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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 mergedinto 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 Plant a caulescent herbaceous vine (acaulescent in its first year, and appearing to be an herb); ultimate leaf segments 5-

10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adlumia2 Plant an acaulescent herb with basal leaves; ultimate leaf segments 1-4 mm wide.

3 Leaves basal only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicentra3 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] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fumaria4 Ovary and fruit elongate, with several to many seeds; [tribe Corydaleae].

5 Flowers pink, the petals tipped with yellow; plant biennial; stem erect, 3-8 (-10) dm tall; capsules erect, 25-35 mmlong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capnoides

5 Flowers yellow; plant 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, & Poggenberg, 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)

The genus is monotypic, an herb, of n. North America. Recent studies have emphasized its distinction from Corydalis, and itscloser 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; plant biennial; stem erect, 3-8 (-10) dm tall; capsules erect, 25-35 mm long . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Capnoides]

1 Flowers yellow; plant annual; stem erect, decumbent, or prostrate, 1-3 (-4) dm tall; capsules erect, ascending, divergent, orpendent, 10-20 (-25) mm long.2 Fruits pendent or divergent; spurred petal 7-9 mm long; seeds 2-2.5 mm wide, with a narrow, acute ring-margin . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. flavula2 Fruits erect or ascending; spurred petal 10-15 mm long; seeds 1.0-2.0 mm wide, without a narrow, acute ring-margin.

3 Capsules ca. 1.0 mm in diameter, strongly constricted between the seeds at maturity; inflorescence long, usually farexceeding the poorly-developed upper leaves; ultimate leaf segments 0.5-1.5 (-3.0) mm wide; seeds less than 1.5mm wide; plant slightly to strongly glaucous; [plants of sandy soils of the outer Coastal Plain] . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. micrantha var. australis3 Capsules 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; seedsmore than 1.5 mm wide; plant green to slightly glaucous; [plants of circumneutral rock outcrops of the upperPiedmont and Mountains] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. micrantha var. micrantha

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Corydalis flavula (Rafinesque) Augustin de Candolle, Short-spurred Corydalis. Pd, Mt, Cp (GA, NC, SC, VA): rich moistforests, especially alluvial forests, glades and outcrops over mafic rocks (such as greenstone); common (rare in NC and SCCoastal 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 micrantha (Engelmann ex A. Gray) A. Gray var. australis (Chapman) Shinners, Southern Corydalis. Cp (GA,NC, SC): sandy roadsides and disturbed areas; uncommon. March-April; May-June. Ssp. australis ranges from e. NC south toFL, west to TX, and inland north to MO and OK. F and S recognized it as a species distinct from C. micrantha; Ownbey reduced itto a subspecies, citing inadequate morphological differences and some alleged intermediates in OK and MO. In our area, the 2taxa appear readily separable on morphological, ecological, and geographical grounds; species status may be warranted. [= C;Corydalis micrantha (Engelmann ex A. Gray) A. Gray ssp. australis (Chapman) G.B. Ownbey -- RAB, FNA, K, Z; Corydalis halei(Small) Fernald & Schubert -- F; Corydalis micrantha -- G, infraspecific taxa not distinguished; Capnoides halei Small -- S]

Corydalis micrantha (Engelmann ex A. Gray) A. Gray var. micrantha, Slender Corydalis. Mt, Pd (NC): circumneutral rockoutcrops and adjacent glades and woodlands; rare (NC Rare). April; June. Ssp. micrantha is primarily midwestern, ranging fromIL, WI, MN, and SD south to AR, TX, and OK, with disjunct outliers in e. TN and w. NC. Ownbey (1947) had no records ofSouthern Appalachian populations of C. micrantha, and considered ssp. micrantha to range no further east than IL and MO; RABincluded montane populations in ssp. australis, stating "this is the only [subspecies] in our range." Morphologically, however, thesepopulations closely resemble ssp. micrantha; their association in the Brushy Mountains with other species disjunct from western orprairie ranges (Anemone berlandieri, Arabis hirsuta, Pellaea wrightiana) provides phytogeographic corroboration. [= C; Corydalismicrantha (Engelmann ex A. Gray) A. Gray ssp. micrantha -- FNA, K, Z; Corydalis micrantha -- F (narrow sense); Corydalismicrantha -- G, infraspecific taxa not distinguished; Capnoides micranthum (Engelmann ex A. Gray) Britton -- S]

Corydalis aurea Willdenow, south and east to MD, WV (?), and PA (Kartesz 1999). [= K] {not yet keyed}

Corydalis crystallina Engelmann, is reported to occur in c. GA (Jones & Coile 1988). [= FNA, K] {not yet keyed}

Dicentra Bernhardi 1833

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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [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 atbase, gradually tapering to the tip.3 Reflexed portions of the outer sepals 4-8 mm long; [plants native and cultivated] . . . . . . . . . . . . . . . . . . . D. eximia3 Reflexed portions of the outer sepals 2-5 mm long; [plants cultivated] . . . . . . . . . . . . . . [D. formosa ssp. formosa]

2 Flowers white or yellowish (very rarely pinkish), in racemes; rootstock with bulblets; ultimate leaf segments not generally3-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. canadensis4 Spurs of the corolla elongate, divergent, 7-9 mm longl bulblets white to pink, tear-shaped (narrowed upwards) . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cucullaria

Dicentra canadensis (Goldie) Walpers, Squirrel Corn. Mt (GA, NC, VA), Pd, Cp (NC, VA): rich, moist forests, especiallyrich 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) (SCRare). 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, rockyslopes, 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 andresembles our native D. eximia. It has the reflexed portion of the outer petals 2-4 mm long. A variety of cultivars, some apparentlyderived 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, &

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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)

Lamprocapnos spectabilis (Linnaeus) Fukuhara, Bleeding Heart, native to e. Siberia, Korea, and n. China, is frequently cultivated. It differs from the native Dicentra eximia and the western American Dicentra formosa ssp. formosa in its leafy stem, theinflorescence 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). This species has recently been considered to constitute amonotypic genus, Lamprocampnos. [= K; Dicentra spectabilis (Linnaeus) Lem.]

GARRYACEAE

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

Aucuba Thunberg (Aucuba, Japanese-laurel)

* 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 plantedcultivars have the dark green leaves prominently speckled with yellow. [= K]

GELSEMIACEAE (Jessamine Family)

A family of 2 genera and about 10 species, shrubs and vines, of tropical and warm temperate America, Africa, and Asia. There ispersuasive evidence that Gelsemium and Mostuea Didr., traditionally treated as part of a heterogeneous Loganiaceae, should beaccorded family status as Gelsemiaceae (Backlund, Oxelman, & Bremer 2000; Struwe, Albert, & Bremer 1994; Sennblad & Bremer1996). 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).

Gelsemium Antoine Laurent de Jussieu (Yellow Jessamine)

A genus of 3 species, vines, our 2 species (distributed in se. North America and Central America) and 1 species in e. Asia. References: Wyatt et al. (1993); Duncan & Dejong (1964); Godfrey (1988); 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 adefinite 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 abruptlynarrowed to a beak 1.5-2 mm long; seeds with a prominent membranous wing sharply differentiated from the body of theseed; 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) andDuncan & Dejong (1964) for extensive discussions of morphology, habitat, pollination, genetics, distribution, and evolutionaryrelationships of our 2 species of Gelsemium. [= RAB, GW, K, S]

Gelsemium sempervirens St. Hilaire, Carolina Jessamine. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (GA): in a widerange of habitats, from swamp forests to dry uplands and thickets, also commonly planted as an ornamental; common. March-early May; September-November. Widespread in the Southeast: VA, se. TN, and AR south to c. peninsular FL and e. TX; disjunctin Guatemala and Mexico (Chiapas, Oaxaca, Puebla, and Veracruz). Jessamine climbs to the tops of trees. [= RAB, F, G, GW, K,S, W]

GENTIANACEAE (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).

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1 Leaves all scale-like, 1-3 (-5) mm long, appressed to the stem; [tribe Gentianeae, subtribe Swertiinae] . . . . . . . . . . . Bartonia1 Leaves larger, spreading or ascending.

2 Stem leaves whorled; plants robust, 1-3 m tall; [tribe Gentianeae, subtribe Swertiinae] . . . . . . . . . . . . . . . . . . . . Frasera2 Stem leaves opposite; plants generally less than 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 thestem;, basal rosette never present; [plants of nutrient-rich, mesic forests]; [tribe Gentianeae, subtribe Swertiinae] . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Obolaria3 Calyx lobes 4-5; stem leaves lanceolate, ovate, elliptic or narrowly elliptic (widest near the middle or towards the

base, the tip acute or acuminate), mostly more than 1.5 cm long, distributed fairly evenly along the stem, basalrosettes sometimes present; [plants of various more-or-less open habitats (except some species of Gentiana, whichcan occur in nutrient-rich, mesic forests].4 Corolla lobes 5-14, much longer than the corolla tube, pink or white; [tribe Chironieae, subtribe Chironiinae] . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sabatia4 Corolla lobes 4-5, shorter than the corolla tube, blue, lavender, pink or white.

5 Corolla tube less than 2 mm wide; [tribe Chironieae, subtribe Chironiinae] . . . . . . . . . . . . . . Centaurium5 Corolla tube more than 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 leavescuneate at the base; plants perennial; [tribe Gentianeae, subtribe Gentianinae] . . . . . . . . . Gentiana

6 Corolla lobes not alternating with corolla appendages; main stem leaves rounded to cordate at thebase; plants biennial or annual; [tribe Gentianeae, subtribe Swertiinae].7 Corolla lobes 4, finely fringed; main stem leaves rounded at base, with lateral veins obscure;

plants biennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gentianopsis7 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 thelower surface); plants 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 thuspresumably partially mycotrophic. References: Gillett (1959)=Z.

1 Corolla lobes white, 5-10 mm long, spreading, spatulate to obovate, rounded at the apex; plants 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 orrounded-mucronate at the apex; plants 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. paniculata2 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 (VA): swamps, bogs, pocosins, pocosin ecotones, sphagnous seepages; 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 northwards to MA(Gillett 1959, Wood & Weaver 1982). [= K, Z; B. paniculata -- RAB, GW, infraspecific taxa not distinguished; B. paniculata -- G, inthe narrow sense; 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 pinesavannas, shores of Coastal Plain depression ponds; uncommon (NC Watch List, VA Rare). February-April (-June). VA (1 siteknown from Virginia Beach City) and se. NC (Carteret County) south to s. FL, west to se. TX. Perhaps not truly congeneric with theother 2 species; Wood & Weaver (1982) commented that "the species [of Bartonia] can be divided into two remarkably distinctgroups." [= RAB, GW, K, S, Z]

Bartonia virginica (Linnaeus) Britton, Sterns, & Poggenberg, 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 Québecwest 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. pulchellum1 Flowers sessile or nearly so (sometimes appearing stalked but with bracteal leaves immediately below the calyx).

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2 Inflorescence a corymbiform cyme (about as broad as long, the central axis poorly developed) . . . . . . . . . . C. erythraea2 Inflorescence a spikelike cyme (distinctly elongate, the central axis straight) . . . . . . . . . . . . . . . . . . . . . . . . C. spicatum

* Centaurium erythraea Rafinesque, Common Centaury, Forking Centaury. Pd (NC, VA), Cp (VA): lawns, disturbed areas;rare, introduced from Europe and w. Asia. July-September. [= C, K; C. minus -- RAB, apparently misapplied; C. umbellatum -- F,G, name not validly published]* Centaurium pulchellum (Swartz) Druce, Lesser Centaury, Branching Centaury. Cp (VA): disturbed areas; rare, introducedfrom Europe. June-September. [= C, F, G, K, S]* Centaurium spicatum (Linnaeus) Fritsch, Spike Centaury. Cp (VA): disturbed areas; rare, introduced from s. Europe. July-August. [= C, F, G, K]

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 ofcove forests, floodplain forests; rare (NC Rare, SC Rare). Late May-June; September-October. W. NY, nw. PA, and s. Ontariowest 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 ofthis species in the Piedmont of SC. [= C, K, S, W, Z; Swertia caroliniensis (Walter) Kuntze -- RAB, F, G]

Gentiana Linnaeus (Gentian)

A genus of about 350-400 species, herbs, primarily temperate and arctic. Even following the removal of Gentianopsis andGentianella, Gentiana is a large and apparently heterogeneous group, perhaps not monophyletic. No satisfactory comprehensivetreatment 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). Key adapted from Z.

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

1 Flower solitary; corolla spotted within; leaves twisted, oblanceolate to oblinear; [subgenus Pneumonanthe, seriesAngustifoliae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. autumnalis

1 Flowers clustered; corolla not spotted within; leaves planar, mostly lanceolate, elliptic, or ovate (rarely linear); subgenusPneumonanthe, 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. alba2 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. linearis4 Margins of leaves and calyx lobes conspicuously ciliate (as seen at10×); 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 theerect one).5 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 narrowersegment often deflexed into the corolla tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. decora

5 Calyx lobes lanceolate, oblanceolate, ovate, or orbicular, 1-5× as long as broad, longer or shorter than theglabrous or puberulent calyx tube; stems glabrous or puberulent; corolla appendages subequally bifid,both segments erect.6 Corollas tightly closed; involucral and upper leaves acuminate; calyx lobes ovate-orbicular.

7 Calyx tubes densely puberulent; stems puberulent; filaments 7-12 mm long; corolla lobes oftentriangular, about 1/2 as wide as the corolla appendages . . . . . . . . . . . . . . . G. austromontana

7 Calyx tubes glabrous; stems glabrous; filaments 10-15 mm long; corolla lobes usually rounded,about as wide as the corolla appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. clausa

6 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. catesbaei8 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

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more than 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. saponaria

Gentiana alba Muhlenberg ex Nuttall, Pale Gentian. Mt (NC): habitat unknown (not collected in NC in this century); rare (NCRare). August-October. Mainly distributed in midwestern United States, from MI west to MN, south to n. AR, G. flavida occurs asscattered disjunctions eastwards to PA, OH, WV, KY, and w. NC. G. alba is the older name; there is controversy, however, overwhether 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 andne. 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]

Gentiana autumnalis Linnaeus, Pinebarren Gentian. Cp (NC, SC, VA): savannas, pine flatwoods, sandhills, in a variety ofsites varying from moist to very xeric, nearly always associated with Pinus palustris and Aristida stricta; uncommon, rare in VA (SCRare, VA Rare). Late September-early January (rarely at other times of the year, such as spring, in response to fire). This speciesis 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 inMexico: G. bicuspidata (G. Don) Briq., G. hooperi Pringle, and G. longicollis Nesom. G. autumnalis is often overlooked, since it isvery inconspicuous except when in flower, it usually flowers at a season when few botanists are about, and sterile plants greatlyoutnumber 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, edgesof moist hardwood forests; common (rare in Piedmont). Late September-November. S. NJ south to n. FL, on the Coastal Plain. [=RAB, C, F, G, GW, K, X, Y, Z; 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 (rarein Piedmont). September-October. Mostly Appalachian: ME south to w. NC and ne. TN, extending east and west to adjacentphysiographic provinces. [= RAB, C, F, G, GW, K, W, X, Y, Z; Pneumonanthe clausa (Raf.) Greene]

Gentiana decora Pollard, Appalachian Gentian. Mt, Pd (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 (NC, VA): openings in spruce-fir forest near the summit of WhitetopMountain, 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 (SevierCounty, TN), G. linearis occurs in thin soils around high elevation outcrops of Anakeesta Slate. [= C, F, G, K, W, X, Y, Z;Pneumonanthe linearis (Frölich) Greene]

Gentiana saponaria Linnaeus, Soapwort Gentian. Cp, Pd, Mt (NC, SC, VA): bogs, marshes, wet hardwood forests, othermoist to wet habitats; uncommon. Late September-November. NY west to n. IL, south to w. FL and e. TX. A peculiar form withvery narrow leaves has been found at several localities in Ashe and Watauga counties, NC and in the South Mountains; it maywarrant taxonomic recognition after further study. [= RAB, C, F, G, GW, K, W, X, Y, Z; 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 (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 southas GA or NC (as by F and G) are apparently based on misidentifications. It will key to the vicinity of G. austromontana and G.clausa, but differs from both in having the corolla lobes reduced to a minute triangular projection, much exceeded in size by theappendages. [= C, K, X, Y; Pneumonanthe andrewsii (Grisebach) W.A. Weber var. andrewsii] {not yet keyed; add refs.}

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] {not yet keyed; add refs.}

Gentianella Moench (Agueweed)

A genus of about 125 species, herbs, temperate. The separation of Gentianella from Gentiana appears to be well warranted; somecharacters 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 onGillett (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 thickenedmargins, and with broadly flattened, frequently nerve-like keels; corolla ca. 20 mm long . . G. quinquefolia var. occidentalis

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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 andvery 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,possibly other habitats as well; rare. Late August-October. Var. occidentalis A. Gray is more western, from OH and s. Ontario westto MN, east and south to w. VA, sc. KY, AR, and se. KS. [= C, G; Gentiana quinquefolia Linnaeus -- RAB, GW, W; Gentianellaquinquefolia ssp. occidentalis (A Gray) J. Gillett -- K, Z; Gentiana quinquefolia var. occidentalis -- F; Gentianella occidentalis (A.Gray) Small -- S]

Gentianella quinquefolia (Linnaeus) Small var. quinquefolia, Eastern Agueweed. Mt (GA, NC, SC, VA): forests, grassybalds; common. Late August-October. Var. quinquefolia is primarily Appalachian, from ME west to w. NY and s. Ontario, south ton. GA and sc. TN. [= C, G; Gentiana quinquefolia Linnaeus -- RAB, GW, W; Gentianella quinquefolia ssp. quinquefolia -- K, Z;Gentiana quinquefolia var. quinquefolia -- F; Gentianella quinquefolia -- S]

Gentianopsis Ma (Fringed-gentian)

A genus of about 20 species, herbs, of north temperate Asia and North America. The reasons for the recognition of Gentianopsisare 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 overcalcareous, mafic, or ultramafic rocks (such as limestone, amphibolite, or serpentinized olivine); rare (GA Threatened, NCEndangered, VA Rare). September-October. ME, s. Ontario, and ND south to NJ, IN, and IA (mostly north of the glacialmaximum) and from PA south to nw. NC and ne. GA in the unglaciated Appalachians. Certainly one of the most beautiful of ournative plants. [= C, K; Gentiana crinita Frölich -- RAB, F, G, GW, W; Anthopogon crinitum (Frölich) Rafinesque -- S; Gentianellacrinita (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 (rarein 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 (reportedfrom TX). The small purplish-green plants are inconspicuous, often nearly hidden under fallen leaves. Obolaria has well-developedmycorrhizae and may be substantially mycotrophic. [= RAB, C, F, G, K, S, W, Z]

Sabatia Adanson (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; [plants of mountain slopes, restricted in our area tosw. NC] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. capitata

3 Basal leaves much broader and shorter than the stem leaves; cauline leaves (15-) 40-80 (-100) mm long, 1-3 mmwide, 20-60× as long as wide; corolla lobes medium rose to deep rose; [plants of bogs and savannas, of the CoastalPlain] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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; [plants of Taxodium ascendens-Nyssa biflora depressions and wet pine flatwoods in se. SC] . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bartramii

4 Upper stem leaves much wider than the diameter of the stem; calyx lobes flat, linear to narrowly oblanceolate;stems 3-12 dm tall; [plants of various habitats, primarily along the shores of blackwater rivers or ponds, or in tidalmarshes].5 Primary branches opposite; terminal flower short-stalked (much shorter than the first internode of the adjacent

branch); stems 5-12 dm tall; [plants 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 theadjacent branch); stems 3-7 dm tall; [plants of brackish marshes or openings along blackwater streams]. 6 Surficial stolons usually absent or poorly developed; internodes commonly much longer than leaves;

[plants of tidal brackish or freshwater marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dodecandra

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6 Surficial stolons usually present and well-developed, elongate; internodes shorter than to equaling theleaves; [plants of openings of blackwater streams] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. foliosa

1 Flowers with 5-6 (-7) corolla lobes.7 Upper branches of main stem alternate.

13 Calyx tube strongly winged; corolla lobes pink; [plant a very rare introduction from c. United States] . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. campestris

13 Calyx tube not winged; corolla lobes pink or white; [plants native, primarily of the Coastal Plain, except S.campanulata which has disjunct occurrences in the Mountains]. 14 Calyx lobes foliaceous, 5-8 mm wide, oblong to oblanceolate, mostly exceeding the corolla lobes . S. calycina14 Calyx lobes linear-setaceous, 0.5-2 mm wide, if equaling the corolla lobes then very narrow and not foliaceous.

15 Calyx lobes (3-) 4-7 (-8) mm long; corolla lobes white; [plants of se. SC southwards] . . . . . . S. brevifolia15 Calyx lobes (4-) 6-17 (-23) mm long; corolla lobes pink (rarely white in individual plants); [plants of the

Coastal Plain of NC, SC, and VA, and rarely the Mountains of NC and VA].16 Plants perennial, often with several stems from a caudex; calyx lobes > 3/4× as long as the corolla

lobes, and sometimes exceeding them; [plants not maritime, of saturated soils from Coastal Plainsavannas to Mountain bogs] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. campanulata

16 Plants annual, solitary; calyx lobes up to 3/4× as long as the corolla lobes; [plants of brackishmarshes and interdune swales] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. stellaris

7 Upper branches of main stem opposite.8 Corolla lobes pink (rarely white); pedicels at least in part > 5 mm long.

9 Lower half of stem winged; leaves ovate, clasping, less than 2× as long as wide; [plants widespread in ourarea] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. angularis9 Lower half of stem not winged; leaves elliptic to lanceolate, more or less tapered to the base, mostly more than

3× as long as wide; [plants of the Coastal Plain or very rarely Piedmont] . . . . . . . . . . . . . . . . . . . S. brachiata8 Corolla lobes white or creamy white; pedicels (above the uppermost bracts or branches) ca. 1-2 (-5) mm long.

10 Lower portion of stem quadrangular, narrowly winged; plants annual or biennial, with 1 (-several) stems arisingfrom a taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. quadrangula

10 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].11 Leaves and upper stem not glaucous; stem terete below, becoming quadrangular or quadrangular-angledabove; corolla lobes (5-) 7-15 (-21) mm long; [widespread in our area] . . . . . . . . . . . . . . . . . . . . . S. difformis11 Leaves and upper stem glaucous; stem terete throughout; corolla lobes (4-) 5-7 (-8) mm long; [from GAsouthwards and westwards].

12 Calyx-lobes erect, (0.1-) 0.2-1.5 (-2.0) mm long, as long as or shorter than the calyx-tube;[sw. GAand n. FL westwards to e. LA] . . . . . . . . . . . . . . . . . . . . . . . . . . . S. macrophylla var. macrophylla

12 Calyx lobes strongly recurved, (1.0-) 1.5-3 mm long, longer than the calyx-tube; [e. and sc. GA southto 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, wetpine 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, of uncertain application]

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 theinterior 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]

Sabatia calycina (Lamarck) Heller, Coastal Rose-pink. Cp (GA, NC, SC, VA): swamp forests, river banks; common (VAWatch 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; scatteredinland as in w. VA, w. NC, c. TN, and KY. [= RAB, C, F, GW, K, S, W, Z; S. campanulata var. gracilis (Michaux) Fernald -- F]* Sabatia campestris Nuttall, Western Marsh-pink, Prairie Rose-gentian, Prairie Sabatia. Mt (NC): roadsides and wodlandedges; 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, oversandstone 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 inPiedmont) (VA Rare). May-September; September-December. S. NJ south to peninsular FL, west to s. AL. [= RAB, C, F, GW, K,S, Z]

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Sabatia dodecandra (Linnaeus) Britton, Sterns, & Poggenberg, 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 ande. GA (Sorrie 1998b). [= F, S; S. dodecandra var. dodecandra -- RAB, in part (see also 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; S. dodecandra var. dodecandra -- RAB, in part; S. dodecandra (Linnaeus) Britton,Sterns, & Poggenberg var. foliosa (Fernald) Wilbur -- GW, K, Z; S. harperi Small -- S]

Sabatia gentianoides Elliott. Cp (GA, NC, SC): pine savannas, bogs; common. July-August; September-October. NCsouth 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 theWaccamaw River (in se. NC and ne. SC), in adjacent ditches and disturbed flats, and very rarely on shores of beaver ponds; rare(NC Threatened, SC Rare, VA Rare). June-August; August-October. This species has a strange, disjunct range, likely related toPleistocene refugia on the (now) Continental shelf, present in s. Nova Scotia; e. MA and RI; ne. VA; se. NC and ne. SC. Thespecies was not believed to be present between se. NC and New England (MA and RI) until discovered in e. VA (Caroline County)(Fleming & Ludwig 1996). Studies underway suggest that the NC/SC 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. GAwest to e. LA. [= K, Z; S. macrophylla -- GW, infraspecific taxa not distinguished; 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, infraspecific taxa not distinguished; S. recurvansSmall -- 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 (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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erodium1 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 60species, herbs, mainly Old World.

1 Primary leaflets sessile or nearly so, sometimes connected by blade tissue; blades of the primary leaflets divided nearly orquite to the base <0.75× to base; apical pits of mericarp lacking sessile glands . . . . . . . . . . . . . . . . . . . . . . . . . E. cicutarium

1 Primary leaflets petiolulate; blades of the primary leaflets divided <0.75× to the base; apical pits of mericarp with sessileglands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [E. moschatum var. moschatum]

* Erodium cicutarium (Linnaeus) L’Héritier, Heron's-bill, Common Stork's-bill, Redstem Filaree, Alfileria, Pin-clover. Mt, Pd,Cp (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, native of Mediterranean Europe, is naturalized southat least to DE and PA; it is reported for SC (Kartesz 1999). {investigate} [= K; E. moschatum – C, infraspecific taxa notdistinguished]

Erodium texanum A. Gray, Texas Stork’s-bill, is reported as an introduction for SC (Kartesz 1999). {investigate} [= K] {not yetkeyed}

Geranium Linnaeus (Geranium, Crane's-bill)

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

1 Petals 12-18 mm long; plant a perennial, from a stout rhizome; anthers >2 mm long . . . . . . . . . . . . . . . . . . . . G. maculatum1 Petals 2-13 mm long; plant an annual or biennial, from a taproot; anthers <1 mm long.

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2 Leaves compound, at least the terminal segment (and often also the two lateral segments) petiolulate, not connected tothe lateral segments by leaf tissue; petals 9-13 (-15) mm long; [plant 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; [plantscollectively 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 mmlong) and short (<0.5 mm long) gland-tipped and eglandular hairs; stamens (all 10) fertile (note that anthersmay fall readily) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. molle

4 Mericarps appressed pubescent, not ridged; stem pubescence of short (<0.3 mm long), gland-tipped andeglandular 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 . . . . . . . . . . . G. dissectum6 Mericarps with long appressed hairs about 1 mm long, these not gland-tipped.

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. 8 Pedicels spreading pubescent, the hairs not glandular; mature stylar beak ca. 1 mm long . [G. sibiricum]8 Pedicels either rerorse-strigose or glandular-villous; mature stylar beak 3-5 mm long.

9 Pedicels glandular-villous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. bicknellii9 Pedicels retrorsely strigose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. columbinum

*? Geranium bicknellii Britton. Mt (VA): {check with VAHP} open woodlands and clearings; rare. July-September. [= C, G, K,W; G. bicknellii var. ?? -- F]

Geranium carolinianum Linnaeus var. carolinianum, Southern Carolina Crane's-bill. Cp, Pd, Mt (NC, SC, VA): fields,roadsides, lawns, pastures, gardens, disturbed areas; common. March-June (and sometimes later). [= C, F, G; G. carolinianum --RAB, W, infraspecific tax not distinguished; G. carolinianum var. carolinianum – K, in part]

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, W, infraspecific tax not distinguished; G. carolinianum var. carolinianum – K, in part]* Geranium columbinum Linnaeus, Long-stalk Crane's-bill. Mt (NC, VA), Pd (VA): roadsides, pastures, disturbed areas;common, introduced from Europe. May-July. [= RAB, C, F, G, K, W]* Geranium dissectum Linnaeus, Cutleaf Crane's-bill. Mt, Pd (NC, SC, VA), Cp (NC, VA): roadsides, pastures, disturbedareas; common, introduced from Europe. April-June. [= RAB, C, F, G, K, W]

Geranium maculatum Linnaeus, Wild Geranium. Mt, Pd, Cp (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. Sometimescultivated. [= RAB, C, F, G, K, W]* Geranium molle Linnaeus, Dove's-foot Crane's-bill. Mt, Pd (NC, SC, VA), Cp (VA): roadsides, pastures, disturbed areas;common, introduced from Europe and w. Asia. April-July. [= RAB, C, F, G, K, 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, W]

Geranium robertianum Linnaus, 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 ibericum Cavanilles has recently been found in Great Smoky Mountains National Park, in both NC and TN (K. Langdonpers. comm.). [= K; G. nepalense – C] {not yet keyed}

Geranium sanguineum Linnaeus is “cultivated and rarely escaped to fields and waste ground” in se. PA (Rhoads & Klein 1993). [=C, K]

Geranium sibiricum Linnaeus, native to Asia, is naturalized south to s. PA (Rhoads & Klein 1993) and is likely to occur in at leastthe northern part of our area. [= C, K]

Geranium thunbergii Siebold & Zuccarini ex Lindley & Paxton, reported for NC (Nesom 2000, Kartesz 1999). [= K; G. nepalenseSweet var. thunbergii (Siebold & Zuccarini ex Lindley & Paxton) Kudo] {add to key, and ‘promote’ to main text}

Geranium versicolor Linnaeus is naturalized south to se. and sw. PA (Rhoads & Miller 1993) and may occur in our area. [= K]

Other Eurasian species are introduced in ne. North America and may be found in our area as well.

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GROSSULARIACEAE (Currant Family)(also see ITEACEAE)

A family of about 24 genera and 330 species, trees and shrubs, cosmopolitan. The familial distinction from the Saxifragaceae issupported by recent molecular data, though the affinities of Grossulariaceae and Saxifragaceae (sensu stricto) appear to be closerthan those of many other groups traditionally included in the Saxifragaceae, such as Parnassia, Lepuropetalon, and Penthorum(Morgan & Soltis 1993).

Ribes Linnaeus (Currant, Gooseberry)

A genus of about 150 species, temperate, of the Northern Hemisphere and montane South America. The genus is separated intodistinctive 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 asmall variety of grape, and have nothing to do with Ribes. A molecular study suggests that recognition of Grossularia as a genusdistinct from Ribes is not warranted, though it does form a monophyletic group nested within Ribes s.l. (Weigend, Mohr, & Motley2002). References: Sinnott (1985)=Z; Weigend, Mohr, & Motley (2002); Spongberg (1972); Schultheis & Donoghue (2004). Keyadapted 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 atmaturirty; 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; [plants of the Mountains] . . . . . . . . . . . . . . . . . . . . . . R. cynosbati3 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; [plants 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. missouriense4 Calyx tube purplish or purplish-green; stamens 6-8 mm long; nodal spines 0-1 (-2) per node, 3-11 mm long, slender

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. rotundifolium1 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. lacustre5 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. glandulosum6 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.7 Leaves with resinous glands beneath; fruits black when mature; [plant native] . . . . . . . . . . . . R. americanum7 Leaves not glandular beneath; fruits red when mature; [plant an introduced species, 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 1-4flowers; 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. cynosbati2 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 fullanthesis exceeding the petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. rotundifolium

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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, IN, NE, and CO. [= C, F, G, K, W]

Ribes cynosbati Linnaeus, Prickly Gooseberry, Dogberry. Mt (GA, NC, VA): moist slopes, periglacial boulderfields, grassybalds, 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, F, G, K, W, Z; 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 McCormickCounty, 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 BritishColumbia, 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 (VARare). May-June. Labrador to AK, south to MA, PA, w. VA, TN (allegedly), n. OH, MI, MN, CO, UT, and CA. The documentationfor 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; rare, probably introduced from further west (VA WatchList). April-June; May-July. Ranging as a native from w. WV, sw. OH, IN, WI, MN, and e. ND south to KY, TN, 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; Grossulariamissouriensis (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 Appalachianendemic: 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 toadjacent 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. UnitedStates, rarely as far south as our area; it may escape. It also is reported as occurring as a native species as far east asMontgomery County in nc. TN (Chester, Wofford, & Kral 1997). It will key to couplet 6, where trouble will be encountered. It hasthe hypanthium long-tubular, the flowers golden yellow, and fruits black (rarely yellow). [= K; R. odoratum H. Wendland -- C]

Ribes curvatum Small, Granite Gooseberry. Pd, Mt (GA): rocky upland forests; rare (GA Special Concern). Native from c. andnw. GA (Jones & Coile 1988) and e. TN, in the Cumberland Plateau (Chester, Wofford, & Kral 1997) westward. Also reported forNC by Sinnott (1985), and his report is backed up by specimens. The specimens, however, indicate that the species was cultivatedin 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 toR. missouriense; it can be distinguished from all our species of gooseberries by its glandular-punctate leaves, and additionally fromR. 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]

Ribes hirtellum Michaux, Northern Gooseberry, ranges south to WV (Tucker County) and may be found in our area. It will key tocouplet 2, where trouble will be encountered. It has glabrous ovaries and fruits, but the stamens are short and included. [= C, K]

Ribes nigrum Linnaeus, Garden Black Currant or Cassis, native to Europe, is cultivated in ne. United States, rarely as far south asour area; it may escape. It will key to R. americanum, from which it can be distinguished by its pedicels 2-8 mm long, much longerthan subtending ovate bracts (vs. pedicels 0-2 mm long, shorter than the subtending lanceolate bracts). [= C, K]

Ribes triste Pallas, Swamp Red Currant, ranges south to WV (Mineral, Pocahontas, and Randolph counties); it may occur in ourarea. 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 obviouslynative habitat, likely in boggy forests or seepage wetlands at high elevations, and probably in VA. [= C, K]

Ribes uva-crispa Linnaeus var. sativum Augustin de Candolle, Garden Gooseberry, native of Europe, is cultivated in ne. UnitedStates, 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, infraspecific taxa not distinguished; R. grossularia Linnaeus]

HALORAGACEAE (Water-milfoil Family)

A family of about 9 genera and 145 species, aquatic and wetland herbs, but also shrubs and trees, cosmopolitan but centered inthe 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)

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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myriophyllum1 Leaves alternate; stamens 3; carpels 3; emersed leaves foliaceous, little if at all reduced . . . . . . . . . . . . . . . . . Proserpinaca

Myriophyllum Linnaeus (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: Aiken (1981). Keys adapted from Aiken (1981), GW, RAB, and C.

Keying Note: {Treatment partial at this time, owing to confusion in various sources about taxa in our area -- material at all areaherbaria needs to be looked at to resolve uncertainties -- neither key includes all species attributed to our area by Aiken, Harvill etal., C, etc.}

Key 1(treating 6 species, not M. humile, M. sibiricum, or M. verticillatum)

1 Submersed leaves entire and scale-like (or absent); plants unbranched, erect from the substrate . . . . . . . . . . . . M. tenellum1 Submersed leaves pinnately divided into filiform segments; plants usually branched, floating in the water column.

2 Leafy vegetative branches considerably emersed, markedly featherlike; emersed leaves mostly 2.5-3.5 cm long; plantsdioecious (ours pistillate), but rarely flowering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aquaticum

2 Leafy vegetative branches mostly submersed (unless stranded by low water), not markedly featherlike (althoughpinnately divided); emersed leaves less than 2 cm long; plants monoecious, the upper flowers staminate.3 Bracts of the inflorescence usually longer than the internodes of the axis, very conspicuous even at a distance.

4 Submersed leaves with a total of 7-17 segments; bracts normally pectinately toothed; fruit mericarps each with with 2 longitudinal ridges, the ridges sharply tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pinnatum

4 Submersed leaves with a total of (15-) 17-31 (-37) segments; bracts normally broad, serrate to entire(sometimes pectinate when stranded); fruit mericarps rounded longitudinally or with 2 low ridges, the ridgesobscurely tuberculate with low, rounded projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. heterophyllum

3 Bracts of the inflorescence shorter than the internodes of the axis, inconspicuous at almost any distance.5 Submersed leaves with a grayish cast, with a total of at least 23 segments neatly disposed in pairs (or

suboppositely), parallel to each other; bracts elliptical to obovate, entire (or nearly so) . . . . . . . . . M. spicatum5 Submersed leaves green or with a reddish cast, with a total of 7-15 (-17) segments disposed alternately,

irregularly, or suboppositely (nt neatly); lower bracts pectinate, above gradually becoming lanceolate and few-tothed, then narrowly obovate and nearly entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. laxum

Key 2(treating 7 species in our area, not M. humile or M. sibiricum)

1 Submersed leaves entire and scale-like (or absent); plants unbranched, erect from the substrate . . . . . . . . . . . . M. tenellum1 Submersed leaves pinnately divided into filiform segments; plants usually branched, floating in the water column.

2 Submersed leaves (at least some of them) alternate or "pseudo-whorled' (some of the leaves separated by very shortinternodes), often some of them also opposite or in true whorls of 3-5.3 Upper bracts subentire, spatulate, shorter than the adjacent flowers; submersed leaves with a total of 11-21 capillary

divisions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. laxum3 Upper bracts pinnately divided, filiform, much longer than the adjacent flowers; submersed leaves with a total of 8-

10 (rarely more) capillary divisions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pinnatum2 Submersed leaves all whorled (sometimes "pseudo-whorled" in M. heterophyllum).

4 Emergent leaves mostly 2.5-3.5 cm long, with 18-36 uniform linear divisions, each 4-8 mm long . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aquaticum

4 Emergent leaves all less than 2.0 cm long, entire, serrate, or pectinate, the teeth or divisions < 4 mm long.5 Upper bracts entire; lower bracts entire, pectinate, or serrate, not more than 2× the length of the adjacent

pistillate flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. spicatum5 Upper bracts pectinate or dentate; lower bracts pectinate, usually > 2× as long as the adjacent pistillate

flowers.6 Bracts laminate, the lower more deeply toothed than the upper; bracteoles 1.0-1.3 mm long . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. heterophyllum6 Bracts without a distinct lamina, the lower pinnatifid, the upper pectinate; bracteoles absent or less than

0.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. verticillatum

* Myriophyllum aquaticum (Vell. Conc.) Verdc., Parrot-feather. Cp, Pd, Mt (NC, VA): ditches, slow-moving rivers, pools,ponds; common (rare in Piedmont and Mountains), native of South America. April-June. An introduced species now widespread inse. United States, north to NY, WV, and MO. [= C, GW, K, W; M. brasiliense Cambessedes -- RAB, F, G; M. proserpinacoidesGillies ex Hooker & Arnott -- S]

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Myriophyllum heterophyllum Michaux, Southern Water-milfoil. Cp (NC, VA): ditches, slow-moving waters of rivers andstreams, pools, ponds; common. April-July. NY west to Ontario and MN, south to FL and TX. [= RAB, C, F, G, GW, K, S]

Myriophyllum humile (Rafinesque) Morong. Cp (VA): floating in an artificial pond; rare (VA Rare). [= K]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 presentdifficulties 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]

Myriophyllum pinnatum (Walter) Britton, Sterns, & Poggenberg, Alternate-leaved Water-milfoil. Cp (NC, VA), Mt (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]* Myriophyllum spicatum Linnaeus, Eurasian Water-milfoil. Cp (SC): ponds and impoundments; uncommon? {habitat andrange in our area uncertain}, native of Eurasia, confused with M. sibiricum, (see below). An introduced species, now widespread ine. United States. Reported for South Carolina by Hill & Horn (1997). [= C, GW, K, W]

Myriophyllum tenellum Bigelow, Leafless Water-milfoil. Cp (NC, VA): natural lakes (Carolina bay lakes), typically growingon the sandy bottoms in water 1-2 meters deep; rare (NC Rare, VA Rare). Newfoundland west to MN, south to PA and NJ, anddisjunct south to a few occurrences in VA and NC. [= C, F, G, K]

Myriophyllum verticillatum Linnaeus. Habitat and range in our area uncertain. A circumboreal species, south in NorthAmerica to FL, MN, and UT. [= C, G, K; M. verticillatum var. pectinatum Wallroth -- F]

Myriophyllum sibiricum Komarov is reported to range south to VA. It is closely related to M. spicatum, but can be distinguished byleaf divisions mostly 10-24 per leaf (vs. mostly 24-40 per leaf), presence of oblanceolate turions (vs. not turion-forming), stem notthickened below inflorescence (vs. stem thickened to nearly 2× the diameter of the lower stem). Given the dispersal of this genusvia waterfowl and boats, other species may be found to occur in our area, at least as vagrants. [= K; M. exalbescens Fernald]

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 mmwide.2 Fruit 2.3-4.0 mm wide, acutely angled, not winged, the sides of the capsule more-or-less planar P. palustris var. crebra2 Fruit (3.5-) 4.0-6.0 mm wide, sharply angled (to somewhat winged), the sides of the capsule concave . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. palustris var. palustris1 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. intermedia3 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): wet places; rare (NC WatchList). July-September. Nova Scotia to SC on the Coastal Plain; disjunct in sc. TN. This taxon is intermediate in morphologybetween P. palustris and P. pectinata; whether it warrants species status is unclear. If merely a rarely produced first-generationhybrid, it should be treated as a hybrid binomial (P. ×intermedia); if it forms independent, self-reproducing populations, it shouldprobably be treated as a species. [= RAB, C, F, G, K, Z; P. palustris -- GW, in part]

Proserpinaca palustris Linnaeus var. crebra Fernald & Griscom, Common Mermaid-weed. Cp (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, infraspecific taxa not distinguished; P. palustris -- GW, infraspecific taxa not distinguished,and also including P. intermedia]

Proserpinaca palustris Linnaeus var. palustris, Coastal Mermaid-weed. Cp (NC, SC, VA), Pd (NC, SC): 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, infraspecific taxa not distinguished; P. palustris -- GW, infraspecific taxa not distinguished, and also including P. intermedia]

Proserpinaca pectinata Lamarck, Feathery Mermaid-weed. Cp (NC, SC, VA): 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. [= K] {not yet keyed}

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

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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 truncate or 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 oblique base, the lateral veinsincluded 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: 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 vegetativecondition; 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 less than 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; [plants of wet savannas,pocosins, and pocosin margins of the Coastal Plain] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. gardenii

1 Leaves glabrous or sparsely stellate-pubescent above, up to 12 cm long and 10 cm wide (the largest more than 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; [plants ofrocky 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). April-May; September-October. Se. NC (and allegedly se. VA) south to panhandle FL and s. AL. Anornamental prized for its small size and attractive fall color. [= RAB, F, FNA, GW, K, S, Z; F. parvifolia Kearney -- S]

Fothergilla major (Sims) Loddiges, Large Witch-alder. Mt, Pd (GA, NC, SC): dry ridgetop forests of middle elevation ridgesin the mountains, especially along the Blue Ridge Escarpment, summits and upper slopes of Piedmont monadnocks, north-facingbluffs in the lower Piedmont; rare (GA Special Concern, NC Rare, SC Rare). April-May; July-October. C. NC west to ne. TN, southto 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 Americanspecies, H. vernalis Sargent, is endemic to the Ozark/Ouachita region of AR, OK, and MO. References: Wen & Shi (1999);Endress in Kubitzki, Rohwer, & Bittrich (1993).

Hamamelis virginiana Linnaeus, 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. Some CoastalPlain populations in SC and GA have very small leaves and may warrant taxonomic recognition; they need additional study. Thebark 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 dropsits leaves. [= RAB, C, F, FNA, G, GW, K, S, W; H. virginiana var. parvifolia Nuttall -- F]

HIPPOCASTANACEAE (Buckeye Family)(see SAPINDACEAE)

HYDRANGEACEAE (Hydrangea Family)

A family of about 17 genera and 190 species, trees, shrubs, vines, and herbs, primarily north temperate. As here interpreted, thefamily Hydrangeaceae includes two well-marked groups, the Hydrangeae (including Decumaria and Hydrangea) and thePhiladelpheae (including Deutzia and Philadelphus). This group has been shown by molecular research to be unrelated to theSaxifragaceae, 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).

1 Plant a woody vine, climbing by aerial rootlets; petals 7-10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decumaria1 Plant a shrub; petals 5 (rarely 10 in the cultivars of the alien Deutzia).

2 Pubescence of leaves and twigs stellate; stamens 10; [plant a cultivated alien, rarely escaped] . . . . . . . . . . . . . . Deutzia2 Pubescence of leaves and twigs simple; stamens 8-10 (in Hydrangea) or 25-90 (in Philadelphus).

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3 Leaf blades 10-30 cm long; inflorescences of 25-many flowers; stamens 8-10 . . . . . . . . . . . . . . . . . . . . Hydrangea3 Leaf blades 3-8 cm long; inflorescences of 1-7 flowers; stamens 25-90 . . . . . . . . . . . . . . . . . . . . . . . Philadelphus

Decumaria Linnaeus (Climbing Hydrangea, Woodvamp)

A genus of 2 species, vines, of e. North America and e. Asia (China).

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

Decumaria barbara Linnaeus, Climbing Hydrangea, Woodvamp. Cp (GA, NC, SC, VA), Pd (GA, SC), Mt (GA, NC): swampforests 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, 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]

Deutzia Thunberg (Deutzia)

A genus of about 60 species, shrubs, mainly Asian. * Deutzia scabra Thunberg, Deutzia, Pride-of-Rochester. Mt (NC, VA), Pd, Cp (VA): fairly commonly cultivated, persistentaround old homesites and escaping to adjacent woodlands; rare, native of Japan and China. [= C, F, K]

Deutzia crenata Siebold & Zuccarini, Chinese Deutzia, is introduced in GA (Kartesz 1999). [= K] {not yet keyed}

Hydrangea Linnaeus (Hydrangea, Sevenbark)

A genus of about 25 species, shrubs, of e. North America and e. Asia. Recent molecular analyses suggest that Hydrangea asusually interpreted is polyphyletic (Soltis, Xiang, & Hufford 1995). Future taxonomic changes are to be expected. References: Pilatowski (1982).

1 Leaves pinnately lobed, the lobes toothed; inflorescence a panicle; large sterile flowers many (more than 20 perinflorescence), 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 (more than 20 per inflorescence), borne throughout the inflorescence;

[plant alien, cultivated and sometimes persistent] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. paniculata2 Inflorescence a corymb; large sterile flowers absent to relatively few (0-15 per inflorescence), borne around the periphery

of the corymb.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 less than 1 cm in diameter . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. arborescens

3 Lower leaf surface variously pubescent, appearing white or gray; trichomes of the lower leaf surface on veins andinterveinal 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); sterileflowers generally very few (0-3 per inflorescence) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. cinerea

4 Lower leaf surface densely floccose-velutinous, felt-like, appearing bright white or silver; trichomes denseenough to entirely mask the green leaf surface; trichomes without tubercles, or with small and inconspicuoustubercles (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, especiallyaround rock outcrops and along streambanks; common (rare in Coastal Plain). May-July. NJ, s. NY, OH, IN, IL, MO, and se. KSsouth to e. NC, c. SC, c. GA, panhandle FL, s. AL, LA, and OK. [= F, K, S, W; H. arborescens ssp. arborescens -- RAB; H.arborescens var. arborescens -- C, G; H. arborescens var. oblonga Torrey & A. Gray -- F]

Hydrangea cinerea Small, Ashy Hydrangea. Mt, Pd (GA, NC, SC): rocky forests and rock outcrops, roadbanks, perhapsstrictly 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; H. arborescens ssp. discolor (Seringe) McClintock -- RAB; H. arborescens var. deamii St. John -- F;H. arborescens var. discolor Seringe -- C, G]* Hydrangea paniculata Siebold, Hydrangea. Mt (NC, VA): persistent after cultivation at old home-sites, sometimes

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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 orescpae from cultivation . Boufford & Wood (1977) describe an apparently native occurrence in nw. SC. This southeastern native isa 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 theFrench Broad River and to its southwest), nw. SC, ne. GA, and se. TN, with outliers (perhaps escaped from cultivation?) in StokesCounty, NC and Calhoun County, SC. This attractive species is especially typical of the escarpment gorge region near the tricornerof 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; H. arborescens ssp. radiata (Walter) McClintock -- RAB]

Philadelphus Linnaeus (Mock-orange)

A genus of 65 (or fewer) species, shrubs, of north temperate areas. The most recent monographer of the genus, Shiu-ying Hu(1954-1955) recognizes many species and varieties on the basis of minor differences in pubescence. Many of the recognized taxaare based only on cultivated material. The native distributions of the varieties have little phytogeographic coherence, and severalvarieties are often reported from the same site, suggesting that they reflect merely variation within a population (if genetically basedat all). For instance, Hu recognizes three varieties in Ph. hirsutus and five in Ph. inodorus, but these seem to be no more thanforms. As Hu writes, "the formerly recognized species, Ph. grandiflorus Willd., and Ph. laxus Schrad., are merely different forms ofa 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 ofvariations exhibited by a large number of specimens collected from the homeland of Ph. inodorus Linn. they appear to be nothingbut a few transitional forms. In this paper, these forms are treated as varieties." Hu's "varieties" should be treated as forms orcultivars, if recognized at all. I have taken a conservative approach, though variation in several of our native species could useadditional 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, sectionDeutzioides] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. hirsutus

1 Axillary buds enclosed; twigs of the current year glabrous; seeds with caudate tails about as long as the embryo; [subgenusPhiladelphus].2 Flowers 1-3 (-9) in a cymule; stamens 60-90; [subgenus Philadelphus, section Pauciflorus] . . . . . . . . . . . . Ph. inodorus2 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. coronarius3 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): 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 mostcommonly 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, rockoutcrops, 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 orless 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 iscultivated and it may escape outside of the range stated. [= RAB, C, F, G, K, S, W, 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 fromnw. GA, is similar to Ph. inodorus, allegedly differing in the pedicels and hypanthium pubescent (vs. glabrous); it is probably only aform of Ph. inodorus. [= RAB, C, F, G, K, S, W, 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. floridus Beadle -- K, S, Z]

Philadelphus pubescens Loiseleur, Ozark Mock-orange, Hairy Mock-orange. Mt (GA): limestone bluffs; rare (GA SpecialConcern). E. TN, KY, nw. GA (Jones & Coile 1988), AL, s. IL, MO, OK, and AR, west of the Blue Ridge. It has been documentedfrom TN counties adjacent to both VA and NC, and is likely to be found in VA, at least. [Ph. intectus Beadle -- S, Z; Ph. latifoliusSchrader ex DC. -- S; Ph. intectus var. pubigerus Hu -- Z; Ph. pubescens var. verrucosus (Schrader) Hu -- Z; Ph. pubescens var.pubescens – K; ; Ph. pubescens var. intectus (Beadle) A.H. Moore – K]

HYDRASTIDACEAE (Golden-seal Family)

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A family of 2 genera and 2 species, perennial herbs, of temperate e. North America and Japan. In chemistry, morphology, andanatomy, Hydrastis shows some relationship to Podophyllum and Diphylleia of the Podophyllaceae (often included in theBerberidaceae). 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 theRanunculaceae, Berberidaceae and probably of Circaeasteraceae, and that Hydrastis has evolved in its own evolutionary lineparallel with other lines leading to the modern representatives of these families." In recent papers on classification of the floweringplants, Thorne (1992) and Reveal (1993) have also accepted Hydrastidaceae as a distinct family. Tobe in Kubitzki & Bayer placesHydrastis with Glaucidium Siebold & Zuccarini in a bigeneric Hydrastidaceae. References: Tamura in Kubitzki, Rohwer, & Bittrich(1993); Tobe in Kubitzki & Bayer (2002).

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 largecolonies after canopy disturbance such as logging; rare (GA Endangered, NC Endangered, VA Watch List). April; May-June. VTand 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 moreremote parts of the mountains, though too rare in our area to support any economically significant wild collection. The root is bitterin taste and contains several alkaloids. Reported for SC (P. McMillan, pers.comm. 2002). [= RAB, C, F, FNA, G, K, S, W)

HYDROLEACEAE

The Hydroleaceae is not closely related to Hydrophyllaceae; recent molecular data confirm the prevailing view through most of the19 century that Hydrolea was in its own family. References: Ferguson (1998); Hilger & Diane (2003); Angiosperm Phylogenyth

Group (1998, 2003).

Hydrolea Linnaeus

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

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 7-8mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. quadrivalvis

1 Flowers in terminal cymes or corymbs; leaves 2-6 cm long, 0.6-2.5 cm wide; axillary spines present or absent; corolla 10-15mm long.2 Leaves elliptic to lanceolate, 2-3 cm long, 0.6-1.0 cm wide; axillary spines absent or rudimentary . . . . . . H. corymbosa2 Leaves ovate to ovate-lanceolate, 3-6 cm long, 1.5-2.5 cm wide; axillary spines preset, wel-developed, to 1.5 cm long . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [H. ovata]

Hydrolea corymbosa J. Macbride ex Elliott, Skyflower. Cp (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 quadrivalvis Walter, Waterpod. Cp (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 ovata Nuttall ex Choisy, ranging from c. GA west to TX, north in the interior to sc. TN and MO. [= C, F, G, GW, K; Namaovata (Nuttall ex Choisy) Britton -- S]

HYDROPHYLLACEAE (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 (1960); 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

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towards 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).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 . . . . . . . . . . . . . . . . . . . . . . . . . . Ellisia4 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemophila3 Flowers all terminal in 3-many-flowered cymes.

5 Inflorescence repeatedly branched subdichotomously; larger leaf blades more than 8 cm wide; stamens wellexserted 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); largerleaf blades less than 5 cm wide (except Ph. bipinnatifida); stamens slightly exserted from the corolla (less than 3mm beyond the corolla) (except well-exserted in Ph. bipinnatifida, included in Ph. covillei); plants annual (biennial inPh. 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 arebadly in need of critical reassessment. References: Constance (1940)=Z.

Identification notes: Ellisia superficially resembles Phacelia ranunculacea and Nemophylla triloba.

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 tosc. VA. Likely to occur in nc. NC. [= C, F, G, GW, K, W, Z; Nyctelea nyctelea (Linnaeus) Britton -- S]

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 7-8mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. quadrivalvis

1 Flowers in terminal cymes or corymbs; leaves 2-6 cm long, 0.6-2.5 cm wide; axillary spines present or absent; corolla 10-15mm long.2 Leaves elliptic to lanceolate, 2-3 cm long, 0.6-1.0 cm wide; axillary spines absent or rudimentary . . . . . . H. corymbosa2 Leaves ovate to ovate-lanceolate, 3-6 cm long, 1.5-2.5 cm wide; axillary spines preset, wel-developed, to 1.5 cm long . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [H. ovata]

Hydrolea corymbosa J. Macbride ex Elliott, Skyflower. Cp (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 quadrivalvis Walter, Waterpod. Cp (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 ovata Nuttall ex Choisy, ranging from c. GA west to TX, north in the interior to sc. TN and MO. [= C, F, G, GW, K; Namaovata (Nuttall ex Choisy) Britton -- S]

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. canadense1 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-) 9-13segments, these toothed but not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. macrophyllum

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3 Inflorescence and upper stem glabrate to strigose with appressed to ascending hairs less than 0.5 mm long; leaves with5-7 (-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;

[plants of low to high elevations in the Mountains] . . . . . . . . . . . . . . . . . . . . . . . . . . H. virginianum var. atranthum4 Corolla white to lavender or pale purple; lower stem slightly to densely pubescent with retrorse to spreading hairs;

[plants of low elevations of the Piedmont, Mountains, and Coastal Plain] . . . . . . . H. virginianum var. virginianum

Hydrophyllum canadense Linnaeus, Mapleleaf Waterleaf, Canada Waterleaf, Broadleaf Waterleaf. Mt (NC, SC, VA), Pd(NC, VA), Cp (VA): cove forests, rocky streambanks, other moist and nutrient-rich forests; common (rare in Coastal Plain and inPiedmont 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, andIL, south to sw. VA, w. NC, n. GA, and n. AL; reports from AR are erroneous, and are based on material of Hydrophyllum browneiKral & 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 hasbeen little consensus about this taxon, some regarding it as merely a color form. Alexander lists numerous characters additional tothat of flower color; they need further investigation. "H. atranthum differs from H. virginianum in the dark-violet flowers, the brownhairs on the appendages, brown filaments, corolla-lobes longer than the tube, stamens shorter [11.5 mm] and more slender, andthe more numerous leaf-lobes. H. virginianum has flowers white to pale lavender or pinkish, white hairs on the appendages, whitefilaments, corolla-lobes and tube equal in length, filaments longer (13.5 mm) and stouter, and 5-7 leaf-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 theSouthern Appalachians do not share the stem pubescence character stated in the key above. The general correlation of flowercolor and lower stem pubescence and the tight geographical range of var. atranthum incline me to accept it provisionally as avariety. It is not, however, limited to high elevations, as stated or implied by some authors. The two varieties provisionallyaccepted 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, infraspecific taxa not distinguished; H. virginicum -- S, orthographic error, infraspecifictaxa not distinguished; 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 closelyrelated vicariant of H. tenuipes Heller of British Columbia south to CA. See Phacelia bipinnatifida for additional suggestions ondistinguishing it from this species. [= C, F, G, Z; H. virginianum -- RAB, K, W, Y, infraspecific taxa not distinguished; H. virginicum -- S, orthographic error, infraspecific taxa not distinguished; H. virginianum -- X, in the narrow sense]

Hydrophyllum appendiculatum Michaux, Biennial Waterleaf, ranges from 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; Decemiumappendiculatum (Michaux) Small -- S]

Nama Linnaeus(also see Hydrolea)

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).

Nemophila aphylla (Linnaeus) Brummitt. Cp, Pd (NC, SC, VA): moist, nutrient-rich floodplain forests; uncommon, thoughoften 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. Nemophila is superficially similar to Phacelia covillei, with which it sometimes co-occurs. Nemophila has white flowers solitaryopposite the leaves and sometimes also in a few-flowered terminal raceme (vs. pale lavender flowers in a terminal raceme only),lateral segments of the leaves often as large as the terminal (vs. lateral segments smaller than the terminal). [= GW, K; N.microcalyx (Nuttall) Fischer & Meyer -- RAB, F, G, S; N. triloba (Rafinesque) Thieret -- C]

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Phacelia Antoine Laurent de Jussieu (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).

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. fimbriata2 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 stem relatively sparse, consisting of appressed to ascending, stiff, pointed hairs; [plants of riversin the Atlantic drainage] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. covillei

4 Pubescence of the stem relatively dense, much of it spreading or even retrorse, most of the hairs weak and twisted,and many of them glandular-tipped; [plants 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 percapsule.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; [plants 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 theleaf 5-15 mm long, 5-9 mm wide; pedicels ascending to spreading in fruit; [plants 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. maculata6 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 1-3pairs of rather broad lobes; [plants of various habitats (including grantic 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 leaveswith 4-5 pairs of lateral leaflets, the terminal leaflet about the same size and unlobed; cauline leaves with2-4 pairs of narrow lobes; [plants associated with granitic flatrocks and domes of the Piedmont of SC andsouthwestwards] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. dubia var. georgiana

Phacelia bipinnatifida Michaux, Fernleaf Phacelia, Forest Phacelia. Mt (GA, NC, SC, VA): cove forests, especially whererocky; 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 and Hydrophyllum virginianum are sometimes confused. Ph. bipinnatifida has the largerand more basal leaves distinctly bipinnatifid, the lower pinnae often stalked (vs. pinnatifid, the basal or terminal pinnae sometimes2-lobed, all the pinnae more-or-less sessile), pubescence of the upper stem and inflorescence in part glandular (pubescencenonglandular), and seeds 4 per capsule, black (vs. 2 per capsule, light brown). 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 stamensand style. These variations are not concomitant, and the distribution of forms showing a complete or partial combination of them issporadic" (Constance 1949). The matter deserves additional study. [= RAB, C, F, G, K, S, W, Z; Ph. bipinnatifida var. plummeriWood -- F; Ph. brevistyla Buckley -- S]

Phacelia covillei S. Watson ex A. Gray, Eastern Buttercup Phacelia. Pd (NC, VA): rich soils of floodplains, and contiguousterraces and slopes; rare (NC Rare, VA Rare). April; May. Ranging in two disjunct areas -- c. NC and sc. VA (in the drainages ofthe Cape Fear, Tar, and Roanoke rivers) and DC, n. VA, and sc. MD (in the drainage of the Potomac River). Most recent authorshave included this taxon within the closely similar Ph. ranunculacea; as thus broadly defined, Ph. ranunculacea was considered tooccur 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), onenear Washington, DC (DC, n. VA, and sc. MD), and a third in c. NC. Chuang & Constance (1977) reported that the westernpopulation center has a chromosome number of n=6, and that the two eastern population centers have n=14. The disparatecytotypes of eastern and western Ph. ranunculacea sensu lato led Chuang & Constance to seek morphological differences thatwould warrant the recognition of separate taxa, but they reported that "no consistent morphological distinction has been foundbetween the two cytotypes." Stem pubescence does, however, show consistent, though subtle, differences between eastern andwestern populations of this complex. Eastern material has the stem pubescence relatively sparse, consisting of appressed toascending, 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 allopatricdistribution 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

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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 Phaceliauncomfortable (see discussion in Chuang & Constance 1977, Constance 1949, Gillett 1968). See Nemophila aphylla forsuggestions on distinguishing these two superficially similar species. [= K; Ph. ranunculacea (Nuttall) Constance -- RAB, C, F, G,Z, in part]

Phacelia dubia (Linnaeus) Trelease var. dubia, Appalachian Phacelia. Mt (GA?, NC, SC, VA), Pd, Cp (NC, SC, VA): floodplain forests, rocky forests, fields, roadsides, granitic flatrocks; common. April-May; June. Var. dubia ranges from PA west toWV, south to nc. SC, sw. NC, and se. TN. In addition to our two varieties, var. interior Fernald is endemic to limestone glades in c.TN. [= K, Y; Ph. dubia -- RAB, C, F, G, S, W, infraspecific taxa not distinguished; Ph. dubia var. fallax (Fernald) Gleason -- G; Ph.dubia var. dubia -- Z, in part only (also including var. interior)]

Phacelia dubia (Linnaeus) Trelease var. georgiana McVaugh, Georgia Phacelia. Pd (GA, SC): granitic flatrocks; rare. April-May; June. Var. georgiana ranges from SC south and west to ec. AL. Var. georgiana is included for our area on the basis ofLevy (1991), who discusses the occurrence of a putative variety ("imitator") from c. SC. This population was morphologicallyindistinguishable from var. georgiana but showed partial sterility when crossed with var. georgiana; see Levy (1991) for furtherdiscussion. [= K, Y, Z; Ph. dubia -- RAB, C, F, G, S, W, infraspecific taxa not distinguished]

Phacelia fimbriata Michaux, Fringed Phacelia, Blue Ridge Phacelia. Mt (GA, NC, VA): moist forests on slopes andfloodplains, at low to high elevations, perhaps mainly over circumneutral soils; uncommon, but locally abundant (GA SpecialConcern, 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): granitic flatrocks; uncommon (NC Watch 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 SpecialConcern, VA Watch List). May-June. S. PA west to s. Ontario, OH and MO, south to nw. SC, nw. GA, and c. AL. Plants "withsmaller flowers, shorter pedicels, and smaller capsules and seeds" are the basis of Ph. boykinii and Ph. bicknellii (Constance1949). A dissertation concluded that these plants warranted taxonomic recognition as "Ph. purshii ssp. boykinii," but the name wasapparently never published. {get dissertation, resolve, both taxa in our area?} [= RAB, C, F, G, K, S, W, Z; 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, infraspecific taxa notdistinguished; Ph. dubia var. dubia -- Z, in part only]

Phacelia ranunculacea (Nuttall) Constance. WV. [= K; Ph. ranunculacea -- RAB, C, F, G, Z, in part]

HYPERICACEAE (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 withPodostemum 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 flesh-colored; stamens fascicled, in 3 fascicles of 3 stamens each; staminodia (hypogynous glands) present, alternatingwith the fascicles of stamens; perianth 5-merous; [tribe Cratoxyleae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triadenum

1 Petals yellow; stamens fascicled or not, if fascicled then not into 3 fascicles of 3 stamens each; staminodia (hypogynousglands) lacking; perianth 4-5-merous; [tribe Hypericeae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypericum

Hypericum Linnaeus (St. John's-wort)

A genus of about 370 species, trees, shrubs, and herbs, primarly temperate. Hypericum in our area is a large, complex, andinteresting genus, with a number of unresolved questions remaining. {The distribution in our area of H. boreale. The distribution inour area of H. ellipticum. The curious bimodal distribution of H. prolificum (more than one taxon?). Basis of Small's H. lobocarpumin Blue Ridge of NC. H. lloydii in VA (it must be)? Overall distribution of H. lloydii. H. nitidum-H. fasciculatum-H. galioides. Andmore....} The species treated in Key B have often been treated in the segregate genus Ascyrum. Evidence from a variety ofdisciplines now suggests that they should be included in Hypericum (Adams & Robson 1961; Calie, Schilling, & Webb 1983;Robson 1996). References: Adams (1973)=Z; Robson (1977, 1981, 1990, 1996, 2001, 2002)=X; 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 texturewhich extends across the petiole; plant a 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 A2 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.

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3 Petals 4; sepals 4 (rarely 2); plant 5-100 cm tall; leaves 2-40 mm long; [subsection Ascyrum] . . . . . . . . . . . . Key B3 Petals 5; sepals 5; plant 50-250 cm tall; leaves (10-) 20-70 mm long; [subsections Centrosperma and Brathydium]

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key C1 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; plant a 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, less than 1 mm wide; inflorescence a compound raceme; [section Brathys] . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key D5 Leaves spreading or ascending, generally multi-nerved, more than 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 astranslucent glandular dots when backlit (except in H. perforatum); sepals and/or petals marked with black glandulardots or lines; [section Hypericum] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key E

6 Capsule 1-locular; stamens separate or connate at the base, but not grouped into fascicles; leaves with translucentglandular dots, without black glandular dots (when backlit); sepals and petals with translucent glandular lines or dotsonly, not marked with black glandular dots or lines.7 Plant a decumbent shrub or suffruticose herb; [section Myriandra, subsections Pseudobrathydium and

Suturosperma] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key F7 Plant an herb; [section Trigynobrathys, and section Myriandra subsection Suturosperma] . . . . . . . . . . . Key G

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

1 Largest leaves 5-14 mm long; sepals 2-5 mm long.2 Capsules 3-4 (-4.5) mm long; seeds brown, 0.6-0.7 mm long, with longitudinal ribbing and alveoli alignment obscure;

largest leaves 11-14 mm long; [plants of ultisol or alfisol pine savannas] . . . . . . . . . . . . . . . . . . . . . . . . . . . H. species 22 Capsules (4-) 6-9 mm long; seeds dark red to black, 0.55-0.6 mm long, with longitudinal ribbing and alveoli alignment

evident; largest leaves 5-11 mm long; [plants of spodosol wet pine flatwoods] . . . . . . . . . . . . . . . . . . . . . H. tenuifolium1 Largest leaves 13-26 mm long; sepals 3-7 mm long.

2 Plant a low shrub to 5 dm tall, forming low rounded clumps; mature capsules ovoid-deltoid, 3-4 mm long, 2-2.5 mm wide,ca. 1.5× as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lloydii

2 Plant an erect shrub to 20 dm tall, with a single main stem, branched above; mature capsules oblong-ovoid, 3-6 mmlong, 1.0-2.5 mm wide, ca. 2-3× as long as broad.3 Mature capsules elliptic-ovoid, usually more than 1.5 mm broad; living leaves with a deep longitudinal groove below

on each side of the midrib, with numerous white papillae present in the groove (visible at 20×); bark somewhatspongy-thickened, soon exfoliating in tissue-thin layers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. fasciculatum

3 Mature capsules oblong, usually 1.5 mm or less broad; living leaves slightly concave below, not grooved, thepapillae inconspicuous, poorly developed; bark thin, even on oldest stems, not spongy, exfoliating in flakes ornarrow strips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. nitidum

[H. brachyphyllum][H. densiflorum var. 3][H. galioides]

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 erectshrub.2 Leaves rounded or subcordate at the base; [plants widespread in our area] . . . . . . . . . . . . . . . . . . . . . . H. crux-andreae2 Leaves cordate-clasping at the base; [plants known from e. GA southwards] . . . . . . . . . . . . . . . . . . . . [H. tetrapetalum]

1 Styles and carpels 2; leaves 1-7 mm wide, mostly cuneate (or if rounded the leaves less than 8 mm long and 3 mm wide);plant an erect or decumbent shrub.3 Pedicels 6-13 mm long, soon reflexed; subtending bractlets located near the last pair of leaves; plant a decumbent shrub,

to 2 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. suffruticosum3 Pedicels 1-5 mm long, erect; subtending bractlets located midway between the base of the flower and the last pair of

leaves; plant an erect or decumbent shrub, mostly 1-15 dm tall.4 Plant an 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; leavesusually variable in size and shape, widest near the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. hypericoides

4 Plant a 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,

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widest above the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. stragulum

[H. microsepalum]

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; [plants of s. SC southwards]; [section Myriandra, subsection Brathydium] . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. myrtifolium

1 Leaves cuneate at the base, oblanceolate, oblong, elliptic, or narrowly elliptic; [plants collectively widespread]; [sectionMyriandra, 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. galioides2 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; [plants introduced in our area, though sometimes appearing native] . . . . . . . . . . . . . . . . . . . H. frondosum3 Flowers (1-) 3-many in terminal cymes; petals 5-10 mm long; sepals 1.5-8 mm long; shrubs to 3 m tall; [plants

native in our area].4 Flowers (1-) 3-7 per inflorescence; capsules (6-) 7-14 mm long; larger leaves (4-) 7-14 mm wide H. prolificum4 Flowers 7-many per inflorescence; capsules (3-) 4.5-6 mm long; larger leaves 3-7 (-11) mm wide.

5 Leaves below the inflorescence only slightly narrower than those of the middle and lower stem; seeds 1.1-1.3 mm long; [plants of Mountain wetlands and uplands] . . . . . . . . . . . . . . . . . . . . . H. densiflorum var. 1

5 Leaves below the inflorescence less than 1/2 as wide as those of the middle and lower stem; seeds 0.8-1.0 mm long; [plants of Coastal Plain wetlands] . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. densiflorum var. 2

[H. lobocarpum][H. densiflorum var. 3]

Key D -- herbaceous St. John's-worts with leaves ascending or appressed, 1-nerved, less than 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 thelateral branches typically much smaller; leaves punctate primarily with translucent glands; [plant alien, usually in disturbedhabitats] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 thelateral branches nearly to quite as large; leaves punctate with black glands; [plant native, in a variety of habitats].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; [plants 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. pseudomaculatum3 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. punctatum2 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; [plants endemic to moderate to high elevations of w. NC, sw. VA, and e. TN].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) mmlong; 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

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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

[H. ascyron ssp. pyramidatum]

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 ofleaves; flowers solitary or in small simple cymes; [plants endemic to rock outcrops at moderate to high elevations in theMountains 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; [plants mostly of the Coastal Plain and Piedmont, very rarely in the Mountains and then at lowelevations]; [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. cistifolium2 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 whwn 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

[H. dolabriforme][H. sphaerocarpum]

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. setosum1 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. adpressum3 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. ellipticum2 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 from 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 leafand toward the midrib; seeds (0.55-) 0.6-0.8 (-0.9) mm long; midstem leaves mostly broadest at or beyond themiddle, the apex usually obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. virgatum

5 Punctate glands frequent on the stem; punctate glands of the leaves and stem large, oval, distributed evenlyand 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); seeds (0.45-) 0.5-0.7 mm long; midstem leaves usuallybroadest at or below the middle, the apex acute to acuminate, or, obtuse (if so then the leaves 5-20 (-24) mmlong, 5-15 mm wide, 1.5-3× as long as wide, ovate to obovate to narrowly elliptic, mostly appressed to thestem, mostly shorter than the internodes).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; seeds 0.55-0.7 mm long; [plants of shallow soilmats 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 pairsof very small bracts; seeds (0.45-) 0.5-0.8 (-0.9) mm long; [plants of Coastal Plain wetlands, very rarelydisjunct 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; seeds (0.45-) 0.5-0.65 mm long; lower stem not spongy-thickened withaerenchymatous tissue; [plants of moist pinelands of the Coastal Plain, very rarely disjunct inland tobog 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-

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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; seeds 0.5-0.8 (-0.9) mm long; lanceolate to linear-lanceolate;upper portion of stem with numerous axillary branches; lower stem usually spongy-thickened withaerenchymatous tissue; [plants of upland depression ponds of the Coastal Plain, growing whereseasonally inundated] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. harperi

4 Styles 0.5-1.5 mm long; stamens 5-22.8 Leaves lanceolate to linear, 6-30 mm long, 0.5-3 mm wide, the leaf base attenuate to cuneate . H. canadense8 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 theinflorescence, giving the inflorescence a naked appearance; [plants of Coastal Plain pinelands] . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gymnanthum9 Sepals broadest near the middle; inflorescence with many normally sized leaves and leaflike bracts, giving

the inflorescence a leafy appearance; [plants 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. boreale10 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 internodebelow; sepals broader above the middle, more-or-less imbricate; [plants of the Coastal Plain]

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. mutilum var. latisepalum11 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 essentiallyabsent; sepals broader below the middle, not imbricate (rarely broader above the middle andimbricate); [plants widespread] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. mutilum var. mutilum

[H. majus][H. ×dissimulatum][H. sp. 3]

Hypericum adpressum Rafinesque ex Barton, Bog St.-John's-wort, Creeping St.-John's-wort. Cp (NC, SC, VA): boggydepressions; 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, X, Z]

Hypericum boreale (Britton) Bicknell, Dwarf St.-John's-wort, Northern St.-John's-wort. Cp (VA), Mt (NC?, VA): sinkholeponds 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. [= C, F, G, K; H. mutilum Linnaeus ssp.boreale (Britton) J.M. Gillett -- Y]

Hypericum buckleyi M.A. Curtis, Granite Dome St.-John's-wort. Mt (GA, NC, SC): thin soil in seasonal seepage aroundrock outcrops, particularly granitic exfoliation domes; rare (GA Special Concern, NC Watch List, SC Rare). June-August. Sw. NCsouth 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, X, Z; H. buckleii -- K, orthographic variant]

Hypericum canadense Linnaeus, Canada St. John's-wort Cp, 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. [= RAB, C, F, G, GW, K, S, W, Y, Z; H.canadense var. galiiforme Fernald -- F]

Hypericum cistifolium Lamarck. Cp (NC, SC): pine savannas, wet pine flatwoods; common. June-August. E. NC south tos. FL, west to e. TX. [= RAB, GW, K, S, X, Y, Z; H. opacum Torrey & A. Gray -- S]

Hypericum crux-andreae (Linnaeus) Crantz, St. Andrew's Cross, St. Peter's-wort. Cp, Pd (NC, SC, VA), Mt (NC, SC): dryforests and woodlands, pine flatwoods; common (rare in Piedmont and Mountains). June-October. NY (Long Island) and NJ southto 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, Z; Ascyrum stansMichaux ex Willdenow -- F, G, S; Ascyrum cuneifolium Chapman -- S]

Hypericum densiflorum Pursh var. 1, Mountain Bushy St. John's-wort. Mt, Pd (NC, VA): bogs, streambanks, dry to moistforests, rock outcrops; common (rare in the VA Piedmont). June-August. Sw. PA south to n. GA and c. AL in and near theMountains. The related H. lobocarpum Gattinger is more western, extending east to TN; the basis for attribution of H. lobocarpumto "Blue Ridge, N.C." by Small (1933) is unknown. [H. densiflorum -- RAB, C, GW, K, S, W, Z, in part; H. densiflorum var.densiflorum -- F, G, in part; H. glomeratum Small -- S]

Hypericum densiflorum Pursh var. 2, Coastal Plain Bushy St. John's-wort. Cp (NC, SC): bogs, moist forests, pinesavannas; common. June-August. NJ south to SC in the Coastal Plain. [H. densiflorum -- RAB, C, GW, K, S, W, X, Z, in part; H.densiflorum var. densiflorum -- F, G, in part]

Hypericum denticulatum Walter, Coppery St.-John's-wort. Cp (GA, NC, SC, VA), Pd, Mt (NC): savannas, wet pineflatwoods, 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,

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and in s. AL. See discussion under H. virgatum. [= K, S; H. denticulatum var. denticulatum -- RAB, C, F, G, Z; H. denticulatum --GW, in part only (also see H. virgatum); H. denticulatum ssp. denticulatum -- Y]

Hypericum drummondii (Greville & Hooker) Torrey & A. Gray, Nits-and-lice, Drummond's St.-John's-wort. Pd, Cp, Mt (NC,SC, VA): dry woodlands, woodland borders, fields; uncommon. July-September. MD west to OH, IL, and se. KS, south topanhandle FL and c. TX. [= RAB, C, F, G, GW, K, W, Y, Z; Sarothra drummondii Greville & Hooker -- S]

Hypericum ellipticum Hooker, Pale St.-John's-wort. Mt (NC?, VA): wet places; rare (NC Watch List, VA Rare). Newfoundland and Nova Scotia west to w. Ontario, south to CT, NY, MI, and MN, and in the mountains to WV, VA, ne. TN(Johnson County) (Chester, Wofford, & Kral 1997), and NC (?). The documentation for C's attribution of H. ellipticum to NC isunknown. [= C, F, G, K, X]

Hypericum fasciculatum Lamarck, Peelbark St.-John's-wort. Cp (NC, SC): wet pine savannas, beaver ponds, uplanddepression ponds; uncommon (NC Watch List). May-September. E. NC south to s. FL, west to s. MS. [= RAB, GW, K, X, Y, Z; H.fasciculatum -- S, in part (also see H. nitidum, and also including H. chapmanii P. Adams of the deep south)]* Hypericum frondosum Michaux. Mt (NC, VA): rock outcrops and rocky woodlands; rare, introduced from further south andwest. Late May-July. This species is native and widespread as far east as e. TN (Chester, Wofford, & Kral 1997); it may occur inour area, especially in sw. VA. [= C, F, G, K, W, Z; H. aureum Bartram -- S; H. splendens Small -- S]

Hypericum galioides Lamarck. Cp (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, S, X, Y, Z; H. ambiguum Elliott -- S]

Hypericum gentianoides (Linnaeus) Britton, Sterns, & Poggenberg, Pineweed, Orange-grass. Mt, Pd, Cp (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, Y, Z; Sarothra gentianoides Linnaeus -- S]

Hypericum graveolens Buckley, Mountain St.-John's-wort. Mt (NC): grassy balds, grassy openings, forests, at highelevations (1200 m or more); rare (NC Watch List). July-August. Nw. NC and ne. TN south to sw. NC, a Southern Appalachianendemic. This and the related H. mitchellianum (another narrow endemic to the Southern Appalachians) hybridize, forming localhybrid populations with intermediate characteristics (Culwell {year}). [= RAB, GW, K, S, W, Z]

Hypericum gymnanthum Engelmann & A. Gray, Clasping-leaf St.-John's-wort. Cp (NC, SC, VA), Pd (VA, Mt): pinesavannas, wet pine flatwoods, sinkhole ponds (Augusta and Rockingham counties, VA), other wet to moist habitats; common (rarein 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, ande. KS; also disjunct in Guatemala (introduced?). [= RAB, C, F, G, GW, K, S, Y, Z]

Hypericum harperi R. Keller, Harper's St.-John's-wort. Cp (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 insc. and se. NC, where it may well occur. This species has generally been considered a part of H. denticulatum or H. virgatum, butWebb (1980) makes a convincing argument for its recognition, including the ecological differentiation and absence of intermediatesor hybrids when growing in proximity to H. denticulatum. See H. virgatum for additional discussion. [= Y; H. denticulatum var.acutifolium -- RAB, Z, in part; H. denticulatum -- GW, in part; H. harperi -- K, in part only; H. acutifolium -- S, in part]

Hypericum hypericoides (Linnaeus) Crantz, St. Andrew's Cross. Cp, Pd, 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 theWest Indies, Mexico, and Central America. [= RAB, C, GW, 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; Ascyrumhypericoides Linnaeus -- S; Ascyrum linifolium Spach -- S]

Hypericum lloydii (Svenson) P. Adams, Lloyd's St.-John's-wort. Pd (NC, SC, VA), Cp (NC, SC): dry woodlands, sandhills,edges of granitic flatrocks, roadbanks; uncommon. June-September. Sc. VA south to c. GA. [= RAB, K, X, Z; Hypericum galioidesLamarck var. lloydii Svenson]

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. [= RAB, C, F, G, GW, K, S, W, Z]

Hypericum mutilum Linnaeus var. mutilum, Common Dwarf St.-John's-wort Mt, Pd, Cp (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. [= F; H. mutilum -- RAB, C,F, G, GW, K, S, W, Z, infraspecific taxa not distinguished; H. mutilum ssp. mutilum -- Y]

Hypericum mutilum Linnaeus var. latisepalum Fernald, Southern Dwarf St.-John's-wort. Cp (SC): marshes and other wethabitats; rare. June-October. Se. SC south to peninsular FL, west to TX (and, according to F, north to s. NJ). [= F; H. mutilum --RAB, F, G, GW, K, S, W, Z, infraspecific taxa not distinguished; H. mutilum ssp. latisepalum (Fernald) N. Robson -- Y]

Hypericum myrtifolium Lamarck, Myrtle-leaf St.-John’s-wort. Cp (SC): ponds; rare. Small (1933) reports this species fromSC; 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, X, Y, Z]

Hypericum nitidum Lamarck. Cp (NC, SC): pine savannas; rare (NC Rare, SC Rare). June-August. Se. NC south topanhandle FL, west to sw. AL. [= RAB, GW, K, Y, Z; H. nitidum ssp. nitidum – X]

Hypericum nudiflorum Michaux ex Willdenow. Pd, Cp (NC, SC, VA), Mt (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 CumberlandPlateau of TN. [= RAB, C, F, G, S, W, X, Y; H. nudiflorum -- GW, K, Z, in part (also see H. apocynifolium of the deep South)]* Hypericum perforatum Linnaeus, European St. John's-wort. Mt, Pd, Cp (NC, SC, VA): fields, pastures, roadsides,woodland borders; common, introduced from Europe. June-September. [= RAB, C, F, G, K, S, W, Z; H. perforatum ssp.perforatum – X]

Hypericum prolificum Linnaeus, Shrubby St-John's-wort. Mt, Pd, Cp (NC, SC, VA): bogs, seepages, rocky forests, rock

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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,X, Z; H. spathulatum (Spach) Steudel -- F]

Hypericum pseudomaculatum Bush. Cp, Pd (SC): wet, moist, or dry forests; rare. June-September. SC south topanhandle FL, west to TX, north in the interior to e. TN, c. IL, s. MO, and c. OK. [= RAB, C, G, K, S, Z; H. punctatum Lamarck var.pseudomaculatum (Bush) Fernald -- F]

Hypericum punctatum Lamarck, Spotted St.-John's-wort. Mt, Pd, Cp (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, S, Z; H. punctatum var. punctatum -- F; H. subpetiolatum Bicknell ex Small -- S]

Hypericum setosum Linnaeus. Cp (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, Y, Z]

Hypericum species 1, Radfords' St. John's-wort. Pd (NC): shallow circumneutral soil mats of granitic domes in the BrushyMountains; 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 themedial and upper nodes (rather than being little if at all branched, and then only from the uppermost nodes), in having leaves withacuminate (rather than acute to obtuse) apices, and electrophoretically (Webb 1980). Additionally, these plants have numerousbracteal 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 andstem (vs. lower leaf surface only). It may be notable that these same outcrops are phytogeographically interesting, with otherdisjunct and weakly differentiated races (see Allium cuthbertii). Further study is planned. [included in concept of H. virgatum (= H.denticulatum var. acutifolium, H. denticulatum ssp. acutifolium) by most earlier authors]

Hypericum species 2. Cp (NC, SC?): wet pine savannas, over ultisols or alfisols; rare? June-September. [included inconcept of H. tenuifolium (=reductum) by earlier authors]

Hypericum stragulum P. Adams & Robson. Mt, Pd (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, W, Z; H. stragalum -- RAB, misspelling; Ascyrumhypericoides 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 (NC, SC): pine savannas and flatwoods; rare(NC Rare). April-June. Se. NC south to c. peninsular FL, west to se. LA. [= RAB, K, Y, Z; Ascyrum pumilum Michaux -- S]

Hypericum tenuifolium Pursh, Sandhill St.-John’s-wort. Cp (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 oldername H. tenuifolium Pursh has now been adequately shown to apply to this taxon. [= X; H. reductum (Svenson) P. Adams -- RAB,GW, K, Y, Z; H. aspalathoides Willdenow -- S, in part (also including H. brachyphyllum)]

Hypericum virgatum Lamarck. Pd, Mt (NC, SC, VA), Cp (SC): woodlands, rock outcrops, woodland borders; common (VAWatch 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, andse. LA. Though treated by most recent authors as a variety of H. denticulatum, H. virgatum is better considered as a distinctspecies. Webb (1980) recognized H. harperi as a separate species (it had previously been considered a part of H. virgatum), andcontinued to recognize this taxon as a variety of H. denticulatum. However, based on the nature of the punctate glands, size ofseeds, inland distribution, etc., it appears that H. virgatum is more distantly related to H. denticulatum and H. harperi than they areto one another; recognition at the species level is warranted for H. virgatum. As pointed out by Webb, H. denticulatum is primarilytetraploid (n = 24), while H. virgatum and H. harperi are (as far as is known) strictly diploid. Additionally, the aberrant populationsfrom granitic outcrops in the Brushy Mountains of Alexander and Wilkes counties, NC referred by Webb (1980) to this taxon aredistinct, 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, in part (also see H. harperi); H. denticulatum var.recognitum Fernald & Schubert -- RAB, F; H. denticulatum -- GW, in part; H. acutifolium Elliott -- S, in part (also see H. harperi); H.denticulatum ssp. acutifolium (Elliott) N. Robson -- Y]

Hypericum apocynifolium Small. c. GA. [= S, X, Y; H. nudiflorum -- GW, K, in part]

Hypericum ascyron Linnaeus ssp. pyramidatum (Aiton) N. Robson, Great St.-John’s-wort. South to s. PA (Rhoads & Klein 1993),MD (Robson 2000), and WV (Kartesz 1999). [H. ascyron Linnaeus -- K; H. pyramidatum Aiton -- C] {not yet keyed} {sectionRoscyna}

Hypericum brachyphyllum (Spach) Steudel. Ranges from e. GA (near SC), and also reported from SC (Kartesz 1999); needsconfirmation. [= GW, K, X, Y]

Hypericum densiflorum Pursh var. 3, Interior Bushy St. John’s-wort. Occurs in e. and c. TN, nw. GA south to c. AL. [H. interiorSmall; H. revolutum Keller] {not yet keyed}

Hypericum ×dissimulatum Bicknell (pro sp.). (NC, VA). [= C; H. dissimulatum Bicknell -- K]

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, K, X]

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Hypericum lobocarpum Gattinger. East to c. TN (Chester, Wofford, & Kral 1997). Credited to SC by Robson (1996), based onspecimen debated and dismissed by Adams. [= C, K, X; H. densiflorum -- GW (in part)]

Hypericum majus (A. Gray) Britton. South to nw. PA (Rhoads & Klein 1993), NJ, DE, and OH (Kartesz 1999). [= C, K]

Hypericum microsepalum (Torrey & A. Gray) A. Gray ex S. Watson. North to se. GA. [= GW, K, X, Y]

Hypericum species 3 J. Allison, Georgia St, John’s-wort. Cp (GA): seepage bogs, roadside ditches; rare (GA Special Concern). {not yet keyed}

Hypericum sphaerocarpum Michaux, Barrens St. John’s-wort. Mt (GA): limestone barrens; rare (GA Special Concern). East toGA (GAHP 2003), e. and c. TN (Chester, Wofford, & Kral 1997), sw. PA (Rhoads & Klein 1993). [= C, K, X; H. turgidum Small] {not yet keyed}

Hypericum tetrapetalum Lamarck, ranges from e. GA (within a few counties of se. SC), south to s. FL, west to panhandle FL. Itoccurs in wet pinelands and in depressional wetlands (open or dominated by Taxodium ascendens). [= GW, K, X, Y; Ascyrumtetrapetalum (Lamarck) Vail]

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 inHypericum, Robson (1977) and others consider it to be more closely related to the tropical Asian shrub Cratoxylum Blume.

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. tubulosum2 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 5-8 mm long at maturity, acute or acuminate; styles 1.8-3 mm long (best seen in fruit) . . . . . . . . . . T. virginicum3 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

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, 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 (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 northin the interior to se. and c. TN, s. IL and s. OH. [= C, GW, K, Z; Hypericum tubulosum Walter -- RAB; Hypericum tubulosum Waltervar. tubulosum -- F; T. longifolium Small -- S]

Triadenum virginicum (Linnaeus) Rafinesque, Marsh St.-John's-wort. Cp, Mt (NC, SC, VA), Pd (VA): bogs, peatywetlands; common (rare in Mountains and Piedmont). July-September. Nova Scotia west to OH and s. Ontario, south to FL andMS, mostly on the Coastal Plain but scattered inland. [= C, GW, K, S; Hypericum virginicum Linnaeus -- RAB; Hypericumvirginicum 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 (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 theinterior to s. MO, s. IL, and OH. [= C, 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)

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).

1 Flowers 2-5-5 cm across; tepals 21-33, red-maroon (rarely white or pinkish); leaves tips acute to acuminate . . . . I. floridanum1 Flowers 0.8-1.2 cm across; tepals 11-16, yellowish green; leaf tips obtuse or rounded . . . . . . . . . . . . . . . . . . . . I. parviflorum

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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). This species occurs in swampy forests, evergreen hammocks, and bayheads and is endemic to scatteredlocalities 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 (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), anarrow Escalloniaceae, or a very narrow (single genus) Iteaceae (S), the relationships of Itea remain problematic. Recentmolecular data suggest that the relationship between Itea and other woody "saxifragaceous" genera (including Escallonia) is onlydistant (Morgan & Soltis 1993). Itea is here conservatively treated in a narrow Iteaceae. The remainder of the genus (about 20species) 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 contrastwith 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 (Walnut Family)

A family of about 8 genera and 60 species, trees and shrubs, mostly temperate. References: Stone in FNA (1997); 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 theterminal or final 2 lateral; nut with shell smooth, ridged, or irregularly wrinkled (but not deeply furrowed); terminal buds withimbricate or valvate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 corrugatedpattern; terminal buds with imbricate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juglans

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

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 twosections, 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 ofCarya. Our three-state area includes 11 of 13 North American species (including the naturalized C. illinoinensis and lacking only amore 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 genusare 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 et al.(1969), it seems possible that reticulate evolution involving extant or extinct diploid species is responsible for some of the difficultiesin 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 beenrecognized".

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 ourarea (in particular the Piedmont) as being characterized by "oak-hickory" or "oak-pine-hickory" forests (e.g. Küchler 1964; Greller

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1988; Schafale & Weakley 1990; Skeen, Doerr, & Van Lear 1993). Ware () 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 greatercation 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 as1820 in an account of the landscape of North Carolina. "The sandy pine barrens, and all the lands on which pine is the exclusivegrowth, are unfriendly to agriculture; but where the pine is intermixed with oak and hickory, the soil is good. Some of our strongestlands 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 asconsisting of between 1 and 10 (or more) taxa. For testing, we present here a plausible middle ground treatment in the key andspecies accounts (4 taxa). References: Stone in FNA (1997); Stone & Hardin in SE (in prep.); 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 Stone & Hardin inSE (in prep.).

Identification notes: Surface vestiture of leaves and bud scales is useful in distinguishing species of Carya. Some use of thesecharacters can be made with a 10× or 20× hand lens; better still is a dissecting microscope. It is important to understand thedifferent trichome types mentioned in the key (terminology follows Hardin 1990 and Hardin & Stone 1984). Short aciculartrichomes are simple, unicellular trichomes tapered to a pointed tip, 0.10-0.35 mm long and with rough walls. Long aciculartrichomes ("solitary" of Hardin & Stone 1984) are similar to short acicular, but are much larger, 0.45-1.6 mm long, and havesmooth walls. Fasciculate trichomes are multicellular and have 2-8 straight or curled rays radiating a clustered base. Multiradiate trichomes are similar to fasciculate, but have 8-17 rays, the inner (and usually more upright) rays attached basallyabove the outer (and usually more spreading) rays. Capitate glandular trichomes are unicellular or multicellular, and aredistinguished by their bulbous or expanded tip; they are usually 0.02-0.1 mm long. Peltate scales are flat or dome-shaped shieldsor disks, slightly to strongly glandular, (sometimes regularly or irregularly lobed) and can be either sessile or stalked (they are oftenreferred to as scales, resin dots, peltate glands, or lepidote scales). On the lower surfaces of leaflets, peltate scales are of twotypes: large peltate scales are 0.08-0.3 mm in diameter and are round, with smooth or slightly irregular margins, while smallpeltate 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 suturesnarrowly 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 ourarea, 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 (-15) leaflets; bark shaggy; lateral petiolules 0-2 mm long; nut flattened in cross-section; kernel

bitter; [native tree of swamp forests, primarily in the Coastal Plain] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aquatica3 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 tree 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 suturesnot winged.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 sloughingwith 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 ondrying); lower surface of leaflets nearly glabrous, except for tufts of trichomes in the main vein axils, and only slightlylepidote 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 mmlong, tomentose, tan to brown (rarely turning black on drying); lower surface of leaflets moderately to densely hirsutewith acicular and fasciculate hairs (sometimes the hairs more or less limited to the main veins), and also moderatelylepidote, the large peltate scales yellow and round, the small peltate scales dark brown and mostly round; terminalleaflet (4-) 6-15 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ovata

4 Bark tight (the ridges typically forming an interlocking diamond pattern), scaly, or shaggy (when shaggy, the separatedsegments normally much less than a meter long); leaves with (3-) 5-9 (-11) leaflets; serrations of the leaflets glabrous orciliate, 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 ofthe 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

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long; nut 2.5-3.5 cm long; [trees common in our area] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. alba7 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 cmlong; nut 3-6 cm long; [trees 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 mostlyglabrous, 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 ofleaflets of various types, 4-lobed and/or irregular scales often more frequent than round scales; fruit husk 2-5 mmthick; 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 theundersurface a reflective, silvery-tan 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; [trees of calcareous swamps, bottomlands and slopes of the CoastalPlain of se. NC and e. SC] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. myristiciformis

9 Lepidote scales silvery-tan; [trees of upland and usually acidic frests and woodlands, widespread in ourarea] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pallida

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-lobedpeltate scales uncommon).10 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. ovalis10 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.11 Rachis (and also often the petiole and lower surfaces of leaflets) densely pubescent . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glabra var. hirsuta11 Rachis, petiole, and lower surfaces of leaflets glabrous or glabrescent.

12 Husk of fruit 1.5-2.5 mm thick; fruit ellipsoidal, subglobose, to obovoid, 1.5-3.5 cm long; terminalleaflet usually 8-17 cm long; [trees primarily of the Piedmont and Mountains] . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glabra var. glabra12 Husk of fruit about 3.5 mm thick; fruit pyriform, 2.5-5 cm long; terminal leaflet usually 20-25 cm

long; [trees primarily of the Coastal Plain] . . . . . . . . . . . . . . . . . . . . C. glabra var. megacarpa

Carya alba (Linnaeus) Nuttall ex Elliott, Mockernut Hickory, White Hickory. Pd, Cp, Mt (NC, SC, VA): forests andwoodlands; common. April-May; October. MA west to IN and IA, south to FL and TX. One of the most common forest trees ofmuch of our area. There has been confusion and controversy for several centuries over the specific epithet. The oldest basionymavailable is Juglans alba Linnaeus, which apparently included disparate elements, including this taxon and C. ovata. Following amore circumscribed typification by Crantz in 1766, the epithet "alba" should have been applied to this taxon, but continued to beapplied in various ways. Rehder (1945) proposed that C. alba should be considered a nomen ambiguum, but agreed that it appliedcorrectly to what has often been called C. tomentosa. He argued that the use of C. alba should be rejected "in order to avoidconfusion and ambiguity." However, C. alba has not been officially rejected, its application now appears to be nomenclaturallyclear, and there is no alternative under the International Code of Botanical Nomenclature (Greuter 1988) to its use. For furtherdiscussion see Rehder (1945), Howard & Staples (1983), and Wunderlin, Hansen, & Hall (1985). [= K; C. tomentosa (Lamarck exPoiret) Nuttall -- RAB, C, F, FNA, G, W; Hicoria alba (Linnaeus) Britton -- S]

Carya aquatica (Michaux f.) Nuttall, Water Hickory, Bitter Pecan. Cp (NC, SC, VA), Pd (SC): swamp forests, where floodedduring 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 (NC, SC,VA): upland flats, especially those weathered from mafic rocks and with shrink-swell soils dominated by montmorillonitic clays, lesstypically on slopes and bottomlands; uncommon (rare in VA). April-May; October. Sc. VA (Halifax County) south to GA, AL, andMS, and inland northwards 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. Itseems to us morphologically and ecologically distinctive. Hardin & Stone (1984) found differences in trichomes, and in a study ofnut oils, Stone, Adrouny, & Flake (1969) found C. ovata "surprisingly distant" from C. carolinae-septentrionalis. There are reportsthat 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 segregated, the two species sometimes occurtogether 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 (NC, SC, VA): forests and woodlands, especiallyin rich, moist alluvial or slope forests; common. April; October. ME and s. Quebec west to MN and NE, south to panhandle FL ande. 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 (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 rangesof the varieties are poorly known. [= F, G; C. glabra -- RAB, C, FNA, GW, K, infraspecific taxa not distinguished; C. glabra var.

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glabra -- W, in part; Hicoria glabra (P. Miller) Britton -- S; Hicoria microcarpa (Nuttall) Britton -- S]Carya glabra (P. Miller) Sweet var. hirsuta (Ashe) Ashe, Hairy Pignut Hickory. Mt, Pd? (NC, SC, VA): forests and

woodlands; common? April-May; October. Var. hirsuta is apparently primarily Appalachian, variously described as being endemicto the Southern Appalachians or ranging north to s. NH. [C. glabra -- RAB, C, FNA, GW, K, infraspecific taxa not distinguished; C.ovalis (Wangenheim) Sargent var. hirsuta (Ashe) Sargent -- F; C. glabra var. glabra -- W, in part]

Carya glabra (P. Miller) Sweet var. megacarpa (Sargent) Sargent, Coastal Pignut Hickory. Cp (NC, SC, VA?): maritimeforests and other forests of the outer Coastal Plain; uncommon? April-May; October. Var. megacarpa is apparently primarily a treeof the se. United States Coastal Plain, ranging from s. NY south to FL, west to TX, and north in the interior to s. IL. [= F, G; C.glabra -- RAB, C, FNA, GW, K, infraspecific taxa not distinguished]* Carya illinoinensis (Wangenheim) K. Koch, Pecan. Cp, Pd (NC, SC, VA): persistent around dwellings and in pecanorchards, 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 ofthe specific epithet has been a source of controversy. [= C, FNA, K; C. illinoensis -- RAB, F, G, GW, orthographic variant; Hicoriapecan (Marshall) Britton -- 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 sometimesplanted, 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 calcareoussubstrates, including calcareous clays and coquina limestone ("marl"), oak flatwoods; rare (GA Special Concern, NC Threatened,SC Rare). April; October. Se. NC south to SC, and from wc. AL west to e. TX and se. OK (apparently absent in GA); and disjunctin Mexico (Nuevo Léon and Tamaulipas). The bronze sheen of the leaflets of this species is diagnostic. First reported for NC byLeonard (1971b). [= FNA, K; C. myristicaeformis -- RAB, GW, orthographic variant; Hicoria myristicaeformis (Michaux f.) Britton --S, orthographic variant; Hicoria myristiciformis (Michaux f.) Britton]

Carya ovalis (Wangenheim) Sargent, Red Hickory. Mt, Pd, Cp (NC, SC, VA): forests and woodlands; common. April-May;October. MA west to WI, south to GA, MS, and MO. [= RAB, C, F, K; C. ovalis var. obcordata (Muhlenberg & Willdenow) Sargent -- F, G; C. ovalis var. obovalis Sargent -- F, G; C. ovalis var. odorata (Marshall) Sargent -- F, G; C. glabra -- FNA, in part; C. glabra(P. Miller) Sweet var. odorata (Marshall) Little -- W]

Carya ovata (P. Miller) K. Koch, Common Shagbark Hickory. Mt, Pd, Cp (NC, VA): rich moist bottomlands, slopes,occasionally on dry upland flats; uncommon. May; October. S. ME and s. Quebec west to MN and NE, south to GA and TX; alsodisjunct in Mexico. [= RAB, C, F, G, GW, K, W; 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 (NC, SC, VA), Mt (NC, SC): dry sandy orrocky 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 is reported to occur as far east as KY, TN, and GA (Kartesz 1999), an eastern extent not reported by FNA. {investigate} [= K; C. texana var. arkansana (Sargent) Little – C] {not yet keyed}

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 distantlyrelated within the genus, with J. nigra most closely related to sw. North American J. microcarpa and J. major, and J. cinerea mostclosely 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).

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 peltatescales 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-richforests; 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, whichnow threatens its continued existence. [= RAB, C, F, FNA, G, K, W; Wallia cinerea (Linnaeus) Alefeld -- S]

Juglans nigra Linnaeus, Black Walnut. Mt, Pd, Cp (NC, SC, VA); moist, nutrient-rich forests; common (uncommon in theCoastal Plain of NC and SC). April; October. MA west to MN, south to nw. FL and TX. The dark brown wood is famous forcabinetry and other uses; it is one of the most prized of North American hardwoods. The nuts, though difficult to crack, are prizedfor their intense flavor. The husk is used as a dye. Country people dehusk the nuts by putting them in dirt or gravel drivewayswhere the passage of car tires removes the husk but does not crack the nut. [= RAB, C, F, FNA, G, K, W; Wallia nigra (Linnaeus)

Flora of the Carolinas, Virginia, and Georgia, Working Draft of March 17, 2004 -- KRAMERIACEAE

390

Alefeld -- S]

KRAMERIACEAE (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 Americato 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 eastwards in the Coastal Plain of FL and GA (east to Bulloch, Bryan, Evans, andEmanuel counties, GA). [= K, Z; K. spathulata Small -- S]