common (VA Rare). Late June-August; September-October. Se. VA south to FL, west to e. TX, and in the Bahama Islands. [= GW, Y; = Cynoctonum sessilifolium J.F. Gmelin – RAB, C, F, G, S; < M. sessilifolia – K, Z (also see M. angustifolia)]

Spigelia Linnaeus (Pinkroot)

A genus of about 50 species, herbs, of tropical and warm temperate America. References: Gould (1996)=Z; Rogers (1986)=Y.

1 Corolla scarlet on the outer surface, yellow on the inner surface...... S. marilandica 1 Corolla light pink to white on the outer and inner surfaces. 2 Corolla 36-50 mm long; pistil 24-27 mm long; sepals 8-11 mm long; inflorescence with 2-4 flowers; [of dolomitic glades in Bibb Co. AL] ...... [S. gentianoides var. alabamensis] 2 Corolla 25-30 mm long; pistil 17-19 mm long; sepals 4-6 mm long; inflorescence with 3-8 flowers; [of pine savannas of Panhandle FL]...... [S. gentianoides var. gentianoides]

Spigelia marilandica (Linnaeus) Linnaeus, Pinkroot, Wormgrass. Mt (GA, NC, SC), Pd, Cp (GA, SC): woodlands and forests, usually on circumneutral soils; common. May-June; late June-July. SC, sw. NC (Cherokee Co. and Macon Co.), and TN west to s. IN and OK, south to FL and TX; some floras allege its occurrence north to MD, NJ, and PA. S. marilandica likely occurs in sw. VA. [= RAB, C, F, G, K, S, W]

Spigelia gentianoides Chapman ex Alphonse de Candolle var. alabamensis K. Gould. Dolostone glades of Bibb County, AL (Gould 1996, Allison & Stevens 2001). [= K, Z] Spigelia gentianoides Chapman ex Alphonse de Candolle var. gentianoides. Pine savannas. FL Panhandle. [= K, Z; = S. gentianoides – S, Y (var. alabamensis not discovered at the time)]

LYTHRACEAE J. St.-Hilaire 1805 (Loosestrife Family)

A family of about 27 genera and 600 species, herbs, shrubs, and trees, primarily tropical (a few warm temperate). References: Graham (1975)=Z. Keys adapted, in large part, from Z.

1 Plant woody or suffrutescent, a shrub or a small tree 1-10 m tall (with peeling, cinnamon-colored bark); petals present, showy, 8-20 mm long. 2 Flowers in axillary clusters; aquatic shrubs with arching suffrutescent or woody stems; leaves opposite or whorled...... Decodon 2 Flowers in terminal panicles; terrestrial shrubs or small trees with erect woody stems; leaves alternate to subopposite...... Lagerstroemia 1 Plant not woody, an herb 0.1-1.2 m tall; petals absent or present, inconspiscuous or showy, 1-10 mm long. 3 Stems pubescent. 4 Floral tube (hypanthium) swollen obliquely at its base; capsule dehiscing longitudinally along the upper surface...... Cuphea 4 Floral tube (hypanthium) symmetrical; capsule dehiscing septicidally at the apex...... Lythrum 3 Stems glabrous. 5 Floral tube cylindric to turbinate, about 2× as long as wide...... Lythrum 5 Floral tube campanulate to globose, about 1× long as wide. 6 Flowers or fruits (1-) 3-10 in the leaf axils (at least some axils with 2 or more flowers or fruits on a given plant) ...... Ammannia 6 Flowers or fruits solitary in the leaf axils (never > 1 per axil). 7 Capsule indehiscent; petals 0; sepals 4, broadly triangular, lacking intersepalary appendages; seeds spatulate or oblanceolate, about 1 mm long, minutely granular on one face and smooth on the other...... Didiplis LYTHRACEAE 483 7 Capsule dehiscing septicidally; petals 4; sepals 4 (-6), triangular, with intersepalary appendages of size about equal to the calyx lobes; seeds hemispheric, about 0.3 mm long, the surface very finely reticulate...... Rotala

{add Punica and Trapa to key}

Ammannia Linnaeus (Toothcup)

A genus of about 25 species, herbs, cosmopolitan. References: Graham (1985)=Y; Graham (1975)=Z. Key based on Y.

1 Style included in fruit, thick, 0.5-1.0 mm long (much shorter than the ovary); calyx lobes obtuse, often with the apices minutely mucronate; petals 0, 1, or 4, pale pink to white, to 1 mm long and 1 mm wide...... A. latifolia 1 Style exserted in fruit, filiform, 1.5-3.0 mm long (equal to or longer than the ovary); calyx lobes triangular, with acute apices; petals 4 (-5), deep rose-purple or pale lavender, ca. 2 mm long and 2 mm wide. 2 Inflorescence usually a short- or long-pedunculate cyme (sometimes reduced); flowers usually > 3 per axil; petals deep rose-purple; fruits 3.5-5 mm in diameter...... A. coccinea 2 Inflorescence sessile; flowers usually 1-3 per axil; petals pale lavender; anthers yellow; fruits 4-6 mm in diameter...... [A. robusta]

Ammannia coccinea Rottböll. Cp, Pd, Mt (GA, NC, SC, VA): marshy areas, ditches, exposed muddy river shores and banks, other wet places; uncommon (VA Watch List). July-October. NJ, OH, IN, IL, IA, and SD south to FL and TX; disjunct in CA; south through Mexico and Central America to n. South America. [= RAB, C, F, GW, K, S, W, Y, Z; > A. coccinea ssp. purpurea (Lamarck) Koehne – G; > A. coccinea – F, GW, S, Z (apparently also including A. robusta)] Ammannia latifolia Linnaeus. Cp (GA, NC, SC, VA): tidal marshes, wet places, ditches; uncommon. July-September. NJ south to s. FL and west to TX (mostly on the coastal plain), and also in the West Indies, Yucatan, Central America, and South America. All plants in North America north of Florida have flowers with petals; most plants from FL south through the West Indies into Central and South America have flowers without petals. Graham (1985) considered these forms; additional study is warranted. The name A. koehnei Britton is available for the petaliferous North American plant should its recognition prove warranted. [= C, GW, K, W, Y, Z; > A. teres Rafinesque – RAB, G; > A. teres var. teres – F; > A. teres var. exauriculata (Fernald) Fernald – F; > A. latifolia – S; > A. koehnei Britton – S]

Ammannia robusta Heer & Regel. OH and British Columbia south to w. KY, w. TN, LA, TX, CA, Mexico, and Belize; West Indies; introduced in NJ and n. South America. [= C, K, Y; < A. coccinea – F, GW, S, Z]

Cuphea P. Browne (Waxweed)

A genus of about 260 species, herbs, of America. References: Graham (1975)=Z.

1 Floral tube green, 4.5-6 mm long, glabrous inside; stamens much shorter than the floral tube; petioles to 8 mm long, often very short ...... C. carthagenensis 1 Floral tube purple-green, 6-10 mm long, villous inside; stamens equal to or exceeding the the floral tube; petioles to 20 mm long...... C. viscosissima

* Cuphea carthagenensis (Jacquin) J.F. Macbride. Cp (GA, NC, SC), Pd (GA): marshes, ditches, wet places; common, native of South America. June-September. [= GW, K, Z; = C. carthagensis – RAB (a misspelling); = Parsonsia balsamona (Chamisso & Schlechtendahl) Standley – S] Cuphea viscosissima Jacquin. Pd, Mt (GA, NC, SC, VA), Cp (VA): dry or wet places; uncommon. July-October. NH west to IA and KS, south to c. GA, LA, and e. OK. [= RAB, C, GW, K, W, Z; = C. petiolata (Linnaeus) Koehne – F, G; = Parsonsia petiolata (Linnaeus) Rusby – S]

* Cuphea procumbens Gómez Ortega is reported for NC by Small (1933). Graham (1975) considers this Mexican species to be represented in se. United States only by "garden escapes that do not persist." {not keyed} [= K, Z; = Parsonsia procumbens (Gómez Ortega) Heller – S]

Decodon J.F. Gmelin (Water-oleander, Water-willow)

A monotypic genus, a weak shrub, endemic to e. North America. References: Graham (1975)=Z.

Decodon verticillatus (Linnaeus) Elliott, Water-oleander, Water-willow, Swamp Loosestrife, Peatweed. Cp (GA, NC, SC, VA), Mt (GA, VA): natural lakes, limesink ponds, peatlands, peaty swamps, not known in the Piedmont or Mountains of NC or SC, but scattered in the Ridge and Valley of VA; common. July-September. Nova Scotia, Ontario, and MN south to c. LYTHRACEAE 484 peninsular FL and e. TX. The lower stems are spongy in texture. [= RAB, GW, K, S, W, Z; > D. verticillatus var. verticillatus – C, F, G; > D. verticillatus var. laevigatus Torrey & Gray – C, F, G]

Didiplis Rafinesque (Water-purslane)

A monotypic genus, an herb, endemic to e. North America. Perhaps better merged into Lythrum. References: Graham (1975)=Z.

Didiplis diandra (Nuttall ex Augustin de Candolle) Wood, Water-purslane. Cp (NC, VA), Pd (SC): stagnant water of pools, streams, and old beaverponds; rare (NC Rare, VA Rare). April-August. VA, IN, and WI south to NC, MS, and LA. [= C, G, GW, K, S, Z; = Peplis diandra Nuttall ex Augustin de Candolle – RAB, F; Lythrum]

Lagerstroemia Linnaeus (Crape-myrtle)

A genus of about 53 species, trees, of tropical se. Asia and Australia. References: Graham (1975)=Z.

* Lagerstroemia indica Linnaeus, Crape-myrtle. Cp, Pd (GA, NC, SC): commonly cultivated, persistent around old plantings, weakly spreading; rare (in the wild), native of Asia. June-September. [= C, K, S, Z]

Lythrum Linnaeus (Loosestrife)

A genus of about 36 species, herbs, cosmopolitan. References: Graham (1975)=Z.

1 Flowers numerous in terminal spike-like thyrses; stamens usually 12; leaves opposite or whorled ...... L. salicaria 1 Flowers solitary or paired in axils; stamens usually (4-) 6; leaves opposite or alternate. 2 Leaves opposite throughout, mostly shorter than to as long as the internodes, 1-4 mm wide...... L. lineare 2 Leaves opposite below, alternate above, mostly longer than the internodes, 2-14 mm wide. 3 Floral tube 3-4 mm long; petals 2-3 mm long; calyx appendages about the same length as the calyx lobes; branch leaves abruptly and definitely reduced in size relative to the stem leaves, and widely spaced; [of sw. GA southward] ...... L. curtissii 3 Floral tube 5-6 mm long; petals 5-6 mm long; calyx appendages about 2× as long as the calyx lobes; branch leaves gradually reduced relative to the stem leaves, dense and overlapping; [collectively widespread in our area]. 4 Leaves ovate to lanceolate, widest at a point 1/6 to 1/2 of the way from the base to the apex, the base rounded to subcordate; stems mostly slender, to 8 dm tall; bracteoles mostly at the base of the pedicel...... L. alatum 4 Leaves lanceolate to linear-lanceolate, widest at a point 1/3 to 2/3 of the way from the base to the apex, the base cuneate, often narrowly so; stems stout, to 13 dm tall; bracteoles mostly on the upper pedicel, immediately below the floral tube...... L. lanceolatum

Lythrum alatum Pursh, Northern Winged Loosestrife. Mt (GA, VA), Pd (VA): calcareous meadows, marl fens, and disturbed wet calcareous places; rare (VA Rare). June-September. ME, NY, MI, and ND south to n. and w. VA, e. TN, nw. GA, n. AL, n.. AR, ne. OK, and CO. [= F, S, W; = L. alatum var. alatum – C, G, GW, K, Z] Lythrum curtissii Fernald, Curtiss's Loosestrife. Cp (GA): calcareous swamps; rare (GA Threatened). June-early September. Sw. GA south to Panhandle FL; the report from Emanuel County, GA (Jones & Coile 1988) is in error. [= GW, K, S, Z] Lythrum lanceolatum Elliott, Southern Winged Loosestrife. Cp, Pd (GA, NC, SC, VA): moist to wet places; uncommon (VA Rare). May-September. Se. VA, se. NC, SC, GA, AL, MS, n. AR, and OK south to s. FL, s. TX, and in the West Indies. Although Graham (1975) argues that L. lanceolatum should be reduced to a variety of L. alatum, her evidence can also be interpreted as warranting specific status. [= RAB, F, S; = L. alatum Pursh var. lanceolatum (Elliott) Torrey & A. Gray ex Rothrock – C, G, GW, K, Z] Lythrum lineare Linnaeus, Narrowleaf Loosestrife. Cp (GA, NC, SC, VA): nearly fresh, brackish, and saline marshes; uncommon. July-October. NJ south to FL and west to TX. [= RAB, C, F, G, GW, K, S, Z] * Lythrum salicaria Linnaeus, Purple Loosestrife. Mt, Pd (NC, VA), Cp (VA): wet places; uncommon, native of Eurasia. June-September. An extremely noxious weed in the ne. United States, aggressively colonizing and coming to dominate a wide variety of freshwater wetlands, sometimes to the near exclusion of native vegetation. [= RAB, C, G, K, W, Z; > L. salicaria var. salicaria – F; > L. salicaria var. gracilior Turczaninow – F; > L. salicaria var. tomentosum (P. Miller) Augustin de Candolle – F]

Punica Linnaeus (Pomegranate)

LYTHRACEAE 485 A genus of 2 species, trees, of Mediterranean Europe and w. Asia. Sometimes treated in the monogeneric family Punicaceae; here included in Lythraceae, following Angiosperm Phylogeny Group (2003). References: Zohary & Hopf (1994).

* Punica granatum Linnaeus, Pomegranate, is reported as cultivated on Hatteras Island (Dare County, NC) (Brown 1959). It is probably not established, but may be persistent. This species has been cultivated in the Old World at least since the 3rd millenium B.C. [= K, S]

Rotala Linnaeus (Toothcup)

A genus of about 44 species, wetland herbs, of temperate to tropical areas. References: Graham (1975)=Z.

Rotala ramosior (Linnaeus) Koehne. Cp, Pd, Mt (GA, NC, SC, VA): marshes, ditches, exposed drawdown muds and silts; common. June-October. Widespread in e. North America, also in the West Indies, Central America, South America, and on the west coast of the United States and Mexico. [= RAB, C, GW, K, S, W, Z; > R. ramosior var. ramosior – F, G; > R. ramosior var. interior Fernald & Griscom – F, G]

Trapa Linnaeus (European Water-chestnut)

A genus of 1 highly polymorphic or up to 15 more narrowly defined species, annual aquatic herbs, native of the Old World. Oftwn placed in a monogeneric family, Trapaceae. References: Angiosperm Phylogeny Group (2003).

* Trapa natans Linnaeus, European Water-chestnut, Water-caltrop. Cp (VA): farm ponds and other stagnant or slow-moving water; rare, introduced from Eurasia and Africa. June-September. [= C, F, G, K]

MAGNOLIACEAE A.L. de Jussieu 1789 (Magnolia Family)

A family of about 7 genera and 223 species, trees and shrubs, tropical and warm temperate, of e. and se. Asia, and from e. North America south through West Indies and Central America to Brazil. References: Hardin (1972); Hardin & Jones (1989)=Z; Meyer in FNA (1997); Figlar & Nooteboom (2004); Frodin & Govaerts (1996); Nooteboom in Kubitzki, Rohwer, & Bittrich (1993); Kim et al. (2001).

1 Leaves about as broad as long, (2-) 4 (-8)-lobed; fruit a 2-seeded, indehiscent samara; [subfamily Liriodendroideae]...... Liriodendron 1 Leaves longer than broad, not lobed (in some species the leaves auriculate-cordate basally); fruit a cone-like aggregate, each follicle dehiscing to reveal the scarlet seed, at first connected to the follicle by a thread-like strand; [subfamily Magnolioideae]...... Magnolia

Liriodendron Linnaeus (Tulip-tree)

A genus of 2 species, trees, relictually distributed, with L. tulipifera in e. North America and L. chinense (Hemsley) Sargent in c. China and n. Vietnam. References: Nooteboom in Kubitzki, Rohwer, & Bittrich (1993); Weakley & Parks (in prep.), abbreviated as Z.

1 Leaves large, 4-8-lobed, the terminal lobes acute; [of the Mountains, Piedmont, and Coastal Plain (especially brownwater rivers and mesic bluffs and slopes)]...... L. tulipifera var. tulipifera 1 Leaves small, 0-4-lobed, the terminal lobes obtuse to broadly rounded; [of the Coastal Plain, especially fire-maintained, acidic, and peaty sites] ...... L. tulipifera var. 1

Liriodendron tulipifera Linnaeus var. tulipifera, Tulip-tree, Yellow Poplar, Whitewood. Mt, Pd, Cp (GA, NC, SC, VA): mesic forests, cove forests in the Mountains to at least 1500m in elevation, bottomland forests and swamps; common. April- June; September-October. Widespread in e. North America. An important timber tree in the Southern Appalachians. [= Z; < L. tulipifera – RAB, C, F, FNA, G, GW, K, S, W, Z] Liriodendron tulipifera Linnaeus var. 1, Coastal Plain Tulip-tree, Southern Yellow Poplar. Cp (GA, NC, SC, VA): blackwater swamps, streamhead pocosins in the fall-line sandhills; uncommon. April-June; September-October. Its occurrence in fire-maintained, acid soil habitats in the Coastal Plain is surprising to people used to Liriodendron as a tree of mesic, rich soil forests. It is, however, a typical species of streamhead pocosins in the fall-line sandhills, growing with Pinus serotina, Nyssa biflora, and Acer rubrum, and often with scorch marks twenty feet up the trunk. [= Z; < L. tulipifera – RAB, C, F, FNA, G, GW, K, S, W, Z]

MAGNOLIACEAE 486 Magnolia Linnaeus 1753 (Magnolia, Cucumber-tree)

A genus of about 130 species, trees and shrubs, of e. Asia (Himalayas and Sri Lanka to Japan and w. Malaysia) and America (e. North America to West Indies, Central America, and South America). Molecular phylogenetics show Magnolia virginiana and M. grandiflora as closely related in a New World primarily subtropical clade, M. macrophylla in a clade with its close relatives, M. fraseri and M. pyramidata together, M. acuminata as basal in a clade that is otherwise Asian (equivalent to subgenus Yulania), and M. tripetala grouped in another clade that is otherwise Asian (Azuma et al. 2001). The sections used follow Figlar & Nooteboom (2004). References: Tobe (1998)=Y; Spongberg (1998)=X; Frodin & Govaerts (1996)=V; Azuma, Thien, & Kawano (1999); Azuma et al. (2001); Figlar & Nooteboom (2004); Nooteboom in Kubitzki, Rohwer, & Bittrich (1993); Kim et al. (2001); Hunt (1998).

1 Leaves cordate-auriculate at base; [subgenus Magnolia]. 2 Leaves glaucous and finely appressed-pubescent beneath; buds and twigs pubescent; [subgenus Magnolia, section Macrophylla] ...... M. macrophylla 2 Leaves green and glabrous beneath; buds and twigs glabrous; [subgenus Magnolia, section Auriculata]. 3 Stamens 8-15 mm long; leaves (most of them) over 25 cm long; conelike aggregate fruit 6.5-11 (-14) cm long...... M. fraseri 3 Stamens 4-8 (-10.5) mm long; leaves (most of them) < 25 cm long; conelike aggregate fruit 3.5-5.5 (-6) cm long...... M. pyramidata 1 Leaves cuneate to rounded (subcordate) at base. 4 Leaves evergreen, coriaceous in texture, glossy dark green above as if varnished, rusty tomentose or green beneath; [subgenus Magnolia, section Magnolia]...... M. grandiflora 4 Leaves variably evergreen to deciduous, herbaceous or subcoriaceous in texture, medium green above with a slightly glossy or dull finish; glaucous or green beneath. 5 Leaves evergreen to deciduous, aromatic when fresh, 8-20 cm long, elliptic, strongly glaucous beneath; [subgenus Magnolia, section Magnolia]. 6 Leaves evergreen; previous year's stems densely pubescent; mature leaves with pubescent midveins; flowers opening near sundown (2-5 hours later than var. virginiana); medium to large tree, to >20 m tall; [of the Gulf Coast and inland, north and east to s. SC (e. NC?)] ...... M. virginiana var. australis 6 Leaves evergreen to deciduous (at least tardily); previous year's stems glabrous; mature leaves with few hairs along the midvein below; flowers opening mid-afternoon; shrub to multi-stemmed small tree, to 10 m (rarely to 15 m) tall; [of the Atlantic Coastal Plain and inland, south and west to s. SC and w. NC] ...... M. virginiana var. virginiana 5 Leaves deciduous, non-aromatic,10-50 cm long, either ovate or lance-obovate, green beneath. 7 Leaf base cuneate-attenuate; leaves 15-50 cm long, broader toward the tip, borne clustered on the end of the twig; buds glabrous; [subgenus Magnolia, section Rhytidospermum, subsection Rhytidospermum] ...... M. tripetala 7 Leaf base rounded to subcordate (often cuneate to widely cuneate in M. acuminata var. subcordata); leaves 10-30 cm long, broader near the middle or toward the base, borne scattered along the twig; buds pubescent; [subgenus Yulania, section Yulania, subsection Tulipastrum]. 8 Twigs of the current year glabrous; petals greenish or greenish-yellow; medium to large tree...... M. acuminata var. acuminata 8 Twigs of the current year pubescent, or at least with persistent hair-bases, petals golden-yellow above, light-yellow below; small tree (rarely larger)...... M. acuminata var. subcordata

Magnolia acuminata (Linnaeus) Linnaeus var. acuminata, Cucumber-tree, Cucumber Magnolia. Mt, Pd (GA, NC, SC, VA): mesic to subxeric forests, especially (but by no means strictly) over mafic or calcareous rocks, up to at least 1550m (where growing with Betula alleghaniensis, Abies fraseri, Picea rubens, and Sorbus americana), ultramafic outcrop barrens (where codominant with Pinus rigida and Quercus alba); common (rare in Piedmont). April-May; July-August. Widespread in e. North America, primarily in or near the Appalachians. The recognition of two varieties is uncertain (see discussion below). [= C, F, G, V, W, X, Y; < M. acuminata – RAB, FNA, K, Z; = Tulipastrum acuminatum (Linnaeus) Small – S] Magnolia acuminata (Linnaeus) Linnaeus var. subcordata (Spach) Dandy, Yellow Cucumber-tree, Showy Cucumber Magnolia. Pd (GA, NC, SC): moist to dry slopes and bottomlands over mafic or calcareous rocks; uncommon. Var. subcordata ranges from sc. NC south to AL, in the lower Piedmont in our area. It has been treated variously as a variety, a species, or merged with M. acuminata. Coker (1943) discusses its history, distribution, and taxonomic status. Additional study is needed. [= V, X, Y; < M. acuminata – RAB, FNA, K, W, Z; = Tulipastrum cordatum (Michaux) Small – S; = Magnolia cordata Michaux] Magnolia fraseri Walter, Fraser Magnolia, Earleaf Umbrella-tree. Mt, Pd (GA, NC, SC, VA): mesic forests; common (uncommon in upper Piedmont only). April-May; July-August. A Southern Appalachian endemic: KY and w. VA south through w. NC and e. TN to nw. SC, n. GA, and ne. AL. [= RAB, C, F, FNA, G, K, S, W, Z; = M. fraseri var. fraseri – V, X; = M. fraseri ssp. fraseri – Y] Magnolia grandiflora Linnaeus, Southern Magnolia, Bull Bay. Cp, Pd (GA, NC, SC, VA): maritime forests, mesic Coastal Plain bluffs and flats, bottomlands, now also widely naturalized, spreading from cultivation into wet to mesic forests; common as an introduction (rare in the Piedmont), rare as a native species in NC (NC Watch List). May-June; October. Curtis (1860) states MAGNOLIACEAE 487 that "the northern limit of this tree is in Brunswick County, south of the Cape Fear; but it flourishes in cultivation through all the lower part of the State." The pre-Columbian range was apparently from se. NC south to FL, west to e. TX, largely on the Coastal Plain, now somewhat expanded northwards and inland by naturalization from centuries of horticultural planting. This is, of course, the classic "southern magnolia," along with live oak (Quercus virginiana), and bald-cypress (Taxodium distichum), one of the totem trees of the Deep South. [= RAB, C, FNA, GW, K, S, V, Y, Z] Magnolia macrophylla Michaux, Bigleaf Magnolia. Pd (GA, NC, SC), Mt (VA), Cp (GA, SC*): mesic forests, primarily over limestone, other calcareous sedimentary rocks (calcareous shales, sandstones, etc.), or mafic rocks (east of the Blue Ridge); rare (NC Rare, SC Rare, VA Rare). May-June; July-August. The range of this species is often stated in such a way as to imply that it is a tree of the southern mountains. Actually, it avoids the Southern Blue Ridge, reaching its greatest abundance in the sedimentary rock Appalachians west of the Blue Ridge, particularly the Cumberland Plateau, and occurs east of the Blue Ridge only as a rare disjunct. M. macrophylla ranges from s. OH and sw. VA south through e. TN to w. GA, west to AL, MS, n. LA, and se. AR (Sundell et al. 1999); disjunct on Crowleys Ridge in ne. AR (population now extirpated), c. and nc. SC, and e. SC (where probably not native). The leaves are up to 1.1 meter long and 3.5 dm wide. See Williams (1999) for additional information about the discovery and nomenclature of this species. The Gulf Coast endemic Magnolia ashei Weatherby is related and is sometimes treated as a variety or subspecies of M. macrophylla. [= RAB, C, F, FNA, G, K, S, W, Z; = M. macrophylla ssp. macrophylla – V, X, Y] Magnolia pyramidata Bartram, Pyramid Magnolia. Cp (GA, SC): mesic forests, especially of bluffs and ravines; rare (GA Watch List, SC Rare). April-May; August. A Southeastern Coastal Plain endemic: c. SC south to panhandle FL, west to e. TX. Sometimes treated as avariety or subspecies of M. fraseri, to which it is clearly closely related, but the distributional and morphological differences are discrete and I therefore prefer specific status. [= RAB, FNA, K, S, Z; = M. fraseri Walter var. pyramidata (Bartram) Pampinini – V, X; = M. fraseri Walter ssp. pyramidata (Bartram) E. Murray – Y] Magnolia tripetala (Linnaeus) Linnaeus, Umbrella Magnolia, Umbrella-tree. Pd, Mt, Cp (GA, NC, SC, VA): mesic forests; common (uncommon in Mountains and Coastal Plain). April-May; July-October. Centered in the Southern Appalachians, but avoiding higher elevations, and therefore occurring primarily "around" the Blue Ridge; ranging from sc. and sw. PA, s. OH, s. IN south to SC, GA, nw. FL, AL, and MS; also disjunct in the Ouachita Mountains of c. AR and e. OK. [= RAB, C, F, FNA, G, K, S, W, V, X, Y, Z] Magnolia virginiana Linnaeus var. australis Sargent, Southern Sweet Bay. Cp (GA, NC?, SC), Pd (GA), Mt (GA): pocosins, bay forests, and swamps in the Coastal Plain, streamhead pocosins, swamps, and sandhill seeps in the Sandhills, bogs and peaty swamps in the Piedmont and Mountains; common (rare in Piedmont and Mountains). April-July; July-October. Primarily a species of the Southeastern Coastal Plain: s. SC (se. NC?) south to s. FL, and west to e. TX, rarely extending into adjacent, more interior provinces. Morphological, molecular, and chemical studies have shown strong variation in M. virginiana, but the patterns are not clear based on the limited current studies (Azuma, Thien, & Kawano 1999). Tobe (1998), McDaniel (1966), and Figlar (pers. comm. 2005) recommend the recognition of two varieties; additional study is needed. [= F, Y; < M. virginiana – RAB, C, FNA, G, GW, K, S, V, W, X, Z] Magnolia virginiana Linnaeus var. virginiana, Northern Sweet Bay. Cp (GA, NC, SC, VA), Pd (NC, VA), Mt (VA): pocosins, bay forests, and swamps in the Coastal Plain, streamhead pocosins, swamps, and sandhill seeps in the Sandhills, bogs and peaty swamps in the Piedmont; common (rare in Piedmont and Mountains). April-July; July-October. Se. MA south to w. NC, s. SC, and e. GA. [= F, Y; < M. virginiana – RAB, C, FNA, G, GW, K, S, V, W, X, Z]

MALVACEAE A.L. de Jussieu 1789 (Mallow Family)

Malvaceae has always been difficult to circumscribe cleanly, relative to members of such families as Sterculiaceae and Tiliaceae. Molecular evidence now adds to morphologic evidence that traditional circumscriptions of these families are highly polyphyletic. Bayer et al. (1999) present a new classification of an expanded Malvaceae, with 9 subfamilies recognized. If circumscribed broadly (as here) to include Sterculiaceae and Tiliaceae, a family of about 243 genera and 4000-4500 species, herbs, shrubs, and trees, of cosmopolitan distribution, but especially diverse in the tropics and subtropics. This family includes several economically important species, including cacao or chocolate, Theobroma cacao Linnaeus, and cola, Cola acuminata R. Brown. References: Whetstone (1983)=Z; Bayer et al. (1999); Bayer & Kubitzki in Kubitzki & Bayer (2003).

Subfamily Byttnerioideae: Melochia

Subfamily Grewioideae: Triumfetta

Subfamily Malvoideae: Tribe Gossypieae: Gossypium Tribe Hibisceae: Abelmoschus, Hibiscus, Kosteletzkya, Malvaviscus, Pavonia, Urena Tribe Malveae: Abutilon, Alcea, Althaea, Anoda, Callirhoe, Iliamna, Malva, Malvastrum, Modiola, Napaea, Sida

Subfamily Sterculioideae: Firmiana

Subfamily Tilioideae: Tilia

MALVACEAE 488 Abelmoschus Medikus 1787 (Okra, Gumbo)

A genus of about 15 species, herbs, of the Old World tropics. Perhaps better included in a broadly circumscribed Hibiscus (Pfeil & Crisp 2005). References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Abelmoschus esculentus (Linnaeus) Moench, Okra, Gumbo. Cp, Pd (GA, NC, SC, VA): frequently cultivated in gardens, rarely persistent or self-seeding the year following; common (in cultivation), rare (as an escape), introduced from Africa. The young capsules are a famous component of southern cooking. [= K, S; = Hibiscus esculentus Linnaeus – F]

Abutilon P. Miller 1754 (Indian-mallow, Indian-hemp)

A genus of about 100-160 species, herbs, of tropical and warm temperate areas. References: Fryxell (2002)=Z; Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Abutilon theophrasti Medikus, Velvetleaf, Pie-marker, Butterprint. Cp (GA, NC, VA), Pd (NC, SC, VA), Mt (GA, NC, SC, VA): fields, roadsides, disturbed areas; common, introduced from s. Asia. June-October. [= F, G, K, W, Z; = A. theophrastii – RAB, orthographic variant; = Abutilon abutilon (Linnaeus) Rusby – S]

Alcea Linnaeus 1753 (Hollyhock)

A genus of about 50-60 species, warm temperate Eurasian (Mediterranean Europe to c. Asia). References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Alcea rosea Linnaeus, Hollyhock. Cp, Pd, Mt (NC, VA) {GA}: roadsides, dumps, frequently cultivated, less commonly escaped or persistent; rare, introduced from Eurasia. Late May-August (rarely later). [= K; = Althaea rosea (Linnaeus) Cavanilles – RAB, C, F, G]

Althaea Linnaeus 1753 (Marsh-mallow)

A genus of about 12 species, herbs, Eurasian. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Althaea officinalis Linnaeus, Marsh-mallow. Cp (VA): marshes; rare, introduced from Europe. The roots of this plant were the original source of the mucilaginous paste used to make marshmallows (which are now made with a synthetic mucilage). [= C, F, G, K]

Anoda Cavanilles 1785 (Anoda)

A genus of about 24 species, herbs, of sw. North America, Central America, and South America. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Anoda cristata (Linnaeus) Schlechtendahl, Spurred Anoda. Cp (VA), Pd (GA, NC, SC, VA): disturbed areas; rare, introduced from sw. United States, Central and South America. July-October. [= G, K; = A. crista – C, orthographic variant; > A. cristata var. cristata – F; > A. cristata var. brachyanthera (Reichenbach) Hochreutiner – F]

Callirhoe Nuttall 1821 (Poppy-mallow)

A genus of about 9 species, herbs, of North America. References: Dorr (1990)=Z; Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Calyx not subtended by an epicalyx. 2 Inflorescence racemose, corymbose, or nearly umbellate; petals white, pink, or mauve; plants ascending, 1.5-8.5 dm tall; mericarps pubescent with simple, appressed hairs...... [C. alcaeoides] 2 Inflorescence paniculate; petals deep red, with a white basal spot; plants erect, 5-20 dm tall; mericarps glabrous ...... C. digitata 1 Calyx subtended by an epicalyx of 3 bractlets. 3 Calyx lobes distinct and divergent in bud; stems decumbent...... C. involucrata var. involucrata 3 Calyx lobes valvate in bud, forming a point; stems erect, ascending, or decumbent. 4 Bractlets of the epicalyx linear, 0.1-1.7 mm wide; peduncles 1-flowered; calyx lobes lanceolate, 7-15.4 mm long; mericarps indehiscent; leaves cordate or ovate in outline, palmately deeply divided into 5-7 lobes...... C. papaver MALVACEAE 489 4 Bractlets of the epicalyx obovate, 2.5-4.6 mm wide; peduncles several-flowered; calyx lobes deltoid, 2-5 (-6.5) mm long; mericarps dehiscent; leaves triangular, not lobed or only slightly so...... C. triangulata

* Callirhoe digitata Nuttall, Finger Poppy-mallow. Mt (GA): occasionally mowed roadside and adjacent powerline right-of- way, with other species of calcareous prairie habitats, one occurrence recorded to date; rare (GA Special Concern), apparently adventive from its native range in prairies of the Ozark region (nw. AR, sw. MO, se. KS, and ne. OK). [= C, F, G, K, Z] * Callirhoe involucrata (Torrey & A. Gray) A. Gray var. involucrata, Purple Poppy-mallow. Pd (VA): disturbed areas; rare, adventive from its native range in the midwestern United States. [= C, G, K, Z; < C. involucrata var. involucrata – F; < C. involucrata – G] Callirhoe papaver (Cavanilles) A. Gray, Woods Poppy-mallow. Cp (GA): forest openings; rare (GA Special Concern). Sw. GA and FL west to e. TX and s. AR (Dorr 1990). [= F, G, K, Z; = Callirrhoë papaver – S, orthographic variant] Callirhoe triangulata (Leavenworth) A. Gray, Sand Poppy-mallow, Clustered Poppy-mallow. Cp (GA, SC, NC): sandhills, sandy scrub, and other dry, open habitats; rare (GA Special Concern). Sc. NC south to GA and west to ec. MA (upper Coastal Plain and lower Piedmont); also sw. WI amd ne. IA south to s. IN and s. IL. [= C, F, G, K, Z; Callirrhoë triangulata – S, orthographic variant]

Callirhoe alcaeoides (Michaux) A. Gray, Pale Poppy-mallow. Calcareous prairies, glades, and other open habitats. East to c. TN (Chester, Wofford, & Kral 1997), IL, KY, and AL (Dorr 1990, Kartesz 1999) from a primary distribution in NE, KS, OK, and TX. [= C, F, G, K, Z; = Callirrhoë alceoides – S, orthographic variant]

Firmiana Marsili (Chinese Parasol-tree, Phoenix Tree)

A genus of about 12 species, trees, of Africa and Asia. References: Brizicky (1966)=Y; Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Firmiana simplex (Linnaeus) W. Wight, Chinese Parasol-tree, Phoenix Tree. Cp, Pd (GA, NC, SC, VA): planted and occasionally naturalized nearby; rare, native of se. Asia, probably China. [= C, K, Y, Z; = F. platanifolia (Linnaeus f.) Schott & Endlicher – RAB, S]

Gossypium Linnaeus 1753 (Cotton)

A genus of about 40-50 species, herbs, shrubs, and trees, of warm temperate to tropical areas. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Gossypium hirsutum Linnaeus, Upland Cotton. Cp (GA, NC, SC, VA): frequently cultivated crop, especially in sandy soils of the Coastal Plain, rarely adventive or a waif; common (as a crop), rare (as an adventive), introduced from tropical America. [= C, G; ? G. hirsutum var. hirsutum – K; ? G. herbaceum Linnaeus – F, S; ? G. barbadense Linnaeus]

Hibiscus Linnaeus 1753 (Hibiscus, Rose-mallow)

A genus of about 200-300 species, trees, shrubs, and herbs, of tropical to warm temperate areas. References: Wise & Menzel (1971); Bayer & Kubitzki in Kubitzki & Bayer (2003). Key based in part on GW.

1 Woody shrub, the stems usually solitary from a creeping rhizome ...... H. syriacus 1 Herb (sometimes robust and to as tall as 3.5 m), often several from ground level, from a crown or taproot. 2 Annual from a taproot, to 0.5 m tall; calyx inflated at maturity; capsule 1.0-1.3 cm long; petals 1.5-4 cm long; leaves 2- 6 cm long, deeply cleft...... H. trionum 2 Perennial from a crown, usually 0.7-3.5 m tall; calyx not inflated at maturity; capsule 1.7-3.5 cm long; leaves 4-25 cm long, deeply cleft, hastate-lobed, or not at all lobed or cleft. 3 Leaves and stems harshly scabrous; calyx lobes each with an elongate purplish nectary on the back; [of pine savannas and dry sandy soils of maritime forest edges, from se. NC southward] ...... H. aculeatus 3 Leaves and stems glabrous, softly pubescent, or slightly scabrous; calyx lobes lacking nectaries; [of marshes and swamps (sometimes cultivated in drier soils), collectively widespread in our area]. 4 Stem glabrous; leaves glabrous; leaves either palmately 3-5-lobed, or prominently halberd-lobed at the base (uncommonly unlobed). 5 Leaves either palmately 3-5-lobed; petals bright scarlet ...... H. coccineus 5 Leaves halberd-lobed at the base (uncommonly unlobed); petals pink or white with a purplish base...... H. laevis 4 Stem pubescent at least when young; leaves pubescent on at least one surface; leaves unlobed or slightly lobed toward the tip. 6 Upper leaf surface densely pubescent; capsule densely pubescent. MALVACEAE 490 7 Leaf blades 3-lobed ...... H. grandiflorus 7 Leaf blades unlobed...... H. moscheutos ssp. incanus 6 Upper leaf surface glabrous or very sparsely pubescent; capsule glabrous. 8 Petals white or creamy, with a red or purple base; flower stalk bearing a leaf well above the base (often about halfway along its length)...... H. moscheutos ssp. moscheutos 8 Petals pink (rarely white), with or without a red or purple base; flower stalk without a leaf, borne in the axil of a leaf...... H. moscheutos ssp. palustris

Hibiscus aculeatus Walter, Savanna Hibiscus, Comfort-root. Cp (GA, NC, SC): pine savannas, dry sandy soils of maritime forest edges; uncommon (rare north of SC) (NC Rare). June-August; July-September. Se. NC south to sc. peninsular FL, west to LA. [= RAB, GW, K, S] Hibiscus coccineus Walter, Scarlet Hibiscus. Cp (GA, *NC, *SC, *VA), *Pd (*NC): marshes, swamp forests, roadside swales, cultivated as an ornamental in yards; rare, presumably introduced from further south, but sometimes appearing native. Native in FL, s. GA, and s. AL. [= GW, K, S] Hibiscus grandiflorus Michaux, Large-flowered Hibiscus. Cp (GA): tidal marshes, lakeshores, wet flatwoods and savannas; rare (GA Special Concern). E. GA (Chatham Co., adjacent to the SC border) (Jones & Coile 1988) south to s. FL, west to e. LA. [= GW, K, S] Hibiscus laevis Allioni, Smooth Rose-mallow, Halberd-leaved Marsh-mallow, Showy Hibiscus. Cp, Pd (GA, NC, SC, VA), Pd (GA), Mt (VA): freshwater marshes, exposed riverbanks, sandbars; common. June-August; August-October. S. PA south to FL, west to TX; north in the interior to around the Great Lakes. [= C, K, W; = H. militaris Cavanilles – RAB, F, G, GW, S] Hibiscus moscheutos Linnaeus ssp. incanus (Wendland f.) Ahles, Eastern Rose-mallow. Cp (GA, NC, SC): marshes; rare. June-September; July-October. S. MD south to c. peninsular FL, west to e. TX. [= RAB, GW; < H. moscheutos var. moscheutos – C; = H. incanus Wendland f. – G, S; < H. moscheutos ssp. moscheutos – K, in part] Hibiscus moscheutos Linnaeus ssp. moscheutos, Eastern Rose-mallow. Cp, Pd, Mt (GA, NC, SC, VA): marshes; common. June-September; July-October. MD west to s. IN, south to n. FL, and se. TX. [= RAB, GW, W; < H. moscheutos var. moscheutos – C; = H. moscheutos Linnaeus – F, G; < H. moscheutos ssp. moscheutos – K; H. oculiroseus Britton – S] Hibiscus moscheutos Linnaeus ssp. palustris (Linnaeus) R.T. Clausen, Eastern Rose-mallow. Cp (NC): marshes; rare. June-September; July-October. E. MA south to e. NC; also around the Great Lakes in NY, OH, IN, MI, IL, and Ontario. [= RAB, GW, W; < H. moscheutos var. moscheutos – C; = H. palustris Linnaeus – F, G; < H. moscheutos ssp. moscheutos – K; ? H. moscheutos – S] * Hibiscus syriacus Linnaeus, Rose-of-Sharon, Althaea. Pd (GA, NC, SC, VA), Cp, Mt (NC, SC, VA): escaped or persistent after cultivation, often spreading by rhizomes; uncommon, introduced from e. Asia. June-September; August-October. [= RAB, C, F, G, K, S, W] * Hibiscus trionum Linnaeus, Flower-of-an-hour. Pd (GA, NC, SC, VA), Mt (NC, SC, VA), Cp (VA): fields, roadsides, disturbed areas; common, introduced from Europe. [= RAB, C, G, K, W; = Trionum trionum (Linnaeus) Wooton & Standley – S]

Hibiscus lasiocarpos Cavanilles, east to c. TN (Chester, Wofford, & Kral 1997), AL, KY, and GA (F, Kartesz 1999). [= F, S; = Hibiscus moscheutos Linnaeus ssp. lasiocarpos (Cavanilles) O.J. Blanchard – K] {not keyed at this time; add to synonymy}

Iliamna Greene 1906 (Globe-mallow)

A genus of 7 species, perennial herbs, of North America. Some authors (such as Mabberley 1997) include Iliamna in Sphaeralcea. References: Bodo Slotta & Porter (2006)=Y; Porter & Wieboldt in Terwilliger (1991)=Z; Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Leaves 5-7-lobed, the lobes narrowly triangular, the sinuses acute; flowers odorless; plant to ca. 1 m in height; [sandstone outcrops on ridgetop] ...... I. corei 1 Leaves 5-7-lobed, the lobes broadly triangular or deltoid, the sinuses obtuse; flowers fragrant; plant to ca. 2.5 m in height; [of river shores and along railroads] ...... I. remota

Iliamna corei Sherff, Peters Mountain Mallow. Mt (VA): in shallow soil in crevices of outcroppings of Clinch sandstone, near the summit of Peters Mountain; rare (US Endangered, VA Endangered). June-August; July-October. Endemic to the summit of Peters Mountain, Giles County, VA. The validity of I. corei as a species distinct from I. remota is supported by Bodo Slotta & Porter (2006). [= F, Y, Z; < I. remota – C, G, W; < I. rivularis (Douglas ex Hooker) Greene var. rivularis – K] Iliamna remota Greene, Kankakee Globe-mallow. Mt (VA): shores and gravel bars along rivers, and along railroad embankments; rare (VA Rare). June-August; July-October. W. VA, nw. IN, and ne. IL. Considered by some to be introduced only in our area, however, the VA populations are genetically different than those in IN and IL (Bodo Slotta & Porter 2006). [= F, Y, Z; < I. remota – C, G, W (also see I. corei); < I. rivularis (Douglas ex Hooker) Greene var. rivularis – K]

Kosteletzkya K. Presl 1835 (Seashore-mallow)

MALVACEAE 491 A genus of about 15-30 species, herbs, of North America, sub-Saharan Africa, and Madagascar. Perhaps better included in a broadly circumscribed Hibiscus (Pfeil & Crisp 2005). References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Hairs of the fruit 0.5-1.0 (-1.5) mm long; flowering calyx mostly 6-10 mm long, the bractlets mostly 2.5-6 mm long; petals mostly 1.8-3 cm long; [of VA northward] ...... K. virginica var. aquilonia 1 Hairs of the fruit 1.2-2.4 mm long; flowering calyx mostly 8-13 mm long, the bractlets mostly 6-10 mm long; petals mostly 3.2-4.5 mm long; [throughout our area]...... K. virginica var. virginica

Kosteletzkya virginica (Linnaeus) K. Presl ex A. Gray var. aquilonia Fernald, Northern Seashore-mallow. Cp (VA): brackish to freshwater tidal marshes; common. July-October. NY (Long Island) south to VA. While geographic trends are readily apparent, the recognition of infraspecific taxa is made problematic by the non-correlation of various characters. [= C, F, G; < Kosteletskya virginica – RAB, orthographic variant; < K. virginica – GW, K; < K. virginica – S] Kosteletzkya virginica (Linnaeus) K. Presl ex A. Gray var. virginica, Southern Seashore-mallow. Cp (GA, NC, SC, VA): brackish to freshwater tidal marshes; common. July-October. DE south to FL, west to TX; also in the West Indies. [= C; < Kosteletskya virginica – RAB, orthographic variant; < K. virginica – GW, K; > K. virginica var. virginica – F, G; > K. virginica var. althaeifolia Chapman – F, G; >< K. virginica – S; > K. althaeifolia (Chapman) Rusby – S]

Malva Linnaeus (Mallow)

A genus of about 40 species, herbs, of temperate Eurasia and montane Africa. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Upper leaves deeply 5-7-lobed, the sinuses cut over halfway to the middle; petals 2-3.5 cm long; erect perennial ...... M. moschata 1 Upper leaves less deeply lobed, rarely to as deep as halfway to the middle; petals 0.5-2.5 cm long; prostrate to erect annual or biennial. 2 Epicalyx of 3 oblong-ovate bractlets; petals reddish purple, 2.0-2.5 cm long; biennial, erect, usually not branched at the base ...... M. sylvestris 2 Epicalyx of 3 linear or narrowly lanceolate bractlets; petals white or pink, 0.6-1.2 cm long; annual, sprawling, usually branched at the base. 3 Petals about 2× as long as the sepals...... M. neglecta 3 Petals about 1× as long as the sepals...... M. rotundifolia

* Malva moschata Linnaeus, Musk Mallow, Rose Mallow. Mt (NC, VA), Pd (VA): pastures, roadsides, barnyards; rare, introduced from Europe. Late May-August. [= RAB, C, F, G, K, W] * Malva neglecta Wallroth, Common Mallow, Cheeses. Mt, Pd (GA, NC, VA), Cp (VA) {SC}: pastures, roadsides, barnyards; rare, introduced from Europe. April-October. [= RAB, C, F, G, K, W; = M. rotundifolia – S, misapplied] * Malva rotundifolia Linnaeus, Small Mallow, Dwarf Mallow, Cheeses. Mt (VA): pastures, roadsides, barnyards; rare, introduced from Europe. [= C, F, G, K, S; = M. pusilla, misapplied] * Malva sylvestris Linnaeus, Common Mallow, High Mallow, Cheeses. Cp, Pd (NC, SC, VA), Mt (VA): pastures, roadsides, barnyards; rare, introduced from Europe. May-July. [= RAB, C, K, S, W; > M. sylvestris var. sylvestris – F, G; > M. sylvestris var. mauritiana (Linnaeus) Boissier – F, G]

* Malva parviflora Linnaeus. SC, MD, FL, etc. [= K] {not keyed at this time; synonymy incomplete} * Malva verticillata Linnaeus is reported as an introduction as far south as s. PA (Rhoads & Klein 1993), MD, and WV (Kartesz 1999). [= K; > M. verticillata var. verticillata – C, F, G; > M. verticillata var. crispa Linnaeus – C, F, G] {not keyed at this time; synonymy incomplete}

Malvastrum A. Gray 1849

A genus of 14 species, herbs, of tropical and warm temperate areas. References: Bates (1967); Bayer & Kubitzki in Kubitzki & Bayer (2003).

Malvastrum hispidum (Pursh) Hochr. Mt (VA): limestone barrens; rare (VA Rare). July-August; August-October. KY, w. VA (Lee Co.), and c. TN, west to IA, KS, and OK. Discovered in our area in 1994 by J.C. Ludwig (Fleming & Ludwig 1996). [= C, K; Malvastrum angustum A. Gray – G, S; Sphaeralcea angusta (A. Gray) Fernald – F; = Sidopsis hispidum (Pursh) Rydberg; = Sida hispida Pursh]

* Malvastrum coromandelianum (Linnaeus) Garcke, introduced, south to se. PA (Rhoads & Klein 1993) and NJ (Kartesz 1999). [= K]

MALVACEAE 492 Malvaviscus Fabricius 1759 (Wax-mallow)

A genus of 3-4 species, herbs, of tropical and subtropical areas. Perhaps better included in a broadly circumscribed Hibiscus (Pfeil & Crisp 2005). References: Turner & Mendenhall (1993)=Z; Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Malvaviscus drummondii Torrey & A. Gray, Wax-mallow, Turk's-cap Mallow. Cp (GA, NC, SC): disturbed areas; rare, native of TX and n. Mexico. July-October. First reported for NC and SC by Leonard (1971b). Although Turner & Mendenhall 91993) cite Leonard’s specimens as M. arboreus var. arboreus, they were correctly determined by Leonard as M. drummondii. Therefore the attribution of M. arboreus var. arboreus to NC by Kartesz (1999) is an error. [= S; = M. arboreus Dillenius ex Cavanilles var. drummondii (Torrey & A. Gray) Schery – K, Z; = Hibiscus drummondii (Torrey & A. Gray) M.J. Young]

Melochia Linnaeus (Chocolate-weed)

A genus of about 54 species, of tropical regions, especially America. References: Brizicky (1966)=Y; Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Petioles >1.5 cm long; pubescence of the stem and leaves sparse, of stellate, forked, and/or simple hairs; cymes terminal on primary and secondary branches...... M. corchorifolia 1 Petioles < 1 cm long; pubescence of the stem and leaves dense (tomentose), of stellate hairs; cymes in upper leaf axils...... M. spicata

* Melochia corchorifolia Linnaeus, Chocolate-weed. Cp (GA, NC, SC): sandy fields, especially in low, wet places; uncommon, introduced from the Old World tropics. [= RAB, GW, K, S, Y, Z] * Melochia spicata (Linnaeus) Fryxell. Cp (GA): disturbed areas; rare, introduced from tropical America. In GA (Kartesz 1999) and FL (Brizicky 1966). [= K; = Riedlea hirsuta (Cavanilles) Alphonse de Candolle – S; = Melochia villosa (P. Miller) Fawcett & Rendle – Y]

Modiola Moench 1794 (Bristly-mallow)

A monotypic genus, an herb, of North America, Central America, and South America. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

Modiola caroliniana (Linnaeus) G. Don, Bristly-mallow. Cp, Pd (GA, NC, SC, VA): lawns, roadsides, disturbed areas, pondshores; uncommon (adventive in part of its range in our area). Late March-June (sometimes later). Ranging as a native from SC south to FL, west to TX, south into the tropics. [= RAB, C, F, G, GW, K, S]

Napaea Linnaeus 1753 (Glade-mallow)

A monotypic genus, an herb, of temperate c. North America. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

Napaea dioica Linnaeus, Glade-mallow. Mt (VA): floodplains; rare (US Species of Concern, VA Rare). June-August. PA and IA south to sw. VA and s. IL. The original distribution of this scarce species is difficult to determine. See the interesting discussion of this species' occurrence in VA in Wieboldt et al. (1998). [= C, F, G, K]

Pavonia Cavanilles 1787

A genus of about 150 species, of tropical and subtropical areas. Perhaps better included in a broadly circumscribed Hibiscus (Pfeil & Crisp 2005). References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

1 Leaves hastate; calyx lobes broadly ovate; carpels un-awned; [introduced species of disturbed habitats] ...... P. hastata 1 Leaves ovate; calyx lobes lanceolate; carpels with 3 apical awns up to 10 mm long; [rare native] ...... P. spinifex

* Pavonia hastata Cavanilles, Swampmallow. Cp (GA): disturbed areas; rare, introduced from tropical America. In se. GA (Jones & Coile 1988). [= K, S] Pavonia spinifex (Linnaeus) Cavanilles, Gingerbush. Cp (SC): hammocks; rare. Reported for the vicinity of Charleston, SC on the basis of a specimen collected by Bachman (Chapman 1878). Small (1933) considers this species as likely native, at least in FL. [= K, S; = Hibiscus spinifex Linnaeus]

Sida Linnaeus 1753 (Sida) MALVACEAE 493

A genus of about 100 species, shrubs and herbs, of tropical, subtropical, and warm temperate areas. References: Fryxell (1985)=Z; Fuertes, Fryxell, & Jansen (2003); Siedo (1999)=Y; Verdcourt (2004)=X; Bayer & Kubitzki in Kubitzki & Bayer (2003). Key adapted in part from Z.

1 Leaves deeply palmately lobed; plants 1-2 (-4) m tall; petals white; [section Pseudonapaea, to be removed from Sida]...... S. hermaphrodita 1 Leaves unlobed; plants 0.2-1 m tall; petals yellow. 2 Mericarps, styles, and stigmas 5; leaves usually truncate to subcordate at the base; [section Spinosae]...... S. spinosa 2 Mericarps, styles, and stigmas (6-) avg. 10 (-13); leaves usually cuneate to rounded at the base. 3 Leaves narrowly elliptic to linear, (3-) 4-20× as long as wide; [section Ellipticifoliae]...... S. elliottii var. elliottii 3 Leaves elliptic-rhombic, mostly 2-3× as long as wide; [section Sidae]. 4 Leaves and branches borne distichously; stipules usually falcate, several-veined ...... S. acuta 4 Leaves and branches borne spirally; stipules linear, 1 (-3)-veined...... S. rhombifolia var. rhombifolia

* Sida acuta Burman f., Broomweed. Cp (GA, SC): disturbed areas; rare, introduced from the Tropics. June-October. [= K, Z; ? S. carpinifolia Linnaeus f. – RAB, S] Sida elliottii Torrey & A. Gray var. elliottii, Coastal Plain Sida. Cp (GA, NC, SC, VA), Pd (NC), Mt (GA): stream banks, sandy openings, pineland pond margins, limestone glades and barrens; uncommon (GA Special Concern). July-October. Var. elliottii ranges from se. VA south to n. FL, west to LA and north in the interior to c. TN and se. MO. A second variety, var. parviflora Chapman, occurs in Peninsular FL, se. TX, and through montane e. Mexico to Guatemala. S. inflexa, of se. VA and ne. NC, is alleged to differ as follows: S. inflexa with calyx 7-10 mm long, leaves elliptic to narrowly elliptic, 4-20 mm wide, (3- ) 4-10× as long as wide (vs. S. elliottii var. elliottii with calyx 5-7 mm long; leaves narrowly lanceolate to linear, 1.5-7 mm wide, 10-20× as long as wide). [= Y; < S. elliottii – RAB, C, F, G, K, S, Z; > S. inflexa Fernald – F, K, Z] Sida hermaphrodita (Linnaeus) Rusby, Virginia Sida, Virginia-mallow. Mt, Pd (VA): sandy or rocky areas along riverbanks; rare. July-August. C. PA and MD west to s. OH, south to DC, WV, w. VA, and ne. TN; disjunct in nw. OH, ne. IN, and s. MI (where presumably native) and with additional collections from e. MA, NY (Long Island), and s. NJ (where probably adventive) (Sp[ooner et al. 1985). Fryxell (1985) comments that this species is so different from the rest of the genus that "one might plausibly argue that it be elevated to generic rank." A molecular phylogenetic analysis suggests that its affinities are not with Sida, but with the South American Sidasodes (Fuertes, Fryxell, & Jansen 2003). Spooner et al. (1985) provide a detailed review of the species. [= C, F, G, K, S, W, Z] * Sida rhombifolia Linnaeus var. rhombifolia, Arrowleaf Sida. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (GA, SC): roadsides, fields, gardens, disturbed areas; common. April-October. Verdcourt (2004) discusses variation in this taxon, and suggests that "studies throughout the entire range of the species will necessitate recognition of more than one species." He recognizes 6 varieties in e. Africa, aside from the Linnaean var. rhombifolia (with type in Jamaica). [= X; < S. rhombifolia – RAB, C, F, G, K, S, W, Z] * Sida spinosa Linnaeus, Prickly Sida, Prickly-mallow, False-mallow. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas, wet fields; common, introduced from the Tropics. June-November. [= RAB, C, F, G, K, S, W, Z]

Sida cordifolia Linnaeus. AL and FL. [= K] {not keyed at this time; synonymy incomplete}

Sphaeralcea St.-Hil. (see Iliamna)

Tilia Linnaeus (Basswood, Whitewood, Linden, Linn)

A genus of about 25-45 species, trees, of temperate regions of North America, Europe and Asia. Hardin's (1990) treatment of American Tilia seems a practical and reasonable approach; it gives taxonomic status to the more distinctive (and geographically based) elements of variation, while recognizing the intergradational nature of the variation. Further investigation of this complex group is, however, warranted. References: Hardin (1990)=Z; Bayer & Kubitzki in Kubitzki & Bayer (2003). Key adapted from Hardin (1990).

Identification notes: While the varieties treated below are broadly distinctive and have definite geographic distributions across e. North America, they are imperfectly distinct in geographic areas of overlap. In our area, their identification is particularly problematic in Virginia, where individuals in many parts of the state show intergradation between the northern var. americana and the Southern and Central Appalachian var. heterophylla.

1 Lower leaf surfaces puberulent with bulbous glands, acicular trichomes, and (rarely) sparsely scattered stellate trichomes; fruiting peduncles and pedicels glabrous or sometimes puberulent; [generally northern, south to VA and w. NC] ...... T. americana var. americana MALVACEAE 494 1 Lower leaf surfaces usually tomentose or becoming puberulent, with bulbous glands, acicular trichomes, and a predominance of stellate or fasciculate trichomes; fruiting peduncles and pedicels stellate-tomentulose (becoming puberulent in age); [collectively widespread in our area]. 2 Lower leaf surfaces pale or whitish, densely stellate tomentose with appressed, sessile-stellate trichomes obscuring the surface (rarely becoming puberulent with age but with some stellate trichomes persisting along major veins, the margin, and/or the apex; lateral buds 5-8 mm long; pericarp 0.8-1.0 mm thick; [widespread in our area] ...... T. americana var. heterophylla 2 Lower leaf surfaces grayish or brownish, loosely but densely tomentose with fasciculate and/or stipitate-stellate trichomes, either remaining tomentose or becoming puberulent, or puberulent from emergence and green beneath; lateral buds 3-5 mm long; pericarp 0.5-0.6 mm thick; [generally southern, Coastal Plain and Piedmont of NC, SC, GA and southward and westward] ...... T. americana var. caroliniana

Tilia americana Linnaeus var. americana, Northern Basswood. Mt (NC, VA), Pd, Cp (VA): rich coves, rocky slopes, metabasalt boulderfileds, rich north-facing river bluffs, calcareous Coastal Plain ravines; common (rare in NC) (NC Watch List). June; August-September. New Brunswick and Manitoba south to e. VA, w. NC, and OK. In VA, var. americana occurs throughout the northern half of the state, with scattered populations southward in the mountains. [= C, K, Z; = T. americana – RAB, G, W; > T. americana – F; > T. neglecta Spach – F, S; > T. glabra Ventenat – S; > T. truncata Spach – S] Tilia americana Linnaeus var. caroliniana (P. Miller) Castig., Southern Basswood, Carolina Basswood. Cp, Pd (GA, NC, SC): mesic forests, in the outer Coastal Plain usually associated with shell deposits, Indian shell middens, or underlying coquina limestone ("marl"); uncommon. June-July; July-August. NC south to FL and west to OK and c. TX. [= K, Z; > T. caroliniana P. Miller – RAB, S; > T. floridana Small – RAB, F, S; > T. georgiana Sargent – S; > T. littoralis Sargent – S] Tilia americana Linnaeus var. heterophylla (Ventenat) Loudon, Mountain Basswood, White Basswood, Linn. Mt, Pd, Cp (GA, NC, SC, VA): rich coves and mesic to dry slopes (the drier sites usually on limestone), often one of the most abundant trees in Southern Appalachian cove forests; common, rare in Coastal Plain. June; July-August. Centered in the Southern Appalachians: sw. PA and WV south to c. NC, wc. GA, FL panhandle, and westward as disjunct populations to the Ozarkian Highlands of s. MO and n. AR. In VA, var. heterophylla dominates the scene in sw. VA and along southern Piedmont river bluffs, with disjunct populations in calcareous ravines in the upper Coastal Plain (Surry County); it also extends less commonly into the northern VA mountains and foothills, where var. americana is more prevalent, but seems to be absent (or very uncommon) in the Potomac valley east of the Blue Ridge. [= C, K, Z; = T. heterophylla Ventenat – RAB, F, W; > T. heterophylla – G, S; > T. monticola Sargent – G; > T. australis Small - S; > T. eburnea Ashe – S; > T. lasioclada Sargent – S; > T. michauxii Nuttall – S; > T. venulosa Sargent]

Several European species are planted as street or yard trees; they differ from the native species in having smaller leaves (usually 4-12 cm long vs. 8-25 cm long) and lacking staminodes. Three of the more commonly planted species are T. platyphyllos Scopoli, T. cordata P. Miller, and T. ×vulgaris Hayne [cordata × platyphyllos]. Planted material should be identified by using appropriate manuals of cultivated species.

Triumfetta Linnaeus

A genus of about 70-150 species, trees, shrubs, and herbs, of tropical regions. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Triumfetta semitriloba Jacquin. Cp (GA): disturbed areas; rare, introduced from further south. In sw. GA (Jones & Coile 1988) and FL. [= K, S]

Urena Linnaeus (Caesarweed)

A genus of about 6 species, of tropical and subtropical regions. References: Bayer & Kubitzki in Kubitzki & Bayer (2003).

* Urena lobata Linnaeus, Caesarweed, Bur Mallow, Congo Jute. Cp (SC): roadsides and vacant lots; rare, introduced from se. Asia. Introduced to se. SC via landscaping plantings, spreading to vacant lots and raodsides (P. McMillan, pers. comm., 2005). [= GW, K, S]

MARTYNIACEAE Stapf 1895 (Martynia Family)

A family of 5 genera and about 16 species, herbs, tropical and subtropical. Bretting & Nilsson (1988) present evidence for maintaining the Martyniaceae as distinct from the Pedaliaceae. References: Ihlenfeldt in Kadereit (2004).

Proboscidea Schmidel 1763 (Unicorn-plant)

MALVACEAE 495 A genus of about 9 species, herbs, of warm temperate to subtropical America. References: Thieret (1977)=Y; Bretting & Nilsson (1988)=Z; Ihlenfeldt in Kadereit (2004).

* Proboscidea louisianica (Miller) Thellung, Unicorn-plant, Devil's-claw, Cow Catcher. Pd (GA, NC, SC), Cp (GA, VA): disturbed areas; rare, introduced from farther west and south (apparently native to the Mississippi valley). The curious fruits are unmistakable. [= RAB, F, GW, Y; = P. louisiana – C, G, orthographic variant; = Martynia louisiana Miller – S; > P. louisianica ssp. louisianica – K, Z]

MELASTOMATACEAE A.L. de Jussieu 1789 (Melastome Family)

A family of about 200 genera and 4500-5000 species, trees, shrubs, vines, and herbs, of tropical, subtropical , and warm temperate areas.

Rhexia Linnaeus 1753 (Meadow-beauty) (by Richard J. LeBlond)

A genus of about 15 species, herbs, of North America. Rhexia is the only genus of the Melastomataceae to occur in North America north of s. FL. References: Kral & Bostick (1969)=Z; Bounds (1987); Wurdack & Kral (1982); Snyder (1996).

Identification notes: measurements of the hypanthium are to the base of the calyx lobes.

1 Anthers straight, ca. 2 mm long. 2 Stem internodes with at least some hairs; leaves oblong, linear, or spatulate; petals yellow ...... Rh. lutea 2 Stem internodes glabrous; leaves ovate, suborbicular, or widely elliptic; petals lavender-rose to pink. 3 Calyx segments blunt to acute; floral tube glandular-pubescent; surface of seeds irregularly ridged ...... Rh. nuttallii 3 Calyx segments acuminate-aristate; floral tube nearly glabrous except along the calyx lobes; surface of seeds pebbled...... Rh. petiolata 1 Anthers curvate, 5-11 mm long. 3 Stem nodes and internodes glabrous; stem and foliage blue-green ...... Rh. alifanus 3 Stem nodes and usually also the internodes hirsute; stem and foliage green. 4 Sepal lobes aristate, the awn-tip 0.5-1.5 mm long, and also with flaring, yellowish, stiff hairs 3-5 mm long ...... Rh. aristosa 4 Sepal lobes obtuse to acuminate, not aristate, the hairs shorter and not yellowish or stiff. 5 Leaves 1-5 (-8) mm wide, linear, linear-elliptic, narrowly oblong, or narrowly spatulate. 6 Petals lavender-rose, (10-) 15-20 (-25) mm long; mature hypanthium 10-14 mm long, with glandular hairs; marginal nerves of leaf abaxial surface either absent or obscure and discontinuous .....Rh. cubensis 6 Petals white to pink (-rose-purple), (7-) 9-14 mm long; mature hypanthium 6-10 mm long, glabrous or sparsely glandular-hairy; marginal nerves of leaf abaxial surface prominent...... Rh. mariana var. exalbida 5 Leaves (5-) 7-20 (-35) mm wide, lanceolate, elliptic, or ovate. 7 Four stem faces at mid-stem about equal, almost flat, the angles sharp or winged. 8 Roots tuberous; stem angles at mid-stem conspicuously winged; hypanthium 7-10 mm long, the neck shorter than the body...... Rh. virginica 8 Roots not tuberous; stem angles sharp to narrowly winged; hypanthium 10-12 mm long, the neck as long as or longer than the body ...... Rh. mariana var. ventricosa 7 Four stem faces at mid-stem markedly unequal, one pair of opposite faces broader, convex, darker green, the narrower pair concave or flat, pale. 9 Mature hypanthium 6-10 (-11) mm long, glandular-setose; petals 12-15 (-18) mm long, glabrous on the lower surface; anthers 5-8 mm long...... Rh. mariana var. mariana 9 Mature hypanthium (9-)10-15 (-20) mm long, glabrous or glabrate; petals (18-) 20-25 mm long, glandular-hairy on the lower surface (best seen in bud); anthers 8-11 mm long ...... Rh. nashii

Alternate Key based largely on vegetative characters

1 Stem internodes glabrous. 2 Stem nodes as well as internodes glabrous, leaf margins entire or remotely low-toothed apically, glabrous...... Rh. alifanus 2 Stem nodes hirsute, leaf margins toothed, the teeth often tipped with hairs. 3 Longest leaves 1.5 (-2) cm long, ovate or suborbicular...... Rh. nuttallii or Rh. petiolata 3 Longest leaves > 2 cm long, lanceolate, elliptic, or ovate. 4 Rhizomes present, roots not tuberiferous or spongy-thickened...... Rh. mariana var. ventricosa MELASTOMATACEAE 496 4 Rhizomes absent, roots tuberiferous or spongy-thickened. 5 Stem leaves gradually reduced upward ...... Rh. virginica 5 Stem leaves gradually lengthening from the base to mid-stem...... Rh. aristosa 1 Stem internodes (and nodes) hirsute or glandular-hairy. 6 Leaves linear, narrowly elliptic, or broadest above the middle. 7 Plant bushy-branched...... Rh. lutea 7 Plant simple below the cymose inflorescence. 8 Mature hypanthium 10-14 mm long, with glandular hairs; petals lavendar-rose, 1.5-2.0 cm long...... Rh. cubensis 8 Mature hypanthium 6-10 mm long, glabrous or sparsely glandular-hairy; petals white, 1.2-1.5 cm long...... Rh. mariana var. exalbida 6 Leaves lanceolate, elliptic, or ovate, broadest at or below the middle. 9 Four stem faces at mid-stem about equal, almost flat, the angles sharp or winged. 10 Rhizomes present, roots not tuberiferous or spongy-thickened...... Rh. mariana var. ventricosa 10 Rhizomes absent, roots tuberiferous or spongy-thickened ...... Rh. virginica 9 Four stem faces at mid-stem markedly unequal, one pair of opposite faces broader, convex, darker green, the narrower pair concave or flat, pale. 11 Mature hypanthium 6-10 mm long, glandular-hairy; petals 1.2-1.5 cm long, glabrous on the lower surface ...... Rh. mariana var. mariana 11 Mature hypanthium 10-15 mm long, glabrous or glabrate; petals 2.0-2.5 cm long, glandular-hairy on lower surface (best seen in bud)...... Rh. nashii

Rhexia alifanus Walter, Smooth Meadow-beauty. Cp (GA, NC, SC): pine flatwoods and savannas, pocosins borders, more able to tolerate merely moist soils than other Rhexia species; common. May-September. A Southeastern Coastal Plain species: e. NC south to n. FL and west to se. Texas. Our tallest and showiest Rhexia: the unbranched (unless injured), wandlike stems, with strongly ascending, bluish-green, generally entire leaves make this species unmistakeable. [= RAB, GW, K, S, Z] Rhexia aristosa Britton, Awned Meadow-beauty, Bristly Meadow-beauty. Cp (GA, NC, SC): clay-based Carolina bays, depression meadows, and limesink ponds (dolines); rare (US Species of Concern, GA Special Concern, NC Threatened, SC Rare, VA Rare). June-September. This species has a very local and disjunct range extending (strictly on the Coastal Plain) from NJ south to AL. The long yellowish bristles at the summit of the calyx/hypanthium are diagnostic. [= RAB, C, F, G, GW, K, S, Z] Rhexia cubensis Grisebach, West Indies Meadow-beauty. Cp (GA, NC, SC): limesink ponds (dolines); rare (NC Rare). June-September. Se. NC south to s. FL and west to sw. MS; also in the West Indies. [= RAB, GW, K, S, Z] Rhexia lutea Walter, Yellow Meadow-beauty, Golden Meadow-beauty. Cp (GA, NC, SC): wet pine flatwoods and savannas, seepage slopes, and bogs; uncommon. April-July (and later in response to growing-season fire). A Southeastern Coastal Plain species: e. NC south to n. FL and west to se. TX. The only yellow-flowered Rhexia and also our bushiest species. [= RAB, GW, K, S, Z] Rhexia mariana Linnaeus var. exalbida Michaux, White Meadow-beauty. Cp, Pd (GA, NC, SC): wet pine flatwoods and savannas, wet meadows, ditches, and wet roadsides; uncommon. June-September. NC south to FL and west to MS. Merging into Rh. mariana var. mariana from FL westward, var. exalbida appears quite distinct in NC, and it here recognized for convenience. The white flowers and linear leaves are diagnostic. [= RAB; Rh. lanceolata – S; < Rh. mariana var. mariana – GW, K, Z] Rhexia mariana Linnaeus var. mariana, Maryland Meadow-beauty, Dull Meadow-beauty, Pale Meadow-beauty. Cp, Pd, Mt (GA, NC, SC, VA): pine flatwoods, wet meadows, bog margins, ditches, wet roadsides, often weedy; common. May- October. E. MA south to FL, west to TX, and north to s. IN and IL. [= RAB, G, W; < Rh. mariana var. mariana – F, GW, K, Z (also see Rh. mariana var. exalbida); > Rh. mariana var. leiosperma Fernald & Griscom – F; ? Rh. delicatula Small – S] Rhexia mariana Linnaeus var. ventricosa (Fernald & Griscom) Kral & Bostick, Swollen Meadow-beauty. Cp (NC, SC, VA), Pd (NC, VA): pine flatwoods and savannas, clearings in cypress-hardwood swamps, ditches, wet roadsides; uncommon. June-September. NJ south to SC. This variety is closely related to Rh. mariana var. interior (Pennell) Kral & Bostick, occurring west of the mountains. Gleason & Cronquist (1991) prefer to retain Rh. interior Pennell at the species level and hedge relative to the distinctiveness of Rh. ventricosa. [= GW, K, W, Z; = Rh. ventricosa Fernald & Griscom – RAB, F; < Rh. interior Pennell – C] Rhexia nashii Small, Hairy Meadow-beauty. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): wet pine flatwoods and savannas; pondshores, bogs, marshes, ditches, wet roadsides; common (uncommon in Piedmont). May-October. Primarily a Southeastern Coastal Plain species: e. VA south to s. FL and west to se. LA. [= GW, K, S, Z; = Rh. mariana var. purpurea Michaux – RAB, F, G] Rhexia nuttallii C.W. James, Nuttall's Meadow-beauty. Cp (GA): pine flatwoods, bogs; rare (GA Special Concern). Coastal Plain of se. GA west to FL Panhandle, south to s. peninsular FL. [= GW, K, Z; = Rh. serrulata Nuttall – S] Rhexia parviflora Chapman, Small-flowered White Meadow-beauty. Cp (GA): limesink pond margins; rare (GA Special Concern). Occurs in sw. GA (Mitchell County) south into Panhandle FL. [= GW, K, S, Z] Rhexia petiolata Walter, Ciliate Meadow-beauty, Short-stemmed Meadow-beauty. Cp (GA, NC, SC, VA), Pd (GA, SC): wet pine flatwoods and savannas, pocosin borders, and ditches; common (VA Rare). June-September. May-October. A Southeastern Coastal Plain endemic: se. VA south to n. FL and west to se. TX. The flowers are sessile, the petals ascending. [= RAB, C, G, GW, K, Z; = Rh. ciliosa Michaux – F, S] MELASTOMATACEAE 497 Rhexia virginica Linnaeus, Virginia Meadow-beauty, Deergrass, Handsome Harry, Wing-stem Meadow-beauty. Mt, Cp, Pd (GA, NC, SC, VA): wet pine flatwoods and savannas, pond shores, bogs, and ditches; common (uncommon in NC Piedmont and NC Coastal Plain). May-October. E. Canada and WI south to n. FL and TX. [= C, G, GW, K, W, Z; > Rh. virginica var. purshii (Sprengel) C.W. James – RAB; > Rh. virginica var. virginica – RAB, F; > Rh. virginica var. septemnervia (Walter) Pursh – F; = Rh. stricta Pursh – S]

Rhexia ×brevibracteata Snyder [Rh. aristosa × virginica] is known as a natural hybrid from the Coastal Plain of NJ (Snyder 1996). {not keyed} Rhexia mariana Linnaeus var. interior (Pennell) Kral & Bostick. Moist to wet areas, ditches, prairies. S. IN, s. IL, s. MO, and se. KS south to c. AL, c, MS, n. LA, and se. OK. [= GW, K, Z; < Rh. interior – C; = Rh. interior Pennell – F, G] {not keyed at this time; synonymy incomplete} Rhexia salicifolia Kral & Bostick, Willowleaf Meadow-beauty. Drawdown zones of Coastal Plain depression ponds and interdune swales. S. AL and FL Panhandle. [= GW, K, Z] {not keyed at this time; synonymy incomplete}

MELIACEAE A.L. de Jussieu 1789 (Mahogany Family)

A family of about 50 genera and 565 species, trees and shrubs, of tropical and subtropical areas. The only native member of the family in e. North America is Swietenia mahogani (Linnaeus) Jacquin (West Indian Mahogany), a very valuable timber tree which ranges north to s. FL. References: Miller (1990)=Z in the synonymy.

Melia Linnaeus (Chinaberry)

A genus of 3 species, trees, of the Old World tropics.

* Melia azedarach Linnaeus, Chinaberry, Carolina Mahogany, Umbrella-tree, Pride-of-India. Cp, Pd (GA, NC, SC, VA), Mt (NC): disturbed areas, abandoned rural yards and fields; common (rare in the Mountains), native to se. Asia, commonly cultivated in our area (mainly in the Coastal Plain) and commonly escaped. April-May; September-October. Rural children often play with the bony drupes, which are poisonous if ingested. [= RAB, C, F, G, K, S, Z]

MENISPERMACEAE A.L. de Jussieu 1789 (Moonseed Family)

A family of about 72 genera and 450 species, vines, shrubs, trees, and herbs, ot tropical, subtropical, and warm temperate areas. References: Rhodes in FNA (1997); Kessler in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves asymmetrically peltate (the stem attached 1-5 mm in from the leaf margin); stamens 12-24; petals 6-9; fruit bluish- black; [tribe Menispermeae] ...... Menispermum 1 Leaves not peltate, usually cordate (the stem attached at the leaf margin); stamens 6 or 12; petals 6 or 0; fruit red or bluish- black. 2 Leaves 3-7-lobed, the sinuses usually deep, the lobes acute; stamens 12; petals 0; fruit bluish-black, 13-25 mm long; [tribe Tinosporeae]...... Calycocarpum 2 Leaves entire to 3-lobed, the sinuses always shallow, the lobes (if present) broadly rounded; stamens 6; petals 6; fruit red, 5-8 mm long; [tribe Menispermeae]...... Cocculus

Calycocarpum (Nuttall) Spach (Cupseed)

A monotypic genus, a woody vine, of e. North America. References: Kessler in Kubitzki, Rohwer, & Bittrich (1993).

Calycocarpum lyonii (Pursh) A. Gray, Cupseed, Lyonia-vine. Cp (GA, SC), Pd, Mt (GA): floodplain forests; rare. May- June. Ranging from nw. GA, s. IN, and MO, south to se. SC, e. GA, Panhandle FL, and LA; some of the easternmost occurrences may be adventive or introduced. [= C, F, FNA, G, K, S]

Cocculus A.P. deCandolle (Coralbeads, Snailseed)

A genus of 8 species, woody vines, shrubs, and trees, of tropical, subtropical, and warm temperate regions of North America, Central America, Africa, Madagascar, India, Malaysia, and the Philippines. References: Kessler in Kubitzki, Rohwer, & Bittrich (1993).

Cocculus carolinus (Linnaeus) A.P. deCandolle, Coralbeads, Carolina Moonseed, Snailseed, Red Moonseed. Cp, Pd, Mt (GA, NC, SC, VA): moist to dry forests and thickets; common (rare in VA and in the Mountains of GA, NC, and SC). June- MELASTOMATACEAE 498 August. VA south to FL, west to TX, north in the interior to s. IN and MO. Its occurrences in VA may be primarily adventive. [= RAB, C, F, FNA, G, K, W; = Epibaterium carolinum (Linnaeus) Britton – S]

Menispermum Linnaeus (Moonseed)

A genus of 2-4 species, vines, of temperate e. North America and temperate e. Asia. References: Kessler in Kubitzki, Rohwer, & Bittrich (1993).

Menispermum canadense Linnaeus, Moonseed, Yellow Parilla. Pd, Cp (GA, NC, SC, VA), Mt (GA, NC, VA): moist nutrient-rich forests, especially on floodplains or lower slopes; common. June-August. Québec west to Manitoba, south to GA and OK. [= RAB, C, F, FNA, G, K, S, W]

MENYANTHACEAE Dumortier 1829 (Buckbean Family)

A family of about 5 genera and 40 species, wetland herbs, of cosmopolitan distribution. References: Wood (1983b)=Z.

1 Leaves trifoliate; inflorescence a raceme...... Menyanthes 1 Leaves simple; inflorescence an umbel...... Nymphoides

Menyanthes Linnaeus (Buckbean, Bogbean)

The genus is monotypic, an herb, circumboreal. References: Wood (1983b)=Z

Menyanthes trifoliata Linnaeus, Buckbean, Bogbean. Mt (NC, VA): mountain bogs at high elevations over amphibolite (in the Blue Ridge), boggy marshes over calcareous rocks (in the Ridge and Valley); rare (NC Threatened, VA Rare). May-June. This circumboreal species is widespread in n. North America and n. Eurasia, ranging south in North America to NJ, VA, IN, MO, and CA, and disjunct to Long Hope Valley, Watauga County, NC. The NC populations are disjunct about 400 km from the next nearest populations in VA. McDowell (1984) reported the first documentation of the species for NC. [= C, G, K, W, Z; > M. trifoliata var. minor Rafinesque – F]

Nymphoides Hill (Floating Heart)

A genus of about 20 species, aquatic herbs, cosmopolitan. References: Wood (1983b)=Z; Burks (2002).

Identification notes: As the scientific name indicates, the leaves of Nymphoides bear a superficial resemblance to those of Nymphaea. The leaves of Nymphoides are more cordate, the two basal lobes more rounded, rather than having a rather sharp corner or angle. Nymphoides cordata has much smaller leaves than Nymphaea, while the thickly pebbled texturing of Nymphoides aquatica is very unlike the glossy smoothness of Nymphaea.

1 Flowers yellow; floating stems usually with multiple leaves; capsules 12-25 mm long ...... N. peltata 1 Flowers white; floating stems with single leaves; capsules 3-14 mm long. 2 Adaxial petal surface bearing a ruffled crest down its length in the middle...... N. cristata 2 Adaxial petal surface not crested. 3 Leaves 5-15 cm wide, roughly pebbled below, thick in texture; stems 1.3-2.5 mm in diameter a few cm below the inflorescence, with conspicuous red spots; tuberous roots of floating clusters stout, blunt-tipped; seeds conspicuously papillate; capsule 10-14 mm long...... N. aquatica 3 Leaves 3-7 cm wide, smooth below, thin in texture; stems 0.6-0.9 mm in diameter a few cm below the inflorescence, rarely spotted with red; tuberous roots of floating clusters slender, with pointed tips; seeds smooth (rarely papillate); capsule 4-5 mm long...... N. cordata

Nymphoides aquatica (Walter ex J.F. Gmelin) Kuntze, Big Floating Heart, Banana Floating Heart. Cp (GA, NC, SC, VA): limesink ponds (dolines), other acidic and nutrient-poor water-filled depressions, sluggish streams, beaverponds, primarily in the Outer and Middle Coastal Plain; uncommon (VA Rare). Late April-September. A Southeastern Coastal Plain endemic: NJ south to FL and west to TX. [= RAB, C, F, GW, K, S, Z; = N. aquaticum – G, orthographic variant] Nymphoides cordata (Elliott) Fernald, Little Floating Heart. Cp (GA, NC, SC): upland depression ponds, sluggish streams, beaverponds, primarily in the fall-line Sandhills; uncommon (NC Watch List). Widespread (though in many parts of its range local) in e. North America, from Newfoundland and Ontario south to FL and LA. [= RAB, C, F, GW, K, Z; = N. cordatum – G, orthographic variant; N. lacunosa (Ventenat) Kuntze – S, misapplied] MELASTOMATACEAE 499 * Nymphoides cristata (Roxburgh) Kuntze, Crested Floating Heart, Water Snowflake. Cp? (SC): ponds and lakes; rare, native of China and India. Apparently first naturalizing in North America in FL in 2000; introduced for water gardens and aquariums. * Nymphoides peltata (S.G. Gmelin) Kuntze, Yellow Floating Heart. Pd (NC, VA): ponds; rare, introduced from Europe. This European native is sparingly naturalized in e. North America; it is sold for cultivation in water gardens, and will likely become more widely naturalized. [= C, F, K; = N. peltatum – G, orthographic variant]

MOLLUGINACEAE Hutchinson 1926 (Carpetweed Family)

A family of about 13-14 genera and 120-125 species, herbs, of tropical and warm temperate areas. References: Vincent in FNA (2003b); Boetsch (2002)=Z; Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

Mollugo Linnaeus (Carpetweed)

A genus of about 35 species, annual herbs, of tropical and subtropical regions of both hemispheres, introduced in temperate regions. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

* Mollugo verticillata Linnaeus, Carpetweed, Indian-chickweed. Cp, Pd, Mt (GA, NC, SC, VA): fields, disturbed areas, drawdown zones on river- and pond-shores; common, introduced from tropical America. May-November. [= RAB, C, F, FNA, G, GW, K, S, W, Z]

MORACEAE Lindley 1847 (Mulberry Family)

A family of about 38 genera and 1100 species, trees, shrubs, vines, and herbs, of tropical, subtropical, and (few) warm temperate areas. References: Wunderlin in FNA (1997); White in SE (in prep.); Rohwer & Berg in Kubitzki, Rohwer, & Bittrich (1993).

1 Herb, 0.3-1.0 m tall; stem without latex; [tribe Moreae] ...... Fatoua 1 Shrub or tree, at maturity over 1 m tall, or woody vine growing appressed to masonry; stem bearing translucent to milky- white latex. 2 Stipules connate, the stipule scar encircling the twig; inflorescence a syconium (the flowers borne on the inner walls of the fleshy receptacle); [tribe Ficeae]...... Ficus 2 Stipules free, the stipule scar not encircling the twig; inflorescence a spike, head, or catkin (the flowers borne exposed on a contracted or elongated axis or receptacle); [tribe Moreae]. 3 Leaves entire, unlobed or shallowly 3-lobed; stems usually thorny. 4 Fruit 2-3 cm in diameter; petioles 5-20 mm long; leaves 3-7 (-10) cm long ...... Cudrania 4 Fruit 10-15 cm in diameter; petioles 30-50 mm long; leaves 6-20 cm long ...... Maclura 3 Leaves serrate, often also 3-15-lobed (the lobes sometimes deep); stems not thorny. 5 Stems and leaves hirsute; leaves alternate, opposite, and whorled...... Broussonetia 5 Stems and leaves glabrous to pubescent; leaves alternate ...... Morus

Broussonetia L'Héritier ex Ventenat 1799 (Paper Mulberry)

A genus of about 8 species, trees, shrubs, and vines, of tropical and subtropical Asia and Madagascar. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

* Broussonetia papyrifera (Linnaeus) L'Héritier ex Ventenat, Paper Mulberry. Pd, Cp, Mt (GA, NC, SC, VA): urban lots, disturbed areas, roadsides; common, native of e. Asia. April. [= RAB, C, F, FNA, G, K, W; = Papyrius papyriferus (Linnaeus) Kuntze – S]

Cudrania Trécul 1847 (Cudrania)

A genus of 1-several species, shrubs, of Asia. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

* Cudrania tricuspidata (Carrière) Bureau ex Lavallée, Cudrania. Pd (NC), Cp (GA): escaped and naturalized from plantings; rare, native of China and Korea, where cultivated as a food for silkworms. July. Naturalized in Orange County, NC, in McIntosh Co. GA (Jones & Coile 1988), and perhaps elsewhere in our area, where recommended as a hedge plant since at least 1940 (Rehder 1940). [= FNA, K; = Maclura tricuspidata Carrière]

MORACEAE 500 Fatoua Gaudichaud-Beaupré 1830 (Crabweed)

A genus of 2-3 species, herbs or weak shrubs, of Asia, Madagascar, and Australia. References: Vincent (2004)=Y; Massey (1975)=Z; Miller & Wood (2003); Kral (1981b); Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

* Fatoua villosa (Thunberg) Nakai, Crabweed, Mulberry-weed, Foolish-weed. Pd (GA, NC, SC, VA), Cp (GA, SC), Mt (NC): disturbed areas, vegetable and flower gardens; rare, native of Asia (apparently se. Asian islands). July-November. As reported by Massey (1975) and Vincent (2004), Fatoua was first reported in the United States (Louisiana) in the early 1960's. As of 2004, its distribution in North America had spread to include 28 states and the District of Columbia, including most states except the Great Plains and Rocky Mountains had spread (Vincent 2004, Sundell et al. 1999, Miller & Wood 2003. Since all early collections seem to be in and around greenhouses and nurseries, it is likely that it has been introduced in horticultural material, perhaps repeatedly (Kral 1981b). Fatoua appears to have become a fairly aggressive weed in eastern North America. It can be expected to continue to spread, and has the potential to become noxious. It has alternate, ovate leaves with cordate bases, borne on long petioles (about as long as the leaf blade), the inflorescences are dense cymes borne on peduncles in the axils of leaves. Pubescence of the stem and foliage is uncinulate, giving the plant a "tacky" feel. An excellent illustration appears in Correll & Correll (1982). [= FNA, K, Y, Z]

Ficus Linnaeus 1753 (Fig)

A genus of about 750 species, trees, shrubs, and vines, of tropical, subtropical, and warm temperate areas. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves lobed, 7-30 cm long; [shrub to small tree] ...... F. carica 1 Leaves unlobed, 1-5 cm long; [vine, climbing appressed to walls]...... F. pumila

* Ficus carica Linnaeus, Edible Fig, Garden Fig. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): grown for its fruits, persistent from plantings, persisting and naturalizing particularly on barrier islands, where it sometimes forms thickets on dunes, or otherwise in the outer Coastal Plain, where proximity to the ocean ameliorates cold winter temperatures; rare, native of w. Asia. May-August; July-October. This is the common cultivated fig, grown for its fruit in the Mid-east for millenia. [= RAB, F, FNA, K, S] * Ficus pumila Linnaeus, Climbing Fig. Cp (GA): walls, disturbed areas; rare, introduced from s. Asia. Locally common in Charleston and Savannah, where grown on walls as an ornamental and certainly persisting. [= FNA, K]

Maclura Nuttall 1818 (Osage-orange)

A monotypic genus (or sometimes broadened to include Cudrania and other genera), a tree, of sc. North America. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

* Maclura pomifera (Rafinesque) C.K. Schneider, Osage-orange, Bow-wood, Bois-d'arc, Hedge-apple. Cp, Pd, Mt (GA, NC, SC, VA): fields, hedgerows, forests, naturalized from extensive planting in the eighteenth and nineteenth centuries, native of TX, OK, AR, and LA. April-May; October. Maclura is a monotypic genus. The large fruits are unmistakeable: yellowish-green, grapefruit-sized, and wrinkled, reminiscent of a giant, spherical mulberry fruit. The wood is extremely heavy, fine-grained, a bright yellow-orange when fresh, but darkening with age, famous for making bows and rarely used in cabinetry. [= RAB, C, F, FNA, G, K, W; = Toxylon pomiferum Rafinesque ex Sargent – S]

Morus Linnaeus 1753 (Mulberry)

A genus of about 10-15 species, trees, of warm temperate, subtropical, and tropical areas. References: Endress & Bittrich in Kubitzki, Rohwer, & Bittrich (1993).

1 Upper leaf surface glossy, glabrous or slightly scabrous; lower leaf surface glabrous, or slightly pubescent on the veins and in the vein axils only; ripe fruits black, purple, red, pink, or white...... M. alba 1 Upper leaf surface dull, scabrous; lower leaf surface pubescent on the veins, veinlets, and the surface between the veins; ripe fruits black or purple...... M. rubra

* Morus alba Linnaeus, White Mulberry, Silkworm Mulberry, Russian Mulberry. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas, vacant lots, roadsides, moist forests; uncommon, native of e. Asia. March-May; May-June. [= RAB, C, F, FNA, G, GW, K, S, W; > M. nigra Linnaeus – S, misapplied as to our material; > M. alba var. tatarica (Linnaeus) Seringe] Morus rubra Linnaeus, Red Mulberry. Cp, Pd, Mt (GA, NC, SC, VA): bottomland forests, mesic slopes, disturbed areas, suburban woodlands; common. April-May; May-June. MA, VT, NY, MI, WI, and se. SD south to s. FL and w. TX, and into MORACEAE 501 Mexico. The fruits are very variable in quality from tree to tree. M. rubra is the only member of the Moraceae native to our area. [= RAB, C, F, G, GW, K, S, W; > M. rubra var. rubra – K]

MYRICACEAE Blume 1829 (Bayberry Family)

A family of about 3-5 genera and 55 species, trees and shrubs, nearly cosmopolitan. See Morella for discussion of our 3 genera. References: Bornstein in FNA (1997); Wilbur (1994)=Z; Elias (1971b)=Y; Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

1 Leaves oblong or linear-lanceolate, pinnatifid, stipulate; fruit in a bristly involucre formed by 8 bractlets ...... Comptonia 1 Leaves mostly obovate or oblanceolate, entire or toothed (especially apically), estipulate; fruit either exposed and densely waxy (Cerothamnus), or partially enclosed in 2 wing-like bractlets (Myrica). 2 Fruit spherical, densely waxy, exposed (the 4-6 bractlets small and inconspicuous); terminal buds present; aments inserted on old wood mainly below the leaves; [in our area, common and in the Coastal Plain, Mountains, and Piedmont]...... Morella 2 Fruit flattened, not waxy, partially enclosed in 2 wing-like bractlets; terminal buds lacking; aments inserted at the summit of the branchlets of the preceding year; [in our area, very rare and restricted to bogs in the Mountains] ...... Myrica

Comptonia L'Heritier ex Aiton 1789 (Sweet-fern)

A monotypic genus, a shrub, of e. North America, known as fossils from a much broader area. References: Bornstein in FNA (1997); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

Comptonia peregrina (Linnaeus) J.M. Coulter, Sweet-fern. Mt (GA, NC, SC, VA), Pd, Cp (NC, VA): in the mountains on xeric ridges at low to medium elevations, usually in fire-maintained habitats, also in xeric and fire-maintained habitats on monadnocks in the upper Piedmont and in dry, sandy sites in the lower Piedmont and fall-line sandhills; common (rare in Piedmont and Coastal Plain) (GA Special Concern, SC Rare). April; August-September. Widespread in ne. North America, south to sc. and w. NC, w. SC, ne. GA, and nc. TN; much more common in the northern parts of its range. [= RAB, C, FNA, K, S, W; > Comptonia peregrina var. asplenifolia (Linnaeus) Fernald – F, Y; > Comptonia peregrina var. peregrina – F, Y; > Myrica asplenifolia Linnaeus var. asplenifolia – G; > Myrica asplenifolia var. tomentosa (Chevallier) Gleason – G]

Morella Loureiro (Bayberry, Wax-myrtle, Candleberry)

Wilbur (1994) makes a compelling case for the recognition of three genera among eastern North American Myricaceae, and for application of the name Myrica to Myrica gale. The typification of the genus Myrica with Myrica gale Linnaeus has been confirmed (Brummitt 1999); thus, the familiar southeastern species placed by many authors in Myrica must take another name. Wilbur (1994) prefers to treat our species as subgenus Cerothamnus (Tidestrom) Wilbur of genus Morella Loureiro; subgenus Morella is restricted to e. Asia, the Philippines, and Malaysia, and differs in a number of ways from subgenus Cerothamnus, including its fleshy and succulent, rather than waxy and hard, berries. Small maintained Cerothamnus at the generic level. Wilbur's inclusion of Cerothamnus in Morella may well be warranted (and is followed here), but I disagree with his provisional decision to include the taxon treated below as Morella pumila in Morella cerifera, and the taxon treated below as Morella pensylvanica in Morella caroliniensis, though their appropriate rank may be questioned. References: Bornstein in FNA (1997); Wilbur (1994)=Z; Wilbur (2002); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

1 Fresh leaves odorless when crushed; flowers unisexual, the staminate and pistillate flowers on different plants; leaves usually entire; [of s. GA south and west]; [subgenus Cerothamnus, series Faya] ...... Morella inodora 1 Fresh leaves aromatic when crushed; flowers bisexual, staminate and pistillate on the same plant; leaves usually serrate, at least near the tip; [collectively widespread in our area]; [subgenus Cerothamnus, series Cerothamnus]. 2 Leaves oblanceolate (generally narrowly so), most of them 0.5-1.5 cm wide, 4-6× as long as wide, evergreen; mature fruits 2.0-3.5 mm in diameter. 3 Medium shrub to small tree (usually 2-10 m tall), not stoloniferous; leaves of fertile branches 4-9 cm long, 8-20 mm wide; [of a wide range of wetland habitats, including wet Coastal Plain pinelands]...... Morella cerifera 3 Small shrub (usually < 1 m tall), strongly stoloniferous; leaves of fertile branches 1.5-4 cm long, 3-8 mm wide; [restricted to Coastal Plain pinelands (or areas formerly so)]...... Morella pumila 2 Leaves elliptic to broadly oblanceolate, most of them 1.5-4 cm wide, 2-4× as long as wide, evergreen to deciduous; mature fruits 3.0-7.0 mm in diameter. 4 Leaves subcoriaceous and more or less evergreen, not revolute (or slightly so if sun-grown), the larger ones usually about 9 cm long and 3.5 cm wide, with punctate glands dense on the lower surface and nearly or entirely absent on the upper surface; fruits 3-4.5 mm in diameter, the fruit wall glabrous or sparsely glandular, the warty protuberances glandular; twigs densely hairy to rarely glabrous; older branches blackish; [of various boggy habitats, widespread in our area]...... Morella caroliniensis MYRICACEAE 502 4 Leaves coriaceous, tardily deciduous, often revolute, the larger ones about 6 cm long and 2 cm wide, with punctate glands fairly dense on both surfaces; fruits 4-6 (-7) mm in diameter, the fruit wall and warty protuberances densely hirsute when young; twigs glabrous to sparsely hairy; older branches whitish gray; [usually of dunes, from Dare County, NC northward]...... Morella pensylvanica

Morella caroliniensis (P. Miller) Small, Pocosin Bayberry, Evergreen Bayberry. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): pocosins, wet savannas and pine flatwoods, sandhill seepage bogs, and other peaty or sandy-peaty wetlands; common. April; August-October. Primarily limited to the Southeastern Coastal Plain, from NJ south to FL and west to TX and AR. [= K; Myrica heterophylla Rafinesque – RAB, C, FNA, W, Y; > Myrica heterophylla var. heterophylla – F; > Myrica heterophylla var. curtissii (Chevallier) Fernald – F; < Myrica pensylvanica – G; < Cerothamnus carolinensis – S (also see Morella pensylvanica); < Morella caroliniensis (P. Miller) Small – Z (also see Morella pensylvanica)] Morella cerifera (Linnaeus) Small, Common Wax-myrtle, Southern Bayberry. Cp (GA, NC, SC, VA): interdune swales (where often dominant), pocosins, brackish marshes, other wet to moist habitats, now also widely planted (including in the Piedmont) as an ornamental or landscaping shrub; common. April; August-October. Widespread in the Coastal Plain of Southeastern United States: NJ south to FL and west to TX. Our most common Morella, and also the largest, sometimes becoming a small tree, to at least 10 m tall and 20 cm DBH. See Morella pumila for a discussion of the controversial taxonomy of Morella cerifera and Morella pumila. [= Myrica cerifera Linnaeus var. cerifera – RAB, Y; < Myrica cerifera – C, FNA, GW (also see C. pumilus); = Myrica cerifera – F, G; = Cerothamnus ceriferus (Linnaeus) Small – S; < Morella cerifera (Linnaeus) Small – K, Z (also see Morella pumila)] Morella inodora (Bartram) Small, Odorless Bayberry. Cp (GA): acid wetlands, especially in wooded, acid, streamhead "bogs" and bayheads, often associated with Magnolia virginiana, Persea palustris, Cyrilla racemiflora, Cliftonia monophylla, and Woodwardia areolata; rare (GA Special Concern). A Southeastern Coastal Plain endemic: se. GA west to s. MS. [= K, Z; = Myrica inodora Bartram – FNA, GW, Y; = Cerothamnus inodorus (Bartram) Small – S] Morella pensylvanica (Mirbel) Kartesz, Northern Bayberry. Cp (NC, VA): dunes, sometimes even on the foredune and stoloniferously colonizing the upper beach, more typically behind the foredune on secondary dunes and sandy flats, often growing intermixed with Morella cerifera, but able to occupy drier sites higher on the dunes, from VA north, also ranging inland in sandy situations; uncommon (restricted in NC to barrier islands of Dare and Currituck counties, but locally common there). April; August-October. This species reaches its southern limit at Avon (Kinnakeet), Dare County, NC. On interdune flats, it often grows intermixed with Morella cerifera, but is readily distinguished (even at a distance) by its stoloniferous growth (appearing as dome-shaped clones 3-20 m in diameter), stouter twigs, and tardily deciduous leaves. The twigs of this species are noticeably stouter than those of Morella cerifera; measured at 10 cm from the twig tips, they are (2-) 3-5 mm in diameter, those of Morella cerifera ca. 1.5-2.5 mm. [= K; = Myrica pensylvanica Loiseleur – RAB, C, F, FNA, GW, Y; < Myrica pensylvanica – G (also see Morella heterophylla); < Cerothamnus carolinensis – S (also see Morella pensylvanica); < Morella caroliniensis (P. Miller) Small – Z; = Cerothamnus pensylvanicus (Mirbel) Moldenke] Morella pumila (Michaux) Small, Dwarf Bayberry, Dwarf Wax-myrtle. Cp (GA, NC, SC, VA): savannas, pine flatwoods, relatively moist to extremely dry sites in sandhills (under Quercus laevis and Q. geminata); common (VA Rare). April; August- October. A Southeastern Coastal Plain endemic: se. VA south to FL and west to LA (or TX). Some authors dismiss the distinction between this taxon and Morella cerifera as merely environmental, while others treat the two as distinct at the varietal or specific level. In our area at least, they appear to be genetically distinct. They often occur in close proximity (though somewhat different habitats), and maintain their distinctiveness. There are some observations that there is a phenologic difference, with Morella pumila peak flowering 3 weeks later than Morella cerifera (J. Townsend, pers. comm. 2002). Though the issue remains unresolved, the stoloniferous growth of Morella pumila is not merely a fire response; I here maintain the two as distinct, pending further research. A varietal distinction might best reflect the relationship of these two taxa, but the appropriate combinations at the varietal level are not available in either Cerothamnus or Morella. [= Myrica cerifera Linnaeus var. pumila Michaux – RAB, Y; < Myrica cerifera – C, FNA, GW; = Myrica pusilla Rafinesque – F, G; = Cerothamnus pumilus (Michaux) Small – S; < Morella cerifera (Linnaeus) Small – K, Z]

Myrica Linnaeus 1753 (Sweet Gale)

A genus of two species, shrubs, of temperate and subarctic regions of North America and Eurasia. References: Bornstein in FNA (1997); Kubitzki in Kubitzki, Rohwer, & Bittrich (1993).

Myrica gale Linnaeus, Sweet Gale. Mt (NC): peaty bogs; rare (NC Endangered Species). April; August-September. A circumboreal species, south in North America to NJ, PA, MI, MN, and OR, disjunct from PA and NJ to Henderson County, NC, where considered extirpated at one time, as a result of the destruction of the famous East Flat Rock Bog. This shrub has been relocated at a single site, where it is abundant in a small area (less than 0.25 hectare). [= C, FNA, G, GW, K, S, W, Y; = Gale palustris Chevallier – RAB; > Myrica gale var. gale – F]

MYRSINACEAE R. Brown 1810 (Myrsine Family)

MYRSINACEAE 503 The traditional families Primulaceae, Myrsinaceae, and Theophrastaceae have been repartitioned by Källersjö, Bergqvist, & Anderberg (2000) in order to create monophyletic groups. References: Källersjö, Bergqvist, and Anderberg (2000); Ståhl & Anderberg in Kubitzki (2004).

1 Leaves alternate (or with some opposite or subopposite); flowers white. 2 Flowers axillary, nearly sessile; leaves 3-10 mm long...... Anagallis 2 Flowers in a terminal raceme, pedicellate, the flowers closely spaced, touching, the inflorescence thus appearing cylindrical, and generally drooping at the tip (reminiscent of Saururus cernuus); leaves longer; [introduced, rarely naturalized in upland situations] ...... Lysimachia clethroides 1 Leaves opposite or whorled; flowers yellow, white, pink, red, or blue. 3 Leaves in a single terminal whorl; petals 7 ...... Trientalis 3 Leaves opposite or whorled (if whorled, with several to many whorls); petals 0 or 5. 4 Leaves > 2 cm long (sometimes less in L. nummularia, and then orbicular, about as wide as long); flowers yellow...... Lysimachia 4 Leaves < 2 cm long (and distinctly longer than wide); flowers red, blue, white, or pink. 5 Flowers on long pedicels; corolla present ...... Anagallis 5 Flowers nearly sessile; corolla absent ...... Glaux

Anagallis Linnaeus 1753 (Pimpernel) (also see Centunculus)

A genus of 20-28 species, herbs, mostly Old World. References: Ståhl & Anderberg in Kubitzki (2004)

1 Leaves alternate; flowers subsessile, on thick pedicels 0.3-1.0 mm long; leaf blades 3-10 mm long ...... A. minima 1 Leaves opposite (occasionally in whorls of 3); flowers pedicellate, on slender pedicels 10-25 mm long; leaf blades 5-30 mm long. 2 Flowers red (rarely white); pedicels usually longer than the leaves...... A. arvensis var. arvensis 2 Flowers blue; pedicels usually shorter than the leaves...... [A. arvensis var. caerulea]

* Anagallis arvensis Linnaeus var. arvensis, Scarlet Pimpernel, Common Pimpernel. Cp (GA, NC, SC, VA), Pd (GA, NC, VA), Mt (VA): lawns, fields, disturbed areas; common, introduced from Europe. April-November. [= C, G; < A. arvensis – RAB, F, GW, W; = A. arvensis ssp. arvensis – K, in the narrow sense; = A. arvensis ssp. arvensis – S] Anagallis minima (Linnaeus) E.H. Krause, Chaffweed, False-pimpernel. Cp, Mt (GA, SC), Pd (GA, VA): ditches, wet disturbed areas, savannas, pond margins; uncommon (rare in Mountains, rare in VA). March-June. This species occurs in widely scattered areas, nearly cosmopolitan. [= GW, K; = Centunculus minimus Linnaeus – RAB, C, F, G, S, W]

* Anagallis arvensis Linnaeus var. caerulea (Schreber) Grenier & Godron, Blue Pimpernel, is reported as introduced in PA, KY, OH, and other scattered states north and west of our area (Kartesz 1999). [= C, G; < A. arvensis – RAB, F, GW, W; = A. arvensis Linnaeus ssp. foemina (P. Miller) Schinz & Thellung – K; = A. arvensis ssp. coerulea Hartman – S] * Anagallis monellii Linnaeus is reported as a waif for Fairfax County, VA by Harvill et al. (1992) and Shetler & Orli (2000). Not keyed. [= K]

Centunculus Linnaeus [see Anagallis]

Glaux Linnaeus 1753 (Sea-milkwort)

A monotypic genus, of north temperate coasts of the Old and New Worlds. Glaux appears to be embedded within Lysimachia and should be merged into that genus (Hao et al. 2004). References: Hao et al. (2004); Ståhl & Anderberg in Kubitzki (2004).

Glaux maritima Linnaeus, Sea-milkwort. Cp (VA): saline coastal habitats; rare. June-July. The species is interruptedly circumboreal, in North America from Québec south to VA on the east coast, and from British Columbia south to OR on the west coast, also inland in w. North America, from Saskatchewan south to NM. G suggests that G. maritima is introduced near its southern limit in the east. [= C, K; > G. maritima var. maritima – F, G]

Lysimachia Linnaeus 1753 (Loosestrife)

A genus of about 150 species, herbs (rarely shrubs), cosmopolitan. Hao et al. (2004) showed that the traditional subgeneric classification of Lysimachia is highly artificial, and that Glaux is embedded within Lysimachia. References: Coffey & Jones (1980)=Z; Hao et al. (2004); Ståhl & Anderberg in Kubitzki (2004). Key partly adapted from Z. MYRSINACEAE 504

1 Leaves alternate; flowers white, in a terminal raceme, the tip often lax ...... L. clethroides 1 Leaves opposite or whorled; flowers yellow, borne variously. 2 Leaves nearly round; plant trailing, rooting at nodes ...... L. nummularia 2 Leaves linear, lanceolate, elliptic, or ovate; plant erect (or trailing and rooting at the nodes in L. radicans, which has lanceolate leaves). 3 Flowers in a terminal raceme or panicle, subtended by bracts much smaller than the stem leaves. 4 Inflorescence a terminal panicle...... L. fraseri 4 Inflorescence a terminal raceme. 5 Leaves narrowly ovate, broadest near the base, with 3 prominent veins ...... L. asperulifolia 5 Leaves linear to lanceolate, broadest near the middle, with 1 prominent vein. 6 Leaves linear to narrowly lanceolate, (1-) 2-4 (-8) mm wide; sepals stipitate-glandular....L. loomisii 6 Leaves lanceolate to elliptic, 7-20 mm wide; sepals glabrous. 7 Flowers in part (the lower) in the axils of well-developed leaves...... L. ×producta 7 Flowers all in the axils of much reduced linear bracts...... L. terrestris 3 Flowers axillary, all or most of them subtended by leaves similar in shape to (though often somewhat smaller than) stem leaves not subtending flowers (or with flowers in axillary, peduncled, densely-flowered racemes in L. thyrsiflora). 8 Flowers in peduncled axillary racemes in the axils of midstem leaves; petals linear to lanceolate, ca. 5 mm long and ca. 1 mm wide, much surpassed by the stamens...... [L. thyrsiflora] 8 Flowers solitary, all or most of them subtended by leaves similar in shape to (though often somewhat smaller than) normal stem leaves; petals lanceolate to ovate, as long or longer than the stamens. 9 Stem leaves whorled (in adult plants); leaves "punctate" with sinuous, elongate markings (visible with the naked eye, but more readily observed with 10 × magnification). 10 Petals yellow, marked with black lines; sepals 2.5-5 mm long...... L. quadrifolia 10 Petals plain yellow, not marked with black lines; sepals 5.5-9 mm long...... L. punctata 9 Stem leaves opposite; leaves not "punctate." 11 Mid-cauline leaves with petioles ciliate their entire length. 12 Mid-cauline leaves 1-2 mm wide; flowers 7-14 mm across; [of ne. AL] ...... [L. graminea] 12 Mid-cauline leaves 4-60 mm wide; flowers 11-26 mm across; [collectively widespread]. 13 Mid-cauline leaves ovate to lanceolate, 17-60 mm wide; sepals with 3 (or 6) usually reddish-brown veins ...... L. ciliata 13 Mid-cauline leaves lanceolate to linear, 4-23 mm wide; sepals without reddish-brown veins. 14 Cilia of the petiole extending onto the base of the leaf blade; leaf blade lanceolate to linear, typically about 8-12× as long as wide, cuneate at the base; sepal venation inconspicuous or apparently absent; capsules 2-4.5 mm in diameter...... L. lanceolata 14 Cilia of the petiole not extending onto the leaf blade; leaf blade lanceolate to ovate, typically 2-4× as long as wide, rounded to cuneate at the base; sepal venation conspicuous; capsules 4-6.5 mm in diameter...... L. hybrida 11 Mid-cauline leaves with petioles pubescent only along basal portion. 15 Rhizomes absent, new shoots arising from crown of rootstock...... L. tonsa 15 Rhizomes present, new shoots arising from the rhizome. 16 Plant reclining or trailing, rooting at the nodes...... L. radicans 16 Plant erect, not rooting at the nodes. 17 Leaf blades linear to narrowly lanceolate, typically 8-14× as long as wide, cuneate to tapering at the base, with a prominent midrib; sepals not conspicuously veined, 1-2 mm wide ...... L. quadriflora 17 Leaf blades ovate to lanceolate, typically 2-4× as long as wide, rounded to cuneate at the base, with the midrib not prominent; sepals conspicuously veined, 0.5-4 mm wide ...... L. hybrida

Lysimachia asperulifolia Poiret, Pocosin Loosestrife, "Roughleaf Loosestrife". Cp (NC, SC): low pocosins, high pocosins, streamhead pocosins, savanna-pocosin ecotones, sandhill-pocosin ecotones; rare (US Endangered, NC Endangered, SC Rare). May-June; August-October. Endemic to the Coastal Plain of NC and SC. L. asperulifolia is a very distinctive species, easily recognized vegetatively by its whorls of sessile, rounded-based, acuminate, bluish-green (to yellowish-green when shaded or otherwise stressed) leaves. The leaves of L. asperulifolia are not rough; the common name "roughleaf loosestrife" is a misnomer, apparently based on a mistranslation of the specific epithet, the translator assuming that "asperulifolia" meant "rough-leaved." The epithet actually refers to the perceived similarity of the leaves to those of the European Asperula odorata (treated in this work as Galium odoratum), Sweet Woodruff, a plant with which Poiret would have been very familiar. The leaves of G. odoratum are similar to those of L. asperulifolia in their whorled disposition. Franklin (2001) studied the biology of this rare species. [= K; = L. asperulaefolia – RAB, GW, S (an orthographic variant)] MYRSINACEAE 505 Lysimachia ciliata Linnaeus, Fringed Loosestrife. Pd, Mt, Cp (GA, NC, SC, VA): mesic forests, especially bottomlands and coves dominated by hardwoods; common. June-August; August-October. Newfoundland west to AK, south to GA, AL, MS, AR, KS, NE, CO, NM, UT, ID, and OR. [= RAB, C, F, GW, K, W, Z; = Steironema ciliatum (Linnaeus) Baudo – G, S] * Lysimachia clethroides Duby. Mt (NC): roadsides (cultivated and rarely persistent or escaped); rare, introduced from Japan. July-August. Collected in the Mountains of NC (Macon County), escaped from cultivation; it is also reported as naturalized in Grundy County, TN (Chester, Wofford, & Kral 1997, Kral 1981). It differs from our other species in its white flowers, in a dense terminal spike (often with secund tip) and alternate leaves. [= C, G, K] Lysimachia fraseri Duby, Fraser's Loosestrife. Mt (GA, NC, SC): hardwood forests, forest edges and roadbanks, thin soils around rock outcrops, usually flowering only when exposed to extra sunlight by a tree-fall light gap or other canopy opening; rare (GA Rare, NC Endangered, SC Rare). June-August; September-October. W. NC and e. TN south to n. SC, n. GA, and AL; disjunct in s. IL and nw. TN (Stewart County) (Chester, Wofford, & Kral 1997). This rare species is limited in NC to the mountains south of the Asheville Basin, especially in the escarpment gorges of Macon and Jackson counties. Potentially the largest and coarsest of our Lysimachia (up to 2 meters tall), L. fraseri usually occurs as much smaller seedlings and non- flowering individuals. When a tree-fall light gap occurs, individuals flower and fruit. Even seedlings can be separated from the more common and widespread L. quadrifolia by the following characteristics (all best observed at 10×): leaves with a narrow, translucent red border, upper internodes of the stem glandular-puberulent, and backlighted leaf without sinuous, translucent lineations (L. quadrifolia: leaves without red border, upper internodes sparsely pubescent with longer, nonglandular hairs, or rarely a few of the hairs with slightly bulbous tips, and backlighted leaf with numerous sinuous, translucent lineations). [= RAB, GW, K, S, W] Lysimachia hybrida Michaux, Lowland Loosestrife. Cp (NC, SC, VA), Mt, Pd (NC, VA): mesic hardwood forests, wet areas; rare (NC Watch List, VA Watch List). June-August; September-October. ME and s. Québec west to Alberta and WA, south irregularly to n. FL, AR, NE, and AZ. [= C, F, K, W, Z; = L. lanceolata var. hybrida (Michaux) A. Gray – RAB, GW; = Steironema hybridum (Michaux) Rafinesque ex B.D. Jackson – G, S] Lysimachia lanceolata Walter, Lanceleaf Loosestrife. Mt, Pd, Cp (GA, NC, SC, VA): mesic to relatively dry forests, forest edges, roadbanks, primarily on circumneutral soils; uncommon. June-August; September-October. NJ, PA, OH, MI, and WI south to GA, panhandle FL, AL, MS, LA, and ne. TX. [= C, F, K, W, Z; = L. lanceolata var. lanceolata – RAB, GW; = Steironema lanceolatum (Walter) Gray – G, S; = Steironema heterophyllum (Michaux) Baudo – S] Lysimachia loomisii Torrey, Carolina Loosestrife. Cp (GA, NC, SC): moist to wet savannas, pocosin ecotones; uncommon (GA Special Concern). May-June; August-October. Endemic to the outer and middle Coastal plain of NC, SC, and e. GA. [= RAB, GW, K, S] * Lysimachia nummularia Linnaeus, Creeping Charlie, Creeping Jenny, Moneywort. Pd (GA, NC, SC, VA), Mt, Cp (NC, SC, VA): lawns, pastures, seepages, other moist, disturbed places; common (uncommon south of VA), introduced from Europe. May-July; August-September. The leaves have many minute, maroon dots. [= RAB, C, F, G, GW, K, S, W] Lysimachia ×producta (A. Gray) Fernald (pro sp.). Mt (NC, VA), Pd (VA), Cp (NC): moist areas; rare. May-July; August-October. This is a fertile hybrid of L. quadrifolia and L. terrestris, sometimes occurring in the apparent absence of one or both parents. [= RAB, C, K; = L. producta (A. Gray) Fernald – G, S] * Lysimachia punctata Linnaeus, Large Loosestrife, Spotted Loosestrife. Mt (NC): disturbed areas; rare, introduced from Eurasia. June-July; August? First found in NC in 1985 (Weakley in prep.). [= C, F, G, K] Lysimachia quadriflora Sims, Smooth Loosestrife, Four-flowered Loosestrife. Mt (GA, VA), Pd? (NC?): wet meadows and calcareous fens, stream banks; rare (NC Watch List, VA Rare). July-September. MA, s. Ontario, MI, and ND south to w. VA, WV, nw. GA, AL, and AR; mainly north and west of the Ohio River, very rare and scattered in or east of the Appalachians. Reported for c. NC by Coffey & Jones (1980). [= C, F, K, W, Z; = Steironema quadriflora (Sims) Hitchcock – G] Lysimachia quadrifolia Linnaeus, Whorled Loosestrife. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): a wide variety of forests and openings, including pine savannas of the outer Coastal Plain, ranging from moist to very dry; common (uncommon in Coastal Plain south of VA). May-August; August-October. ME west to WI and MN, south to SC, c. GA, AL, and TN. Although the species normally has whorled leaves, immature and small plants often have opposite leaves only. See L. fraseri for discussion of vegetative features useful in distinguishing the two species. [= RAB, C, F, G, GW, K, S, W] Lysimachia radicans Hooker, Trailing Loosestrife. Mt, Cp (VA): moist forests, swamps; rare (VA Rare). June-August. The main distribution of this species is in the Mississippi Embayment, from MO and w. TN south to s. AL, MS, AR, LA, and e. TX; disjunct occurrences in VA and (allegedly) e. NC are curious. The report for NC is from a species list for Nags Head Woods, Dare County; it is unpublished, apparently not documented by an herbarium specimen, and rejected unless additional documentation is found. [= C, F, K, W, Z; = Steironema radicans (Hooker) A. Gray – G, S] Lysimachia terrestris (Linnaeus) Britton, Sterns, & Poggenburg, Bog-candles, Swamp-candles. Mt (GA, NC, SC, VA), Pd, Cp (NC, SC, VA): bogs, wet meadows, and swamp forests; uncommon. May-July; August-October. Newfoundland west to MN and Saskatchewan, south to SC, GA, e. TN, and sc. TN. [= RAB, C, G, GW, K, S, W; L. terrestris var. terrestris – F] Lysimachia tonsa (Wood) Wood ex Pax & R. Knuth, Southern Loosestrife, Appalachian Loosestrife. Pd (GA, NC, SC, VA), Mt (GA, VA): upland forests, especially over calcareous or mafic rocks; rare (NC Watch List, VA Watch List). May-July; August-October. Sc. VA and KY south to SC, wc. GA, and e. TN. The range is centered on the Southern Appalachians, but the species is essentially absent from the higher mountains – a "doughnut range." [= RAB, C, F, K, W, Z; =? Steironema intermedium Kearney – G; = Steironema tonsum (Wood) Bicknell ex Britton – S]

* Lysimachia barystachys Bunge. Reported from a single county in nc. GA (Jones & Coile 1988). [= K] {investigate; not keyed at this time} MYRSINACEAE 506 Lysimachia graminea (Greene) Handel-Mazzetti, Grassleaf Yellow-loosestrife. Endemic to ne. AL (Little River Canyon area). [= K, Z; = Steironema gramineum Greene – S] * Lysimachia japonica Thunberg, native of Japan and China. Reported for WV (Kartesz 1999). [= K] {investigate; not keyed at this time} Lysimachia thyrsiflora Linnaeus, Tufted Loosestrife, ranges south to NJ, PA, OH, and MO (Kartesz 1999), and MD (from Big Marsh, Kent County) (Steury, Tyndall, & Cooley (1996). [= C, K; = Naumburgia thyrsiflora (Linnaeus) Duby] {not keyed at this time; synonymy incomplete} * Lysimachia vulgaris Linnaeus, Garden Loosestrife. Pd (VA): disturbed bottomland, native of Europe. Introduced and naturalized south at least to se. and sc. PA (Rhoads & Klein 1993), WV, KY, MD, and NJ (Kartesz 1999). [= C, K] {not keyed at this time; synonymy incomplete}

Another hybrid has been reported: L. ×radfordii Ahles, a hybrid of L. loomisii × quadrifolia. It is intermediate between its parents.

Trientalis Linnaeus 1753 (Starflower)

A genus of 2 species, herbs, north temperate. References: Ståhl & Anderberg in Kubitzki (2004).

Identification notes: Trientalis can be recognized by its terminal whorl of leaves (4-10 cm long), the one to several white flowers borne on terminal, slender pedicels, each flower typically with 7 petals (inconspicuously united at the bases), each petal acuminate. The plant is reminiscent of a white-flowered Lysimachia with only one whorl of leaves.

Trientalis borealis Rafinesque ssp. borealis, Starflower. Mt (GA, NC, VA): northern hardwood forests, rich slope forests, often in second-growth areas; uncommon in VA, rare farther south (GA Endangered, NC Rare). May-June. This northern species, widespread in the mountains of VA, and known from a few locations in n. GA and ne. TN (Chester, Wofford, & Kral 1997), was first located in NC only in 1988 (Dellinger 1989). "The attractive white corollas, usually with 7 petals united only at the very base, are open in the late spring and they drop intact – like fallen stars" (Voss 1996). [= K; < T. borealis – C, F, G, W]

NELUMBONACEAE Dumortier 1829 (Lotus-lily Family)

A family of 1 genus and 2 species, aquatic herbs, of temperate and subtropical e. North America and e. Asia. References: Wiersema in FNA (1997); Williamson & Schneider in Kubitzki, Rohwer, & Bittrich (1993).

Nelumbo Adanson 1763 (Lotus-lily, Lotus, Sacred-lotus, Sacred-bean)

A genus of 2 species, aquatic herbs, of temperate and subtropical e. North America and e. Asia. References: Williamson & Schneider in Kubitzki, Rohwer, & Bittrich (1993).

Identification notes: Nelumbo can be immediately distinguished in vegetative condition from the other "pads" (Nymphaea, Nuphar, and Nymphoides) by its peltate leaves, and from the peltate Brasenia by the much larger size and roundness of the leaves.

1 Petals yellow; mature fruits (“nuts”) usually < 1.25× as long as wide...... N. lutea 1 Petals pink or white; mature fruits (“nuts”) usually > 1.5× as long as wide...... N. nucifera

Nelumbo lutea Willdenow, Yonkapin, American Lotus-lily, Yellow Lotus, Yockernut, Water-chinquapin, Pond-nuts. Cp (GA, NC, SC, VA), Pd (VA), Mt (GA, VA): ponds, natural lakes; uncommon. June-September. NY and s. Ontario west to MN and IA, south to s. FL and e. TX, and south into the West Indies and Mexico. [= RAB, C, F, FNA, G, GW, K, S, W; N. pentapetala (Walter) Fernald] * Nelumbo nucifera Gaertner, Sacred-lotus, Oriental Lotus-lily, Pink Lotus. Cp (NC, SC), Pd (NC), Mt (GA): ponds and lakes; rare, introduced from Asia. June-September. [= RAB, C, F, FNA, G, GW, K]

NYCTAGINACEAE A.L. de Jussieu 1789 (Four-o'clock Family)

A family of about 31 genera and 400 species, trees, shrubs, vines, and herbs, of tropical, subtropical, and (less commonly) warm temperate regions, especially diverse in the New World. References: Bogle (1974)=Z; Spellenberg in FNA (2003b); Bittrich & Kühn in Kubitzki, Rohwer, & Bittrich (1993).

1 Flowers < 3 mm long, lacking involucral bracts subtending the petaloid calyx...... Boerhavia 1 Flowers > 10 mm long, with involucral bracts (simulating a calyx) subtending the petaloid calyx...... Mirabilis NYCTAGINACEAE 507

Boerhavia Linnaeus 1753 (Spiderling)

A genus of about 20-40 species, annual and perennial herbs, of tropical, subtropical, and warm temperate regions of the Old and New World. References: Spellenberg in FNA (2003b); Bogle (1974)=Z; Bittrich & Kühn in Kubitzki, Rohwer, & Bittrich (1993).

1 Fruit rounded at apex, stipitate-glandular; perennial...... B. coccinea 1 Fruit flat at the apex, glabrous; annual...... B. erecta

* Boerhavia coccinea P. Miller, Wineflower. Cp (NC, SC, VA): disturbed areas, adventive on ballast; rare, introduced from tropical America. June-September. Contrary to the statement in RAB that this species is "apparently not established," it is well established on the Wilmington (New Hanover County, NC) waterfront. [= RAB, FNA, K; = Boerhaavia coccinea – S, orthographic variant] Boerhavia diffusa Linnaeus, Red Spiderling, Spreading Hogweed. Cp (SC): [= FNA, K, Z] {not keyed at this time} {disentangle coccinea and diffusa; rewrite key} Boerhavia erecta Linnaeus, Erect Spiderling. Cp, Pd (GA, NC, SC): sandy fields, roadsides, disturbed areas; uncommon. May-October. NC south to FL, west to TX and AZ, perhaps only introduced in our area. [= RAB, FNA, K, Z; = Boerhaavia erecta – G, S, orthographic variant]

Mirabilis Linnaeus 1753 (Umbrella-wort, Four-o'clock)

A genus of about 55-60 species, annual and perennial herbs, of warm temperate America and s. Asia. References: Spellenberg in FNA (2003b); Le Duc (1995); Bittrich & Kühn in Kubitzki, Rohwer, & Bittrich (1993).

1 Petaloid calyx with a narrow tube 3-4 cm long, the spreading portion to 5 cm in diameter; involucre with 1 flower, not expanding in fruit; [section Mirabilis] ...... M. jalapa 1 Petaloid calyx with a broad tube < 0.5 cm long, the spreading portion < 1.5 cm in diameter; involucre with 3-5 flowers, expanding in fruit; [section Oxybaphus]. 2 Leaves cuneate at the base, 2.5-6× as long as wide; [native, of dry sandy habitats in s. SC southward] ...... M. albida 2 Leaves cordate at the base, 1-2× as long as wide; [introduced, of disturbed habitats] ...... M. nyctaginea

Mirabilis albida (Walter) Heimerl, Wild Four-o'clock, Pale Umbrella-wort. Cp (GA, SC): sandhills, adjacent disturbed sandy soils; rare (GA Special Concern). May-August. S. SC south to GA, west to TX, north in the interior to c. TN, IA, and KS. [= RAB, C, F, K, Z; = Oxybaphus albidus (Walter) Sweet – G; = Allionia albida Walter – S] * Mirabilis jalapa Linnaeus, Garden Four-o'clock, Marvel-of-Peru. Cp (GA, NC, SC, VA), Pd (NC, SC, VA): disturbed areas, or persistent at former garden sites; uncommon, introduced from tropical America. June-November. [= RAB, C, F, G, K, S, Z] * Mirabilis nyctaginea (Michaux) MacMillan, Heart-leaved Umbrella-wort, Wild Four-o'clock. Mt, Pd (NC, VA), Cp (VA): railroad embankments, other disturbed areas; rare, introduced from c. North America. June-October. [= RAB, C, F, K, W, Z; = Oxybaphus nyctagineus (Michaux) Sweet – G; = Allionia nyctaginea Michaux – S]

Mirabilis linearis (Pursh) Heimerl var. linearis. Manitoba and Alberta, south to TN, MS, and CA; scattered elsewhere in e. North America by introduction. [= FNA; < M. linearis – C, F, K; < Oxybaphus linearis (Pursh) B.L. Robinson – G] {not keyed at this time}

NYMPHAEACEAE R.A. Salisbury 1805 (Water-lily Family)

A family of 6 genera and about 75 species, aquatic herbs, cosmopolitan. References: Wiersema & Hellquist in FNA (1997); Schneider & Williamson in Kubitzki, Rohwer, & Bittrich (1993); Les et al. (1999).

1 Flowers nearly spherical, 2-5 cm in diameter; sepals 6 (in our species), petaloid, green to yellow, incurved; petals many, inconspicuous, scalelike or staminodial; leaves often of 2 types, the submersed leaves (when present) thinner in texture than the floating or emersed leaves; floating or emersed leaves having 60-90% of their surface area with vasculature derived from the midrib; rhizome with triangular or winged leaf scars; [subfamily Nupharoideae]...... Nuphar 1 Flowers hemispheric, 4-20 cm across; sepals 4, greenish, inconspicuous; petals spreading and ascending, white or yellow, showy; leaves of 1 type, floating; floating leaves having 25-40 % of their surface area with vasculature derived from the midrib; rhizome with circular leaf scars; [subfamily Nymphaeoideae]...... Nymphaea

Nuphar J.E. Smith 1809 (Spatterdock, Yellow Pondlily) NYMPHAEACEAE 508

A genus of about 16 species, aquatic herbs, of north temperate areas. Beal (1956) recognized 8 taxa of Nuphar in North America, which he treated as subspecies of the European N. lutea. Voss's (1985) statement (about the genus in Michigan) "our plants are quite easily distinguished ... and they are treated here as closely related species" applies equally (or better!) in our area. References: Beal (1956)=Z; Wiersema & Hellquist in FNA (1997); Padgett (1999)=Y; Schneider & Williamson in Kubitzki, Rohwer, & Bittrich (1993). Key based in large part on FNA.

1 Sepals 5 (or 5-6 in N. rubrodisca); stigmatic disc red; fruit deeply contricted below the stigmatic disc; leaf blades 3.5-25 cm long; [section Nuphar]. 2 Anthers 1-3 mm long; stigmatic disc with 6-10 deep crenations; stigmatic rays terminating 0-0.2 mm from the margin of the disc; constriction below disc 1.5-5 mm in diameter; leaf sinus 2/3 or more the length of the midrib; leaf blades 3.5-10 (-13) cm long...... [N. microphylla] 2 Anthers (2-) 3-6 mm long; stigmatic disc with 8-15 shallow crenations; stigmatic rays terminating 0-1.6 mm from the margin of the disc; constriction below disc 5-10 mm in diameter; leaf sinus ca. 1/2 the length of the midrib; leaf blades 5-25 cm long ...... [N. rubrodisca] 1 Sepals 6-9 (-12); stigmatic disc yellow, green, or sometimes reddish; fruit slightly or not at all constricted below the stigmatic disc; leaf blades 7-50 cm long; [section Astylus]. 3 Floating leaf blades 2-6× as long as wide, the sinus < 1/4 as long as the midrib; thin-textured submersed leaves often more abundant than the floating leaves; [of blackwater or tidal streams, rivers, and lakes of the Coastal Plain, se. VA, e. NC, e. SC, Panhandle FL, s. AL]. 4 Floating leaf blades 3-6× as long as wide; stigmatic rays elliptic, terminating < 1 mm from the edge of the disk; [of blackwater or tidal streams, rivers, and lakes of the Coastal Plain of se. VA to e. SC] ...... N. sagittifolia 4 Floating leaf blades 2-3× as long as wide; stigmatic rays linear, mostly terminating 1-2 mm from the edge of the disk; [blackwater streams and rivers, Panhandle FL and s. AL] ...... [N. ulvacea] 3 Floating leaf blades 1-2× as long as wide, the sinus > 1/3 as long as the midrib; floating or emersed leaves more abundant than submersed leaves; [collectively of various habitats and distributions, but not as above]. 5 Leaf petiole flattened on the upper (adaxial) surface and winged along the margins; fruit usualy purplish; sepals red or maroon at the base adaxially...... [N. variegata] 5 Leaf petiole terete or slightly flattened, not winged; fruit usually greenish or yellowish; sepals yellow or red at the base adaxially. 6 Lower leaf surface glabrous to sparsely pubescent; leaves 7-30 cm wide, the lobes acute to broadly rounded; leaves mostly emersed...... N. advena 6 Lower leaf surface densely pubescent; leaves 20-45 cm wide, the lobes broadly rounded; leaves mostly floating ...... N. orbiculata

Nuphar advena (Aiton) R. Brown ex Aiton f., Broadleaf Pondlily. Cp, Pd, Mt (GA, NC, SC, VA): lakes, ponds, natural depression ponds, old millponds, slow-flowing rivers (blackwater and brownwater); common. April-October. The most widespread and common Nuphar in e. North America, ranging from ME west to WI, south to s. FL, Cuba, TX, and n. Mexico. See N. sagittifolia for discussion of the two taxa. [= C, FNA; = Nuphar luteum (Linnaeus) Sibthorp & J.E. Smith ssp. macrophyllum (Small) E.O. Beal – RAB, GW, W, Z; > Nuphar advena – F, G; > Nuphar fluviatile (Harper) Standley – F, G; > Nuphar puteorum Fernald – F; = Nuphar lutea J.E. Smith ssp. advena (Aiton) Kartesz & Gandhi – K; > Nymphaea advena Aiton – S; > Nymphaea chartacea Miller & Standley – S; > Nymphaea fluviatilis Harper – S; = Nuphar advena ssp. advena – Y] Nuphar orbiculata (Small) Standley. Cp (GA): quiet waters in blackwater swamps; uncommon (GA Watch List). A Southeastern Coastal Plain endemic: FL north to ne. GA. [= FNA; = Nuphar lutea ssp. orbiculata (Small) E.O. Beal – K; > Nymphaea orbiculata Small – S; > Nymphaea bombycina (Miller & Standley) Standley – S; = Nuphar advena (Aiton) Aiton f. ssp. orbiculata (Small) D. Padgett – Y; = Nuphar luteum ssp. orbiculatum (Small) E.O. Beal – Z] Nuphar sagittifolia (Walter) Pursh, Narrowleaf Pondlily, Bonnets. Cp (NC, SC, VA): blackwater streams, rivers, and lakes, in swift, sluggish, or stagnant water, extending downriver into freshwater tidal areas; uncommon (NC Rare, VA Rare). April-October. Endemic to our area: e. VA south to ne. SC, very conspicuous and locally abundant on shallow bars along rivers such as the Northeast Cape Fear, Black, and Waccamaw, and forming dense colonies in Lake Waccamaw. Apparent hybrids with N. advena have been named Nuphar ×interfluitans Fernald. The submersed leaves have somewhat the texture and appearance of a thin leaf lettuce or the marine alga Ulva. This species appears to be closely related to N. ulvacea (Miller & Standley) Standley of blackwater rivers of panhandle FL, another phytogeographic connection between se. NC and panhandle FL. DePoe & Beal (1969) and Beal & Southall (1977) argue that this taxon and N. advena intergrade clinally, with N. advena inland and N. sagittifolia in the outer Coastal Plain, and that the two taxa are maintained by water temperatures. This ignores the fact that the two taxa often occur in close proximity to one another in both the inner and outer Coastal Plain. The frequency of so-called intermediates has also been exaggerated; few populations will present any difficulties in identification. I prefer to treat these taxa as species, with rare hybridization or introgression. [= C, FNA; = Nuphar luteum (Linnaeus) Sibthorp & J.E. Smith ssp. sagittifolium (Walter) E.O. Beal – RAB, GW, Z; = Nuphar sagittifolium – F, G, an orthographic variant; = Nuphar lutea J.E. Smith ssp. sagittifolia (Walter) E.O. Beal – K; = Nymphaea sagittifolia Walter – S]

Nuphar microphylla (Persoon) Fernald. Nova Scotia, Québec, and Manitoba south to s. NJ, se. PA, MI, IL, and MN. [= C, FNA, Y; = Nuphar microphyllum – F, G; = Nuphar lutea ssp. pumila (Timm) E.O. Beal – K; = Nuphar pumila Timm; = Nuphar luteum ssp. pumilum (Timm) E.O. Beal – Z] NYMPHAEACEAE 509 Nuphar rubrodisca Morong. New Brunswick, Québec, and Ontario south to NJ, PA, MI, and MN. [= FNA; = Nuphar ×rubrodisca Morong – C; = Nuphar ×rubrodiscum Morong – F; = Nuphar rubrodiscum – G; = Nuphar lutea J.E. Smith ssp. rubrodisca (Morong) Hellquist & Wiersema – K] Nuphar ulvacea (G.S. Miller & Standley) Standley, Sea-lettuce Pondlily. Blackwater streams of Panhandle FL and s. AL. [= FNA; = Nuphar luteum (Linnaeus) Sibthorp & J.E. Smith ssp. ulvaceum (G.S. Miller & Standley) E.O. Beal – GW, K; = Nymphaea ulvacea G.S. Miller & Standley – S] Nuphar variegata Durand in G.W. Clinton. Widespread in ne. North America, south to DE, PA, OH, IN, IL, IA, and NE. [= C, FNA; = Nuphar variegatum – F, G; = Nuphar lutea ssp. variegata (Durand) E.O. Beal – K; = Nuphar luteum ssp. variegatum (Durand) E.O. Beal – Z]

Nymphaea Linnaeus 1753 (Water-lily)

A genus of about 50 species, aquatic herbs, cosmopolitan. References: Wiersema in FNA (1997); Woods et al. (2005a, 2005b)=Z; Schneider & Williamson in Kubitzki, Rohwer, & Bittrich (1993).

1 Petals yellow; plants producing stolons ...... N. mexicana 1 Petals white (to pink); plants not producing stolons. 2 Petiole solid-colored; leaf length/width ratio (length measured from petiole attachment to tip of leaf, along midvein) (0.44-) avg. 0.56 (-0.71); two leaf lobes with rounded lobe tips; lower leaf surface reddish-purple...... N. odorata ssp. odorata 2 Petiole striped; leaf length/width ratio (-.55-) 0.63 (-0.73); leaf lobes with pointed tips; lower leaf surface green ...... [N. odorata ssp. tuberosa]

Nymphaea mexicana Zuccarini, Banana Water-lily, Yellow Water-lily. Cp (GA, NC, SC): sluggish or stagnant waters; rare, scattered in occurrence and possibly introduced from further south, but the introduction agents may well be wild ducks, such as canvasbacks. June-September. Ne. NC south to s. FL, west to TX, also in sw. US and the New World tropics. [= RAB, FNA, K, Z; = Castalia flava (Leitner) Greene – S] Nymphaea odorata W.T. Aiton ssp. odorata, White Water-lily. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): ponds, sluggish waters; common (uncommon in Piedmont and Mountains). June-September. Newfoundland west to Manitoba, south to FL and TX; also scattered in the w. United States. N. odorata is polymorphic, leading to the naming of numerous species, subspecies, and varieties (see synonymy for a few of the named entities). Wiersema in FNA (1997) recognize ssp. odorata (all of our plants) and ssp. tuberosa (Paine) Wiersema & Hellquist, more western and northern, but approaching our area (see below). Other named entities warrant further evaluation. N. odorata var. gigantea [= Castalia lekophylla Small] occurs on the Coastal Plain, and is considered to differ from var. odorata in its larger leaves (1.5-6 dm in diameter vs. 0.5-2.5 dm), larger flowers (mostly > 15 cm wide vs. mostly < 10 cm), and leaves upturned at the margins (vs. flat). N. odorata var. minor [= Castalia minor (Sims) Nyar] is considered to differ from var. odorata in its generally smaller size, leaves 5-11 cm in diameter, flowers mostly < 8 cm wide (vs. mostly > 9 cm wide); it may be merely a dwarfed form of extremely nutrient-limited waters of the Coastal Plain. [= FNA, K, Z; < N. odorata – RAB; >< N. odorata var. odorata – C; > N. odorata var. odorata – F, G; > N. odorata var. gigantea Tricker – C, F, G; > N. odorata var. stenopetala Fernald – F; > Castalia odorata (W.T. Aiton) Wood – S; > Castalia minor (Sims) Nyar – S; > Castalia lekophylla Small – S]

Nymphaea odorata W.T. Aiton ssp. tuberosa (Paine) Wiersema & Hellquist. South to MD, DE, NJ, PA. [= FNA, K, Z; < N. odorata var. odorata – C; = N. tuberosa Paine – F, G]

NYSSACEAE A.L. de Jussieu ex Dumortier 1829 (Tupelo Family)

A family of 1-3 genera and 8-12 species, trees and shrubs, of e. Asia, se. Asia, e. North America, and Central America. The circumscription and recognition of this family has been controversial; Nyssaceae has sometimes been included in a broadly circumscribed Cornaceae, but this appears to be phylogenetically incorrect (Xiang et al. 2002). References: Xiang et al. (2002).

Nyssa Linnaeus (Tupelo, Sour Gum, Black Gum)

A genus of about 8-10 species, trees and shrubs, of e. North America, e. Asia, se. Asia, and Central America. The only taxon other than those in our area is N. ursina Small [N. sylvatica Marshall var. ursina (Small) Wen & Stuessy], a 2-5 m tall shrub or small tree related to N. biflora and endemic to panhandle FL. The only other members of the genus are 2-4 e. and se. Asian species and a recently discovered species of Costa Rica (Hammel & Zamora 1990, Wen & Stuessy 1993). References: Burckhalter (1992)=Z; Wen & Stuessy (1993)=Y; Eyde (1966)=X.

Identification notes: Nyssa sylvatica is often mistaken (especially as seedlings, saplings, or fire-sprouts) for Diospyros virginiana, because of their similar, alternate, glossy-green, acuminate leaves. Nyssa can be distinguished by its three vascular bundle scars per leaf scar (vs. one Diospyros), leaves often with a few irregular teeth (vs. never toothed), leaves pale to medium NYSSACEAE 510 green beneath (vs whitish-green beneath), leaves lacking dark glands on the midrib above and the outer petiole (vs. present), and leaves glabrous or nearly so below (vs. glabrate to tomentose with curly hairs) (McKenney 1967).

1 Petioles of mature leaves 3-6 cm long; leaves to 30 cm long and 15 cm wide, at least the larger on a tree normally > 8 cm wide, often with a few irregular teeth, these typically located near the widest part of the blade...... N. aquatica 1 Petioles of mature leaves 0.5-2.0 (-2.5 cm) long; leaves to 18 cm long and 10 cm wide, the largest leaves on a tree rarely > 7 cm wide, generally entire, rarely with a few irregular teeth, these typically located toward the leaf apex. 2 Fruits 20-40 mm long, yellow, orange, or red when mature, the stone winged; pistillate flowers and fruits 1 per peduncle; trees often multiple-trunked, the trunks crooked; mature leaves densely pubescent beneath ...... N. ogeche 2 Fruits 6-15 mm long, blue-black when mature, the stone slightly ridged to nearly smooth; pistillate flowers (1-) 2-5 per peduncle; trees typically single-trunked, the trunk fairly straight; mature leaves glabrous to pubescent beneath. 3 Pistillate flowers and fruits (1-) 2 (-3) per peduncle; leaves with thick texture, rather stiff, typically widest beyond the middle, the apex typically obtuse, the margins entire (rarely with a few teeth on vigorous sprouts); trunk swollen or buttressed at base; bark of large trees rough, a vertical ridge-furrow pattern most prominent; [trees of swamps with periodic or seasonal flooding; mostly on the Coastal Plain]...... N. biflora 3 Pistillate flowers and fruits (2-) 3-5 (-8) per peduncle; leaves with thin texture, pliable, typically widest near the middle, the apex typically acuminate, the margins often with a few irregular teeth near the apex (though sometimes an entire tree with no toothed leaves); trunk not swollen or buttressed at base (even when growing in moist or wet habitats); bark of large trees rough, divided by deep vertical and horizontal furrows into a pattern of squarish checks; [trees of dry to mesic upland forests, less commonly in bottomlands or other wetlands, where flooding occurs at most occasionally and is of short duration; throughout our area] ...... N. sylvatica

Nyssa aquatica Linnaeus, WaterTupelo, Tupelo Gum, Cotton Gum. Cp (GA, NC, SC, VA): river swamps, where inundated for substantial periods of time; common. April-May; September-October. Se. VA south to panhandle FL, west to se. TX, north in the Mississippi Embayment to se. MO, s. IL, and e. KY, primarily on the Coastal Plain, but with scattered locations in other physiographic provinces, such as in sc. TN. [= RAB, C, F, GW, K, S, X, Y, Z; = N. uniflora Wangenheim – G] Nyssa biflora Walter, Swamp Tupelo, Water Gum, Swamp Black Gum. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): blackwater river swamps, depressions in pinelands, pocosins, either where inundated for substantial periods of time or in more- or-less permanently saturated organic peaty soils; common (rare in Piedmont). April-June; August-October. NJ south to s. FL, west to e. TX, primarily on the Coastal Plain, but scattered inland to c. NC, w. SC, c. TN, w. KY (Clark et al. 2005), se. MO, and c. AR. [= G, K, S, Z; = N. sylvatica Marshall var. biflora (Walter) Sargent – RAB, C, F, GW, X, Y] Nyssa ogeche Bartram ex Marshall, Ogeechee Lime, Ogeechee Tupelo, Ogeechee Plum. Cp (GA, SC): river swamps and wet forests with peaty soils, also in upland depression ponds; common, rare north of GA (SC Rare). April; August-October. A Southeastern Coastal Plain endemic: se. SC south to c. peninsular FL, west to s. AL. [= RAB, GW, K, X, Y, Z; > N. acuminata Small – S; > N. ogeche – S] Nyssa sylvatica Marshall, Sour Gum, Black Gum, Pepperidge. Mt, Pd, Cp (GA, NC, SC, VA): dry or mesic upland forests, less commonly in bottomlands, pine savannas, or upland depressions, where occasionally inundated briefly; common. April- June; August-October. S. ME west to MI and se. WI, south to c. peninsular FL, west to e. TX and e. OK. N. sylvatica is quite variable in morphology and ecology, at least some of the morphologic variation correlated with geography and ecology. The status of varieties recognized by previous authors (such as Fernald 1950) needs reassessment. In the Mountains of our area N. sylvatica is typically found in dry woodlands, such as pine-oak/heath, with xerophytic species such as Pinus virginiana and Quercus montana. In the outer Coastal Plain of the Carolinas, N. sylvatica often occurs in wet savannas with Pinus serotina, where often mistaken (because of the wetland habitat) for N. biflora. The leaves turn a brilliant orange-red in fall (often a few on any tree turning prematurely in July or August). [= G, K, S, Z; = N. sylvatica var. sylvatica – RAB, C, GW, X, Y; > N. sylvatica var. sylvatica – F; > N. sylvatica var. dilatata Fernald – F; > N. sylvatica var. caroliniana (Poiret) Fernald – F]

OLEACEAE Hoffmansegg & Link 1813 (Olive Family)

A family of about 25 genera and 600-615 species, trees and shrubs, nearly cosmopolitan, but centered in Asia. References: Hardin (1974)=Z; Green in Kadereit (2004).

1 Leaves compound. 2 Leaves pinnately compound with > 5 leaflets; petals absent; fruit a samara; small to large tree; [tribe Oleeae, subtribe Fraxininae] ...... Fraxinus 2 Leaves trifoliolate; petals 6-10, yellow, conspicuous; fruit a deeply 2-lobed dryish berry; [tribe Jasmineae]...... [Jasminum] 1 Leaves simple. 3 Flowers bright yellow, showy; fruit a many-seeded capsule; [tribe Forsythieae]...... Forsythia 3 Flowers white, lilac, or purplish; fruit a drupe or 4-seeded capsule. 4 Leaves cordate or truncate at the base; fruit a 4-seeded capsule; corolla lobes shorter than the tube; flowers lilac or white, in terminal panicles; [tribe Oleeae, subtribe Ligustrinae] ...... Syringa 4 Leaves cuneate to rounded at the base; fruit a drupe; corolla lobes either shorter or longer than the tube; flowers white or greenish-white, in terminal or lateral panicles or fascicles. OLEACEAE 511 5 Corolla absent; calyx minute or lacking; flowers in axillary fascicles; [tribe Oleeae, subtribe Oleinae] ...... Forestiera 5 Corolla present (often conspicuous and showy); calyx present; flowers in lateral or terminal panicles or in terminal subumbellate clusters. 6 Corolla lobes 5-12; flowers in terminal subumbellate clusters; [tribe Jasmineae] ...... [Jasminum] 6 Corolla lobes 4; flowers in lateral or terminal panicles 7 Corolla lobes elongate, much longer than the corolla tube; [tribe Oleeae, subtribe Oleinae]...... Chionanthus 7 Corolla lobes short, no longer than the corolla tube. 8 Inflorescence a many-flowered terminal panicle; leaves generally ovate, elliptic or lanceolate (widest below or at the middle); [tribe Oleeae, subtribe Ligustrinae] ...... Ligustrum 8 Inflorescence a few-flowered axillary panicle; leaves generally oblanceolate or obovate (widest above the middle); [tribe Oleeae, subtribe Oleinae] ...... Osmanthus

Chionanthus Linnaeus 1753 (Fringe-tree, Old Man's Beard)

A genus of controversial circumscription, either of only 3 species, limited to se. North America and e. Asia, or (if including Linociera) of about 60-100 species, primarily tropical. Ch. pygmaeus Small is endemic to scrub in peninsular FL. References: Hardin (1974)=Z.

Chionanthus virginicus Linnaeus, Fringe-tree, Old Man's Beard. Pd, Mt, Cp (GA, NC, SC, VA): dry, mesic, or wet forests and woodlands, granitic flatrocks and domes, glades and barrens over various rocks (including granite, greenstone, etc.), swamp forests in the Coastal Plain, rarely pocosins; common. April-May; July-September. NJ, s. PA, s. OH, and MO south to n. FL and e. TX. Ch. virginicus in our area shows a diversity of morphology and correlated habitat that suggests the possible presence of two taxa. Swamp- and pocosin-inhabiting populations in the outer Coastal Plain have leaves 4-8× as long as wide and seem very different than Piedmont dry woodland populations with leaves 1-2× as long as wide; further and more careful study is needed. Ch. virginicus is a traditional southern yard plant, often used as a "specimen plant," very showy in spring, particularly when grown to its full size. [= RAB, C, F, G, GW, K, S, W, Z]

Forestiera Poiret 1812 (Forestiera)

A genus of about 15-20 species, shrubs, of sw. and se. North America, Central America, and the West Indies. References: Anderson (1985)=Y; Godfrey (1988)=X; Hardin (1974)=Z; Johnston (1957)=Q; Green in Kadereit (2004).

1 Leaves (6-) 7-8 (-9) cm long, long-acuminate or acuminate (rarely acute) at the apex, the tip sharply pointed; [of swamp forests, sloughs, and ponds] ...... F. acuminata 1 Leaves 1.5-7 (-8) cm long, obtuse at the apex, or if short-acuminate the ultimate tip blunt; [of shell middens and calcareous bluffs]. 2 Leaves evergreen, glabrous above, glabrous and punctate below; leaf margins entire...... F. segregata var. segregata 2 Leaves deciduous, at least sparsely pubescent on the midrib above, pubescent and non-punctate below. 3 First-year twigs pubescent, the pubescence evenly distributed (not in 2 lines); petioles moderately pubescent; flowering in early spring from buds on twigs of the previous season; leaves 5-7 (-8) cm long ...... F. godfreyi 3 First-year twigs pubescent, the pubescence in 2 lines on either side of the twig; petioles glabrous (or with a very few hairs; flowering in mid-late summer, the flowers in leaf axils; leaves mostly 2-5 cm long...... F. ligustrina

Forestiera acuminata (Michaux) Poiret, Swamp-privet. Cp (GA, SC): swamp forests, especially over calcareous substrates; uncommon. March; May-June. SC south to n. FL, west to TX, north in the interior to KY, e. and c. TN, IN, IL, MO, and KS. [= RAB, C, F, G, GW, K, S, Q, X, Y, Z] Forestiera godfreyi L.C. Anderson, Godfrey's Forestiera. Cp (GA, SC): shell middens, maritime forests over shell substrate; rare (GA Special Concern). Mid January-February; April-May. Se. SC (Beaufort and Charleston counties) to e. GA and n. FL. [= K, X, Y; < F. pubescens Nuttall – S, in part (apparently)] Forestiera ligustrina (Michaux) Poiret in Lamarck, Southern-privet. Cp, Pd (GA, SC): upland forests and slopes along streams, mostly on shell middens or calcareous rocks; rare (SC Rare). E. SC south to n. FL, west to se. TX, north in the interior to c. TN and KY. [= K, S, Q, X, Z] Forestiera segregata (Jacquin) Krug & Urban var. segregata, Florida-privet. Cp (GA, SC): calcareous scrub, shell middens, maritime forests and thickets; rare (GA Special Concern, SC Rare). Se. SC south to s. FL, and in the West Indies. Var. pinetorum (Small) M.C. Johnston is restricted to s. FL. [= K, Q, Z; > F. porulosa (Michaux) Poiret – S; > F. globularis Small – S; < F. segregata – X]

Forsythia Vahl 1804 (Forsythia, Golden-bells)

OLEACEAE 512 A genus of about 7-9 species, shrubs, of e. Asia and se. Europe. References: Hardin (1974)=Z; Green in Kadereit (2004).

1 Mature branches hollow or irregularly pith-filled between the nodes; leaves oblong-ovate, toothed or 3-parted; branches arching when well-developed ...... F. suspensa 1 Mature branches cross-septate (chambered) between the nodes; leaves oblong-lanceolate, toothed; branches upright...... F. viridissima

* Forsythia suspensa (Thunberg) Vahl, Weeping Forsythia. Pd (GA, NC, VA), Mt (VA): waste places, vacant lots, suburban woodlands; commonly planted and persistent, rarely escaped (native of China). [= C, G, K, Z] * Forsythia viridissima Lindley, Greenstem Forsythia. Pd (GA, NC, VA), Cp, Mt (VA): waste places, vacant lots, suburban woodlands; commonly planted and persistent, rarely escaped (native of China). [= C, G, K, W, Z]

Fraxinus Linnaeus 1753 (Ash)

A genus of about 45-65 species, trees, mostly north temperate (Asia, North America, Europe). References: Hardin & Beckmann (1982)=Z; Miller (1955)=Y; Green in Kadereit (2004).

1 Young twigs 4-angled or narrowly 4-winged; [trees of calcareous woodlands in the Mountains of sw. VA and northward and westward]...... F. quadrangulata 1 Young twigs terete (rounded in cross-section); [trees of various habitats, collectively widespread in our area]. 2 Lateral leaflets sessile; calyx absent...... F. nigra 2 Lateral leaflets with petiolules (1-) 3-20 mm long; calyx present, persisting as a minute cup at the base of the fruits. 3 Leaves minutely papillose beneath (best seen at magnification of 40× or more), and sometimes also pubescent, more-or-less strongly whitened; wing of the samara decurrent only onto the upper 1/3 (or less) of the samara body ...... F. americana 3 Leaves glabrous to pubescent beneath (never papillose), green; wing of the samara decurrent onto 1/2 (or more) of the samara body. 4 Petiolules of the lowermost leaflets 1-9 mm long, all but 1-2 mm narrowly winged; samara mostly < 7 mm wide; calyx 0.5-1.5 mm long...... F. pennsylvanica 4 Petiolules of the lowermost leaflets 3-20 mm long, not winged (except F. caroliniana); samara mostly > 7 mm wide; calyx 1-6 mm long. 5 Body of samara flattened, winged the full length of the samara body; calyx 1 mm long; leaf scars slightly notched; small tree, often multi-trunked...... F. caroliniana 5 Body of samara terete or subterete, winged about 1/2 the length of the samara body; calyx 2.5-6 mm long; leaf scars deeply notched; medium to large tree, typically single-trunked ...... F. profunda

Fraxinus americana, White Ash, American Ash. Mt, Pd, Cp (GA, NC, SC, VA): mesic slopes, rich cove forests, dryish calcareous or mafic glades and woodlands (with Juniperus virginiana var. virginiana and Carya glabra); common (rare in Coastal Plain of NC, SC, and GA). April-May; August-October. Nova Scotia west to MN, south to FL and TX. A valuable timber tree. The division into 2 taxa, var. americana and var. biltmoreana, needs further study. [= C, K, W, Z; > F. americana Linnaeus var. americana – RAB, F, G; > F. americana Linnaeus var. biltmoreana (Beadle) J. Wright ex Fernald – RAB, F, G; > F. americana var. microcarpa A. Gray – F; > F. americana – S, Y; > F. biltmoreana Beadle – S, Y] Fraxinus caroliniana P. Miller, Water Ash, Pop Ash, Carolina Ash. Cp, Pd (GA, NC, SC, VA): deeply to shallowly flooded swamps; common (rare in Piedmont). May; July-October. Se. VA south to FL, west to TX, primarily on the Coastal Plain. A small tree, sometimes very abundant (and nearly the only subcanopy species) as the understory in Taxodium-Nyssa swamps. [= RAB, C, G, GW, K, Y, Z; > F. caroliniana var. caroliniana – F; > F. caroliniana var. oblanceolata (M.A. Curtis) Fernald & Schubert – F; > F. caroliniana var. cubensis (Grisebach) Lingelsh. – F; > F. caroliniana – S; > F. pauciflora Nuttall – S] Fraxinus nigra Marshall, Black Ash. Mt, Pd (VA): seepage swamps and mountain streambanks; rare (VA Watch List). April-May; August-October. Newfoundland and Québec west to Manitoba, south to DE, VA, IN, and IA. [= C, F, G, K, W, Y, Z] Fraxinus pennsylvanica Marshall, Green Ash, Red Ash. Cp, Pd, Mt (GA, NC, SC, VA): bottomlands and swamps, especially along brownwater rivers and streams; common. April-May; August-October. Nova Scotia west to Alberta, south to FL and TX. Variation in this species (see synonymy) needs further study. [= C, GW, K, W, Z; > F. pennsylvanica var. subintegerrima (Vahl) Fernald – RAB, F, G; > F. pennsylvanica var. pennsylvanica – RAB, F, G; > F. pennsylvanica var. austinii Fernald – F; > F. darlingtonii Britton – S; > F. pennsylvanica – S; > F. smallii Britton – S; ? F. pennsylvanica ssp. pennsylvanica – Y] Fraxinus profunda (Bush) Bush, Pumpkin Ash. Cp, Pd (GA, NC, SC, VA), Mt (NC): swamps, especially along blackwater rivers and streams and in freshwater tidal wetlands (as along the James, Pamunkey, Mattaponi, and Rappahannock rivers in e. VA), also in brownwater bottomlands; common (rare in Piedmont and Mountains). April-May; August-October. S. NJ south to n. FL, west to LA, mostly on the Coastal Plain, north in the interior to w. NC, sc. TN, e. AR, se. MO, s. IL, IN, OH, sc. MI, ne. PA, and w. NY. This species has a peculiar distribution; see McCormac, Bissell, & Stine (1995) for additional OLEACEAE 513 discussion. The nomenclature is controversial. [= C, GW, K, W, Z; = F. tomentosa Michaux f. – RAB, F, G, Y; > F. profunda – S; > F. michauxii Britton – S] Fraxinus quadrangulata Michaux, Blue Ash. Mt (GA, VA): mesic to dry calcareous woodlands and forests; rare (GA Special Concern, VA Watch List). April; July-October. S. Ontario west to s. MI and e. KS, south to sw. VA, e. TN, nw. GA, n. AL, and OK. [= C, F, G, K, S, Y, Z]

Jasminum Linnaeus 1753 (Winter Jasmine)

A genus of about 200 species, shrubs and woody vines, of tropical (and rarely temperate) Eurasia. References: Green in Kadereit (2004)

1 Leaves simple; flowers white...... [J. multiflorum] 1 Leaves trifoliolate; flowers yellow. 2 Leaflets 2.5-7 cm long; flowers 3.5-5 cm across...... [J. mesnyi] 2 Leaflets 1-3 cm long; flowers ca. 2.5 cm across ...... [J. nudiflorum]

* Jasminum mesnyi Hance, Japanese Jasmine, Primrose Jasmine, native of w. China, is planted and often persists. Reported for GA (K). [= K] * Jasminum multiflorum (Burmann f.) Andrews, Star Jasmine, is planted and naturalized at least as far north as Jacksonville, Duval County, FL (Wunderlin & Hansen 2004). [= K] * Jasminum nudiflorum Lindley, Winter Jasmine, native of China, is commonly planted and often persists. It has green stems and yellow flowers. Reported for GA (K). [= K]

Ligustrum Linnaeus 1753 (Privet)

A genus of about 40 species, shrubs and trees, of the Old World. I have here largely followed Hardin (1974), though, as he points out, "the taxonomy and nomenclature of our plants seem uncertain in a few cases" and "it is difficult to determine which are really naturalized in our area." It is possible that not all the species treated below are truly naturalized, and that taxonomic changes will be needed. References: Hardin (1974)=Z; Green in Kadereit (2004). The key is based closely on Hardin (1974).

1 Twigs glabrous. 2 Corolla tube equalling or shorter than the corolla lobes. 3 Leaves persistent or tardily deciduous, 6-15 cm long ...... L. lucidum 3 Leaves deciduous, 3-6 cm long...... L. vulgare 2 Corolla tube slightly longer than or up to 3× as long as the corolla lobes. 4 Leaves persistent and glossy, rounded or broadly cuneate at the base; corolla tube slightly longer than the corolla lobes...... L. japonicum 4 Leaves deciduous or semi-evergreen, cuneate at the base; corolla tube ca. 3× as long as the corolla lobes...... L. ovalifolium 1 Twigs pubescent. 5 Corolla tube equalling or shorter than the corolla lobes. 6 Flowers sessile or subsessile ...... L. quihoui 6 Flowers pedicellate. 7 Twigs densely pubescent; leaves pubescent on the midrib beneath ...... L. sinense 7 Twigs minutely puberulent; leaves glabrous ...... L. vulgare 5 Corolla tube slightly longer than or up to 3× as long as the corolla lobes. 8 Pedicels pubescent; calyx pubescent...... L. obtusifolium 8 Pedicels glabrous; calyx glabrous or slightly pubescent at the base. 9 Leaves 2-6 cm long; twigs conspicuously pubescent ...... L. amurense 9 Leaves 4-10 cm long; twigs minutely puberulent...... L. japonicum

* Ligustrum amurense Carrière, Amur Privet. Pd (NC, VA), Cp, Mt (VA {SC}: disturbed places; rare, native of n. China. [= RAB, C, F, G, K, Z] * Ligustrum japonicum Thunberg, Japanese Privet. Cp (GA, NC, SC, VA), Pd (NC, VA): disturbed places; rare, native of Japan and Korea. [= RAB, K, Z] * Ligustrum lucidum Aiton f., Glossy Privet. Pd, Cp (NC): disturbed places; rare, native of China, Japan, and Korea. This species is superficially similar to L. japonicum; the lateral leaf veins are translucent in this species. [= K, S, Z] * Ligustrum obtusifolium Siebold & Zuccarini. Cp, Pd (NC, VA), Mt (VA): disturbed places; uncommon, native of Japan. [= C, F, G, K, Z] * Ligustrum ovalifolium Hasskarl, California Privet. Cp, Pd (NC, VA): disturbed places; rare, native of Japan. [= RAB, C, F, G, K, S, Z] * Ligustrum quihoui Carrière, Wax-leaved Privet. Cp (NC, VA): disturbed places; rare, native of China. [= K, Z] OLEACEAE 514 * Ligustrum sinense Loureiro, Chinese Privet. Cp, Pd, Mt (GA, NC, SC, VA): moist forests, especially alluvial bottomlands; common, native of China. This species is one of the most noxious of our weeds, choking out native vegetation in hundreds of square kilometers of land in our area. The rapidity with which it has engulfed southern wetlands is hinted at by Small's (1933) mention of it only as "an escape in southern Louisiana." [= RAB, C, G, GW, K, S, W, Z] * Ligustrum vulgare Linnaeus, Common Privet. Cp, Pd (NC, VA): disturbed places; rare, native of Europe and n. Africa. [= C, F, G, K, S, Z]

Osmanthus Loureiro 1790 (Wild Olive, Devilwood)

A genus of about 15-32 species, shrubs and trees, of se. Asia (most species) and se. North America. References: Hardin (1974)=Z; Green in Kadereit (2004).

1 Leaf margins entire; leaves usually >7 cm long; [native tree of Coastal plain forests] ...... O. americanus 1 Leaf margins spiny-toothed; leaves < 10 cm long; [horticulturally planted, rarely naturalizing]...... O. ×fortunei

Osmanthus americanus (Linnaeus) Bentham & Hooker f. ex A. Gray, Wild Olive, Devilwood. Cp (GA, NC, SC, VA): maritime forests and (in GA, SC, and extreme s. NC) dry, sandy forests well inland, and reported for wet habitats as well further south; uncommon (VA Rare). April-May; August-October. Se. VA south to FL, west to LA; also in Mexico. O. megacarpus Small, sometimes treated as a variety, O. americanus var. megacarpus (Small) P.S. Greene, is endemic to pine scrub in peninsular FL and differs primarily in having a larger fruit. The very hard, tough, and unsplittable wood is the inspiration for the common name "Devilwood." O. americanus is a conspicuous element of maritime forests in most of our area, readily recognizable by the flattened twigs characteristic of the family, and the opposite (often a few subopposite), glossy, evergreen leaves. [= RAB, F, G, GW; = O. americanus var. americanus – C, K, Z; = Amarolea americana (Linnaeus) Small – S] * Osmanthus ×fortunei Carrière [= O. fragrans × heterophyllus], Fortune’s Sweet Olive. Pd (NC): suburban woodlands, escaped from horticultural plantings; rare, hybrid originating in Japan of two species native to Japan.

Syringa Linnaeus 1753 (Lilac)

A genus of about 20-23 species, shrubs, from s. Europe to se. Asia. References: Hardin (1974)=Z; Green in Kadereit (2004).

* Syringa vulgaris Linnaeus, Lilac. Mt (NC, VA): commonly planted, persistent and naturalizing around old farms; rare, native of se. Europe. [= C, F, G, K, Z]

ONAGRACEAE A.L. de Jussieu 1789 (Evening-primrose Family)

A family of about 18 genera and 650 species, herbs, shrubs, and rarely trees, cosmopolitan (especially of temperate and subtropical America). References: Munz (1965)=X; Crisci et al. (1990).

1 Flowers 2-merous, the petals white; fruits with uncinate trichomes; leaves opposite, decussate, borne spreading at right angles to the stem, mostly ovate, on petioles mostly 0.5-8 cm long ...... Circaea 1 Flowers (3-) 4 (-7)-merous, the petals yellow, pink, or white (or absent); fruits lacking uncinate trichomes; leaves alternate (rarely opposite). not decussate, usually ascending or appressed (rarely spreading at right angles to the stem), mostly lanceolate, mostly sessile or subsessile. 2 Fruit indehiscent; seeds 1-6 per capsule, 1.5-3.5 mm long ...... Gaura 2 Fruit dehiscent; seeds (10-) 50-many per capsule, 0.3-2 mm long. 3 Seeds with an elongate coma at one end (wind-dispersed); petals pink or white. 4 Leaves all alternate; flowers numerous in a terminal raceme (with small bracts); flower buds reflexed, the flowers held horizontally or ascending; petals 10-20 mm long; stigma 4-lobed; plants 10-30 dm tall...... Chamerion 4 Leaves all or at least the lowermost opposite; flowers few, axillary, or in poorly developed, leafy racemes; flower buds not reflexed, the flowers ascending; petals 2-8 mm long (except 10-15 mm long in E. hirsutum); stigma capitate (except 4-lobed in E. hirsutum); plants 1-20 dm tall ...... Epilobium 3 Seeds not comose (gravity-dispersed); petals yellow or absent (rarely white or pink). 5 Calyx tube not extended beyond the summit of the ovary; sepals persistent on the capsule (rarely deciduous); stamens 4, 8, or 10-14; petals yellow or absent; [primarily of wetlands] ...... Ludwigia 5 Calyx tube extended beyond the summit of the ovary; sepals deciduous; stamens 8; petals yellow (rarely pink or white); [primarily of uplands] ...... Oenothera

Calylophus Spach

A genus of 6 species, of North America. ONAGRACEAE 515

Calylophus serrulatus (Nuttall) Raven, east to w. KY. [= K; = Oenothera serrulata Nuttall] {not keyed at this time; synonymy incomplete}

Chamerion (Rafinesque) Rafinesque (Fireweed)

There is increasingly strong evidence for the recognition of this group of plants as a genus separate from Epilobium. References: Mosquin (1966)=Z; Holub (1972)=Y; Munz (1965)=X.

Chamerion platyphyllum (Daniels) Löve & Löve, Great Willow-herb, Fireweed. Mt (NC, VA): grassy balds, roadsides, disturbed areas; uncommon (NC Rare). July-September. Chamerion platyphyllum has a circumboreal distribution; it is a member of a circumboreal complex, consisting of several related taxa that differ in chromosome number, a variety of morphological characters, and distribution. The tetraploid Chamerion platyphyllum is generally more southern, extending south in North America to NJ, montane w. NC and ne. TN, n. IN, MN, SD, AZ, NM, and CA; it may be more appropriately treated as a variety or subspecies of Ch. angustifolium. The diploid Chamerion angustifolium (Linnaeus) Holub is arctic and boreal, extending south in North America to New Brunswick, Québec, Ontario, alpine WY, and British Columbia. The hexaploid is Chamerion danielsii D. Löve. [< E. angustifolium – RAB, G, GW, W; = E. angustifolium var. canescens A. Wood – C; >< E. angustifolium var. angustifolium – F, X; > E. angustifolium var. platyphyllum (Daniels) Fernald – F; = Chamerion angustifolium (Linnaeus) Holub ssp. circumvagum (Mosquin) Kartesz – K; < Chamaenerion angustifolium (Linnaeus) Scopoli – S; < Chamerion angustifolium (Linnaeus) Holub – Y; = Epilobium angustifolium Linnaeus ssp. circumvagum Mosquin – Z]

Circaea Linnaeus (Enchanter's-nightshade)

A genus of 7-8 species, herbs, of temperate and boreal regions of the Northern Hemisphere. References: Boufford (1983)=Z; Boufford (2005)=Y; Munz (1965)=X; Averett & Boufford (1985); Skvortsov (1979). Key based on Z.

Identification notes: Sometimes confused in vegetative condition with Phryma.

1 Flowers opening before elongation of the raceme axis, therefore clustered and corymbiform at the apex of the raceme, borne on erect or ascending pedicels; plant 5-25 (-30) cm tall; fruits clavate, 2.0-2.5 mm long, 0.7-1.2 mm thick, 1-locular ...... C. alpina ssp. alpina 1 Flowers opening after elongation of the raceme axis, more or less loosely spaced, borne on spreading pedicels; plants (12-) 20-100 cm tall; fruits obovoid to pyriform, 2.8-3.9 (-4.5) mm long, 1.5-3.6 mm thick, 2-locular, or the fruits sterile and aborting shortly after anthesis, 1-2-locular when present. 2 All, or nearly all, ovaries developing to maturity; fruit with corky-thickened ribs separated by deep grooves...... C. canadensis ssp. canadensis 2 All ovaries aborting shortly after anthesis (very rarely a few persistent after anthesis); fruit (when somewhat persistent) with low ribs and shallow grooves...... C. ×sterilis

Circaea alpina Linnaeus ssp. alpina, Alpine Enchanter's-nightshade. Mt (GA?, NC, VA): moist organic soil at high elevations (especially in spruce-fir and northern hardwood forests), rocky seepages, in spray behind waterfalls, at dripping cliff bases; uncommon. June-September. C. alpina is treated by Z as a circumboreal complex of six subspecies. Ssp. alpina is itself circumboreal, in North America ranging from Newfoundland and Labrador, west to AK, south to MD, w. NC, e. TN, n. GA (?), KY, n. IL, MN, MT, and WA, disjunct in montane sites southward in the w. United States, such as the Black Hills of SD, and isolated montane sites in CO, AZ, and NM. Another subspecies occurs in w. North America, and four subspecies occur in humid and montane parts of Asia. [= K, X, Z; < C. alpina – RAB, F, G, GW, S, W; = C. alpina var. alpina – C] Circaea canadensis (Linnaeus) Hill ssp. canadensis, Canada Enchanter's-nightshade. Mt (GA, NC, VA), Pd (GA, NC, SC, VA), Cp (GA, NC, VA): mesic, nutrient-rich forests; common (SC Rare). June-August. Nova Scotia and New Brunswick west to se. Manitoba and ND, south to e. NC, c. SC, s. GA, LA, OK, and NE. The systematics of this taxon is controversial, and the best treatment is still unclear. Most recently, Boufford (2005) has treated the complex as 2 species, C. canadensis and C. lutetiana, the former with 2 subspecies, ssp. canadensis of eastern North America and ssp. quadrisulcata of Asia. Previously, Boufford (1983) treated the complex as a circumboreal complex of 3 subspecies of C. lutetiana, including the North American ssp. canadensis (Linnaeus) Ascherson & Magnus, the primarily Asian ssp. quadrisulcata (Maximowicz) Ascherson & Magnus, and the European ssp. lutetiana. Other authors have preferred varietal status for the 3 entities, full species status, no formal status at all (C. lutetiana as a polymorphic complex), or associating the more similar pair (North American and Asian) as 2 subspecies separate from the European at specific rank. Boufford (1983) and Averett & Boufford (1985) show convincingly that separate taxonomic status for the three entities is warranted, and that ssp. canadensis is more closely related to ssp. quadrisulcata. The question of the appropriate taxonomic level remains. Boufford (1983) states that "although subspp. canadensis and quadrisulcata are placed in C. lutetiana, this might not ultimately prove to be the best treatment." Later, flavonoid data showed strong differences between the three taxa, stronger than the differences between many of the other species in the genus (Averett & Boufford 1985). Morphologic differences between the three taxa are fairly subtle but appear to be consistent. The complicated synonymy is perhaps an example of a too-zealous attempt to have nomenclature reflect subtleties of relationship and ONAGRACEAE 516 evolutionary divergence, our understanding of which is unclear and changeable. [=Y; = C. lutetiana Linnaeus ssp. canadensis (Linnaeus) Ascherson & Magnus – RAB, K, W, X, Z; = C. lutetiana var. canadensis Linnaeus – C; = C. quadrisulcata (Maximowicz) Franchet & Savatier var. canadensis (Linnaeus) Hara – G; > C. canadensis var. canadensis – F; > C. canadensis var. virginiana Fernald – F; C. latifolia Hill – S; = C. quadrisulcata ssp. canadensis (Linnaeus) Löve & Löve] Circaea ×sterilis Boufford, Hybrid Enchanter's-nightshade. Mt (NC, VA): mesic, nutrient-rich forests; rare. June-August. C. ×sterilis is reported to occur frequently in the absence of one or both of its parents (Z, Skvortsov 1979), and is therefore treated separately and keyed here. It ranges from Newfoundland west to Ontario and MN, south to w. NC, OH, and WI. It appears to be rare in our area, but should be sought more carefully. Recognition of C. canadensis (Linnaeus) Hill as distinct from C. lutetiana renders the hybrid binomial name C. ×intermedia inappropriate for North American plants, since it is the hybrid of C. alpina ssp. alpina and the European C. lutetiana. [= C. ×sterilis Boufford – Y; = C. ×intermedia Ehrhart (pro sp.) – RAB, C, K, W, X, Z (but misapplied as to our material if C. canadensis is accepted as a species); > C. canadensis var. canadensis – F, misapplied; > C. canadensis var. virginiana Fernald – F; = C. canadensis (Linnaeus) Hill – G, misapplied]

Epilobium Linnaeus (Willow-herb) (also see Chamerion)

Epilobium is a large genus (ca. 200 species), distributed primarily in boreal and alpine latitudes and elevations. All five of the species in our area reach or approach their southern limits in eastern North America here. There is increasing opinion that E. angustifolium and its relatives should be distinguished at the generic level from Epilobium, as Chamerion. References: Munz (1965)=Z.

1 Leaves all alternate; flowers numerous in a terminal raceme (with small bracts); flower buds reflexed, the flowers held horizontally or ascending; petals 10-20 mm long; stigma 4-lobed; plants 10-30 dm tall...... [see Chamerion] 1 Leaves all or at least the lowermost opposite; flowers few, axillary, or in poorly developed, leafy racemes; flower buds not reflexed, the flowers ascending; petals 2-8 mm long (except 10-15 mm long in E. hirsutum); stigma capitate (except 4-lobed in E. hirsutum); plants 1-20 dm tall; [section Lysimachion]. 2 Stigma 4-cleft; petals 10-15 mm long ...... [E. hirsutum] 2 Stigma capitate; petals 2-8 mm long. 3 Leaves lanceolate, distinctly broader below the middle, flat, the larger generally at least 10 mm wide, toothed. 4 Principal leaves 3-7 cm long, with obscure marginal teeth, the apices merely acute; internodes (below the inflorescence) glabrous, glabrescent, or with pubescence scattered over the surface; mature coma (attached to plump seeds) nearly white; plants often strict or sparingly branched; seeds striate (with well-developed papillae arranged conspicuously in lines)...... E. ciliatum ssp. ciliatum 4 Principal leaves 5-15 cm long, with conspicuous and often irregular marginal teeth, the apices acuminate to attenuate; internodes (below the inflorescence) with lines of pubescence (some internodes on a given plant sometimes with scattered pubescence or glabrous); mature coma cinnamon (attached to plump seeds) brown (pale when immature); plants generally well-branched, with a bushy habit; seeds papillose (the papillae sometimes forming weak lines)...... E. coloratum 3 Leaves linear to narrowly lanceolate, broadest near the middle, revolute, the larger generally < 10 mm wide, not toothed. 5 Pubescence spreading...... E. strictum 5 Pubescence appressed. 6 Upper leaf surface finely and rather densely pubescent ...... E. leptophyllum 6 Upper leaf surface glabrous or with a few scattered hairs near the midrib ...... [E. palustre]

Epilobium ciliatum Rafinesque ssp. ciliatum, American Willow-herb. Mt (NC, VA): bogs, seeps, disturbed wet places (such as moist edges of logging roads); uncommon (NC Rare). June-September. Newfoundland and Labrador west to AK, south to VA, w. NC, ne. TN, IN, IA, CA, TX, Mexico, Central America; disjunct in Chile and Argentina. [= K; < E. ciliatum – RAB, W; = E. ciliatum var. ciliatum – C; > E. ciliatum – F, X, in a narrower sense; > E. glandulosum Lehm. var. adenocaulon (Hausskn.) Fernald – F; > E. adenocaulon Hausskn. var. adenocaulon – G, Z] Epilobium coloratum Biehler, Bronze Willow-herb, Eastern Willow-herb. Mt (GA, NC, SC, VA), Pd, Cp (NC, VA): seepages, moist open places; common (rare in Coastal Plain of NC). June-September. ME west to MN, south to NC, n. GA, AL, AR, and TX. There are some difficulties in distinguishing this species and E. ciliatum in our area. [= RAB, C, F, G, GW, K, S, W, Z] Epilobium leptophyllum Rafinesque, Narrowleaf Willow-herb, American Marsh Willow-herb. Mt (NC, VA): bogs, seepages, and boggy meadows; rare (NC Watch List, VA Rare). July-October. Newfoundland and Mackenzie west to British Columbia, south to w. NC, ne. TN, KS, and CA. [= RAB, C, F, G, GW, K, W, Z] Epilobium strictum Muhlenberg ex Sprengel, Northeastern Willow-herb, Downy Willow-herb, Soft Willow-herb. Pd (VA): moist places; rare (VA Rare). Québec west to MN, south to n. VA, OH, and n. IL. The single record is regarded as questionable. [= C, F, G, K, Z]

* Epilobium hirsutum Linnaeus, Hairy Willow-herb, introduced from Eurasia, ranges south to s. PA (Rhoads & Klein 1993) and WV (Kartesz 1999). [= C, F, G, K, Z] ONAGRACEAE 517 Epilobium palustre Linnaeus, Marsh Willow-herb, ranges south to DE and ne. PA (Rhoads & Klein 1993). [= C, K; > E. palustre var. palustre – F, G, Z]

Gaura Linnaeus (Gaura)

A genus of about 21 species, herbs, of North America. The flowers of all our species open about sunset and wither early the following morning. The genus is rather weedy; other western species may be expected to turn up in our area as adventive weeds. References: Raven & Gregory (1972)=Z; Munz (1965)=X.

1 Pedicels 2-4 mm long; fruit with a stipe at maturity; clumped or matted perennials from woody rhizomes or rootstocks; [of sandy habitats of SC and GA southward]. 2 Clumped perennial; petals 4-10 mm long; body of the fruit 5-10 mm long; stipe of the fruit 0.5-4.5 mm long ...... G. filipes 2 Mat-forming perennial; petals 7-15 mm long; body of the fruit 8-15 mm long; stipe of the fruit 2-8 mm long ...... G. sinuata 1 Pedicels 0-1 mm long; fruit without a stipe; annual, winter annual, or biennial; [collectively of various habitats and more widespread in our area. 3 Sepals 2-3.5 mm long; petals 1.5-3 mm long...... G. parviflora 3 Sepals 2.5-12 mm long; petals 2.5-9 mm long. 4 Sepals 2.5-8 mm long; leaves 0.1-1.3 cm wide, the widest rarely over 1 cm wide; flowers 3-4-merous (often mixed on a plant); fruits 3-4-angled (often mixed on a plant); [of the outer Coastal Plain of GA, NC, and SC]...... G. angustifolia 4 Sepals 8-13 mm long; leaves 0.3-2.5 cm wide, the larger nearly always > 1 cm wide; flowers 4-merous; fruits 4- angled; [primarily of the Mountains and Piedmont of NC, SC, and VA, extending to the Coastal Plain of GA and SC] ...... G. biennis

Gaura angustifolia Michaux, Southeastern Gaura. Cp (GA, NC, SC): open woodlands, sandy fields, roadsides, primarily in the outer Coastal Plain; common. May-September. E. NC (Dare County) south to s. FL, west to e. TX, endemic to the Coastal Plain. [= RAB, K, S, Z; > G. angustifolia var. angustifolia – X] Gaura biennis Linnaeus, Biennial Gaura, Northeastern Gaura. Mt (NC, SC, VA), Pd (GA, NC, VA), Cp (GA, SC): roadsides, woodlands, streambanks, disturbed areas; common. June-October. MA and NY west to WI, se. MN, and IA, south to sw. NC, c. GA (Jones & Coile 1988), sc. TN, and c. IL. [= RAB, K, S, W, Z; G. biennis var. biennis – C, F, G, X] Gaura filipes Spach, Threadstalk Gaura. Cp (GA, SC), Pd, Mt (GA): sandy fields, disturbed areas, and clearings; common. April-July. SC west to n. TN and s. IN, south to ne. FL and e. LA. [= RAB, C, G, K, W, Z; > G. filipes var. filipes – F, X; > G. filipes var. major Torrey & A. Gray – F, X; = G. michauxii Spach – S] * Gaura parviflora Douglas ex Lehmann, Small-flowered Gaura. Cp (GA, SC), Pd (GA): sandy fields, disturbed areas, and clearings; rare, introduced from further west. May-July. IN and IL west to WA, south to MS, and Mexico; apparently introduced eastward to MA, TN, GA, and SC. Kartesz's (1999) adoption of G. mollis as the name for this taxon has been rejected (Wagner & Hoch 2000, Brummitt 2001). [= RAB, F, G, S, Z; = G. mollis James – K; > G. parviflora var. parviflora – X; > G. parviflora var. lachnocarpa Weatherby – X] * Gaura sinuata Nuttall ex Seringe, Texas Gaura. Cp (GA, SC), Pd (GA): sandy fields, disturbed areas, and clearings; uncommon, introduced from further west. April-June. AR and OK south to s. TX, introduced eastward to SC and FL. [= RAB, K, X, Z]

Gaura drummondii (Spach) Torrey & A. Gray. Disjunct eastward in GA (Kartesz 1999). [= K] {not keyed at this time} Gaura longiflora Spach. East to MD, PA, KY, TN, and AL (Kartesz 1999). [= K; = G. biennis Linnaeus var. pitcheri Torrey & A. Gray – C, F, G, X; > G. filiformis Small – S; > G. longiflora – S] {not keyed at this time}

Ludwigia Linnaeus 1753 (Seedbox, Water-primrose, Water-purslane)

A genus of about 82 species, herbs and shrubs, cosmopolitan. References: Peng (1989)=Z; Munz (1965)=X; Nesom & Kartesz (2000)=Q; Zardini, Gu, and Raven (1991)=V; Peng (1984, 1986, 1988); Peng & Tobe (1987); Raven (1963); Munz (1938, 1944); Eyde (1977, 1978, 1981); Raven & Tai (1979); Duke (1955). Key based in part on GW, Z, and Q.

1 Leaves opposite; plants creeping (rooting at the nodes); [section Dantia]...... Key A 1 Leaves alternate; plants erect or ascending (not rooting at the nodes), or creeping (rooting at the nodes). 2 Stamens 8-14; sepals 4-7; petals 4-7; [of various habits, including annual and perennial herbs and shrubs, variously erect, ascending, creeping, or forming floating mats] ...... Key B 2 Stamens 4; sepals 4; petals 0-4; [perennial herbs, with erect ascending flowering stems]...... Key C

Key A – Ludwigia with opposite leaves ONAGRACEAE 518

1 Pedicels of flowers and fruits 5-35 mm long. 2 Petals 7-11 mm long; pedicels of capsules 15-35 mm long, longer than the leaves...... L. arcuata 2 Petals 4-5 mm long; pedicels of capsules 5-16 mm long, shorter than to equalling the leaves ...... L. brevipes 1 Pedicels of flowers and fruits 0-3 mm long. 3 Stems, leaves, capsules, and calyx densely hirsute; seeds dark reddish-brown, 0.3-0.4 mm long ...... L. spathulata 3 Stems, leaves, capsules, and calyx glabrous to sparsely puberulent; seeds tan, 0.4-0.8 mm long. 4 Petals 0; floral tubes and capsules with 4 longitudinal dark green bands; bractlets (borne at or near base of floral tube) absent or present, if present then 0-1 mm long ...... L. palustris 4 Petals 4; floral tubes and capsules lacking green banding; bractlets (borne at or near base of floral tube) present, 2- 4 mm long ...... L. repens

Key B – Ludwigia with alternate leaves, 8-14 stamens, 4-7sepals, and 4-7 petals

1 Sepals 4; stamens 8; seeds in 2-several vertical series in each locule, free of endocarp tissue. 2 Internodes of the stem conspicuously winged on the angles by 2 decurrent wings running down from each leaf base; petals 0.6-1.2 cm long; capsule 1.0-2.0 cm long, 4-angled or 4-winged; [section Myrtocarpus]...... L. decurrens 2 Internodes of the stem not winged on the angles (or very faintly so); petals 1.0-5.0 cm long; capsule (1.5-) 2-5 cm long, obtusely 4-angled; [section Macrocarpon]. 3 Petals (1.5-) 3-5 cm long; sepals ca. 10 mm wide at base ...... L. bonariensis 3 Petals 1-2 cm long; sepals 3-5 mm wide at base ...... L. octovalvis 1 Sepals 5 (-7); stamens 10 (-14); seeds in 1 vertical series in each locule, loosely embraced or embedded in endocarp tissue. 4 Stems erect; floral tube much longer than the pedicel; seeds loosely embraced by a corky, horseshoe-shaped segment of endocarp; [section Seminuda] ...... L. leptocarpa 4 Stems (at least the lower portions) decumbent, creeping, or floating in mats (the flowering stems more-or-less erect in L. grandiflora and L. hexapetala); floral tube much shorter than the pedicel; seeds embedded in the woody endocarp; [section Oligospermum]. 5 Flowering stems decumbent, floating, or creeping; stem and leaves glabrous or glabrescent; petals mostly 1-1.5 cm long; anthers 1-1.7 mm long ...... L. peploides var. glabrescens 5 Flowering stems more-or-less erect; stem and leaves sparsely to densely pubescent with long soft hairs; petals (1.2-) 1.6-3 cm long; anthers 2.5-3.5 mm long. 6 Sepals (6-) 8-11 (-14) mm long; primary leaves 5-8.5 cm long, 7-11 mm wide, usually linear-lanceolate, usually widest below the middle; petals (1.2-) 1.6-2.0 (-2.6) cm long; style 4.7-6.7 (-8.2) mm long; stems densely villous...... L. grandiflora ssp. grandiflora 6 Sepals (8-) 12-19 mm long; primary leaves 5.5-13 cm long, 9-18 mm wide, usually narrowly elliptic to oblanceolate, usually widest above the middle; petals (1.5-) 2.0-2.9 (-3) cm long; style (5.8-) 6-10 mm long; stems sparsely to densely villous (rarely glabrous) ...... L. grandiflora ssp. hexapetala

Key C – Ludwigia with alternate leaves, 4 stamens, 4 sepals, and 0-4 petals

1 Pedicels 2-15 mm long; capsules subglobose to spheric or cubic, about as long as wide, box-like, 4-angled, dehiscence by an apical pore (later sometimes also irregularly loculicidal); petals present, 4-15 mm long, persistent or caducous; roots fascicled, fusiform, tuberous; plants lacking basal, stoloniform shoots; [section Ludwigia]. 2 Leaves cuneate at base; pedicels 2-5 mm long; nectary discs at base of style flattish, inconspicuous; [widespread in our area, in a wide variety of habitats] ...... L. alternifolia 2 Leaves rounded or truncate at base; pedicels 4-15 mm long; nectary discs at base of style domed, prominent; [nearly restricted to the Coastal Plain, primarily of pinelands]. 3 Styles 6-10 mm long; plants glabrous, glabrescent, or pubescent with very short hairs; sepals strongly reflexed in fruit ...... L. virgata 3 Styles 1.5-3 mm long; plants glabrescent or pubescent with short to long, spreading to shaggy hairs; sepals strongly reflexed, spreading, or ascending in fruit. 4 Sepals narrowly deltoid, broadest at or near the base, 3-4× as long as wide, ascending or spreading in fruit; plants glabrescent to hirtellous with long spreading hairs...... L. hirtella 4 Sepals ovate, broadest near the middle, ca. 2× as long as wide, strongly reflexed in fruit; plants pubescent with relatively short, appressed to spreading hairs...... L. maritima 1 Pedicels 0-1 (-5) mm long; capsules subglobose, obconic, or obpyramidal, about as long as wide or longer than wide, circular to quadrangular in cross-section, dehiscence irregularly loculicidal; petals absent or present, if present (L. linearis, L. linifolia) then 0-6 mm long and caducous; roots fibrous or rhizomatous; plants frequently with basal, stoloniform shoots; [section Microcarpium]. 5 Capsules cylindrical, narrowly obconical, or narrowly obpyramidal, at least 2.5-5× as long as broad; petals present or absent. 6 Primary leaves of the flowering stems narrowly elliptical, 6-12 (-20) mm wide; petals absent...... L. glandulosa ONAGRACEAE 519 6 Primary leaves of the flowering stems linear, 1.5-5 mm wide; petals present. 7 Sepals (3.3-) 4-7 mm long; lateral and marginal veins obscure on lower leaf surface; seeds reddish brown; capsules cylindric, parallel-sided through most of their length, not grooved; anthers 0.5-1.1 mm long...... L. linifolia 7 Sepals 2.3-5 (-5.6) mm long; lateral and marginal veins distinct on lower leaf surface; seeds yellowish; capsules elongate obpyramidal, tapering through most or all of their length, with a shallow longitudinal groove on each face; anthers 1.1-2 mm long. 8 Sepals 2.3-4 mm long, acuminate, the surfaces densely and minutely papillose, the papillae 0.02-0.05 mm long and appressed; capsules 5-8.5 (-10) mm long, 2-4 (-5) mm in diameter; pedicels 0-0.4 mm long; seed surface cells elongate parallel to the seed length (as seen at 20× or more); anthers 1.1-1.6 mm long ...... L. linearis var. linearis 8 Sepals 3-5 (-5.6) mm long, elongate-acuminate to cuspidate, the surfaces densely minutely strigillose, the hairs 0.06-0.10 mm long and appressed to ascending; capsules 5-10 (-12) mm long, 3-5.5 mm in diameter; pedicels 0-3.5 (-5) mm long; seed surface cells elongate transverse to the seed length, or irregular (as seen at 20× or more); anthers (1.1-) 1.3-2 mm long...... L. linearis var. puberula 5 Capsules subglobose, obovoid, or broadly obpyramidal, 1-1.5 × as long as broad; petals absent. 9 Flowers in compact, headlike or elongate spikes, the inflorescence lacking well-developed leaves; stems rarely branched; rhizomes often present...... L. suffruticosa 9 Flowers axillary in the axils of well-developed leaves; stems usually much branched; rhizomes absent. 10 Plants densely pubescent throughout. 11 Sepal apex elongate-acuminate or subcuspidate, reflexed; pubescence of stems and leaves hirtellous (the hairs spreading); seed surface cells suborbicular (as seen at 20× or more); anthers 0.6-0.9 (-1.3 mm long; style 1-2 mm long...... L. pilosa 11 Sepal apex acuminate, ascending; pubescence of stems and leaves strigillose (the hairs appressed) or hirtellous (the hairs spreading); seed surface cells elongate; anthers 0.3-0.8 mm long; style 0.25-1 (-1.25) mm long. 12 Plants hirtellous; capsules oblong-obovoid; sepals greenish on the upper surface; bracteoles (1.5-) 2-4.3 mm long, borne at or near the base of the capsule; seed surface cells elongate transverse to the seed length; anthers 0.3-0.35 mm long; style 0.25-0.5 mm long ...... L. ravenii 12 Plants strigillose; capsules subglobose; sepals yellowish on the upper surface; bracteoles 0.5-1.5 mm long, usually borne on the short pedicel; seed surface cells in patches, some patches with cells elongate parallel to seed length, others with cells transverse to seed length, others with cells diagonal (rather resembling a badly laid-out parque floor); anthers 0.5-0.8 mm long; style 0.55-1 (- 1.25) mm long ...... L. sphaerocarpa 10 Plants glabrous or subglabrous throughout. 13 Primary leaves of the flowering stems 4-17 mm long, 1.5-10 mm wide, mostly obovate-spatulate and 1.5-3× as long as wide; capsules 1-1.5 (-2) mm long, containing 10-20 dark reddish-brown seeds; plants typically 1-4 dm tall ...... L. microcarpa 13 Primary leaves of the flowering stems (18-) 30-110 mm long, 2-10 (-20) mm wide, mostly elliptic, lanceolate, oblanceolate, or linear and 4-20× as long as wide; capsules 1.8- 2-7 mm long, containing 40- 500 light brown, yellowish, or tan seeds; plants typically 3-10 dm tall. 14 Capsules obpyramidal, the corners narrowly winged with wings 0.3-0.9 mm wide; bracteoles 1.5- 4.7 mm long. 15 Stems often distinctly ridged or winged; sepals creamy-white, nearly as long as the capsule; capsule wall bulging out longitudinally between the wings; seed surface cells elongate parallel to the seed length...... L. alata 15 Stems nearly smooth or slightly ridged; sepals greenish, about 1/2 as long as the capsule; capsule wall flat between the wings; seed surface cells suborbicular ...... L. lanceolata 14 Capsules oblong-ovoid or subglobose, the corners not winged; bracteoles either 0.5-1.5 mm or 3.5- 6.5 (-8) mm long. 16 Bracteoles 3.5-6.5 (-8) mm long; sepals green, the apex long-acuminate, reflexed; capsules oblong-obovoid; seed surface cells elongate parallel to the seed length; [known from our area only in the Piedmont of VA] ...... L. polycarpa 16 Bracteoles 0.5-1.5 mm long; sepals yellowish, the apex acuminate, ascending capsules subglobose; seed surface cells in patches, some patches with cells elongate parallel to seed length, others with cells transverse to seed length, others with cells diagonal (rather resembling a badly laid-out parque floor); [of the Coastal Plain of GA, NC, SC, and VA in our area]...... L. sphaerocarpa

Ludwigia alata Elliott, Winged Seedbox. Cp (GA, NC, SC, VA): interdune ponds, freshwater to slightly brackish (oligohaline) marshes; rare (NC Rare, VA Rare). June-September. Se. VA south to s. FL, west to se. LA; disjunct in Jamaica. This species is a hexaploid (n = 24). One third of the genome of L. alata is apparently derived from L. microcarpa or its ancestor (Peng 1988). [= RAB, C, F, G, K, Z; > L. alata – GW, in part only (also see L. lanceolata); > L. alata – S, in a narrower sense; > L. simulata Small – S] ONAGRACEAE 520 Ludwigia alternifolia Linnaeus, Alternate-leaf Seedbox. Mt, Pd, Cp (GA, NC, SC, VA): ditches, marshes, open wet places, disturbed wet places; common. May-October. MA west to s. Ontario, s. MI, IA, and KS, south to n. FL and e. TX. [= RAB, G, GW, K, S, W; > L. alternifolia var. alternifolia – C, F; > L. alternifolia var. pubescens E.J. Palmer & Steyermark – C, F] Ludwigia arcuata Walter. Cp (GA, SC): marshes or submerged in water of natural Coastal Plain ponds; rare. June- September. SC south to s. FL, west to panhandle FL and s. AL. [= RAB, GW, K; = Ludwigiantha arcuata (Walter) Small – S] * Ludwigia bonariensis (M. Micheli) Hara. Cp (NC, SC): freshwater tidal marshes and adjacent disturbed areas; rare, apparently introduced from tropical America. June-September. Locally abundant in disturbed edges of freshwater tidal marshes near Wilmington, NC, perhaps introduced on ship's ballast. Material from Wilmington apparently has larger flowers than material of L. bonariensis elsewhere; its source and appropriate taxonomic treatment uncertain and needing further study. First reported for SC by Leonard (1971b). [= RAB, GW, K; = Jussiaea neglecta Small – S] Ludwigia brevipes (B.H. Long ex Britton, A. Braun, & Small) Eames, Long Beach Seedbox, Coastal Plain Water-purslane. Cp (GA, NC, SC, VA): pondshores, blackwater rivers, interdunal swales, borrow ponds, ditches, impoundments, marshes; rare (NC Rare, VA Rare). July-October. NJ south to e. GA (Jones & Coile 1988), in the Coastal Plain. [= RAB, C, F, G, GW, K] Ludwigia decurrens Walter, Wingstem Water-primrose. Cp, Pd, Mt (GA, NC, SC, VA): swamp forests, ditches; common (rare in VA Mountains). June-October. MD, w. VA, WV, s. IN, s. IL, and MO, south to s. FL and TX; also in tropical America. [= RAB, C, GW, K, W; = Jussiaea decurrens (Walter) Augustin de Candolle – F, G, S] Ludwigia glandulosa Walter, Small-flowered Seedbox. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): low forests, marshes, ditches; common (VA Watch List). June-September. E. MD south to n. FL, west to e. TX, north in the interior to c. TN, w. KY, s. IN, s. IL, se. MO, c. AR, and se. OK, primarily on the Southeastern Coastal Plain. A related species, treated by Peng as L. glandulosa ssp. brachycarpa (Torrey & A. Gray) Peng, ranges from sw. LA north and west to s. OK and c. TX. This species is tetraploid (n = 16). [< L. glandulosa – RAB, C, F, G, GW, S, in a broader sense; = L. glandulosa ssp. glandulosa – K, Z] * Ludwigia grandiflora (Michaux) Greuter & Burdet ssp. grandiflora, Showy Water-primrose. Cp (GA, SC): ponds, lakes, sluggish waters of ditches or streams; rare. May-September. Se. SC south to FL, west to TX; disjunct in MO, Guatemala, and in s. South America. This taxon is hexaploid (n= 24). See Zardini, Gu, and Raven (1991) and Nesom & Kartesz (2000) for additional information. [= Q; < L. uruguayensis (Cambessedes) Hara – RAB, C, GW, K (also see L. hexapetala); = L. grandiflora (Michaux) Zardini, Gu, & Raven – V] * Ludwigia grandiflora (Michaux) Greuter & Burdet ssp. hexapetala (Hooker & Arnott) Nesom & Kartesz, Common Water- primrose. Cp (NC, SC), Pd (GA, NC, VA), Mt (VA): ponds, lakes, sluggish waters of ditches or streams; uncommon (but often locally abundant). May-September. NC south to FL, west to OK and TX; also in CA, Europe, South America, Mexico; also introduced farther north in North America. This taxon is decaploid (n = 40). See Zardini, Gu, and Raven (1991) and Nesom & Kartesz (2000) for additional information. [= Q; < L. uruguayensis (Cambessedes) Hara – RAB, C, GW, K, W, in part (also see L. grandiflora); < Jussiaea uruguayensis Cambessedes – F, G; Jussiaea michauxiana Fernald – F; = L. hexapetala (Hooker & Arnott) Zardini, Gu, & Raven – V] Ludwigia hirtella Rafinesque, Rafinesque's Seedbox. Cp (GA, NC, SC, VA), Pd (GA), Mt (GA, NC): savannas, rarely in mountain bogs; common (VA Rare). June-September. S. NJ south to panhandle FL, west to e. TX, north in the interior to KY, c. TN, AR, and se. OK. [= RAB, C, F, G, GW, K, S, W] Ludwigia lanceolata Elliott, Lanceleaf Seedbox. Cp (GA, NC, SC): interdune ponds, open wet areas; rare (NC Rare). August-September. Se. NC south to c. peninsular FL, west to panhandle FL. This species is tetraploid (n = 16). [= RAB, K, S, Z; < L. alata – GW, in part] Ludwigia leptocarpa (Nuttall) Hara, Water-willow. Cp (GA, NC, SC, VA), Pd (GA, VA), Mt (GA): riverbanks, marshes, and ditches; common, uncommon in VA (VA Watch List). June-September. VA south to c. peninsular FL, west to e. TX, north in the interior to se. MO and s. IL; and in tropical America. [= RAB, C, GW, K, W; = Jussiaea leptocarpa Nuttall – F, G, S] Ludwigia linearis Walter var. linearis, Eastern Narrowleaf Seedbox. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): savannas; common. June-September. Var. linearis ranges from s. NJ south to c. peninsular FL, west to se. LA, extending inland to the Cumberland Plateau of nc. AL and c. TN. Var. linearis is here interpreted to be equivalent to Peng's subglabrous morph. Peng (1989) declines to recognize infraspecific taxa in L. linearis, but his discussion makes clear that 2 distinctive entities are present, as characterized by orientation of seed surface cells and characters of leaves, bracteoles, pedicels, sepals, stigmas, and styles (see key). The orientation of seed surface cells, recognized as a distinctive character in other difficult species pairs (such as L. alata and L. lanceolata) is the most reliable character separating the 2 varieties. This species is diploid (n = 8). [< L. linearis – RAB, C, F, G, GW, K, S, W, Z] Ludwigia linearis Walter var. puberula Engelmann & A. Gray, Western Narrowleaf Seedbox. Cp (GA, NC, SC, VA), Pd (GA, SC): savannas, interdunal swales; uncommon. June-September. Var. puberula ranges primarily from c. AL west to c. AR, south to e. TX, with intergradational material extending as far north and east as n. FL and e. NC. Var. puberula is here interpreted to include Peng's intermediate morph, densely strigillose morph, and completely glabrous morph (Peng 1989). As pointed out by Peng (1989), the glabrous morph is exactly like the densely strigillose morph except for the absence of pubescence. They often grow together, have essentially the same distribution, and may differ only at a single allele. Peng's intermediate morph is heterogeneous; some likely being truly intermediate between (and possibly hybrid derivatives of) the two varieties here recognized, while others clearly belong to var. puberula (based on surface cell orientation and floral characteristics) and merely have an amount of pubescence intermediate between the densely strigillose and completely glabrous morphs. [< L. linearis – RAB, C, F, G, GW, K, S, W, Z] Ludwigia linifolia Poiret in Lamarck, Flaxleaf Seedbox. Cp (GA, NC, SC), Pd (NC): limesink ponds (dolines) and Taxodium ascendens savannas; rare (NC Rare). June-September. Nc. NC south to s. FL, west to s. MS; disjunct in Tabasco, Mexico. This species is diploid (n = 8). [= RAB, GW, K, S, Z] ONAGRACEAE 521 Ludwigia maritima Harper, Harper's Seedbox. Cp (GA, NC, SC): savannas; common. June-September. E. NC south to s. peninsular FL, west to e. LA. [= RAB, GW, K, S] Ludwigia microcarpa Michaux, Small-fruited Seedbox. Cp (GA, NC, SC), Pd (GA, NC), Mt (GA): in circumneutral or alkaline soils of moist places, over calcareous rock, mafic rock, shell hash, or brackish sands, such as in maritime wet grasslands, savannas and adjacent ditches over coquina limestone ("marl"), and wet clay flats over diabase, often in roadside ditches; uncommon. July-October. Ne. NC south to s. FL, west to se. TX (Brown & Marcus 1998); disjunct inland on calcareous or mafic rocks in nc. NC, n. GA, n. AL, c. TN, and sc. MO; also in the Bahamas, Cuba, and Jamaica. This species is diploid (n = 8). [= RAB, F, GW, K, S, W, Z] Ludwigia octovalvis (Jacquin) Raven. Cp (GA, NC, SC): marshes, disturbed areas; rare. May-September. Se. NC south to s. FL, west to TX; and widespread in tropical America. [= GW; > L. octovalvis spp. octovalvis – K; > L. octovalvis ssp. sessiliflora (M. Micheli) Raven – K; > Jussiaea angustifolia Lamarck – S; > Jussiaea scabra Willdenow – S] Ludwigia palustris (Linnaeus) Elliott, Common Water-purslane. Cp, Pd, Mt (GA, NC, SC, VA): moist to wet disturbed areas; common. May-October. Widespread in North America, Eurasia, and Africa. [= RAB, C, GW, K, W; > L. palustris var. americana (Augustin de Candolle) Fernald & Griscom – F, G; > L. palustris var. nana Fernald & Griscom – F; = Isnardia palustris Linnaeus – S] *? Ludwigia peploides (Humboldt, Bompland, & Kunth) Raven var. glabrescens (Kuntze) Shinners. Cp (GA, NC, VA), Mt (VA), Pd (GA, SC, VA): pools, ditches, disturbed places; rare. May-September. PA, VA and NC south and west to FL and AZ, widespread in the West Indies, Central and South America. Doubtfully native in all or part of our area. [= RAB, C; L. peploides ssp. peploides – GW; Jussiaea diffusa Forskl. – S; Jussiaea grandiflora Michaux – S; = Jussiaea repens Linnaeus var. glabrescens Kuntze – F, misapplied; Jussiaea repens – G, misapplied; = L. peploides ssp. glabrescens (Kuntze) Raven – K; < L. peploides – W, infraspecific taxa not distinguished] Ludwigia pilosa Walter, Hairy Seedbox. Cp (GA, NC, SC, VA), Pd (NC, VA): ditches, wet places; common, rare in VA (VA Rare). June-October. Se. VA south to n. FL, west to se. TX, restricted to the Coastal Plain except for disjunct occurrences inland in NC, VA, and n. AL. This species is tetraploid (n= 16). [= K, W, Z; < L. pilosa – RAB, C, F, G, GW, S (also see L. ravenii)] Ludwigia polycarpa Short & Peter. Pd (VA): {habitat}; rare (VA Watch List). June-September; July-October. MA, CT, and w. VT west to s. Ontario, MI, WI, MN, and c. NE, south to c. VA, KY, s. IL, s. MO, and e. KS. This species is tetraploid (n = 16). [= C, F, G, GW, K, S, Z] Ludwigia ravenii Peng, Raven's Seedbox. Cp (NC, SC, VA): savannas, swamps, marshes, wet open places; rare (NC Watch List, VA Rare). June-October. Se. VA south to ne. FL (no known records for GA), restricted to the Coastal Plain. For further information, see Peng (1984, 1988, 1989). This species is tetraploid (n = 16). [= K, Z; < L. pilosa – RAB, C, F, G, GW, S (included within concept of L. pilosa by most earlier authors)] Ludwigia repens Forster, Creeping Seedbox. Cp (GA, NC, SC, VA), Pd (GA): ditches, pools, and streams; uncommon (VA Rare). June-September. Se. VA south to s. FL, west to TX and n. Mexico, north in the interior to TN, MO, and OK; also in CA, Bermuda, and the West Indies. Reveal et al. (2003) propose the name L. repens for nomenclatural conservation with a conserved type; if this proposal is not accepted, L. natans Elliott will become the name of this species. [= RAB, GW, K; Ludwigia natans Elliott – F, G; Isnardia repens – S] Ludwigia spathulata Torrey & A. Gray, Southern Water-purslane. Cp (GA, SC), Pd (GA): sinkhole ponds, cypress-gum ponds, depression meadows, boggy shores; rare (GA Special Concern, NC Rare, SC Rare). June-October. SC south to panhandle FL and s. AL. [= RAB, GW, K; Isnardia spathulata (Torrey & A. Gray) Small – S] Ludwigia sphaerocarpa Elliott, Globe-fruited Seedbox. Cp (GA, NC, SC, VA): boggy areas, pools, ditches, river marshes, interdune swales, river and pondshores; rare (NC Watch List, VA Rare). June-September. E. MA south to n. FL, west to e. TX, primarily on the Coastal Plain, spottily distributed in that range, and also disjunct in w. NY, sc. TN, s. IN, and nw. IN and ne. IL. This species is tetraploid (n = 16). Peng (1989) considers it likely that L. sphaerocarpa is of allopolyploid origin, one or both of its parents now extinct. [= RAB, C, GW, K, S, Z; > L. sphaerocarpa var. sphaerocarpa – F, G; > L. sphaerocarpa var. jungens Fernald & Griscom – F, G] Ludwigia suffruticosa Walter, Shrubby Seedbox. Cp (GA, NC, SC): periodically to seasonally flooded portions of limesink ponds (dolines) and clay-based Carolina bays; rare north of SC (NC Rare). June-October. Se. NC south to s. peninsular FL, west to panhandle FL and se AL. This species is tetraploid (n = 16). Peng (1989) reports that "with its whitish creamy sepals, which are very showy in the dense flower aggregates, the cross-pollinating L. suffruticosa successfully attracts many insects, mostly bumblebees, honeybees, and wasps." [= RAB, GW, K, S, Z; = L. capitata Michaux] Ludwigia virgata Michaux, Savanna Seedbox. Cp (GA, NC, SC, VA): wet savannas; common (VA Rare). June- September. Se. VA south to s. peninsular FL, west to panhandle FL and se. AL. [= RAB, C, F, GW, K, S]

Ludwigia peruviana (Linnaeus) Hara, in s. GA (Jones & Coile 1988). Reported for NC (Kartesz 1999). {investigate} [= K; Jussiaea peruviana Linnaeus]

The following natural hybrids are known, not necessarily in our area. Hybrids are generally recognizable from their intermediate morphology and usual association with their two parents. However some hybrids resemble one parent much more than the other, and some hybrids are found in populations independent (and even disjunct) from one or both parents. Allopolyploidy may have had a major role in the evolution of this genus, especially section Microcarpium, which has a majority of polyploid species.

L. alata × pilosa. Pentaploid, sterile. L. alata × suffruticosa. Pentaploid, sterile. ONAGRACEAE 522 L. arcuata × pilosa. L. glandulosa × linearis. Triploid, sterile. L. glandulosa × palustris. Triploid, sterile. L. glandulosa × pilosa. Tetraploid, fertile. L. glandulosa × sphaerocarpa. Tetraploid, fertile. L. lanceolata × pilosa [= L. ×simulata Small (pro sp.)]. Tetraploid, fertile. L. lanceolata × suffruticosa. Tetraploid, fertile. Frequent south of our area. L. linearis × sphaerocarpa. Triploid, sterile. L. microcarpa × palustris. L. pilosa × ravenii. Tetraploid, fertile. L. pilosa × sphaerocarpa. Tetraploid, fertile. Frequent. L. pilosa × suffruticosa. Tetraploid, fertile. L. polycarpa × sphaerocarpa. Tetraploid, fertile.

Oenothera Linnaeus 1753 (Evening-primrose) (also see Calylophus)

A genus of about 124 species, herbs, of America (especially temperate regions). This treatment provisional, with further revision likely, especially in the Oe. fruticosa-Oe. tetragona-Oe. pilosella complex. References: Dietrich, Wagner, & Raven (1997)=Z; Dietrich & Wagner (1988)=Y; Munz (1965)=X; Straley (1977)=V. Keys adapted in part from those references.

1 Ovary essentially terete; fruit terete or with 4 rounded ridges; stamens equal in length (except in Oe. speciosa). 2 Flowers white or pink; flower buds nodding; [section Hartmannia] ...... Oe. speciosa 2 Flowers yellow; flower buds erect; [section Oenothera] 3 Fruit linear, nearly isodiametric through its length; seeds borne ascending in the locules, rounded or fusiform, more or less regularly pitted; [section Oenothera, subsection Raimannia]. 4 Petals acute to rounded at the apex. 5 Inflorescence dense, with > 2 flowers per spike opening each day; leaves gray-green ...... [Oe. clelandii] 5 Inflorescence lax, 1-2 flowers per spike opening on each day; leaves green...... Oe. curtissii 4 Petals truncate to emarginate at the apex. 6 Nonflowering portion of stems stiff, densely strigillose or sometimes also villous; leaves gray-green, densely strigillose, usually subentire to shallowly dentate (rarely lyrate); [in maritime situations]. 7 Sepals 2.0-3.3 cm long; petals 2.5-4.5 cm long; stigma elevated above the anthers at anthesis; capsule 2.5-5.5 cm long; rosette leaves 5-14 cm long, 1-2 cm wide...... Oe. drummondii ssp. drummondii 7 Sepals 0.3-1.1 cm long; petals 0.45-1.6 cm long; stigma surrounded by the anthers at anthesis; capsule 1.5-4.5 cm long; rosette leaves 4-8 cm long, 0.7-1.0 cm wide...... Oe. humifusa 6 Nonflowering portion of stem not stiff, moderately to sparsely strigillose to sometimes densely villous, and also more orless glandular puberulent; leaves green, sparsely to moderately strigillose and usually villous, deeply lobed to dentate (rarely some of them subentire); [in inland disturbed situations]. 8 Petals 2.5-4 cm long; style 4-7.5 cm long; stigma lobes well elevated above the anthers at anthesis...... Oe. grandis 8 Petals 0.5-2.2 cm long; style 2-5 cm long; stigma lobes surrounded by the anthers at anthesis ...... Oe. laciniata 3 Fruit thickest near the base, tapering to the apex; seeds borne horizontally in the locules, angled-prismatic, not regularly pitted; [section Oenothera, subsection Oenothera]. 9 Stigma elevated above the anthers at anthesis; petals 2.5-5 cm long. 10 Cauline leaves 0.4-1.0 cm wide; apex of the inflorescence curved; free sepal tips subterminal, usually spreading; capsules spreading at nearly right angles to the stem, long-attenuate toward apex, usually conspicuously arcuate...... Oe. argillicola 10 Cauline leaves 1.5-6 cm wide; apex of the inflorescence erect; free sepal tips terminal, erect; capsules erect or slightly spreading, gradually attenuate toward the apex. 11 Upper stem, ovary, floral tube, and sepals always conspicuously pubescent, usually with at least some red-pustulate hairs; bracts green, persistent; sepals often flushed with red, or red-striped...... Oe. glazioviana 11 Upper stem, ovary, floral tube, and sepals often apparently glabrous without magnification; pustulate hairs absent, or if present not red (in fresh material); bracts often pale-green and deciduous; sepals yellowish green, or flushed with some red...... Oe. grandiflora 9 Stigma surrounded by or below the anthers at anthesis; petals 0.7-2.5 (-3) cm long. 12 Plant appearing exclusively appressed-pubescent (as seen without magnification). 13 Apex of the inflorescence curved; free sepal tips subterminal in bud, erect to spreadin; dry capsules usually rusty brown ...... Oe. oakesiana ONAGRACEAE 523 13 Apex of the inflorescence erect; free sepal tips erect in bud; dry capsules gray-green or dull green. 14 Leaves green to pale green; stems, ovary, floral tube, and sepals sparsely appressed-pubescent ...... Oe. biennis 14 Leaves dull green to gray-green; stems, ovary, floral tube, and sepals densely appressed- pubescent...... Oe. villosa ssp. villosa 12 Plant appearing either glabrous or with a mixture of long pustular hairs and appressed pubescence (as seen without magnification). 15 Apex of inflorescence curved; free sepal tips subterminal in bud. 16 Plant (at least the lower portions) predominantly strigillose; leaves dull green to gray-green; dry capsules rusty brown ...... Oe. oakesiana 16 Plant predominantly erect-pubescent or appearing glabrous (as seen without magnification); leaves usually bright green; dry capsules usually dark green or black...... Oe. parviflora 15 Apex of inflorescence erect; free sepal tips terminal or subterminal in bud. 17 Inflorescence conspicuously pubescent ...... Oe. biennis 17 Inflorescence glabrous (or appearing so without magnification). 18 Free sepal tips terminal in bud; petals 1.4-2.5 (-3) cm long; bracts caducous, pale green; capsules dull green when dry; petals fading yellowish-white to translucent...... Oe. nutans 18 Free sepal tips subterminal in bud; petals 0.8-1.5 (-2) cm long; bracts persistent, green; capsules usually black or dark green when dry; petals fading pale yellow, usually opaque ...... Oe. parviflora 1 Ovary 4-angled or 4-winged (at least near its tip); fruit sharply 4-angled or 4-winged; stamens of two lengths (except Oe. triloba and Oe. macrocarpa ssp. macrocarpa). 19 Leaves all basal, pinnatifid; [section Lavauxia] ...... Oe. triloba 19 Leaves in part cauline, entire or toothed. 20 Petals 50-70 mm long; flowers opening in the evening; wings of the fruit 10-25 mm wide; [section Megapterium]...... [Oe. macrocarpa ssp. macrocarpa] 20 Petals 3-30 mm long; flowers opening in the day; wings of the fruit <3 mm wide; [section Kneiffia] 21 Cauline leaves linear, < 1 mm wide; petals 3-5 (-7) mm long; floral bracts shorter than the subtended ovaries; mature fruits ellipsoid-rhomboid, 4-6 mm long; annual; [section Kneiffia, subsection Peniophyllum]...... Oe. linifolia 21 Cauline leaves lanceolate to ovate, > 1 mm wide; petals 5-30 mm long; floral bracts longer than the subtended ovaries; mature fruits clavate to oblong-elliptic, 8-20 mm long; perennial; [section Kneiffia, subsection Kneiffia]. 22 Petals 5-10 mm long; inflorescence usually nodding ...... Oe. perennis 22 Petals 15-30 mm long; inflorescence usually erect. 23 Plant conspicuously pilose-hirsute with hairs 1-3 mm long; free sepal tips 1-4 mm long, divergent ...... Oe. pilosella 23 Plant either with shorter or appressed pubescence, of glandular or nonglandular hairs; free sepal tips 0-2 (-6) mm long, divergent or not. 24 Capsules oblong, widest near the middle, usually abruptly tapered to a stipe 0.1-3 (-7) mm long; hairs of the ovary and capsule predominantly glandular (or the ovary glabrous); leaves subglabrous or sparsely pubescent, more or less dentate. 25 Petals (20-) 25-35 mm long; cauline leaves lanceolate to ovate, 2-7 cm long, 1-3 cm wide, often glaucous beneath...... Oe. tetragona var. fraseri 25 Petals 12-20 (-25) mm long; cauline leaves linear to lanceolate, 2-7 cm long, 0.5-1.0 (- 1.5) cm wide ...... Oe. tetragona var. tetragona 24 Capsules clavate, widest above the middle, gradually tapered to a stipe 3-10 mm long; hairs of the ovary and capsule nonglandular (or with a mixture of glandular and nonglandular hairs); leaves generally pubescent, subentire. 26 Petals 15-30 mm long; stems 7-12 dm tall, freely branched, slightly pubescent; cauline leaves lanceolate, 5-12 cm long, 0.5-1.5 cm wide; [of tidal marshes, usually with spongy lower stems and adventitious roots where regularly submerged] ...... Oe. riparia 26 Petals (8-) 15-22 mm long; stems 1-8 dm tall, less branched (unless mowed, grazed, or otherwise damaged), more pubescent; cauline leaves 2-6 (-8) cm long, 0.2-1.0 (-1.2) cm wide. 27 Capsule vestiture a mixture of glandular and nonglandular hairs. 28 Cauline leaves not velutinous, 5-10× as long as wide...... Oe. tetragona var. brevistipata 28 Cauline leaves velutinous, 2-4× as long as wide. 29 Petals 7-12 mm long; leaves lance-oblong, obtuse; [of barrens of TN, KY, and AL]...... [Oe. tetragona var. sharpii] 29 Petals 15-20 mm long; leaves lanceolate, acute; [of the Atlantic Coastal Plain]...... Oe. tetragona var. velutina 27 Capsule vestiture strictly nonglandular. ONAGRACEAE 524 30 Free sepal tips 1-3 mm long, often arching; calyx strigose...... Oe. fruticosa var. unguiculata 30 Free sepal tips < 1 mm long; calyx various. 31 Capsule body 6-11 mm long, the pubescence rather coarse...... Oe. fruticosa var. fruticosa 31 Capsule body 3-5 mm long, the pubescence very fine. 32 Capsule body 3.5-4 mm long, strigose-pilose; [of Coastal Plain bogs] ...... Oe. fruticosa var. microcarpa 32 Capsule body 4-5 mm long; very finely strigillose; [of Piedmont rock outcrops] ...... Oe. fruticosa var. subglobosa

Oenothera argillicola Mackenzie, Shale-barren Evening-primrose. Mt (VA): shale barrens and woodlands; uncommon. Sc. PA south through MD to e. WN and w. VA (south to Montgomery County). [= C, F, G, H, K, W, Z; > Oe. argillicola var. argillicola – X; > Oe. argillicola var. pubescens Core & Davis – X] Oenothera biennis Linnaeus, Common Evening-primrose. Mt, Pd, Cp (GA, NC, SC, VA): fields, pastures, roadsides, disturbed areas; common. June-October. Ranging widely in e. North America and Europe, and scattered in w. North America. [= H, K, W, Z; < Oe. biennis – RAB, G, S (also see Oe. nutans); = Oe. biennis var. biennis – C, F; > Oe. biennis var. biennis – F, in a narrower sense; > Oe. biennis var. pycnocarpa (Atkinson & Bartlett) Wiegand – F; > Oe. biennis ssp. caeciarum Munz – X; > Oe. biennis ssp. centralis Munz – X] Oenothera curtissii Small. Cp (GA, SC): sandhills, sandy fields; rare. May-September. Se. SC south to n. peninsular FL, west to s. AL. Closely related to Oe. rhombipetala, which is restricted to the Great Plains, with scattered occurrences east to AR, IL, and MI. [= K, Y; < Oenothera rhombipetala Nuttall ex Torrey & A. Gray – RAB, F, X, misapplied; = Raimannia curtissii Rose – S] *? Oenothera drummondii Hooker ssp. drummondii, Drummond's Evening-primrose. Cp (NC, SC): sandy ocean beaches; rare, perhaps only introduced or adventive from the Gulf Coast. April-October. Ssp. drummondii ranges from se. NC south to s. FL, west to se. TX, and south to Tamaulipas and Vera Cruz. Ssp. thalassiphila (Brandegee) W. Dietrich & W.L. Wagner is restricted to the southern tip of Baja California. [= Y; < Oe. drummondii – RAB, K; < Raimannia drummondii (Hooker) Rose ex Sprague & Riley – S; = Oe. drummondii var. drummondii – X] Oenothera fruticosa Linnaeus var. fruticosa, Southern Sundrops. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, VA): dry forests and woodlands, glades, and rock outcrops; common. April-August. MA west to IN, south to FL and LA. [= F, G, X; < Oe. fruticosa – RAB, C; < Oe. fruticosa ssp. fruticosa – H, K, V, W; > Oe. fruticosa var. linearis (Michaux) S. Watson – F; > Oe. fruticosa var. humifusa Allen – F, G, X; > Kneiffia fruticosa (Linnaeus) Raimann – S; > Kneiffia arenicola Small – S; > Kneiffia semiglandulosa Pennell – S] Oenothera fruticosa Linnaeus var. microcarpa Fernald, Small-fruited Sundrops. Cp (NC, SC, VA): boggy depressions. April-August. E. MD south to e. SC. [= F, X; < Oe. fruticosa – RAB, C; < Oe. fruticosa ssp. fruticosa – H, K, V] Oenothera fruticosa Linnaeus var. subglobosa (Small) Munz, Flatrock Sundrops. Pd (GA): granite flatrocks and domes; rare. GA to AL. [= X; < Oe. fruticosa – RAB, C; < Oe. fruticosa ssp. fruticosa – H, K, V; = Kneiffia subglobosa Small – S] Oenothera fruticosa Linnaeus var. unguiculata Fernald, Southern Sundrops. Cp (NC, SC, VA): sandhills, moist to wet loamy savannas; uncommon. April-August. Se. VA south to e. SC. [= F, X; < Oe. fruticosa – RAB, C; < Oe. fruticosa ssp. fruticosa – H, K, V] * Oenothera glazioviana Micheli in Martius, Garden Evening-primrose. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas; uncommon. This species apparently arose as a garden hybrid, and has been widely cultivated and naturalized nearly worldwide. [= K, Z; = Oe. erythrosepala Borbás – X] Oenothera grandiflora L'Héritier ex Aiton. Cp, Pd, Mt (GA?, NC, SC): disturbed areas; uncommon. June-October. VT west to KY, south to c. peninsular FL and s. MS. [= F, K, X, S, Z] * Oenothera grandis (Britton) Smyth. Cp (NC): roadsides; rare, introduced from further west. March-July. The native range of this species is centered in KS, OK, and TX. [= K, X, Y; = Oe. laciniata Hill var. grandiflora (S. Watson) B.L. Robinson – RAB, F, G] Oenothera humifusa Nuttall, Seabeach Evening-primrose, Spreading Evening-primrose. Cp (GA, NC, SC, VA): coastal sand dunes; common. Early May-October. S. NJ south to s. FL, west to s. LA, along the coast. [= RAB, C, F, G, H, K, X, Y; = Raimannia humifusa (Nuttall) Rose – S] Oenothera laciniata Hill, Cutleaf Evening-primrose. Cp, Pd, Mt (GA, NC, SC, VA): disturbed areas; common. February- October. ME west to ND, south to s. FL and TX; also in CA. [= K, W; = Oe. laciniata var. laciniata – RAB, C, F, G; = Raimannia laciniata (Hill) Rose – S; = Oe. laciniata ssp. laciniata – X] Oenothera linifolia Nuttall, Threadleaf Sundrops, Flaxleaf Sundrops. Pd (GA, NC, SC, VA*), Cp (GA, SC), Mt (GA): dry openings and fields; rare (SC Rare, VA Watch List). C. VA west to s. IL and se. KS, south to panhandle FL and se. TX. Occurrences east of the Mississippi River may be mainly or entirely adventive. Belden et al. (2004) discuss the Virginia occurrence. [= RAB, C, F, G, K, W, V, X; = Peniophyllum linifolium (Nuttall) Pennell – S] Oenothera nutans Atkinson & Bartlett. Mt, Pd (GA?, NC, SC?, VA): roadsides, openings, forest edges, pastures; common (NC Watch List, VA Watch List). July-October. ME west to MI, south to n. FL, s. AL, and s. MO. [= K, Z; < Oe. biennis – RAB, G, S; = Oe. biennis Linnaeus var. austromontana (Munz) Cronquist – C; = Oe. biennis var. nutans (Atkinson & Bartlett) Wiegand – F; = Oe. austromontana (Munz) Raven, Dietrich, & Stubbe – H, W; = Oe. biennis ssp. austromontana Munz – X] Oenothera oakesiana (A. Gray) Robbins ex S. Watson & Coulter. Cp (NC, VA), Pd, Mt (VA): disturbed areas, roadsides; uncommon (VA Watch List). Nova Scotia west to Manitoba, south to e. NC, sc. VA, PA, n. IN, n. IL, and s. MN. [= G, K, Z; = ONAGRACEAE 525 Oe. parviflora Linnaeus var. oakesiana (A. Gray) Fernald – C, F; = Oe. parviflora ssp. parviflora var. oakesiana (A. Gray) Fernald – X] Oenothera parviflora Linnaeus, Small-flowered Evening-primrose. Mt, Pd (NC, VA), Cp (NC, SC, VA): fields, disturbed areas; uncommon. May-September. Nova Scotia west to Manitoba, south to NC, TN, KY, and MO. Reported for GA (GANHP). [= RAB, G, K, W, Z; > Oe. parviflora var. parviflora – C, F; > Oe. parviflora var. angustissima (R.R. Gates) Wiegand – F; > Oe. parviflora ssp. parviflora var. parviflora – X; > Oe. parviflora ssp. angustissima (R.R. Gates) Munz – X] Oenothera perennis Linnaeus, Little Sundrops. Mt (NC, SC, VA), Pd (NC, VA), Cp (VA): bogs, sphagnous seeps; uncommon (rare south of VA) (NC Rare, SC Rare). May-August. Nova Scotia west to Manitoba, south to w. NC, nw. SC, KY, and MO. [= RAB, C, G, K, W, X; > Oe. perennis var. perennis – F; = Kneiffia perennis (Linnaeus) Pennell – S] Oenothera pilosella Rafinesque, Midwestern Evening-primrose. Mt, Pd, Cp (VA): disturbed areas; rare. NH west to Ontario, south to s. VA, KY, n. AL, c. MS, and c. LA. Oe. sessilis (Pennell) Munz, treated by Straley (1977) as Oe. pilosella ssp. sessilis (Pennell) Straley, seems sufficiently distinct to be recognized as a species; it is restricted to Mississippi River Alluvial Plain. [= F, G, X; = Oe. pilosella ssp. pilosella – C, K, V; > Kneiffia pratensis Small – S; = Oe. fruticosa Linnaeus var. hirsuta Nuttall ex Torrey & A. Gray] Oenothera riparia Nuttall, Riverbank Evening-primrose. Cp (NC, SC): tidal marshes; rare (NC Rare). June-July. Se. VA (?) south to se. NC and e. SC. Distinct from Oe. fruticosa. Present in the freshwater tidal portions of the Waccamaw, Northeast Cape Fear, Black, Greater Pee Dee, and Cape Fear (?) rivers. [< Oe. fruticosa – RAB; < Oe. fruticosa ssp. fruticosa – K, V; = Kneiffia riparia (Nuttall) Small – S; = Oe. tetragona Roth ssp. glauca (Michaux) Munz var. riparia (Nuttall) Munz – X] * Oenothera speciosa Nuttall, White Evening-primrose. Cp, Pd, Mt (GA, NC, SC, VA): roadsides and fields, also cultivated as an ornamental; common (rare in Mountains), introduced from further west. May-August. [= RAB, C, F, G, K, W, X; = Hartmannia speciosa (Nuttall) Small – S] Oenothera tetragona Roth var. brevistipata (Pennell) Munz. Mt, Pd (GA, NC, SC, VA), Cp (VA): dry forests and woodlands, roadsides; common. May-August. SC and KY, south to GA and MS. Should perhaps be considered more closely related to Oe. fruticosa (where placed in synonymy by Straley), if it is determined to be valid. [= G; < Oe. tetragona – RAB, C; < Oe. fruticosa Linnaeus ssp. fruticosa – H, K, V, W; = Kneiffia brevistipata Pennell – S; = Oe. tetragona ssp. tetragona var. brevistipata – X] Oenothera tetragona Roth var. fraseri (Pursh) Munz, Appalachian Sundrops. Mt (GA, NC, SC, VA), Pd (NC, SC, VA), Cp (VA): dry to moist forests and woodlands, roadsides; common. May-August. NY and PA, south to nw. SC and n. GA. This is the more montane and high elevation variant of Oe. tetragona. [= F, G, X; < Oe. tetragona – RAB, C; > Oe. tetragona var. hybrida (Michaux) Fernald – F; > Oe. tetragona var. latifolia (Rydberg) Fernald – F; < Oe. fruticosa Linnaeus ssp. glauca (Michaux) Straley – H, K, V, W; > Kneiffia glauca (Michaux) Spach – S; > Kneiffia hybrida (Michaux) Small – S; > Kneiffia latifolia Rydberg – S; = Oe. tetragona ssp. glauca var. glauca – X] Oenothera tetragona Roth var. tetragona, Northern Sundrops. Mt, Pd (NC, SC, VA), Cp (VA): dry forests and woodlands, roadsides; common. May-August. Newfoundland west to MI, south to e. VA and MO. [= F; < Oe. tetragona – RAB, C; < Oe. tetragona var. tetragona – G; < Oe. fruticosa Linnaeus ssp. glauca (Michaux) Straley – H, K, V, W; = Kneiffia tetragona (Roth) Pennell – S; = Oe. tetragona ssp. tetragona var. tetragona – X] Oenothera tetragona Roth var. velutina (Pennell) Munz. Cp (VA): dry sandy soils. Se. NY (Long Island) south to se. VA. Should perhaps be considered more closely related to Oe. fruticosa (where placed in synonymy by Straley), if it is determined to be valid. [= F, G; < Oe. tetragona – RAB, C; < Oe. fruticosa Linnaeus ssp. fruticosa – H, K, V, W; = Kneiffia velutina Pennell – S; = Oe. tetragona ssp. tetragona var. velutina – X] Oenothera triloba Nuttall, Stemless Evening-primrose. Mt (GA, VA): limestone glades, {habitat not known in VA}; rare, perhaps only introduced, though native into eastern KY and TN (GA Watch List). [= C, F, G, H, K, X; = Lavauxia triloba (Nuttall) Spach – S] * Oenothera villosa Thunberg ssp. villosa. Mt, Pd (VA): disturbed areas; uncommon, apparently naturalized in our area from an original distribution in the Great Plains. [= K, Z; ? Oe. strigosa (Rydberg) Mackenzie & Bush – G; ? Oe. biennis var. canescens Torrey & A. Gray – C, F; ? Oe. strigosa (Rydberg) Mackenzie & Bush ssp. canovirens (Steele) Munz – X]

* Oenothera clelandii W. Dietrich, Raven, & W.L. Wagner. Reported for SC (Kartesz 1999). {investigate} Centered in IL and WI, ranging east, probably mostly as introductions, to NJ, WV, KY. [= C, K, Y; < Oe. rhombipetala, misapplied] Oenothera macrocarpa Nuttall ssp. macrocarpa, Wingfruit Evening-primrose, occurs as a disjunct in c. TN. [= K; < Oe. missouriensis Sims – F; < Oe. macrocarpa Nuttall – C, G; = Oe. missouriensis Sims var. missouriensis – X; ? Megapterium missouriense (Sims) Spach] Oenothera tetragona Roth var. sharpii Munz. Known from the Eastern Highland Rim of TN, AL, and KY. [< Oe. tetragona – RAB, C; < Oe. tetragona var. tetragona – G; < Oe. fruticosa Linnaeus ssp. fruticosa – H, K, V, W; = Oe. tetragona ssp. tetragona var. sharpii – X]

Many hybrids are known.

OROBANCHACEAE Ventenat 1799 (Broomrape Family)

A family of about 96 genera and 2060 species, root-parasitic herbs lacking chlorophyll (Orobanchaceae sensu stricto) and chlorophyllose hemi-parasites (formerly placed in the Scrophulariaceae), of temperate and subtropical regions of the Northern Hemisphere (Manen et al. 2004). References: Thieret (1971); Olmstead et al. (2001); Fischer in Kadereit (2004).