Part 7 - UNC Herbarium

ORCHIDACEAE 836 

Zeuxine Lindley 1826 (Soldier Orchid) 

A genus of about 26 species, of tropical and subtropical Old World (introduced elsewhere). References: Ackerman in FNA 

(2002a). 

* Zeuxine strateumatica (Linnaeus) Lindley, Lawn Orchid, Soldier Orchid. Cp (GA): lawns, rare, introduced from Asia. [= 

FNA, GW, K, L] 

POACEAE (R. Brown) Barnhart 1895 or GRAMINEAE A.L. de Jussieu 1789 (Grass Family) 

A family of about 670 genera and 10,000 species, cosmopolitan. References: Flora of North America Editorial Committee 

(2003a)=FNA; Hitchcock and Chase (1950)=HC; Blomquist (1948). 

[note: only a small portion of the key to genera complete] 

Key A – tribe Andropogoneae 

1 Leaves ovate-lanceolate, 2-10 cm long, 2.5-7× as long as wide; plants weak-stemmed annuals, branching, decumbent, 

rooting at the lower nodes; [alien weeds]. 

2 Leaves cordate-clasping at base; spikelets not paired, unaccompanied by a vestige.......................................... Arthraxon 

2 Leaves tapering to a broadly cuneate base; spikelets paired (one of the pair sometimes vestigial)...............Microstegium 

1 Leaves lanceolate to linear, either longer or proportionately narrower; plants either perennial or coarse annuals with erect 

and mostly unbranched culms. 

3 Spikelets embedded in the thickened rachis (the inflorescence thus like an ear of corn), or fitting into grooves in the 

thickened rachis (the inflorescence thus cylindrical and resembling a rat's tail), or the pistillate inflorescences enclosed 

in a hard, bead-like, pearly-white, modified bract. 

4 Spikelets unisexual, with male and female spikelets in separate inflorescences or in different parts of the same 

inflorescence. 

5 Internode narrower than and more-or-less enclosed by the female spikelet .................................................Coix 

5 Internode broader than and more-or-less enclosing the female spikelet. 

6 Racemes of mixed sex, female below, male above ....................................................................Tripsacum 

6 Racemes of single sex ........................................................................................................................... Zea 

4 Spikelets, or at least one of each pair, bisexual. 

7 Pedicels fused to the internode; [coarse alien grass of disturbed habitats] .........................................Rottboellia 

7 Pedicels free from the internodes; [either a native coarse grass of pinelands or prairie-like areas, or a short 

alien grass of lawns and disturbed areas]. 

8 Sessile spikelet smooth or pitted; culms 50-200 cm tall; [native grass of pinelands or prairie-like areas] .. 

.................................................................................................................................................Coelorachis 

8 Sessile spikelet with pectinate margins; culms 5-40 cm tall; [alien grass of lawns and disturbed areas] .... 

................................................................................................................................................ Eremochloa 

3 Spikelets not embedded or fitting into grooves in the rachis, the rachis slender (the spikelets visibly separate and often 

pedicelled). 

9 Pedicelled spikelet similar to the sessile spikelet, both fertile. 

10 Spikelets falling in pairs together with sections of the disarticulating rachis ....................................Saccharum 

10 Spikelets falling separately from the persistent rachis. 

11 Panicle contracted, spikelike; glumes membranous ..................................................................... Imperata 

11 Panicle loose; glumes cartilaginous or coriaceous.................................................................... Miscanthus 

9 Pedicelled spikelet differing from the sessile in shape and sex (sometimes represented only by a pedicel). 

12 Spikelets awned, the awn 10-20 cm long. 

13 First glume lacking glands; panicle open, the branches 5-8 cm long .....................................Chrysopogon 

13 First glume with a row of punctate, concave glands; panicle contracted, spikelike................ Heteropogon 

12 Spikelets awned or not, if awned the awn < 5 cm long. 

14 Inflorescence a panicle, the branches not subtended by sheaths. 

15 Pedicelled spikelet represented by pedicel only; apex of sheath bearing 2 auricles 1-10 mm long; 

[native] ........................................................................................................................... Sorghastrum 

15 Pedicelled spikelet present, staminate; apex of sheath truncate; [alien] ............................... Sorghum 

14 Inflorescence of 1-13 digitate (whorled) racemes borne at the summit of a peduncle, the peduncle 

subtended by a raceme sheath. 

16 Racemes 1 per peduncle and raceme sheath................................................................ Schizachyrium 

16 Racemes 2-13 per peduncle and raceme sheath.

POACEAE 837 

17 Pedicels of the pedicelled (reduced or absent) spikelets terete or slightly flattened and grooved 

on one side only....................................................................................................... Andropogon 

17 Pedicels of the pedicelled (reduced or absent) spikelets strongly flattened and grooved on both 

sides, the central portion thin or membranous ........................................................Bothriochloa 

References: Tucker (1996)=Z. 

Aegilops Linnaeus 1753 (Goat Grass) 

1 Spikelets cylindric; glumes with 4 awns; rachis disarticulating at maturity..........................................................Ae. cylindrica 

1 Spikelets nearly ovate; glumes with 1 awn; rachis not disarticulating at maturity.................................................. Ae. neglecta 

* Aegilops cylindrica Host, Jointed Goat Grass. Mt, Pd (VA): disturbed areas; uncommon, introduced from s. Europe. [= C, 

F, G, HC, K, Z] 

* Aegilops neglecta Req. ex Bertoloni, Small Goat Grass. Cp (VA): disturbed areas; rare, introduced from s. Europe. 

Reported from Arlington County, VA. [= Z; Ae. ovata Linnaeus – C, G, HC, apparently misapplied; Ae. geniculata Roth – K, 

apparently misapplied] 

* Aegilops triuncialis Linnaeus. {MD} [= K] {not keyed at this time; synonymy incomplete} 

Agropyron Gaertner 1770 

(see Elymus, Elytrigia, Pascopyrum) 

Agrostis Linnaeus 1753 (Bentgrass) 

(also see Lachnagrostis and Polypogon) 

A genus of about 220 species, primarily temperate. References: Tucker (1996)=Z. 

1 Palea 1/2-3/4 as long as the lemma, 0.6-1.2 mm long; plants introduced, often (though not always) in disturbed habitats; 

plants flowering (collectively) June-October; [subgenus Agrostis]. 

2 Ligule mostly 0.5-2 mm long, truncate; panicle branches naked towards the base, diffuse when in fruit, the spikelets 

well-separated ..................................................................................................................................................A. capillaris 

2 Ligule mostly 2.5-6 mm long, acute, rounded, or truncate; panicle branches (some of them) with spikelets to near the 

base, the spikelets usually agglomerated. 

3 Leaves 3-8 mm wide; inflorescence triangular-ovoid, the branches widely spreading at maturity, usually reddish; 

plant with rhizomes, without stolons.........................................................................................................A. gigantea 

3 Leaves mostly 1-3 mm wide; inflorescence narrowly ovoid, the branches ascending at maturity, usually tan; plant 

without rhizomes, with or without stolons. 

4 Stolons well developed; leaves mostly < 5 cm long................................................A. stolonifera var. palustris 

4 Stolons poorly developed or absent; leaves mostly > 7 cm long......................... A. stolonifera var. stolonifera 

1 Palea < 2/5 as long as the lemma, 0-0.5 mm long; plants native, typically in more or less natural habitats; plants flowering 

(collectively) March-November; [subgenus Vilfa]. 

5 Lemma usually awned (sometimes unawned), the awn inserted near the apex, 4-10 mm long, straight, very delicate and 

flexuous; annual, flowering April-June.......................................................................................................... A. elliottiana 

5 Lemma awned or not, the awn (when present) inserted either near the middle of the lemma or near the apex, 0-6 mm 

long, straight or bent, neither delicate nor flexuous; perennial, flowering (collectively) March-November. 

6 Lemma with a 3-6 mm long, geniculate awn inserted near the middle; [of high elevation rock outcrops] ................. 

.................................................................................................................................................................A. mertensii 

6 Lemma awnless or with a 0-3 mm long, straight awn inserted near the tip; [of various habitats]. 

7 Spikelets 1.2-2 mm long; anthers 0.3-0.6 mm long; plants flowering March-July........................... A. hyemalis 

7 Spikelets (1.8-) 2.2-3.5 (-3.7) mm long; anthers (collectively) 0.3-1.2 mm long; plants flowering June- 

November. 

8 Leaves mostly involute, 1-2 (-3) mm wide; panicle branches mostly forking well beyond the middle....... 

.....................................................................................................................................................A. scabra 

8 Leaves flat, 2-6 mm wide; panicle branches mostly forking at or below the middle. 

9 Lemma 1.8-3 mm long, minutely but copiously scabrous (at 20× or more); anthers 0.7-1.2 mm 

long; spikelets (2.3-) 2.7-3.5 (-3.7) mm long, usually clustered near the tips of the branchlets; 

panicle branches scabrous; culms to 15 dm tall; [of wet savannas and other wet habitats of the 

Coastal Plain] ................................................................................................................... A. altissima 

9 Lemma 1.4-2 mm long, glabrous; anthers 0.3-0.6 mm long; spikelets (1.8-) 2.2-2.7 (-3.2) mm long, 

usually not clustered near the tips of the branchlets; panicle branches glabrous to scabrous; culms to 

10 dm tall; [of various habitats, nearly throughout our area]..........................................A. perennans

POACEAE 838 

Agrostis altissima (Walter) Tuckerman, Coastal Bog Bentgrass. Cp (GA, NC, SC, VA), Mt (VA): wet savannas, sinkhole 

ponds, edges of swamp forests; rare (NC Watch List, VA Watch List). October-November. MA (?) and NJ south to se. LA, 

primarily on the Coastal Plain. [= F, HC, Z; < A. perennans – RAB, FNA GW, K; = A. perennans var. elata (Pursh) A. 

Hitchcock – C, G, S] 

* Agrostis capillaris Linnaeus, Rhode Island Bentgrass, Colonial Bentgrass, Browntop. Mt (NC, SC, VA), Pd, Cp (VA): 

meadows, roadsides, disturbed areas; uncommon, introduced from Europe (and possibly n. North America). June-August. [= C, 

FNA, K, Z; = A. tenuis Sibthorp – RAB, G, HC, S, W; > A. tenuis var. tenuis – F] 

Agrostis elliottiana J.A. Schultes, Elliott's Bentgrass, Southern Bentgrass. Pd, Cp (GA, NC, SC, VA), Mt (GA, NC, SC): 

dry soils of barrens, fields, and rock outcrops; uncommon (VA Watch List). April-June. MD west to s. OH, and e. KS, south to 

panhandle FL and c. TX. [= RAB, C, F, FNA, G, HC, K, S, W, Z] 

* Agrostis gigantea Roth, Redtop, Black Bentgrass. June-October. Mt, Pd (GA), {provinces} (SC, VA). [= C, F, FNA, K, 

W, Z; < A. stolonifera – RAB, GW; = A. stolonifera Linnaeus var. major (Gaudin) Farwell – G; = A. alba – HC, misapplied; >< 

A. alba – S, misapplied] 

Agrostis hyemalis (Walter) Britton, Sterns, & Poggenburg, Ticklegrass, Small Bentgrass. Mt, Pd, Cp (GA, NC, SC, VA: 

roadsides, other disturbed habitats; common. March-July. [= F, FNA, K, Z; < A. hyemalis – RAB (also see A. scabra); = A. 

hyemalis var. hyemalis – C, G; = A. hiemalis – GW, HC, orthographic variant; < A. hiemalis – S, W, orthographic variant (also 

see A. scabra var. scabra] 

Agrostis mertensii Trinius, Arctic Bentgrass. Mt (NC, VA): in thin soil of high elevation rocky summits; rare (NC Rare). 

July-August. Circumboreal, in North America south to ME (Mt. Katahdin), NH (White Mountains), VT, NY (Adirondack 

Mountains), WV (Spruce Knob), TN (Roan Mountain, Mt. Leconte), NC (Roan Mountain, Big Yellow Mountain, Black 

Mountains), Québec, British Columbia, CO, UT (?), and AK. [= C, FNA, K, W, Z; > A. borealis Hartman – RAB, HC, S; > A. 

borealis Hartman var. americana (Scribner) Fernald – F, G] 

Agrostis perennans (Walter) Tuckerman, Upland Bent, Autumn Bentgrass. Mt, Pd, Cp (GA, NC, SC, VA): . August- 

October. [= HC, Z; < A. perennans – RAB, FNA, GW, K, W (also see A. altissima); = A. perennans var. perennans – C, G, S; > 

A. perennans var. perennans – F; > A. perennans var. aestivalis Vasey – F] 

Agrostis scabra Willdenow, Fly-away Grass, Rough Bentgrass. (GA, NC, SC, VA). (VA Watch List). June-November. [= 

FNA, GW, K, Z; < A. hyemalis – RAB, W; = A. hyemalis (Walter) Britton, Sterns, & Poggenburg var. scabra (Willdenow) 

Blomquist – C; > A. scabra var. scabra – F; = A. hyemalis (Walter) Britton, Sterns, & Poggenburg var. tenuis (Tuckerman) 

Gleason – G; = A. scabra var. scabra – HC] 

* Agrostis stolonifera Linnaeus var. palustris (Hudson) Farwell, Creeping Bentgrass. June-October. [= C; < A. stolonifera – 

RAB, FNA GW, W (also see A. gigantea); = A. alba Linnaeus var. palustris (Hudson) Persoon – F, misapplied; = A. stolonifera 

var. compacta Hartman – G; = A. palustris Hudson – HC, Z; < A. stolonifera – K; < A. alba – S, misapplied] 

* Agrostis stolonifera Linnaeus var. stolonifera. June-October. [= C, G; < A. stolonifera – RAB, FNA, GW, W (also see A. 

gigantea); = A. alba Linnaeus var. alba – F, misapplied; = A. stolonifera – HC, Z; < A. stolonifera – K; >< A. alba – S, 

misapplied] 

* Agrostis canina Linnaeus, Brown Bentgrass, Velvet Bentgrass, ranges south to DE, se. PA (Rhoads & Klein 1993), WV, 

and TN (Kartesz 1999). [= C, FNA, K] {not keyed at this time; synonymy incomplete} 


Aira Linnaeus (Hair Grass) 

1 Panicle dense and spike-like, the branches short and appressed to ascending ...........................................................A. praecox 

1 Panicle open, the branches elongate, diffusely spreading or ascending. 

2 Lemma of both the lower floret and the upper floret with an awn 2-4 mm long........................................A. caryophyllea 

2 Lemma of upper floret with an awn 1.5-2.5 mm long, lemma of the lower floret awnless or with a minute awn < 1 mm 

long ...........................................................................................................................................................A. elegantissima 

* Aira caryophyllea Linnaeus, Silver Hair Grass. Pd (GA, NC, SC, VA), Cp (GA, NC, VA), Mt (NC): fields, roadsides, 

disturbed areas; uncommon, introduced from Europe. May. [= RAB, C, G, HC, K, Z; = Aspris caryophyllea (Linnaeus) Nash – 

S] 

* Aira elegantissima Schur, Elegant Hair Grass. Pd, Cp (GA, NC, SC, VA), Mt (GA, SC): fields, roadsides, disturbed areas; 

common, introduced from Europe. May-June. [= C, Z; ? A. elegans Willdenow ex Kunth – RAB, G, HC, K; = Aspris capillaris 

(Host) A.S. Hitchcock – S] 

* Aira praecox Linnaeus, Early Hair Grass. Cp (NC, VA): fields, roadsides, disturbed areas; uncommon, introduced from 

Europe. Reported for NC by Burk (1961), and recently collected in the NC Sandhills (B.Sorrie, pers.comm. 2004). [= C, G, HC, 

K, Z] 

Alopecurus Linnaeus (Foxtail Grass)

POACEAE 839 

A genus of about 36 species, north temperate and temperate South America. References: Tucker (1996)=Z. 

1 Glumes 4-6 mm long, acute or acuminate. 

2 Glumes with hairs < 1.0 mm long on the keel, merely scabrous towards the tip ....................................... A. myosuroides 

2 Glumes with hairs 1.0-1.5 mm long on the keel, including towards the tip ..................................................... A. pratensis 

1 Glumes 2-3.2 mm long, obtuse or truncate. 

3 Awn about as long as the glumes (at most exceeding the glumes by 1 mm)............................... A. aequalis var. aequalis 

3 Awn longer than the glumes, exceeding the glumes by 1.5-3.5 mm. 

4 Anthers 0.4-0.7 mm long; annual....................................................................................................... A. carolinianus 

4 Anthers 1.3-2 mm long; perennial........................................................................................................A. geniculatus 

Alopecurus aequalis Sobolewski var. aequalis, Short-awn Foxtail Grass. Mt (VA): [habitat]; rare (VA Watch List). 

Circumboreal, south in North America to NJ, w. VA, IN, MO, and CA. [= F, K; < A. aequalis – C, G, HC] 

Alopecurus carolinianus Walter, Carolina Foxtail Grass. Cp, Pd (GA, NC, SC, VA), Mt (VA): moist fields, ditches, 

forests; common (rare in Mountains). April-May. MA west to British Columbia, south to n. FL and CA. [= RAB, C, F, G, GW, 

HC, K, Z; = A. ramosus Poiret – S] 

* Alopecurus geniculatus Linnaeus, Water Foxtail Grass. Mt, Pd, Cp (VA): disturbed areas; rare, introduced from Eurasia. 

[= C, F, G, HC; > A. geniculatus var. geniculatus – K] 

* Alopecurus myosuroides Hudson, Slender Foxtail Grass. Pd (NC, VA), Cp (VA): moist fields; uncommon, introduced 

from Europe. April-May. [= RAB, C, F, G, HC, K, S, Z] 

* Alopecurus pratensis Linnaeus, Meadow Foxtail. Mt (NC, VA), Pd (GA): roadsides, fields; rare, native of Eurasia. May- 

July. Reported for Piedmont of nc. GA (Jones & Coile 1988), for scattered locations in PA (Rhoads & Klein 1993), and for VA, 

KY, WV, MD, and DE (Kartesz 1999). [= C, F, G, HC, K] 

Ammophila Host (Beach-grass) 

A genus of 2 species, north temperate. References: Tucker (1996)=Z. 

1 Ligule 10-30 mm long ........................................................................................................................................... [A. arenaria] 

1 Ligule 1-3 mm long ........................................................................................................................................... A. breviligulata 

Ammophila breviligulata Fernald, American Beach-grass. Cp (NC, *SC, VA): dunes; common. August-September. 

Newfoundland south to about Cape Hatteras, Dare County, NC, and on shores around the Great Lakes; planted further south. As 

a native grass, Ammophila ranged south only to NC, where it was rare; it is now commonly planted ("sprigged") in the Carolinas 

as a sand-binder and is now common south into SC. [= RAB, C, F, G, HC, K, S, Z] 

* Ammophila arenaria (Linnaeus) Link, European Beach-grass, is introduced in MD and PA (Kartesz 1999). [= C, F, HC, K] 

Amphicarpum Kunth (Peanut-grass, Goober-grass) 

The genus consists only of the two species treated here, remarkable for their dimorphic spikelets, some of them cleistogamous 

and subterranean ("goobers"), others aerial and chasmogamous. A series of publications over the past century make 

Amphicarpum one of the best studied "useless" grasses anywhere (Holm 1896; Weatherwax 1934; Gray & Fairbrothers 1971; 

McNamara & Quinn 1977; Cheplick & Quinn 1982, 1983, 1986, 1987, 1988a, 1988b; Cheplick 1989). References: Wipff in 

FNA (2003a). 

1 Leaf blades hirsute with pustular-based hairs on both surfaces, the margins ciliate (and also slightly cartilaginous-thickened); 

[of moist to wet, peaty or sandy-peaty soils]..................................................................................................... A. amphicarpon 

1 Leaf blades glabrous, the margins cartilaginous-thickened; [of seasonally flooded natural ponds]......... A. muhlenbergianum 

Amphicarpum amphicarpon (Pursh) Nash, Pinebarrens Peanut-grass, Pinebarrens Goober-grass. Cp (GA, NC, SC, VA), 

Pd (VA): wet, peaty, open soils, especially peat-burns in pocosin edges, primarily in the outer Coastal Plain, responding strongly 

to fire; uncommon, rare in VA (VA Rare). August-October. An Atlantic Coastal Plain endemic, scattered and rather rare, from 

e. MA to GA. If one carefully excavates young plants in spring or summer, they will generally be found to be connected to the 

remnants of the previous year's subterranean spikelet. [= FNA; = Amphicarpum purshii Kunth – RAB, C, F, G, GW, HC, K; = 

Amphicarpon amphicarpon (Pursh) Nash – S] 

Amphicarpum muhlenbergianum (J.A. Schultes) Hitchcock, Florida Peanut-grass, Florida Goober-grass, Blue Maidencane. 

Cp (GA, NC, SC): clay-based Carolina bays in the inner Coastal Plain; rare (NC Rare, SC Rare). August-October. A 

Southeastern Coastal Plain endemic: FL and s. AL north to se. NC, rare north of s. GA. First found in NC in the late 1980's by 

M. Boyer. [= RAB, FNA, GW, HC; = A. muehlenbergianum – K, orthographic variant; = Amphicarpon floridanum Chapman – 

S]

POACEAE 840 

A genus of about 100-110 species, mainly tropical. 

Andropogon Linnaeus 1753 (Broomsedge, Bluestem) 

(also see Bothriochloa and Schizachyrium) 

The difference between this treatment and that in RAB may cause some users to react with skepticism, dismay, or alarm, but I am 

confident that it represents a much truer description of the genus. Campbell's work (1983, et seq.) has greatly clarified the 

taxonomy of Andropogon in e. North America. Great confusion and disagreement were previously the rule in dealing with the A. 

virginicus-A. glomeratus complex. Campbell's careful morphologic work has provided workable technical characters which 

distinguish the taxa he recognizes. 

I have generally followed Campbell (1983, et seq.) in his circumscriptions of taxa. I disagree, however, with his strongly 

morphologic species concept and the basis for his decisions regarding the rank of the taxa (species, variety, and "variant," an 

informal, English name for a subvarietal entity). Campbell bases the rank recognition of taxa on their "morphological distance" 

from one another, as determined by the sum of non-matching characters out of 33 characters analyzed. In general, he regards 

species as separated by a morphologic distance of 9 or more, varieties by 6 or more, and variants by 3 or more. Such an approach 

fails to take into account additional evidence of the ecological preferences, geographic distributions, reproductive isolation, 

evolutionary pathways, and population biology of the taxa. 

Evidence presented by Campbell (1983) is useful in determining a more meaningful assignment of taxonomic rank. For instance, 

he states that "there are ample opportunities for gene flow between taxa because they frequently grow together and flower at the 

same time of day and (mostly) at the same time of year. I have observed two taxa growing within one to three meters of one 

another over our hundred times. In only five of these opportunities for hybridization were there plants whose intermediate 

morphology suggested that they were hybrids. In the rare instances where hybridization does take place, there are few mature 

hybrid individuals. I have found only twelve putative hybrid individuals in the five localities where hybridization is suspected. 

The parents outnumber these hybrids by between five and one hundred or more to one." In discussing A. virginicus var. glaucus 

(here treated as A. capillipes) he states "the drylands variant ... produces generally shorter raceme sheaths, racemes, and spikelets; 

its flowers are more frequently chasmogamous, and unlike the wetlands variant, it has no hairs below the raceme sheath. In 

addition, it grows in better-drained soil and has a narrower geographic range ... I have seen these taxa growing within one to three 

meters of one another at three localities in northwestern Florida. At only one of these was there difficulty in classifying any 

individual: a single plant on a slope between a bog inhabited by the wetlands variant and a roadside lined with the drylands 

variant..." 

Some additional examples would be Campbell's discussion of several closely related taxa in the A. glomeratus complex. 

"Andropogon glomeratus var. glomeratus and the robust variant of var. pumilus have inflorescences so similar in shape that most 

previous workers have united them and have overlooked the differences between them. The robust variant is taller, usually with 

rather smooth sheaths and with shorter, more ciliate, and darker ligules, narrower raceme sheaths, and lower glume keels that are 

scabrous below the middle. Although both taxa grow in wet sites, the robust variant is weedier, shows a greater tolerance for 

drier conditions and various soil types, and has a wider geographic range." His discussion of A. glomeratus var. glaucopsis and 

var. hirsutior is also worth repeating. "Often the two grow together in populations of thousands of individuals. Because they 

grow together so frequently and are morphologically so alike, the possibility that they are not distinct taxa but merely genetic 

segregates of one another has been carefully considered. Based on observations of several hundred seedlings grown from seeds 

from both taxa (growing together in nature), there is no evidence for [mere] genetic segregation. The glaucousness/greenness and 

pubescence/glabrousness of the stem sheaths are discernible in the seedlings within a few weeks of germination." 

Taxa differing in numerous morphologic characters, with different (though overlapping) geographic ranges, with different 

ecological preferences (often rather narrowly segregated by hydrology), and (when they do occur in proximity to one another) 

showing little or no sign of introgression or hybridization are probably better treated as biological species. Thus, I have treated a 

number of Campbell's varieties as species. Several of his "variants" also seem to warrant taxonomic recognition, at varietal or 

specific rank; in fact, he subsequently elevated several (Campbell 1986). References: Campbell (1983)=Z; Campbell in FNA 

(2003a). Key adapted in part from Z. 

Identification notes: A thorough understanding of the architecture of the inflorescences of Andropogon is necessary in order to 

identify them successfully. The parts will be described, beginning from the apex of a branch of the inflorescence. Spikelets 

occur in pairs, the sessile spikelet (usually just referred to as the spikelet) and the pedicelled spikelet, which is usually vestigial 

or absent (except in A. gerardii) and sterile (except in A. gerardii, where it is staminate). The first or lower glume of the sessile 

spikelet has two keels, and the presence and location of antrorse prickle hairs (scabrousness) is an important character in the A. 

glomeratus complex. The length of the sessile spikelet is an important character; it should be measured exclusive of the awn, 

borne at the apex of the lemma. Awn length is also a useful taxonomic character. The pedicelled spikelet is borne on the 

pedicel, which is attached at the base of the sessile spikelet and typically angles away from it at about a 45 degree angle. The 

rachis internode extends from the base of one sessile spikelet to the next sessile spikelet above, breaking apart (upon 

dehiscence) just below the next spikelet and remaining attached to the sessile spikelet below. The dispersal unit consists of a 

sessile spikelet sitting in the V shape formed by (on one side) the pedicel and pedicelled spikelet and (on the other side) the rachis

POACEAE 841 

internode. Both the pedicel and the rachis internode are usually pubescent with long hairs, and the color of those hairs and their 

distribution are useful characters. 

While the dispersal units are still attached to one another, the rachis internodes form a continuous and more-or-less straight 

rachis. The dispersal units attached together in an unbranched sequence are termed a raceme, whose length is a useful character. 

Two or more racemes are attached digitately at the summit of the peduncle (in Schizachyrium only a single raceme is found). 

The number of racemes attached is an important character. A raceme sheath subtends the peduncle, often more or less 

surrounding the peduncle and the racemes. The length of the peduncle (distance between the points of attachment of the raceme 

sheath and the racemes) is an important character. The length and width (at its widest point) of the raceme sheath are very useful 

characters, used throughout the key. The racemes, peduncle and subtending raceme sheath make up an inflorescence unit. The 

overall inflorescence is more-or-less complexly branched; its overall size and shape are very useful in recognizing the various 

taxa, but variation in such a subjective (and environmentally plastic) character has added to the taxonomic confusion in 

Andropogon. The use of inflorescence shape in the key has been minimized, but is often mentioned in the discussion of each 

species. The number of inflorescence units per plant varies from species to species, in some species rarely exceeding 10, in 

others ranging upwards to 500 or 600. The absence or presence of hairs immediately below the raceme sheath is useful in some 

groups. 

There are several important characters of the foliage. A. capillipes and A. glaucopsis have culm sheaths and leaf blades that are 

strongly glaucous; this is usually very obvious, but can be tested for by running the finger along the surface of the leaf (a white 

coating of wax will come off on the finger). The key often calls for the ligule length; measure the longest portion of the undivided 

portion of the ligule. The ligule often has an erose or ciliate upper margin; measure the length of the ciliations. The length of 

leaf blade is measured from the ligule to the leaf apex; do not include the leaf sheath, which is often long and (especially late in 

the year) only loosely sheathing the culm or even divergent it. Whether the culm is antrorsely scabrous or smooth is better 

determined by touch than by sight. Choose several mid-culm sheaths, run one's finger downwards and upwards along the sheath 

surface (near the collar is best). If the sheath is antrorsely scabrous one will feel a somewhat greater resistance to moving the 

finger downwards than upwards. 

1 Pedicellate spikelet staminate, as large as the sessile, fertile spikelet; sessile spikelets > 7 mm long; [section Andropogon] .... 

...................................................................................................................................................................................A. gerardii 

1 Pedicellate spikelet sterile, vestigial or absent; sessile spikelets < 7 mm long; [section Leptopogon]. 

2 Leaves strongly glaucous (often nearly white with a powdery wax that can be rubbed off on the fingers), glabrous. 

3 Ligules (0.9-) 1.5 (-2.0) mm long, with ciliations 0-0.2 mm long; leaf blades usually (33-) avg. 40 (-75) cm long; 

pubescence beneath raceme sheaths moderate to dense; raceme sheaths (2.0-) 2.4-3.6 (-4.4) cm long, (1.3-) 2.0-2.5 

(-3.0) mm wide.......................................................................................................................................A. glaucopsis 

3 Ligules (0.2-) 0.4 (-0.5) mm long, with ciliations 0.3-1.2 mm long; leaf blades (12-) avg. 19 (-38) cm long; 

pubescence beneath raceme sheaths absent to dense; raceme sheaths (2.1-) 2.9-4.3 (-6.0) cm long, (2.7-) 3.1-3.8 (- 

5.5) mm wide. 

4 Summit of branchlet below attachment of raceme sheath pubescent with hairs 2-4 mm long; raceme sheaths 

(2.4-) 3.2-4.8 (-6.0) cm long; spikelets (3.0-) 3.5-3.9 (-4.4) mm long; racemes (1.5-) 2.0-3.0 (-4.0) cm long; 

leaves 2.5-6.5 mm wide, averaging 5 mm; upper floret lemma awn 0.9-2.1 mm long, averaging 1.4 mm ......... 

.............................................................................................................A. capillipes var. 1 ["wetland variant"] 

4 Summit of branchlet below attachment of raceme sheath glabrous; raceme sheaths (2.1-) 2.6-3.8 (-4.9) cm 

long; spikelets (2.6-) 3.2-3.5 (-3.9) mm long; racemes (1.4-) 1.7-2.4 (-3.2) cm long; leaves 2-5 mm wide, 

averaging 3.5 mm; upper floret lemma awn 0.6-1.5 mm long, averaging 1.1 mm.............................................. 

............................................................................................................. A. capillipes var. 2 ["dryland variant"} 

2 Leaves green (to somewhat glaucous, but never powdery white), pubescent or glabrous. 

5 Upper culm sheaths distinctly broadened and strongly overlapping, often largely hiding the raceme sheaths before 

senescence (but in some forms with the raceme sheaths strongly exserted); culms mostly < 1 m tall (to 1.4 m tall) . 

.................................................................................................................................................................... A. elliottii 

5 Upper culm sheaths reduced, not strongly overlapping, not hiding the raceme sheaths after anthesis; culms mostly 

> 1 m tall (except A. perangustatus, A. tracyi, and small forms of A. virginicus). 

6 Many or all peduncles longer than the subtending raceme sheaths at maturity, racemes then fully exserted 

above the apex of the raceme sheath. 

7 Inflorescence branches arching outwards in pronounced curves; racemes (1.2-) 1.5-2.1 (-2.6) cm long; 

awn (0.2-) avg. 0.7 (-1.1) cm long; spikelets (4.1-) 4.4-4.6 (-5.0) mm long...................A. brachystachyus 

7 Inflorescence branches erect; racemes (2.2-) 2.6-6 cm long; awn 0.5-2.0 cm long; spikelets (4.3-) 4.9-6.5 

(-7.5) mm long. 

8 Lower glumes more or less folded; stamen 1; racemes (2.2-) 2.6-4.3 (-5.3) cm long; awn (0.5-) avg. 

0.8 (-1.6) cm long; spikelets (4.3-) 4.9-5.4 (-6.1) mm long ................................................A. arctatus 

8 Lower glumes flat; stamens 3; racemes 3-6 cm long; awn 1-2 cm long; spikelets (4.5-) 5-6.5 (-7.5) 

mm long ....................................................................................................A. ternarius var. ternarius 

6 Peduncles all shorter than the subtending raceme sheaths at maturity, at least the bases of the racemes not 

exserted above the apex of the raceme sheath.

POACEAE 842 

9 Inflorescence units with (2-) 4-7 (-13) racemes; raceme sheaths (4.1-) 5.3-8.0 (-10-1) mm wide; hairs of 

the rachis internode and pedicel yellow-tawny when dry ............................................................A. mohrii 

9 Inflorescence units with 2-5 (-7) racemes; raceme sheaths (1.5-) 2.0-4.8 (-6.3) mm wide; hairs of the 

rachis internode and pedicel gray to whitish when dry. 

10 Postflowering peduncles < 10 mm long. 

11 Culm sheaths antrorsely scabrous (often hirsute as well); leaf blades usually > 35 cm long. 

12 Ligules (0.6-) 0.8 (-1.3) mm long (usually < 1 mm long), with ciliations 0.2-0.9 mm long; 

raceme sheaths (1.5-) 2.0-2.5 (-3.0) mm wide (usually < 2.5 mm wide); keels of first 

glume often scabrous below the middle.................................................... A. tenuispatheus 

12 Ligules (0.7-) 1.2 (-2.2) mm long (usually > 1 mm long), with ciliations 0.0-0.3 mm long; 

raceme sheaths (2.0-) 2.4-3.4 (-4.7) mm wide (usually > 2.5 mm wide); keels of first 

glume scabrous only above the middle, smooth below. 

13 Inflorescences oblong to obpyramidal; spikelets (3.8-) 4.1-4.4 (-5.0) mm long; 

anthers usually not marcescent within spikelet; mature peduncles (4 -) 11-35 (-60) 

mm long (usually some of them > 10 mm long)......... A. glomeratus var. glomeratus 

13 Inflorescences (linear to) oblong; spikelets (3.4-) 3.6-3.8 (-4.6) mm log; anthers 

usually marcescent within spikelets; peduncles (2-) 3-5 (-8) mm long........................ 

........................................................................................ A. glomeratus var. hirsutior 

11 Culm sheaths not scabrous (often hirsute); leaf blades < 35 cm long (except in A. glomeratus 

var. pumilus). 

14 Leaves glabrous. 

15 Ligules (0.8-) 1.1 (-1.5) mm long, with ciliations 0-0.1 mm long; basal leaves often 

filiform, < 1.5 mm wide, strongly erect ............................................A. perangustatus 

15 Ligules (0.2-) 0.5 (-0.8) mm long, with ciliations 0.2-1.3 mm long; basal leaves 

usually > 2 mm wide, soon arching. 

16 Culm internodes green (or glaucous just below the node only); raceme sheaths 

(2.2-) 2.5-3.8) (-4.5) cm long, (1.7-) 2.4-3.1) (-4.0) mm wide; peduncles (1-) 4- 

9 (-30) mm long; racemes 2 (-3) per inflorescence unit....................................... 

................................................................................. A. virginicus var. decipiens 

16 Culm internodes glaucous; raceme sheaths (2.8-) 3.3-4.7 (-6.7) cm long, (3.0-) 

3.2-3.8 (-5.2) mm wide; peduncles (2-) 3-4 (-6) mm long; racemes 2-4 (-7) per 

inflorescence unit, at least some inflorescence units (especially at culm and 

branch apices) with 3 or more racemes................................................................ 

.................................................. A. virginicus var. virginicus ['smooth variant'] 

14 Leaves pubescent, at least on the margin near the collar. 

17 Keels of first glume often scabrous below the middle; leaves usually > 44 cm long... 

.......................................................................................................... A. tenuispatheus 

17 Keels of first glume scabrous only above middle; leaves usually < 31 cm long. 

18 Pubescence of young culm sheaths appressed; spikelets usually > 4 mm long; 

hairs on rachis internode and sterile pedicel dense, long; callus hairs 1.5-5 mm 

long................................................................................................A. longiberbis 

18 Pubescence of young culm sheaths spreading; spikelets mostly < 4 mm long; 

hairs on rachis internode and sterile pedicel rather sparse and short; callus hairs 

1-2.5 mm long. 

19 Raceme sheaths (2.2-) 2.5-3.8 (-4.5) cm long, (1.7-) 2.4-3.1 (-4.0) mm 

wide; racemes 2 (3) per inflorescence unit; spikelets (3.0-) 3.3-3.6 (-4.0) 

mm long........................................................... A. virginicus var. decipiens 

19 Raceme sheaths (2.3-) 3.4-5.2 (-6.7) cm long, (2.7-) 3.3-4.0 (-5.5) mm 

wide; racemes 2-5 (-7) per inflorescence unit; spikelets (2.9-) 3.7-3.9 (- 

4.7) mm long................................................... A. virginicus var. virginicus 

10 Postflowering peduncles > 15 mm long. 

20 Culm sheaths antrorsely scabrous (often hirsute as well). 

21 Ligules (1.0-) 1.2 (-2.0) mm long, with ciliations 0-0.3 mm long; keels of first glume 

scabrous only above middle................................................ A. glomeratus var. glomeratus 

21 Ligules (0.6-) 0.8 (-1.3) mm long, with ciliations 0.2-0.9 mm long; keels of first glume 

often scabrous below middle .................................................................... A. tenuispatheus 

20 Culm sheaths not scabrous (often hirsute). 

22 Culms < 1.2 m tall; leaf blades < 30 cm long and < 3 mm wide; inflorescence units rarely 

> 20/culm. 

23 Raceme sheaths (2.2-) 2.5-3.8 (-4.5) cm long; spikelets (3.0-) 3.3-3.6 (-4.0) mm 

long; leaf blades (2.5-) 3.6 (-5.5) mm wide ..................... A. virginicus var. decipiens 

23 Raceme sheaths (2.6-) 4.1-6.6 (-8.5) cm long; spikelets (3.0-) 3.4-5.1 (-5.5) mm 

long; leaf blades (0.8-) 1.8 (-3.0) mm wide. 

24 Ligules (0.8-) 1.1 (-1.5) mm long, with ciliations 0-0.1 mm long........................

POACEAE 843 

..................................................................................................A. perangustatus 

24 Ligules (0.2-) 0.4 (-0.5) mm long, with ciliations (0.1-) 0.2-0.8 mm long........... 

................................................................................................................A. tracyi 

22 Culms usually > 1.2 m tall; leaf blades often > 30 cm long and > 3 mm wide; 

inflorescence units usually > 20/culm. 

25 Inflorescence branches arching outwards in pronounced curves; awn mostly < 1 cm 

long; spikelets (4.1-) 4.4-4.6 (-5.0) mm long; anther > 1.7 mm long........................... 

........................................................................................................A. brachystachyus 

25 Inflorescence branches erect; awn mostly >1 cm long; spikelets (3.0-) 3.3-3.8 (-4.5) 

mm long; anther < 1.5 mm long. 

26 Raceme sheaths (1.5-) 2.0-2.5 (-3.0) mm wide; keels of first glume often 

scabrous below middle; culms to 2.5 m tall; leaves to 109 cm long and 9.5 mm 

wide .......................................................................................... A. tenuispatheus 

26 Raceme sheaths (1.7-) 2.4-3.1 (-4.0) mm wide; keels of first glume scabrous 

only above middle; culms < 1.7 m tall; leaves < 35 cm long and 5.5 mm wide... 

................................................................................. A. virginicus var. decipiens 

* Andropogon arctatus Chapman, Florida Bluestem. Cp (NC): moist disturbed ground; rare, apparently introduced from 

farther south (NC Watch List). This curious record (the specimen at GH, collected by Randolph and Randolph in 1922 in 

Pamlico County, NC, annotated as A. arctatus by Campbell) is likely a waif. The species is native to pinelands from n. FL west 

to w. panhandle of FL and adjacent s. AL, south to s. FL. [= FNA, HC, K, S, Z] 

Andropogon brachystachyus Chapman, Shortspike Bluestem. Cp (GA, SC): moist to wet pinelands, natural pond margins, 

bogs, disturbed roadsides; rare (GA Special Concern). Se. SC (McMillan et al. 2002) south to FL, south to s. FL, west to e. FL 

Panhandle. A. brachystachyus is considered by some to range north to NC. [= FNA, K, Z; = A. brachystachys – GW, HC, S, 

orthographic variant] 

Andropogon capillipes Nash var. 1, Wetland White Bluestem. Cp (GA, NC, SC): wet savannas, ditches adjacent to 

savannas, depressional wetlands; common. September-October. S. NJ south to s. FL and west to e. TX; also in the Bahamas 

(Sorrie & LeBlond 1997). Campbell (1983) informally describes two "variants" of this species (which he treats at the varietal 

level, as A. virginicus var. glaucus). A. capillipes is clearly a species distinct from A. virginicus; moreover, the substantial 

morphological and ecological differences between Campbell's two "variants" (which he describes as nearly always sharply 

distinct, even when growing in close proximity) warrant recognition as good species, or at least as varieties. [< A. virginicus – 

RAB; < A. virginicus var. glaucus Hackel – F, FNA; < A. capillipes – GW, HC, K, S; = A. virginicus var. glaucus "wetlands 

variant" – Z; = A. virginicus var. dealbatus Mohr ex Hackel] 

Andropogon capillipes Nash var. 2, Dryland White Bluestem. Cp (GA, NC, SC): dry to mesic pine flatwoods, sandhills, 

adjacent roadbanks; uncommon. September-October. Se. NC south to s. FL and west to s. AL. See A. capillipes var. 1 for 

discussion of these two taxa. The type of A. capillipes (collected by A.H. Curtiss in FL) is of this taxon; Nash (1900) states that 

it occurs "in dry soil, North Carolina to Florida" and emphasizes that it is "abundantly distinct from A. virginicus, to which it is 

related." [< A. virginicus – RAB; < A. virginicus var. glaucus Hackel – F, FNA; < A. capillipes – GW, HC, K, S; = A. virginicus 

var. glaucus "drylands variant" – Z] 

Andropogon elliottii Chapman. Cp, Pd, Mt (GA, NC, SC, VA): dry to moist forests, woodlands, fields, and disturbed 

areas; common (uncommon in Mountains). September-October. Widespread in se. United States, from s. NJ west to s. IN, s. IL, 

s. MO, south to s. FL and TX. Campbell (1983) argued that the name A. elliottii should be replaced by A. gyrans; Ward (2004c) 

argues for retention of the traditional A. elliottii. [= HC; > A. elliottii – RAB, S; > A. campyloracheus Nash – RAB, S; = A. 

gyrans Ashe – C, W; = A. gyrans var. gyrans – FNA, K, Z; > A. elliottii var. elliottii – F, G; > A. elliottii var. gracilior Hackel – 

F, G; > A. elliottii var. projectus Fernald & Griscom – G] 

Andropogon floridanus Scribner, Florida Bluestem. Cp (GA): longleaf pine sandhills; rare. September-October. S. GA 

west to FL Panhandle, south to s. FL. Reported for e. and s. GA (FNA, Jones & Coile 1988). [= FNA, HC, K, S] {not keyed at 

this time} 

Andropogon gerardii Vitman, Big Bluestem, Turkeyfoot. Cp, Pd, Mt (GA, NC, SC, VA): in a wide variety of habitats, 

usually rather dry, such as sandhills, glades, cliffs, and rock outcrops, in the Piedmont in woodlands, former prairie-like sites, 

woodlands, open forests, and river-scour grasslands, in the Mountains in glades, riverside scour areas, and rarely in grassy balds, 

ascending to at least 1600 m over mafic rocks (on Old Field Bald, Watauga and Ashe counties, NC); common. July-October. 

Québec west to Saskatchewan, south to FL and AZ. Some favor treating A. hallii Hackel as a subspecies of A. gerardii (Wipff 

1996c). I do not agree, but if that course is followed, then our eastern taxon should be known as A. gerardii ssp. gerardii. [= 

RAB, C, FNA, G, GW, HC, K, W; > A. gerardii var. gerardii – F; = A. provincialis Lamarck – S] 

Andropogon glaucopsis Elliott, Chalky Bluestem. Cp (GA, NC, SC, VA): wet savannas, pine flatwoods, ditches, wet 

disturbed sites; uncommon (VA Watch List). September-October. Se. VA south to c. peninsular FL and west to e. TX. The 

extent of the western Gulf Coastal Plain distribution (to the West Gulf Coastal Plain of w. LA and e. TX) is based on specimens 

(at BRIT) and sight records (B. Sorrie, pers. comm.). Although sometimes included in the past in either A. glomeratus or A. 

virginicus, this species is distinctive and easily recognized in the field (even from a car at 60 m.p.h.) by the combination of blue 

color, height of well over 1 m (taller than the other glaucous bluestems), and semi-bushy inflorescence. [= GW, K; < A. 

virginicus – RAB; = A. virginicus var. glaucopsis (Elliott) A.S. Hitchcock – F, HC; = A. glomeratus var. glaucopsis (Elliott) A.S. 

Hitchcock – FNA, Z; < A. glomeratus – S]

POACEAE 844 

Andropogon glomeratus (Walter) Britton, Sterns, & Poggenburg var. glomeratus. Cp, Pd, Mt (GA, NC, SC, VA): 

swamps, wet savannas, pine flatwoods, wet disturbed sites; common. September-October. S. MA south to c. peninsular FL and 

west to s. MS, primarily on the Coastal Plain, but scattered inland to w. PA, WV, c. KY, c. TN and AR. [= FNA, K, Z; < A. 

virginicus – RAB; = A. virginicus var. abbreviatus (Hackel) Fernald & Griscom – C, F, G, GW; < A. glomeratus – HC, S, W] 

Andropogon glomeratus (Walter) Britton, Sterns, & Poggenburg var. hirsutior (Hackel) C. Mohr. Cp (GA, NC, SC, VA): 

wet savannas, pine flatwoods, adjacent ditches, other wet disturbed sites; common. September-October. E. MD south to c. 

peninsular FL west to se. LA. This taxon should be recognized at the specific level, but the appropriate combination has not been 

made. [= FNA, K, Z; < A. virginicus – RAB; ? A. virginicus var. glaucopsis (Elliott) A.S. Hitchcock – G, misapplied; = A. 

virginicus var. hirsutior (Hackel) A.S. Hitchcock; < A. glomeratus – HC, S] 

Andropogon longiberbis Hackel, Longbeard Bluestem. Cp (GA, NC, SC): dry sandy soils of sandhills and dunes; rare (GA 

Special Concern, NC Watch List). September-October. Se. NC south to s. and w. FL, and in the Bahamas. [= FNA, HC, K, S, 

Z] 

Andropogon mohrii (Hackel) Hackel ex Vasey, Tawny Bluestem, Bog Bluestem. Cp (GA, NC, SC, VA): wet savannas, 

sphagnous bogs; rare (GA Special Concern, NC Rare, VA Rare). September-October. Se. VA south to n. FL, west to LA. [= 

RAB, C, F, G, GW, HC, K, S; = A. liebmannii Hackel var. pungensis (Ashe) C.S. Campbell – FNA, Z] 

Andropogon perangustatus Nash, Narrow-leaved Bluestem. Cp (GA, NC, SC, VA): clay-based Carolina bays and boggy 

wetlands; rare (NC Watch List, VA Watch List). August-October. E. VA south to c. peninsular FL, east to e. TX. Growth form, 

general appearance, and habitat (dense bluish tussocks with very narrow leaves and long ligules, growing in wet areas such as 

clay-based Carolina bays) make A. perangustatus readily recognizable. [= HC, S; = A. gyrans Ashe var. stenophyllus (Hackel) 

C.S. Campbell – FNA, K, Z; = A. elliottii Chapman var. stenophyllus (Hackel) D.B. Ward] 

Andropogon tenuispatheus (Nash) Nash. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): maritime wet grasslands, brackish 

marsh edges, moist disturbed sites; common (VA Watch List). September-October. Se. VA and c. OK south to s. FL and w. TX, 

also south into Central America and the Caribbean. [< A. virginicus – RAB; = A. glomeratus (Walter) Britton, Sterns, & 

Poggenburg var. pumilus Vasey ex Dewey – FNA, K, Z ("robust variant"); < A. glomeratus – HC, S] 

Andropogon ternarius Michaux var. ternarius, Splitbeard Bluestem. Cp, Pd, Mt (GA, NC, SC, VA): dry to moist soils; 

common (uncommon in Mountains). September-October. Var. ternarius ranges from DE west to KY and s. MO, south to FL 

and TX. Var. cabanisii (Hackel) Fernald & Griscom is endemic in s. and c. peninsular FL. [= FNA, K, Z; < A. ternarius – RAB, 

C, G, W; > A. ternarius var. ternarius – F; > A. ternarius var. glaucescens (Scribner) Fernald & Griscom – F; = A. ternarius – 

HC, S] 

Andropogon tracyi Nash, Tracy's Bluestem. Cp, Pd (GA, NC, SC): dry sandy or clayey soils of sandhills, disturbed sites; 

rare (NC Watch List). September-October. E. NC south to s. FL and west to MS. [= FNA, HC, K, S, Z] 

Andropogon virginicus Linnaeus var. decipiens C.S. Campbell, Deceptive Bluestem. Cp (GA, NC, SC, VA): savannas, 

flatwoods, maritime wet grasslands, disturbed pinelands; uncommon (VA Watch List). September-October. Se. VA south to s. 

FL and west to w. FL; also in the Bahamas (Sorrie & LeBlond (1997). [= FNA, K, Z (1986); < A. virginicus – RAB, S; < A. 

virginicus var. virginicus – F, G, HC; = A. virginicus var. virginicus – Z (1983 – "deceptive variant")] 

Andropogon virginicus Linnaeus var. virginicus, Old-field Broomstraw, Broomsedge, "Sedge Grass", "Sage Grass". Cp, 

Pd, Mt (GA, NC, SC, VA): old fields, roadbanks, disturbed sites; common. September-October. Widespread, from MA west to 

MI and e. KA, south to FL and e. TX, and in the Caribbean and Central America. Campbell (1983) recognized 3 "variants" 

within A. virginicus var. virginicus; the "deceptive variant" he later (1986) described formally as var. decipiens (see above). The 

"old-field variant" is the common "variant" in our area, occurring abundantly throughout the state. It has green stem internodes 

and the leaves usually pubescent, at least on the margins near the collar. The "smooth variant" is known only from the Coastal 

Plain and is apparently rare in our area, known from NC and SC (Berkeley and Marion counties; P. McMillan, pers. comm.). It 

has glaucous stem internodes and glabrous leaves. It is unclear whether the "smooth variant" warrants taxonomic recognition. [= 

FNA, K, Z ("oldfield variant" and "smooth variant"); < A. virginicus – RAB, S, W; < A. virginicus var. virginicus – C; < A. 

virginicus var. virginicus – G, HC (also see var. decipiens); >< A. virginicus var. virginicus – F; > A. virginicus var. 

tetrastachyus (Elliott) Hackel – F] 

Anthenantia Palisot de Beauvois (Silkyscale) 

A genus of 3 species, of se. North America (or 4-5 species of se. North America and tropical America, if Leptocoryphium is 

included in Anthenantia). Clayton & Renvoize (1986) state that "Anthenantia is the etymologically correct version of three 

alternative spellings given by Beauvois." References: Wipff in FNA (2003a); Crins (1991)=Z; Kral (2004)=Y; Clayton & 

Renvoize (1986). 

1 Leaves mostly 3-5 mm wide, ascending to erect, not squarrose (lacking a sharp bend outward at the summit of the sheath), 

medium green, with a very short taper to a blunt or rounded apex, the lower sheaths crowded and keeled (therefore 

distichous), generally suffused with purple; spikelets usually purple (fading tan); leaf margins scaberulous .................A. rufa 

1 Leaves mostly 5-10 mm wide, spreading, usually squarrose (with a sharp bend outward at the summit of the sheath), 

yellowish green, with a long taper to a sharp apex, the lower sheaths not crowded, keeled, or distichous, green; spikelets 

usually green (fading yellow); leaf margins papillose-ciliate towards the base ........................................................... A. villosa 

Anthenantia rufa (Nuttall) J.A. Schultes, Purple Silkyscale. Cp (GA, NC, SC): wet savannas in the outer Coastal Plain, 

seepage bogs and moist sandhill-pocosin ecotones in the fall-line sandhills; rare (NC Watch List, SC Rare). September-October.

POACEAE 845 

Se. NC south to n. FL and west to w. LA. A. rufa inhabits much wetter habitats than the similar A. villosa, and is more typical of 

the outer Coastal Plain. Plants without culms are reminiscent of the Liliaceae. [= FNA, Y; = Anthaenantia rufa – RAB, GW, 

HC, K, S, Z, orthographic variant] 

Anthenantia villosa (Michaux) Palisot de Beauvois, Green Silkyscale. Cp (GA, NC, SC): sandhills, especially in submesic 

swales; uncommon (rare in the outer Coastal Plain). September-October. Se. NC south to s. FL and west to e. TX. A. villosa is 

found in drier habitats than A. rufa, most typically in upland swales in the sandhills. Kral (2004) has segregated a new species, 

A. texana Kral, of the w. Gulf Coastal plain, previously confused with A. villosa. [= Y; < Anthaenantia villosa – RAB, HC, K, S, 

Z, orthographic variant; < Anthenantia villosa – FNA] 


Anthoxanthum Linnaeus (Vernal Grass) 

1 Annual, geniculate; ligules 0.5-2 mm long; glumes glabrous; leaves 1-2 mm wide ............................................... A. aristatum 

1 Perennial, erect; ligules (1-) 2-3 mm long; glumes villous throughout or at least on the keel; leaves 2-5 mm wide .................. 

................................................................................................................................................................................A. odoratum 

* Anthoxanthum aristatum Boissier, Annual Vernal Grass. Cp (NC, SC, VA), Pd, Mt (VA): roadsides, disturbed areas; rare, 

introduced from Europe. April-June. [= RAB, C, G, HC, K, S, Z; = A. puelii Lecoq & Lamotte – F] 

* Anthoxanthum odoratum Linnaeus, Sweet Vernal Grass Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): lawns, roadsides, 

disturbed areas; common, introduced from Europe. April-June. A. odoratum is a familiar grass of suburban areas and roadsides, 

and its pollen is known as a major cause of spring hay fever. From a letter from Charles Darwin to J.D. Hooker, in June 1855: 

"Have just made out my first grass, hurrah! hurrah! I must confess that fortune favours the bold, for, as good luck would have it, 

it was the easy Anthoxanthum odoratum: nevertheless it is a great discovery; I never expected to make out a grass in all my life, 

so hurrah! It has done my stomach surprising good..." [= RAB, C, F, G, HC, S, W, Z; = A. odoratum ssp. odoratum – K] 

Apera Adanson 

* Apera spica-venti (Linnaeus) Palisot de Beauvois, reported for se. PA (Rhoads & Klein 1993), MD, and KY (Kartesz 1999). 

[= K] 

Aristida Linnaeus 1753 (Three-awn Grass) 

A genus of about 250-300 species, widespread in the tropics, subtropics, and warm temperate zones. References: Allred in FNA 

(2003a); Allred (1986)=Z; Allred (1984, 1985); Peet (1993)=Y; Ward (2001)=X; Henrard (1929)=Q; Kesler, Anderson, & 

Hermann (2003)=V. Key adapted, in part, from Z. 

Identification notes: the awns must be dry and relatively mature to assume their characteristic positions (immature awns and 

moist mature awns are erect and parallel). It is sometimes useful to dry a collection unpressed. Beware, however, that drying 

followed by dispersal can take place very quickly under the right conditions (such as the dashboard of a hot car)! 

1 Plant a perennial, forming dense tussocks, the leaves primarily basal, usually very numerous, mostly > 3 dm long, 0.5-1.5 

mm wide, almost always tightly involute; flowering only in the growing season following fire. 

2 Base of blade and collar (and often the upper sheath) with conspicuous tuft or bearding of woolly to villous pubescence 

(sometimes deciduous on foliage more than a year old); leaves usually glabrous above the basal 2 cm of the blade; [of 

s. SC south]................................................................................................................................................. A. beyrichiana 

2 Base of blade, collar, and upper sheath lacking a conspicuous tuft of woolly to villous pubescence; leaves with 2 lines 

of villous pubescence on either side of the midrib on the lower surface extending nearly or entirely the length of the 

blade (sometimes deciduous on foliage more than a year old); [of n. SC and NC]...............................................A. stricta 

1 Plant an annual or perennial, forming small tufts (or solitary), the leaves primarily cauline, usually few, mostly < 3 dm long 

(if as long as 3 dm then > 2 mm wide), flat to slightly folded, but not wiry; flowering not strongly triggered by fire. 

3 First glume 3-7 nerved .................................................................................................................................... A. oligantha 

3 First glume 1-2-nerved. 

4 Lateral awns < 8 mm long; plant an annual. 

5 Central awn 1-27 mm long, not spirally coiled at its base (above the awn column), either straight, curving, or 

contorted (when dry); lateral 2 awns 0-18 mm long, contorted at base and more-or-less divergent. 

6 Central awn (8-) 12-27 mm long; lateral awns (1-) 6-18 mm long............... A. longespica var. geniculata 

6 Central awn mostly 1-10 (-14) mm long; lateral awns 0-5 (-8) mm long.....A. longespica var. longespica 

5 Central awn 3-8 mm long, spirally coiled at its base (above the awn column) like a corkscrew, 1/2 to 3 full 

turns (when dry); lateral 2 awns 0.7-4 mm long, straight, erect. 

7 First glume 1/2 to 2/3 as long as the second glume; lemma 6-11 mm long, glabrous to scaberulous..........

POACEAE 846 

...................................................................................................................................................A. curtissii 

7 First glume as long as or nearly as long as the second glume; lemma 3-8 mm long, sparsely appressedpubescent................................................................................................................................A. 

dichotoma 

4 Lateral awns > 8 mm long; plant an annual or perennial. 

8 Sheaths lanose or floccose (the hairs kinked and intertwined); nodes of the panicle axis with tufts of lanose or 

floccose hairs........................................................................................................................................A. lanosa 

8 Sheaths glabrous to pilose (the hairs straight and usually appressed, not intertwined); nodes of the panicle 

axis glabrous or pilose. 

9 Awn column (the connivent awns twisted together) or lemma beak (slender, narrowed, and twisted 

portion of lemma body below the awns) 7-30 mm long; lemma body (including the beak, if present) 

separated from the awns (or awn column) by an articulation zone, the awns (or awn column) 

disarticulating at maturity from the lemma. 

10 Panicle spiciform, broadest near the middle, dense, the spikelets overlapping strongly; awns (10-) 

20-30 mm long, borne at the summit of a twisted lemma beak 7-30 mm long; culms simple or with 

very few branches; plants perennial ..............................................................................A. spiciformis 

10 Panicle almost corymbiform, broadest above the middle, open, the spikelets overlapping only 

slightly; awns 30-40 mm long, not including the 8-15 mm long column formed by the twisting 

together of the 3 awn bases; culms often much-branched; plants annual .....................A. tuberculosa 

9 Awn column or lemma beak absent or < 7 mm long; lemma body not separated from the awns by an 

articulation zone. 

11 Spikelets borne singly at each node of the main axis, the inflorescence thus a spike or raceme ......... 

..............................................................................................................................................A. mohrii 

11 Spikelets 2 or more per node of the main axis at most nodes (a few nodes may have single 

spikelets), often with side branches present as well, the inflorescence thus a panicle or raceme. 

12 First glume 1/3 to 3/4 as long as the second glume; awns 40-100 mm long................................ 

...........................................................................................................A. purpurea var. longiseta 

12 First glume > 3/4 as long as the second glume; awns 8-40 mm long. 

13 Annual .................................................................................. A. longespica var. geniculata 

13 Perennial. 

14 Central awn 15-40 mm long; first glume prominently 2-keeled, (8-) 9-14 mm long 

when mature ..............................................................................................A. palustris 

14 Central awn 8-25 mm long; first glume either 1-keeled and 6-14 mm long, or 

weakly 2-keeled and 5.5-9 (-10) mm long when mature. 

15 Central awn about 2× as thick as the lateral awns, divergent to reflexed; first 

glume 1-keeled or weakly 2-keeled; [plants of moist to wet habitats]. 

16 Basal internode of the culm 0.3-0.6 mm wide; most nodes of the 

inflorescence with 1-2 spikelets; all awns spreading, the central spirally 

twisted basally and often contorted by as much as 180 degrees (best seen 

in fresh material); central awn 15-20 mm long, lateral awns 11-16 mm 

long, the ratio of the lateral:central awn length 0.69-0.80; lemma callus 

beard 0.6-1.0 mm long........................................................ A. simpliciflora 

16 Basal internode of the culm 0.7-1.2 mm wide; most nodes of the 

inflorescence with 3 or more spikelets; central awn spreading to slightly 

deflexed, not spirally twisted basally, the lateral awns ascending to erect 

(best seen in fresh material); central awn 13-22 mm long, lateral awns 8- 

15 mm long, the ratio of the lateral:central awn length 0.55-0.69; lemma 

callus beard 0.2-0.6 mm long....................................................... A. virgata 

15 Central awn < 1.5× as thick as the lateral awns, erect to divergent; first glume 

1-keeled (rarely weakly 2-keeled); [plants of dry habitats]. 

17 Culms mostly > 10 dm tall and 3-6 mm in diameter near the base; awns 8- 

15 mm long; panicle branches > 4 cm long; callus ca. 1.0 mm long............ 

.............................................................................................. A. condensata 

17 Culms 5-8 (-10) dm tall and 1-4 mm in diameter near the base; awns 12-25 

mm long; panicle branches 1-4 cm long; callus 0.4-0.8 mm long. 

18 First glume 1-4 mm longer than the second glume (rarely about equal 

to it); awns 15-25 mm long, straight or slightly contorted at the base; 

leaf blades 1-3 mm wide, usually curling...................A. purpurascens 

18 First glume shorter than or about equal to the second glume; awns 12- 

18 mm long, spirally contorted at the base; leaf blades about 1 mm 

wide, usually not curling................................................. A. tenuispica 

Aristida beyrichiana Trinius & Ruprecht, Southern Wiregrass. Cp (GA, SC): sandhills, savannas, from very dry to 

seasonally saturated soils; common. September-November. S. SC south to s. FL, west to s. MS. See Peet (1993) for discussion 

of the taxonomy and ecology of this species; also see comments under A. stricta, which also apply here. Ward (2001) proposes

POACEAE 847 

varietal status for A. stricta and A. beyrichiana. [= K, Y; < A. stricta – RAB, FNA, GW, HC, S, V, Z; = A. stricta Michaux var. 

beyrichiana (Trinius & Ruprecht) D.B. Ward – X] 

Aristida condensata Chapman, Big Three-awn. Cp (GA, NC, SC): dry sandy soils of sandhills; rare (NC Watch List, SC 

Rare). August-October. Sc. NC south to s. FL, west to s. MS (Sorrie & Leonard 1999). [= RAB, FNA, HC, K, S, Z] 

Aristida curtissii (A. Gray ex S. Watson & Coulter) Nash, Curtiss's Three-awn. Cp, Pd (GA, NC, SC, VA): roadsides, 

disturbed areas, bare eroding soil; uncommon. August-October. ME west to WY, south to FL, AR, OK, and CO, perhaps largely 

or entirely adventive in our area. See Z for a discussion of the rationale for reducing A. curtissii to a variety of A. dichotoma. C 

reduces it to a variety of the more western A. basiramea Engelmann ex Vasey. For now, and for simplicity, I prefer to retain the 

two as species. [= RAB, G, HC, S; = A. basiramea Engelmann ex Vasey var. curtissii (A. Gray ex S. Watson & Coulter) 

Shinners – C; = A. dichotoma Michaux var. curtissii A. Gray – F, FNA, K, W, Z] 

Aristida dichotoma Michaux, Fork-tip Three-awn. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, fields, disturbed areas, bare 

eroding soil; common. August-October. ME west to WI, south to FL and TX. See A. curtissii for comments. [= RAB, C, G, 

HC, S; = A. dichotoma var. dichotoma – F, FNA, K, W, Z] 

Aristida lanosa Muhlenberg ex Elliott, Woollysheath Three-awn. Cp, Pd (GA, NC, SC, VA): dry sandy soils of sandhills 

and fields; common, rare in Piedmont (VA Watch List). August-October. NJ south to FL, west to TX, north in the interior to 

MO and OK. [= RAB, C, FNA, G, HC, K, S, Z; > A. lanosa var. lanosa – F; A. lanosa var. macera Fernald & Griscom – F] 

Aristida longespica Poiret var. geniculata (Rafinesque) Fernald, Eastern Slim-spike Three-awn. Cp, Pd (GA, NC, SC, VA), 

Mt (NC, SC, VA): disturbed areas; common? August-October. The distribution and habitats of the 2 varieties in our area are 

poorly known, pending further field and herbarium investigation. [= C, F, FNA, HC, K, Z; < A. longespica – RAB, W; > A. 

intermedia Scribner & Ball – F, G, S; > A. longespica – G] 

Aristida longespica Poiret var. longespica, Eastern Slim-spike Three-awn. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): 

disturbed areas; uncommon? August-October. The distribution and habitats of the 2 varieties in our area are poorly known, 

pending further field and herbarium investigation. [= C, F, FNA, HC, K, Z; < A. longespica – RAB, G, W; = A. longespica – S] 

Aristida mohrii Nash, Mohr's Three-awn. Cp (GA, SC): sandhills; rare. August-October. Panhandle FL and sw. GA west 

to s. AL; apparently disjunct in SC (Chesterfield and Richland counties). [= FNA, HC, K, S, Z] 

Aristida oligantha Michaux, Prairie Three-awn. Pd, Cp, Mt (GA, NC, SC, VA): roadsides, fields, disturbed areas; 

common. August-October. VT west to SD, south to FL and TX, scattered elsewhere as a weed. [= RAB, C, F, FNA, G, HC, K, 

W, S, Z] 

Aristida palustris (Chapman) Vasey, Longleaf Three-awn. Cp (GA, NC, SC): wet pine savannas, limesink depressions; 

uncommon. August-October. Se. NC south to FL, west to TX; apparently disjunct on the Cumberland Plateau of KY. [= C, 

FNA, K, S, Z; = A. affinis (Schultes) Kunth – RAB, F, G, GW, HC, misapplied] 

Aristida purpurascens Poiret, Arrowfeather. Cp, Pd, Mt (GA, NC, SC, VA): dry habitats, especially in dry sandy soils; 

common. August-October. MA west to WI and KS, south to FL and TX. In the Sandhills occurring in two forms, one green, the 

other strongly glaucous-blue. [= RAB, C, G, HC, S, W; > A. purpurascens var. purpurascens – F; > A. purpurascens var. minor 

Vasey – F; = A. purpurascens var. purpurascens – FNA, K, Z] 

* Aristida purpurea Nuttall var. longiseta (Steudel) Vasey, Red Three-awn. Cp (SC): disturbed areas; rare, adventive from 

further west. August-October. Also reported from NC, but the collection is from a Soil Conservation Service test nursery, and 

there is no evidence of naturalization. [= C, FNA, K, Z; > A. longiseta var. robusta Merrill – F; = A. longiseta Steudel – G, HC] 

Aristida simpliciflora Chapman, Southern Three-awn, Chapman's Three-awn. Cp (GA, NC): wet pine savannas; rare (GA 

Special Concern, NC Rare). Sw. GA west through the FL Panhandle and c. AL to s. MS (Sorrie & Leonard 1999), and south into 

central Peninsular Florida; also in se. NC, where apparently disjunct (it should be searched for in SC). A. simpliciflora was 

believed to be a Gulf Coastal Plain endemic until found by R. LeBlond in 1999 in wet savannas in se. NC (Green Swamp 

savannas, Brunswick County; Old Dock Savanna, Columbus County; and The Neck Savanna, Pender County). It is reported for 

sw. GA (Jones & Coile 1988, Kartesz 1999). Harper also reports it for c. GA. [= FNA, HC, K, S, Z] 

Aristida spiciformis Elliott, Bottlebrush Three-awn, Spike Three-awn. Cp (GA, NC?, SC): wet pine savannas and seepage 

areas; rare (NC Watch List). August-October. E. SC (McMillan et al. 2002) south to FL, west to MS. Allred (1986) also reports 

this species from NC, but the documentation is unknown to me. [= RAB, FNA, GW, HC, K, S, Z] 

Aristida stricta Michaux, Carolina Wiregrass, Pineland Three-awn. Cp (NC, SC), Pd (NC): Coastal Plain pinelands of 

nearly all sorts, ranging from the driest white-sand sandhills to seasonally saturated pine savannas dominated by a mixture of 

longleaf pine and pond pine, largely or entirely replaced in the wettest savannas by Sporobolus teretifolius, Sporobolus 

pinetorum, Muhlenbergia expansa, Ctenium aromaticum, and Calamovilfa brevipilis; also in Piedmont areas adjacent to the 

Coastal Plain and formerly supporting fire-maintained longleaf pine woodlands; common, rare in Piedmont. September- 

November. Ne. NC (south of Albemarle Sound and the Roanoke River), south to ne. SC (Lee and Kershaw counties). A. stricta 

was the keystone species of much of the upland Coastal Plain of the Carolinas. Its flammable foliage facilitated the spread of 

lightning-set fires that maintained the biologically rich pine savanna, sandhill, and pine flatwood ecosystems once widespread in 

our area. Though still locally common in parts of the Sandhill region and in portions of Brunswick, Pender, Onslow, and Carteret 

counties, NC, A. stricta is much rarer than formerly. The conversion of vast acreages of former pinelands to agriculture, pine tree 

farms, and developed areas has taken its toll over the years. In the twentieth century, suppression of fire has also led to the 

destruction of A. stricta. More recently, pine-straw raking is leading to the serious decline of A. stricta in its few remaining 

strongholds on public lands. A. stricta has little tolerance for ground disturbance. See Peet (1993) for discussion of the 

taxonomy and ecology of this species. Ward (2001) proposes varietal status for A. stricta and A. beyrichiana. [= K, Y; < A. 

stricta – RAB, FNA, GW, HC, S, V, Z (also see A. beyrichiana); = A. stricta var. stricta – X]

POACEAE 848 

Aristida tenuispica A.S. Hitchcock, Southern Arrowfeather. Cp (GA, NC, SC): sandy habitats in the Coastal Plain; 

uncommon? August-October. NC south to FL and west to MS. [= HC, S; = A. purpurascens Poiret var. tenuispica (A.S. 

Hitchcock) Allred – FNA, K, Z] 

Aristida tuberculosa Nuttall, Seabeach Needlegrass. Cp (GA, NC, SC, VA): sandhills, coastal dunes (in VA), other dry, 

sandy habitats such as sandy roadsides; common, rare in VA (VA Watch List). August-October. Se. NH south to NJ and 

disjunct in e. VA in the outer Coastal Plain; from sc. NC south to Panhandle FL and west to s. MS (Sorrie & Leonard 1999), 

mostly in the inner Coastal Plain; and also near the Great Lakes in sw. MI, n. IN, n. IL, s. WI, se. MN, and e. IA. The curious 

trimodal distribution is unexplained. [= RAB, C, F, FNA, G, HC, K, S, Z] 

Aristida virgata Trinius. Cp (GA, NC, SC, VA), Pd (GA), Mt (NC): moist to wet savannas, mountain bogs (Henderson 

Co., NC), other moist habitats; common. August-October. S. NJ south to FL, west to TX, primarily on the Coastal Plain. [= 

RAB, C, F, G, GW, HC, S; = A. purpurascens Poiret var. virgata (Trinius) Allred – FNA, K, Z] 

* Aristida adscensionis Linnaeus, Sixweeks Three-awn, has been reported as an introduction (from western United States) in 

SC (FNA, Kartesz 1999). {investigate} [= F, FNA, G, K] {not keyed at this time; synonymy incomplete} 

Aristida basiramea Engelmann ex Vasey, Forktip Three-awn. Occurs in VA, SC, etc. (FNA). [= F, FNA, G, K; = A. 

basiramea var. basiramea – C] {not keyed at this time; synonymy incomplete} 

Aristida gyrans Chapman, Corkscrew Three-awn. Cp (GA): dry pinelands; rare (GA Special Concern). E. GA and w. 

Panhandle FL, south to s. FL. In Bryan, Long, and Montgomery counties in e. GA (Sorrie 1998b), and in wc. GA (J. Allison, 

pers. comm.). [= FNA, K, S] {not keyed at this time; synonymy incomplete} 

Aristida ramosissima Engelmann ex A. Gray. East to Panhandle FL, c. TN, and e. KY (FNA) and might occur in our area. 

It is similar to A. oligantha and will key to it; it differs in having the central awn of the lemma 9-30 mm long (vs. 30-70 mm 

long), and the awn of the second glume 3-7 mm long (vs. 7.5-17 mm long). [= C, F, FNA, G, K, S] {not keyed at this time; 

synonymy incomplete} 

Allred (1986) reports the collection of several additional non-native species from our area, including A. divaricata Willdenow 

from sw. United States (from a Soil Conservation Service test nursery in Chapel Hill, NC) and uncertainly identified material of 

an Australian species (from a wool-combing mill at Jamestown, SC). There is no evidence that either are naturalized. 


Arrhenatherum Palisot de Beauvois (False Oatgrass) 

1 Base of culm consisting of a series of adjacent (moniliform) corms ca. 1 cm in diameter...................A. elatius var. bulbosum 

1 Base of culm not swollen or cormose ....................................................................................................... A. elatius var. elatius 

* Arrhenatherum elatius (Linnaeus) J. & K. Presl var. bulbosum (Willdenow) Spenner, Tuber Oatgrass, Onion Couch. 

(VA): habitat in our area not known; abundance not known, introduced from Europe. This variety was apparently cultivated for 

the edible tubers in Bronze Age Europe (Tucker 1996). Cited for VA in HC. [= C, F, G, HC, K, Z; < A. elatius – GW, W; = A. 

elatius var. tuberosum Thiel. – S] 

* Arrhenatherum elatius (Linnaeus) J. & K. Presl var. elatius, Tall Oatgrass. Pd (GA, NC, VA), Mt, Cp (NC, VA): 

meadows, fields, roadsides; common, introduced from Europe. May-June. [= C, F, G, HC, K, S, Z; < A. elatius – RAB, GW, 

W] 

Arthraxon Palisot de Beauvois (Basket Grass) 

References: van Welzen (1981)=Y; Thieret in FNA (2003a); Kiger (1971)=Z. 

Identification notes: Sometimes confused (especially before flowering) with Microstegium, but Arthraxon has distinctly 

cordate-clasping leaves, which Microstegium lacks. Also vegetatively similar to Oplismenus. 

* Arthraxon hispidus (Thunberg) Makino var. hispidus, Basket Grass. Cp, Pd, Mt (GA, NC, SC, VA): moist ditches, 

bottomlands, disturbed areas; common, native of se. Asia. September-October. Like Microstegium, Arthraxon appears to be 

steadily increasing its abundance in our area. [= FNA, Y; < A. hispidus – C, GW, K, Z; > A. hispidus var. cryptatherus (Hackel) 

Honda – RAB, F, G, HC, W] 

Arundinaria Michaux (Cane) 

Both species of Arundinaria were much reduced by the foraging of free-range livestock in the eighteenth and early nineteenth 

centuries and by fire suppression in the late nineteenth century and throughout the twentieth century. "Canebrakes," large areas 

dominated by cane, were described in many historical accounts and apparently occupied large parts of the landscape of the

POACEAE 849 

Coastal Plain, also occurring in the Piedmont and low Mountains. References: Tucker (1988)=Y; McClure (1973)=Z; McClure 

(1963); Judziewicz et al. (2000)=X; Triplett, Weakley, & Clark (2006)=Q. The key adapted from Y and Z. 

1 Primary branches with 0-1 compressed basal internodes; culm internodes usually sulcate; culm leaves deciduous; culms to 

10 m tall; rhizomes lacking air canals.......................................................................................................................A. gigantea 

1 Primary branches with 2-5 compressed basal internodes; culm internodes usually terete; culm leaves persistent to tardily 

deciduous; culms to 4 m tall; rhizomes with or without longitudinal air canals (visible in cross-section as a cylinder of 

hollow canals 1 mm or less from the outer surface). 

2 Foliage blades chartaceous, deciduous, abaxial surfaces pilose or glabrous, weakly tessellate; primary branches usually 

< 35 cm long, basal nodes not developing secondary branches; top knot blades 12-22.5 cm long; rhizomes with or 

without air canals ......................................................................................................................................A. appalachiana 

2 Foliage blades coriaceous, persistent, abaxial surfaces densely pubescent or glabrous, strongly tessellate; primary 

branches usually > 50 cm long, basal nodes developing secondary branches; top knot blades 20-30 cm long; rhizomes 

with air canals ......................................................................................................................................................... A. tecta 

Arundinaria appalachiana,Triplett, Weakley, & L.G. Clark, Hill Cane. Mt, Pd (GA, NC, SC): dry to moist forests on 

slopes; common. The short plants (often only knee-high, though sometimes head-high) on mountain slopes south of Asheville 

are autumn-deciduous, whereas both "rivercane" appears to be evergreen. [= Q; < A. gigantea (Walter) Walter – RAB, GW; < A. 

gigantea ssp. tecta (Walter) McClure – K, X, Z; < A. tecta – HC, S, Y; = A. tecta var. decidua Beadle in L.H. Bailey; apparently 

and implicitly included in the concept of either A. tecta or A. gigantea by earlier authors] 

Arundinaria gigantea (Walter) Walter, Giant Cane, River Cane. Mt, Pd, Cp (GA, NC, SC, VA): swamps, floodplains; 

common. April-July. Widespread in se. North America, ranging from s. DE s. OH, south to FL and e. TX. There has been much 

disagreement over the recognition of one, two, or several taxa of cane in the Southeastern United States. This species reaches 

heights of 6-7 (-10) m and is supposed to flower only once every 40-50 years. A. macrosperma Michaux is controversial, 

sometimes considered to be a synonym of A. gigantea or to represent hybridization or introgression between A. gigantea and A. 

tecta. [= F, HC, Q, S, Y; < A. gigantea – RAB, C, GW (also see A. tecta); = A. gigantea ssp. gigantea – K, Z; > A. gigantea ssp. 

gigantea – X; > A. gigantea (Walter) Walter ssp. macrosperma (Michaux) McClure – X; > A. macrosperma Michaux] 

Arundinaria tecta Walter, Switch Cane, Small Cane. Cp, Pd (GA, NC, SC, VA): savannas, pocosins, canebrakes, generally 

(but not solely) in wetlands; common. April-July. Primarily a Southeastern Coastal Plain endemic: e. VA to FL and s. AL. A. 

tecta is a smaller plant than A. gigantea (normally 1-2 m tall, but reaching heights of up to 4 m where fire-suppressed), and 

flowers more frequently, supposedly every 3-4 years (Tucker 1988), probably actually in response to fire. [= Q; < A. gigantea – 

RAB, C, GW; < A. tecta – F, HC, S, Y; < A. gigantea ssp. tecta (Walter) McClure – K, X, Z] 

Arundo Linnaeus (Giant Reed) 

A genus of 3 species, widespread in the tropics, subtropics and warm-temperate areas. References: Allred in FNA (2003a). 

* Arundo donax Linnaeus, Giant Reed. Cp, Pd (GA, NC, SC, VA), Mt (NC, VA): disturbed areas; uncommon, introduced 

from the Old World. September-October. Var. versicolor, with leaves transversely striped white and green, is better considered 

only as a form or cultivar. [= RAB, F, FNA, K, S; > A. donax var. donax – HC; > A. donax var. versicolor (P. Miller) Stokes – 

HC] 


Avena Linnaeus 1753 (Oats) 

1 Lemmas pubescent with brown hairs; lemmas with long bent awns.............................................................................. A. fatua 

1 Lemmas glabrous or scabrous; lemmas unawned or with relatively straight awns ....................................................... A. sativa 

* Avena fatua Linnaeus, Wild Oats. (VA). {needs herbarium checks}. [= C, F, G, HC, K] 

* Avena sativa Linnaeus, Oats. Mt (NC, SC, VA), Pd, Cp (GA, NC, SC, VA): fields and disturbed areas; commonly 

cultivated, uncommonly escaping. May-June. An important crop, but apparently only a weed until transported from the Middle 

East to the moister central Europe, where cultivated beginning about 3000 BP (Hancock 2004). [= RAB, G, HC, K, S, W, Z; > 

A. sativa var. orientalis (Schreber) Alefeld – F; > A. sativa var. sativa – F] 

Axonopus Palisot de Beauvois (Carpet Grass) 

A genus of ca. 100 species, primarily tropical and subtropical. Phylogenetic studies suggest that Axonopus may be included in 

Paspalum. References: Barkworth in FNA (2003a). 

1 Spikelets 4-6 mm long .............................................................................................................................................. A. furcatus

POACEAE 850 

1 Spikelets 1.5-2.8 mm long. 

2 Spikelets 1.5-2.2 mm long; leaf blades mostly 8-10 mm wide.....................................................................A. compressus 

2 Spikelets 2.2-2.8 mm long; leaf blades mostly 2-4 (-6) mm wide...................................................................A. fissifolius 

Axonopus compressus (Swartz) Palisot de Beauvois, Southern Carpet Grass. Cp (GA, SC, VA?): lawns; rare, probably 

introduced. Reported for VA by HC. Sometimes used as a lawn grass in the deep South. [= FNA, HC, K, S; Paspalum] 

Axonopus fissifolius (Raddi) Kuhlm., Common Carpet Grass. Cp, Pd (GA, NC, SC, VA): sandy forests, roadsides, lawns; 

common. June-October. VA south to FL, west to TX and OK, and extending into tropical America. [= FNA, K; ? A. affinis 

Chase – RAB, GW, HC, W; = Paspalum fissifolium Raddi] 

Axonopus furcatus (Flügge) A.S. Hitchcock, Big Carpetgrass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): sandy forests, 

bottomlands, roadsides, lawns; common. July-October. Se. VA south to FL, west to TX and AR. [= RAB, C, F, FNA, G, GW, 

HC, K, S; = Paspalum furcatum Flügge] 

Bambusa Schreber (Bamboo) 

* Bambusa oldhamii Munro is reported for NC (Kartesz 1999). {investigate} [=Sinocalamus latiflorus (Munro) McClure – 

K, misapplied] 

* Bambusa vulgaris Schrader ex J.C. Wendland, Common Bamboo, is reported for SC (Kartesz 1999). {investigate} [= K] 

Bothriochloa Kuntze (Beardgrass, Cane Bluestem) 

A genus of ca. 35 species, widespread in tropical and subtropical regions of the Old and new World. References: Allred in FNA 

(2003a); Vega (2000)=Z; Allred & Gould (1983)=Y. Key adapted from Allred in FNA (2003a). 

1 Sessile spikelets 4.5-8.5 mm long......................................................................................................................... B. barbinodis 

1 Sessile spikelets 3-4.5 mm long. 

2 Pedicellate spikelets about as long as the sessile spikelets. 

3 Rachises longer than the branches.............................................................................................................[B. bladhii] 

3 Rachises shorter than the branches...............................................................................B. ischaemum var. songarica 

2 Pedicellate spikelets much shorter than the sessile spikelets. 

4 Panicles reddish when mature; hairs below the sessile spikelets sparse and ca. 1/4 as long as the spikelets, not 

obscuring the spikelets ..............................................................................................................................[B. bladhii] 

4 Panicles silvery-white or tannish when mature, hairs below the sessile spikelets dense and > ½ as long as the 

spikelets, somewhat obscuring the spikelets ..................................................................B. laguroides ssp. torreyana 

* Bothriochloa barbinodis (Lagasca y Segura) Herter, Cane Bluestem, Pinhole Bluestem. Cp, Pd (SC): disturbed areas; rare, 

introduced from w. United States. [= FNA, K; > Bothriochloa perforata (Trinius ex E. Fourn.) Herter – Z; = Andropogon 

barbinodis Lagasca y Segura – HC; > Bothriochloa barbinodis (Lagasca y Segura) Herter var. perforata (Trinius ex E. Fourn.) 

Gould; > Andropogon perforatus Trinius ex E. Fourn.] 

* Bothriochloa ischaemum (Linnaeus) Keng var. songarica (Ruprecht ex Fischer & C.A. Meyer) Celarier & Harlan, King 

Ranch Bluestem. Cp (SC): disturbed places; rare, introduced from western North America. Reported for SC (Kartesz 1999). [= 

K, Z; < B. ischaemum – FNA] 

* Bothriochloa laguroides (Augustin de Candolle) Herter ssp. torreyana (Steudel) Allred & Gould, Silver Bluestem. Cp 

(GA, SC), Pd (GA): disturbed areas; rare, introduced from {}. Reported for SC (Kartesz 1999), ne. GA (Jones & Coile 1988; 

Allred & Gould 1983), e. TN, and c. TN (Chester et al. 1993), in some cases as B. saccharoides var. torreyana. [= FNA, K, Y, Z; 

= B. saccharoides (Sw.) Rydberg var. torreyana (Steudel) Gould] 

* Bothriochloa bladhii (Retzius) S.T. Blake, Australian Bluestem, is reported from e. TN (according to specimen cited by 

FNA and Z) and FL. [= FNA, K, Z] {synonymy incomplete} 

* Bothriochloa pertusa (Linnaeus) A. Camus, Pitted Bluestem. Introduced at scattered sites in e. North America, including 

FL, LA, MD, and MS (FNA, Kartesz 1999). [= FNA, K, Z] {not keyed at this time; synonymy incomplete} 

Bouteloua Lagasca y Segura (Grama) 

A genus of about 40 species, of the Western Hemisphere. References: Herrera Arrieta, Peterson, & de la Cerda Lemus 

(2004)=X; Columbus (1999)=Z; Gould (1979)=Y; Wipff in FNA (2003a); Snow in FNA (2003a). Key based in part on Wipff in 

FNA (2003a) 

1 All spikelets unisexual, plants usually dioecious; [introduced species] ................................................................B. dactyloides 

1 Lowest floret in each spikelet bisexual, the upper staminate or sterile; [introduced or native species].

POACEAE 851 

2 Panicle branches deciduous; disarticulation occurring at the base of the branch (the branch therefore falling whole); 

spikelets 2-3 per branch, appressed to the branch; [native species of limestone habitats, also with introduced 

populations]; [subgenus Bouteloua]..............................................................................B. curtipendula var. curtipendula 

2 Panicle branches persistent; disarticulation occurring above the glumes (the individual florets therefore falling); 

spikelets >6 per branch, pectinately disposed; [rare introductions]; [subgenus Chondrosum]. 

3 Panicle branches terminating in a spikelet ..................................................................................................B. gracilis 

3 Panicle branches extending beyond the base of the terminal spikelets.................................... B. hirsuta var. hirsuta 

Bouteloua curtipendula (Michaux) Torrey var. curtipendula, Side-oats Grama. Mt (GA, VA), Pd (GA): dry rocky slopes 

and bluffs over limestone or serpentine, limestone glades; uncommon. July-September. S. CT west to MT, south to VA, e. TN, 

nw. GA, AL, panhandle FL, TX, AZ, and CA; also in Central and South America. The older literature refers to B. curtipendula 

as introduced in SC, but the single specimen documenting its occurrence there appears to be from experimental plantings at 

Clemson University; there is apparently no evidence of its establishment. B. curtipendula occurs on serpentine in the Piedmont 

of Georgia (Allison, pers. comm.). Var. caespitosa Gould & Kapadia is cespitose rather than rhizomatous and occurs in sw. 

United States. [= C, FNA, K, Y; < B. curtipendula – RAB, F, G, HC, S, W] 

* Bouteloua dactyloides (Nuttall) J.T. Columbus, Buffalo Grass. Mt (VA), Pd, Cp (GA): disturbed areas; rare, introduced 

from w. North America. [= Z; = Buchloe dactyloides (Nuttall) Engelmann – C, F, FNA, G, HC, K] 

* Bouteloua gracilis (Willdenow ex Kunth) Lagasca y Segura ex Griffiths, Blue Grama. Cp (SC): disturbed areas; rare, 

introduced from western North America. Reported for SC (Gould 1979). [= F, FNA, K, Y; > Bouteloua gracilis var. gracilis – 

HC] 

* Bouteloua hirsuta Lagasca y Segura var. hirsuta, Hairy Grama. Cp (GA, SC): disturbed areas; rare, introduced from 

western North America. Reported for SC and GA (Kartesz 1999). [= K, Y; < Bouteloua hirsuta – F, HC; = Bouteloua hirsuta 

ssp. hirsuta – FNA] 

Brachiaria 

(see Urochloa) 

Brachyelytrum Palisot de Beauvois (Shorthusk) 

The only other species of the genus is B. japonicum Hackel, of s. Japan, Korea, and ec. China (Saarela et al. 2003, Tucker 1988). 

References: Saarela et al. (2003)=Z; Tucker (1988)=Y; Stephenson (1971); Voss (1972); Campbell, Garwood, & Specht (1986). 

Key based in part on Saarela et al. (2003). 

1 Lemmas hirsutulous or minutely scabrous, the longest hairs (0.06-) 0.08-0.14 (-0.2) mm long (not evident at 10×); lemma 

(0.7-) 0.8-1.2 (-1.4) mm wide; widest leaf blade (8-) 10-14 (-16) mm wide; second glume (0.6-) avg. 1.2 (-3.0) mm long; 

[plants of the Mountains]........................................................................................................................................B. aristosum 

1 Lemmas hirsute with hairs (0.2-) 0.4-0.8 (0.9) mm long (easily seen at 10×); lemma (0.8-) 1.1-1.5 (-1.8) mm wide; widest 

leaf blade (9-) 11-17 (-20) mm wide; second glume (0.2-) avg. 2.2 (7.0) mm long; [plants widely distributed in our area] ...... 

...................................................................................................................................................................................B. erectum 

Brachyelytrum aristosum (Michaux) Trelease in Branner & Coville, Northern Shorthusk. Mt (GA, NC, VA): moist forests, 

mostly at moderate to high elevations, such as northern hardwoods and spruce-fir; rare (GA Special Concern, NC Watch List). 

July-August. Fairly widespread in ne. North America, south in the mountains to sw. NC and n. GA. In MI, B. septentrionale 

flowers about 10 days before co-occurring B. erectum, with strongly synchronized anthesis of each species occurring on a single 

day (Stephenson 1971)). Reputed intermediates and hybrids between the two taxa are apparently based on the use of ambiguous 

characters. [= Z; = Brachyelytrum septentrionale (Babel) G. Tucker – K, Y; < B. erectum – RAB, G, HC, S, W; = B. erectum 

var. septentrionale Babel – F; = B. erectum var. glabratum (Vasey ex Millspaugh) Koyama & Kawano – C] 

Brachyelytrum erectum (Schreber ex Sprengel) Palisot de Beauvois, Common Shorthusk. Mt, Pd (GA, NC, SC, VA), Cp 

(NC, SC, VA): mesic forests, in the Mountains at lower elevations than B. septentrionale); common (uncommon in Piedmont, 

rare in Coastal Plain). June-August. MA, NY, OH, MI, and s. WI south to FL and TX. [= K, Y, Z; < B. erectum – RAB, G, HC, 

S, W (also see B. septentrionale); = B. erectum var. erectum – C, F] 

Brachypodium Palisot de Beauvois 

* Brachypodium sylvaticum (Hudson) Palisot de Beauvois ssp. sylvaticum, Slender False Brome. Pd (VA): roadsides and 

yards; rare, introduced from Europe. [= FNA; < B. sylvaticum – HC, K] 


Briza Linnaeus (Quaking Grass)

POACEAE 852 

* Briza maxima Linnaeus, Greater Quaking Grass. Cp (GA): disturbed areas; rare, introduced. Reported in e. GA (Jones & 

Coile 1988). [= K] {not keyed at this time; synonymy incomplete} 

* Briza minor Linnaeus, Lesser Quaking Grass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): fields, disturbed areas; common, 

introduced from Europe. April-June. [= RAB, C, F, G, GW, HC, K, S, Z] 

* Briza media Linnaeus, Perennial Quaking Grass, reported for scattered locations in PA (Rhoads & Klein 1993), MD, DE, 

and AL (Kartesz 1999). [= K] {not keyed at this time; synonymy incomplete} 

Bromus Linnaeus 1753 (Brome-grass) 

A genus of about 150 species, north temperate and South American. References: McNeill (1976); Sales (1993, 1994)=Z; Tucker 

(1996)=Y; Pavlick (1995)=X; McKenzie & Ladd (1995); Pavlick & Anderton in FNA (in prep.). 

1 Lemmas compressed and strongly keeled (the whole spikelet thus strongly laterally flattened); first glume 3-9-nerved; 

[section Ceratochloa] ...........................................................................................................................................B. catharticus 

1 Lemmas rounded or weakly keeled (the whole spikelet therefore terete to somewhat laterally flattened); first glume either 3- 

5-nerved or 1-3-nerved. 

2 First glume 3-5 nerved (at least 3 nerves well-developed). 

3 Lemma awn 2-3 mm long; plant perennial; [native species of dry woodlands]; [section Pnigma] .............. B. kalmii 

3 Lemma awn 3-12 mm long (or 0-6 mm long in B. secalinus); plant annual; [introduced species of disturbed 

habitats]; [section Bromus]. 

4 Panicle compact, the lateral branches erect or ascending, the pedicels < 10 mm long (shorter than the 

spikelets) ........................................................................................................... B. hordeaceus ssp. hordeaceus 

4 Panicle relatively open, the lateral branches erect, ascending, or spreading, the pedicels > 15 mm long (longer 

than the spikelets). 

5 Margins of the lemmas involute in fruit, wrapping around the grain, exposing the rachilla.....B. secalinus 

5 Margins of the lemmas gaping, overlapping in fruit. 

6 Panicle branches erect or ascending, relatively stiff and straight .................................. B. racemosus 

6 Panicle branches spreading (at least the lower), either relatively stiff and straight, or flexuous and 

lax. 

7 Panicle branches stiff; lemma awns 5-12 mm long, straight................................B. commutatus 

7 Panicle branches flexuous and lax; lemma awns 7-15 mm long, flexuous ..............B. japonicus 

2 First glume 1 (-3) nerved (only 1 nerve well-developed). 

8 Longer lemma awns 10-60 mm long; plants annual; [introduced species of disturbed habitats]; [section Genea]. 

9 Panicle dense, spikelike........................................................................................................................B. rubens 

9 Panicle open, not spikelike. 

10 First glume 13-20 mm long; second glume 20-30 mm long; lemma awns 35-60 mm long......... B. rigidus 

10 First glume 5-14 mm long; second glume 8-17 mm long; lemma awns 10-30 mm long. 

11 First glume 7-14 mm long; second glume 9-17 mm long; lemma awns 18-30 mm long......B. sterilis 

11 First glume 5-7 mm long; second glume 8-11 mm long; lemma awn (7-) 10-17 mm long ................. 

.......................................................................................................................................... B. tectorum 

8 Longer lemma awns 1-6 (-8) mm long; plants perennial; [native and introduced species, collectively of disturbed 

and natural habitats]. 

12 Plants with creeping rhizomes, forming clonal colonies; both surfaces of leaves glabrous or glabrescent; 

[section Bromopsis]............................................................................................................................ B. inermis 

12 Plants not strongly rhizomatous, the stems solitary or tufted; surfaces of leaf blades usually pubescent 

(sometimes sparsely so). 

13 Pedicels erect or ascending, mostly shorter than the spikelet; leaves 2-3 mm wide; [introduced, of 

disturbed habitats]; [section Bromopsis] ..................................................................................... B. erectus 

13 Pedicels ascending at first, later arching-drooping, mostly longer than the spikelet; leaves 4-15 mm 

wide; [native, mostly of forests]; [section Pnigma]. 

14 Lemmas glabrous (or very minutely pubescent) on the back, hairy along the lower margins with 

long hairs.............................................................................................................................B. ciliatus 

14 Lemmas uniformly hairy over the entire back-surface (or rarely entirely glabrous). 

15 Culms with 10-20 leaves, often weak and leaning or reclining; junction of sheaths and base of 

leaf blades with 2 well-developed flanges prolonged into auricles or divergent spurs; second 

glume primarily 5-nerved; flowering late, with anthesis August-October...............B. latiglumis 

15 Culms with 6-10 leaves, erect; junction of sheaths and base of leaf blades lacking flanges or 

auricles; second glume primarily 3-nerved; flowering earlier, anthesis from May-August. 

16 Underleaf surfaces with a conspicuous satiny sheen (when fresh); summit of sheath 

opposite the ligule with a conspicuous tuft of hairs.................................. B. nottowayanus

POACEAE 853 

16 Underleaf surfaces lacking a conspicuous satiny sheen; summit of sheath opposite the 

ligule lacking a conspicuous tuft of hairs........................................................ B. pubescens 

* Bromus catharticus Vahl, Rescue Grass. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): disturbed areas; common, native 

of South America. April-June. [= RAB, F, G, HC, K, W, X, Y; ? Bromus willdenowii Kunth – C; = Bromus unioloides Kunth – 

S] 

Bromus ciliatus Linnaeus, Fringed Brome. Mt (NC, VA): seepage areas, edges of fens, moist areas near high elevation 

creeks, grassy balds, high elevation woodlands, mostly over amphibolite or other mafic rocks; rare (NC Rare, VA Rare). July- 

August. Widespread in n. North America: Labrador to AK, south in the east to PA, and in the mountains to NC. Known in NC 

only from Bluff Mountain and Long Hope Valley, Ashe and Watauga counties, and Roan Mountain, Mitchell County. [= RAB, 

C, G, HC, S, W, X, Y; > Bromus ciliatus var. ciliatus – F, K; = Bromopsis ciliata (Linnaeus) Holub] 

* Bromus commutatus Schrader, Hairy Chess, Meadow Brome. Pd, Mt (GA, NC, SC, VA), Cp (NC, SC, VA): disturbed 

areas; common, native of Europe. May-June. The relationship and relative distribution of this species and Bromus racemosus is 

poorly known for our area. See Bromus racemosus for further comments. [= C, F, HC, K, S, X, Y; < Bromus commutatus – 

RAB (also see Bromus racemosus); < Bromus racemosus – G, W] 

* Bromus erectus Hudson, Short-branched Brome. Mt, Pd (VA): disturbed areas; rare, native of Europe. [= C, F, G, HC, K, 

S, X; = Bromopsis erecta (Hudson) Fourr.] 

* Bromus hordeaceus Linnaeus ssp. hordeaceus, Soft Chess, Lopgrass. Mt (NC, VA), Pd (VA), Cp (SC, VA): disturbed 

areas; rare, native of Europe. July. [= K, X; ? Bromus mollis Linnaeus – RAB, F, G, HC, misapplied; < Bromus hordeaceus – 

C, Y] 

* Bromus inermis Leysser, Smooth Brome, Hungarian Brome. Cp (NC, SC, VA), Mt (NC, VA), Pd (VA): disturbed areas; 

rare, native of Europe. June-July. [= RAB, C, G, HC, S, W, X, Y; > Bromus inermis var. inermis – F; > Bromus inermis ssp. 

inermis var. inermis – K; = Bromopsis inermis (Leysser) Holub] 

* Bromus japonicus Thunberg ex Murray, Japanese Chess. Mt, Pd, Cp (GA, NC, SC, VA): disturbed areas; common, native 

of Asia. May-June. [= RAB, C, G, K, S, W, X, Y; > Bromus japonicus var. japonicus – F, HC; > Bromus japonicus var. 

porrectus Hackel – F, HC] 

Bromus kalmii A. Gray, Kalm Brome. Mt (VA): shale woodlands and barrens; rare (VA Rare). ME west to SD, south to 

MD, w. VA, and IA. Distinctive for its few leaves (usually 3-4) clustered near the base, the spikelets large and approximate to 

one another in a narrow, nodding panicle. [= C, F, G, HC, K, X] 

Bromus latiglumis (Shear) A.S. Hitchcock, Riverbank Brome, Auricled Brome, Hairy Woodbrome, Flanged Brome. Mt 

(NC, VA), Pd (VA): alluvial soils along rivers; rare (NC Watch List). August-October. Widespread in ne. North America, from 

ME to MT, south to NC and OK. Flowering many weeks later than co-occurring B. pubescens. In NC apparently only along 

large rivers flowing west through the Appalachians into the Mississippi River drainage, notably the New and the French Broad. 

[= F, G, HC, K, X; < Bromus purgans Linnaeus – RAB; = Bromus altissimus Pursh – C; < Bromus latiglumis – Y (also see 

Bromus nottowayanus)] 

Bromus nottowayanus Fernald, Satin Brome, Nottoway River Brome, Virginia Brome. Pd (NC, VA), Cp, Mt (VA): moist 

forests, especially along small stream bottoms; rare (NC Watch List). June-August. The range of this species is poorly known, 

owing to confusion between it, B. pubescens and B. latiglumis. It is apparently known from MD, VA, and NC, west to TN, IL, 

IN, MO, and AR. McKenzie & Ladd (1995) report on the biology and taxonomy of this species. [= C, F, HC, K, X; < Bromus 

purgans Linnaeus – RAB; < Bromus latiglumis – Y; = Bromopsis nottowayana (Fernald) Holub] 

Bromus pubescens Muhlenberg ex Willdenow, Common Eastern Brome, Canada Brome. Mt, Pd (GA, NC, SC, VA), Cp 

(GA, VA): mesic forests, generally on rocky slopes, common. May-August. Widespread in e. North America: s. Ontario west 

to Alberta, south to FL and AZ. [= C, K, W, X, Y; < Bromus purgans Linnaeus – RAB, S, misapplied (also see Bromus 

latiglumis and Bromus nottowayanus); = Bromus purgans Linnaeus – F, G, misapplied; > Bromus purgans var. purgans – HC; > 

Bromus purgans var. laeviglumis (Scribner ex Shear) Swallen – HC; = Bromus laeviglumis – S, misapplied (?); = Bromopsis 

pubescens (Muhlenberg ex Willdenow) Holub] 

* Bromus racemosus Linnaeus, Smooth Brome. Pd, Cp, Mt? (NC, SC, VA): disturbed areas, native of Europe. May-June. 

The relative distribution, abundance, and habitats in our area of this species and B. commutatus poorly understood. Additional 

characters are as follows (from Stace 1997): lemmas 6.5-8 mm long (vs. 8-11 mm long in B. commutatus), anthers mostly 1.5-3 

mm long (vs. mostly 1-1.5 mm long), spikelets 10-16 mm long (vs. 15-28 mm long), lowest rachilla segment mostly 0.7-1 mm 

long (vs. mostly 1.3-1.7 mm). [= C, F, HC, K, X; < Bromus commutatus – RAB; < Bromus racemosus – G, W (also see Bromus 

commutatus)] 

* Bromus rigidus Roth, Ripgut Brome, Ripgut Grass. Cp (NC, SC, VA), Pd (GA, NC): disturbed areas; rare, native of 

Mediterranean Europe. April. [= RAB, C, F, G, HC, K; ? Bromus diandrus Roth – Y; ? Bromus diandrus var. ?? – Z] 

* Bromus rubens Linnaeus, Foxtail Chess, Red Brome. Cp (SC, VA): disturbed areas; rare, introduced from Mediterranean 

Europe. Specimens in our area come from areas around wool-combing plants, and were likely introduced on wool from w. 

United States, where this European species is well-established. [= C, G, K, X; ? Bromus madritensis Linnaeus – F, misapplied; = 

Bromus madritensis ssp. rubens (Linnaeus) Husnot] 

* Bromus secalinus Linnaeus, Cheat, Common Chess, Ryebrome. Pd, Cp, Mt (GA, NC, SC, VA): disturbed areas; common, 

native of Europe. May-June. [= RAB, C, F, G, HC, K, S, W, X, Y] 

* Bromus sterilis Linnaeus, Barren Brome, Poverty Brome, Cheatgrass. Pd (NC, VA), Mt, Cp (VA): disturbed areas; rare, 

native of southern Europe. May-June. [= RAB, C, F, G, HC, K, S, W, X, Y, Z]

POACEAE 854 

* Bromus tectorum Linnaeus, Downy Brome, Downy Chess, Downy Cheat, Junegrass, Cheatgrass. Mt, Pd (GA, NC, SC, 

VA), Cp (NC, SC, VA): disturbed areas; common, native of Europe. April-June. [= RAB, C, F, G, HC, K, S, W, X, Y; ? 

Bromus tectorum ssp. tectorum – Z] 

* Bromus arvensis Linnaeus has been reported as introduced for nc. GA (Jones & Coile 1988) and for VA, MD, PA, and NJ 

(Kartesz 1999). {investigate} [= C, K] {not keyed at this time} 

* Bromus briziformis Fischer & C.A. Meyer. Reported as an introduction in ne. North America, south to MD, NJ, PA, DE 

(Kartesz 1999). [= K; = B. brizaeformis – C, orthographic variant] {not keyed at this time} 

* Bromus carinatus Hooker & Arnott. Reported by Jones & Coile (1988) for nc. GA. [= C, K] {not keyed at this time} 

* Bromus madritensis Linnaeus. Reported introduced in VA and MD (Kartesz 1999). {investigate} [= K] {not keyed at this 

time} 

* Bromus ramosus Hudson. Introduced. Reported for DC and MS (Kartesz 1999). [= K] {not keyed at this time} 

* Bromus squarrosus Linnaeus. Introduced. Reported for KY and NJ (Kartesz 1999). [= K] {not keyed at this time} 

Buchloe Engelmann (Buffalo Grass) 

(see Bouteloua) 

Calamagrostis Adanson (Reed-grass) 

A genus of about 230 species, north and south temperate. References: Marr, Hebda, & Greene in FNA (in prep.); Tucker 

(1996)=Z; Greene (1980). 

1 Awn sharply bent; callus hairs 1/2 or less the length of the lemma; [subgenus Ankylatherae]. 

2 Plant densely tufted, delicate, the culms 30-60 cm tall, with 2-3 nodes; leaves 1-2 mm wide, involute; callus hairs about 

1 mm long (< ½ the length of the lemma)..............................................................................................................C. cainii 

2 Plant rhizomatous or loosely tufted, coarse, the culms 60-120 cm tall, with 3-5 nodes; leaves 4-8 mm wide, flat; callus 

hairs about 2 mm long (< ½ the length of the lemma) .....................................................................C. porteri ssp. porteri 

1 Awn straight; callus hairs 3/4 as long as to equal to the lemma. 

3 Panicle loose, spreading, 3.5-6 cm wide; callus hairs 2-3 mm long (about ¾ the length of the lemma); [subgenus 

Calamagrostis]....................................................................................................................C. canadensis var. canadensis 

3 Panicle dense, erect, 1.5-4 cm wide; callus hairs 3-4 mm long (about equal to the lemma); [subgenus Orthatherae]........ 

........................................................................................................................................................................ C. cinnoides 

Calamagrostis cainii A.S. Hitchcock, Cain's Reed-grass. Mt (NC): high elevation rocky summits; rare (US Species of 

Concern, NC Endangered). July-September. Endemic to a few mountain-tops in the Southern Appalachians, C. cainii, once 

thought to be endemic to Mount LeConte, TN, was discovered at two sites in NC in 1989 and 1990 – Mount Craig, Yancey 

County, and Craggy Pinnacle, Buncombe County (Wiser 1991). This species is more likely to be mistaken (especially 

superficially) for an Agrostis than for any of the other Calamagrostis in our area, but is distinguishable by its larger spikelets (5-6 

mm long, rather than 1.3-2 mm) and the presence of a callus beard. [= FNA, HC, K, W, Z] 

Calamagrostis canadensis (Michaux) Palisot de Beauvois var. canadensis, Bluejoint, Canada Reed-grass. Mt (GA, NC, 

VA): wet meadows along streams, high elevation openings, such as grassy balds and cliff bases; uncommon (NC Rare). August. 

Widespread and common across n. North America, reaching its southern limit in the east in w. NC, e. TN (Chester et al. 1993), 

and ne. GA (Rabun Bald, Rabun County). [= FNA, G, HC, K; < C. canadensis – RAB, C, S, W, Z; > C. canadensis var. robusta 

Vasey – F] 

Calamagrostis cinnoides (Muhlenberg) W.P.C. Barton, Nuttall's Reed-grass. Mt (GA, NC, SC, VA), Cp, Pd (NC, SC, VA): 

savannas, bogs, and other wet sites; common. July-October. ME and NY south to n. GA (Jones & Coile 1988), AL, and LA, 

primarily on the Coastal Plain. The replacement of the familiar C. cinnoides is suggested for nomenclatural reasons (Kartesz 

1999). [= RAB, C, F, G, GW, HC, S, W, Z; = C. coarctata Torrey ex Eaton – K] 

Calamagrostis porteri A. Gray ssp. porteri, Porter's Reed-grass. Mt (GA, NC, SC, VA), Pd (VA): dry to dry-mesic forests, 

forest edges, cliff bases; uncommon, rare in NC (GA Special Concern, NC Rare). NY to AL, in the Appalachians, with disjunct 

populations s. MO and w. AR; it was first reported from NC by Ware (1973). In addition to the key characters above, it can be 

distinguished from C. canadensis by its having leaf sheaths pubescent at the summit (Matthews & Radford 1985). [= K; = C. 

porteri – C, HC, W; ? C. porteri – G, Z] 

Calamagrostis canadensis (Michaux) Palisot de Beauvois var. macouniana (Vasey) Stebbins. Reported for VA (FNA). 

Reported south to NJ and KY only (Kartesz 1999). [= F, FNA, G, HC, K; < C. canadensis – C, Z; = C. macouniana (Vasey) 

Vasey] {not keyed at this time} 

Calamagrostis porteri A. Gray ssp. insperata (Swallen) C.W. Greene, ranges east to KY and TN (Kartesz 1999). [= K; = C. 

insperata Swallen – C, HC] {not keyed at this time} 

Calamagrostis stricta (Timm) Koeler ssp. inexpansa (A. Gray) C.W. Greene ranges south to n. WV (Preston and Randolph 

counties). [= K; < C. stricta – C; ? C. neglecta (Ehrhart) Gaertner, Mey., & Scherb. var. neglecta – F] {not keyed at this time}

POACEAE 855 

Calamovilfa (A. Gray) Hackel ex Scribner & Southworth (Sandreed) 

A genus of 5 species, of e. and c. North America. Reeder & Ellington (1960) studied various anatomic features of Calamovilfa, 

and determined that its closest relative was Sporobolus. A molecular phylogenetic study of Sporobolus and closely related 

genera suggests that Calamovilfa should be included in Sporobolus (Ortiz-Diaz & Culham 2000). References: Thieret (1966)=Z. 

Key based in part on Thieret in FNA (in prep.). 

Identification notes: Superficially somewhat similar to Sporobolus pinetorum, S. floridanus, and S. curtissii (herbarium 

specimens of the two species have been regularly confused), Calamovilfa is distinguished by its leaves tapered to either end and 

long-acuminate (vs. parallel-margined and abruptly acute in Sporobolus) and tendency to form larger, clonal patches (Sporobolus 

forms wiregrass-like bunches or clumps). In flower or fruit, the Calamovilfa can be distinguished by characters of the spikelet, 

by vegetative characters, or by its coarser, generally taller culms, with the panicle branches usually spreading (rather than always 

ascending in Sporobolus). The three have very similar bases, unlike any other grasses in our area – the lower leaf sheaths are 

indurated and shiny, forming a hard, polished, knotty, and fire-proof covering over the short-creeping rhizome. Aristida stricta 

has a somewhat similar base, but less indurated, less creeping, and with an unpolished appearance. Calamovilfa brevipilis also 

has a cartilaginous, pale yellow annulum surrounding the outer (abaxial) surface of the juncture of the sheath and leaf, a structure 

not visible in the other species. Positive identification in sterile condition is not difficult. 

1 Panicles narrow, the branches appressed-ascending; [of the Coastal Plain of FL].................................................. [C. curtissii] 

1 Panicles broad, the bracnches ascending-spreading; [either of the Coastal Plain of SC northwards, or of the interior]. 

2 Spikelets 6-7.4 mm long; glumes acute to acuminate, usually arcuate; lemmas 5.5-7 mm long, usually arcuate; [of river 

scour areas in the rocky inland parts of the South]...........................................................................................[C. arcuata] 

2 Spikelets 4-5.8 mm long; glumes acute, straight; lemmas 4-5.4 mm long, straight; [of pineland habitats of the Coastal 

Plain of South Carolina northwards .................................................................................................................C. brevipilis 

Calamovilfa brevipilis (Torrey) Scribner, Pinebarren Sandreed. Cp (NC, SC, VA), Pd (NC): savanna-pocosin ecotones, 

sandhill seepage bogs, pocosins; rare (NC Watch List, SC Rare, VA Rare). June-October. A "bimodal endemic", with two areas 

of distribution: Pine Barrens of NJ and the Coastal Plain (very rarely lower Piedmont) of e. NC, n. SC, and s. VA. Var. 

heterolepis Fernald, no longer considered valid, refers to the NC-SC material; var. calvipes Fernald, no longer considered valid, 

refers to the VA material. Like Aristida stricta, this grass is dependent on fire for flowering (it will also sometimes flower in 

response to mowing or other disturbance). Suppression of the natural fire regime has led to its substantial decline and the severe 

contraction of its range in the Southeast, since fire exclusion in its seepage or ecotone habitat leads to rapid invasion by shrubs 

and competitive elimination of Calamovilfa and many other herbs. [= RAB, C, FNA, G, GW, K, S, Z; > C. brevipilis var. 

brevipilis – F, HC; > C. brevipilis var. heterolepis Fernald – HC; > C. brevipilis var. calvipes Fernald – F, HC] 

Calamovilfa arcuata K.E. Rogers occurs in Morgan and Cumberland counties, TN, in the Cumberland Plateau, and in AL. 

[= FNA, K] 

Calamovilfa curtissii (Vasey) Scribner of FL is a closely related sibling species of C. brevipilis. It occurs in the FL 

Panhandle and e. peninsular FL. [= FNA, GW, HC, K, S, Z] 

References: Soreng et al. (2003)=Z. 

Catapodium Link 1827 

* Catapodium rigidum (Linnaeus) Dony, native to Europe, is known from collections from wool-combing mills in South 

Carolina; it is probably not established. [= Z; = Desmazeria rigida (Linnaeus) Tutin – K; = Scleropoa rigida (Linnaeus) 

Grisebach] 

Cenchrus Linnaeus (Burgrass, Sandspur) 

A genus of about 16 species, primarily tropical and subtropical. References: Stieber & Wipff in FNA (2003a); Crins (1991)=Z. 

{vegetative characters} {VA distribution of C. tribuloides} 

Identifications note: Spikelets of Cenchrus are subtended by an involucre of spines and/or bristles which are (in most of our 

species) fused into a bur. Bristles are narrow-based and terete. Spines are broad-based, and somewhat flattened (not terete) in 

cross-section, at least basally. 

1 Involucre of bristles only, these not fused into a bur; perennial, to 2 m tall....................................................... C. myosuroides 

1 Involucre of spines fused into a coherent bur, sometimes also with bristles; annual or perennials, to 1 m tall. 

2 Spines in a single whorl, subtended by numerous smaller, narrower bristles..................................................C. echinatus

POACEAE 856 

2 Spines in multiple whorls or irregular in their disposition (if few and in a single whorl, then not subtended by smaller, 

narrower bristles). 

3 Burs (excluding the spines) 9-16 mm long, 4-6 mm wide, the spines 4-8 mm long; spikelets 1 (-2) per bur, 

concealed; leaf blades 3-14 mm wide ................................................................................................... C. tribuloides 

3 Burs (excluding the spines) 5.5-12 mm long, 2.5-6 mm wide, the spines 2-7 mm long; spikelets 2-4 per bur, 

exserted at the tip; leaf blades 1-5 (-7) mm wide. 

4 Spines slender, 45-75, 3.5-7 mm long; spikelets 6-8 mm long.....................................................C. longispinus 

4 Spines stout, 6-10 (-40), 2-5 mm long; spikelets 3.5-6 mm long .......................................................C. spinifex 

Cenchrus echinatus Linnaeus, Southern Sandspur, Bristly Sandspur, Hedgehog Grass. Cp (GA, NC, SC), Pd (GA, SC): 

fields, roadsides, disturbed areas; common (uncommon in NC). June-October. NC (and DC?) south to FL, west to CA, south 

into the tropical America. [= RAB, C, HC, K, S, Z] 

Cenchrus longispinus (Hackel) Fernald, Northern Sandspur, Common Sandspur. Cp, Pd, Mt (GA, NC, SC, VA): fields, 

roadsides, disturbed areas, lawns; common (uncommon in Mountains). June-October. ME west to OR, south to FL, TX, and 

CA. [= RAB, C, F, FNA, K, W, Z; = C. pauciflorus Bentham – G, HC, S, misapplied] 

* Cenchrus myosuroides Kunth. Cp (SC): roadsides, disturbed areas; rare, introduced from further south. December. SC 

south to FL, west to TX, south into the West Indies and other parts of tropical America. [= RAB, FNA, HC, K, S, Z] 

Cenchrus spinifex Cavanilles, Coastal Sandspur. Cp (GA, NC, SC, VA), Pd (GA, SC, VA), Mt (VA): fields, roadsides, 

disturbed areas; common (rare in NC and VA). July-October. VA south to FL, west to AR and KS, south into tropical America. 

[= FNA, K; > C. incertus M.A. Curtis – RAB, C, F, G, HC, S, Z] 

Cenchrus tribuloides Linnaeus, Dune Sandspur. Cp (GA, NC, SC, VA), Pd*, Mt* (VA): dunes, sandy fields, sandy 

woodlands in the outer Coastal Plain; common. August-October. NY (Long Island) south to FL, west to TX, south into tropical 

America. This is the sandspur so familiar and disliked by beach-goers in our area. [= RAB, C, F, FNA, HC, K, S, W, Z] 

Cenchrus brownii Roemer & J.A. Schultes. Reported for NC (Kartesz 1999) and GA (FNA). {investigate} [= FNA, K] 

{not keyed at this time; add to synonymy} 

Cenchrus gracillimus Nash. Reported for sc. GA by Jones & Coile (1988) and FNA. [= FNA, K] {not keyed at this time; 

add to synonymy} 

Chasmanthium Link (Spanglegrass, Spikegrass) 

A genus of 5 species endemic to se. North America. References: Sánchez-Ken & Clark in FNA (2003a); Yates (1966a, 

1966c)=Z. 

1 Panicle branches elongate, pendulous; spikelets (15-) 20-40 mm long, with 6-20 flowers...................................Ch. latifolium 

1 Panicle branches short, erect or ascending; spikelets 5-18 mm long, with 2-8 (-11) flowers. 

2 Fully-developed spikelets 12-18 mm long, 8-12 mm wide. 

3 Axils of the spikelets and panicle branches glabrous; empty lemmas 9 (-12); [se. NC south to c. peninsular FL and 

e. FL panhandle] .....................................................................................................................................Ch. nitidum 

3 Axils of the spikelets and panicle branches with a tuft of long hairs; empty lemmas 2-4; [w. FL panhandle west to 

e. LA (Florida parishes)].......................................................................................................[Ch. ornithorhynchum] 

2 Fully-developed spikelets 4-9 mm long, 3-7 mm wide. 

4 Collar (junction of leaf and sheath) glabrous or nearly so; leaves 3-7 mm wide........................................ Ch. laxum 

4 Collar (junction of leaf and sheath) pilose; leaves 6-12 mm wide. 

5 Inflorescence with divergent branches; [in outer Coastal Plain calcareous sites from SC southwards] .............. 

...................................................................................................................................... Ch. sessiliflorum var. 1 

5 Inflorescence with appressed branches; [more widespread in our area]......Ch. sessiliflorum var. sessiliflorum 

Chasmanthium latifolium (Michaux) Yates, River Oats, Fish-on-a-pole. Pd, Mt, Cp (GA, NC, SC, VA): riverbanks, 

streambanks, bottomland forests, seepages and glades over mafic or calcareous rock, usually in nutrient-rich soils; common 

(uncommon in Coastal Plain and Mountains). June-October. Widespread in se. North America, north to NJ, OH, IL, and KS. [= 

C, FNA, GW, K, W, Z; = Uniola latifolia Michaux – RAB, F, G, HC, S] 

Chasmanthium laxum (Linnaeus) Yates, Slender Spikegrass. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): savannapocosin 

ecotones, sandhill-pocosin ecotones, moist hardwood swamps, other moist habitats; common (rare in Mountains). June- 

October. Widespread in se. North America, north to s. NY, KY, and OK. See Ch. sessiliflorum for comments on the suggestion 

that these two taxa are only varietally distinct. [= C, FNA, GW, K, W, Z; = Uniola laxa (Linnaeus) Britton, Sterns, & 

Poggenburg – RAB, F, G, HC, S; = Chasmanthium laxum var. laxum] 

Chasmanthium nitidum (Baldwin) Yates, Shiny Spanglegrass. Cp (GA, NC, SC): blackwater swamp forests; rare (NC 

Rare, SC Rare). September-November. A Southeastern Coastal Plain endemic: se. NC south to c. FL and west to se. AL. [= 

FNA, GW, K, Z; = Uniola nitida Baldwin – RAB, HC, S] 

Chasmanthium sessiliflorum (Poiret) Yates var. 1, Coastal Hammock Longleaf Spikegrass. Cp (GA, SC): calcareous 

hammocks; rare. August-October. An additional taxon warrants recognition: it is characterized by divergent panicle branches 

and occurs in outer Coastal Plain calcareous sites (J. Allison, pers. comm.). [< Chasmanthium sessiliflorum (Poiret) Yates – C,

POACEAE 857 

FNA, GW, K, Z; < Uniola sessiliflora Poiret – RAB, F, G, HC; < Uniola longifolia Scribner – S; < Chasmanthium laxum 

(Linnaeus) Yates var. sessiliflorum (Poiret) L. Clark] 

Chasmanthium sessiliflorum (Poiret) Yates var. sessiliflorum, Longleaf Spikegrass. Cp (GA, NC, SC, VA), Pd (GA, SC): 

moist hardwood forests, swamps, other moist habitats; rare (NC Watch List, VA Watch List). August-October. Widespread in 

se. North America, north to se. VA, TN, AR, and OK. This species and Ch. laxum are morphologically somewhat similar, but 

their treatment as varieties of a single species is completely unwarranted. They frequently co-occur (especially on the Gulf 

Coastal Plain), growing side by side, and show no sign of intergradation. [< Chasmanthium sessiliflorum – C, FNA, GW, K, W, 

Z; < Uniola sessiliflora Poiret – RAB, F, G, HC; < Uniola longifolia Scribner – S; < Chasmanthium laxum (Linnaeus) Yates var. 

sessiliflorum (Poiret) L. Clark] 

Chasmanthium ornithorhynchum (Steudel) Yates, Birdbill Spikegrass. S. AL and w. FL panhandle west to e. LA (Florida 

Parishes). Also reported for NC and SC (FNA 2003a). {investigate} [= FNA, GW, K, Z; = Uniola ornithorhyncha Steudel – S] 

Key based partly on C. 

Chloris Swartz (Finger-grass, Chloris) 

(also see Eustachys) 

1 Inflorescence verticillate, typically the panicle branches in 2-5 verticels; perennial; fertile lemma inconspicuously appressedpilose; 

spikelets not imbricate............................................................................................................................. Ch. verticillata 

1 Inflorescence digitate, the panicle branches in a single verticel at the apex of the culm; annual; lemma conspicuously longciliate; 

spikelets imbricate......................................................................................................................................... Ch. virgata 

* Chloris verticillata Nuttall, Windmill-grass. Mt (VA), {SC}: disturbed areas, bottomland fields; rare, introduced from 

further west. [= C, F, G, HC, K] 

* Chloris virgata Swartz, Feather Finger-grass, Showy Chloris. Pd (GA, NC, SC, VA), Cp (GA): disturbed areas; rare, 

introduced from tropical America. [= RAB, C, F, G, HC, K] 

* Chloris barbata Sw. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, introduced from {}. [= K] 

{not keyed} 

* Chloris canterei Arech. var. grandiflora (Roseng. & Izag.) D.E. Anderson. Cp (SC): waste areas near wool-combing 

mills; rare, perhaps only a waif, introduced from Paraguay. [= K; < Ch. cantérai – HC] {not keyed} 

* Chloris cucullata Bisch. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, introduced from {}. [= 

K] {not keyed} 

* Chloris divaricata R. Brown. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, introduced from 

{}. [= K] {not keyed} 

* Chloris gayana Kunth, Rhodes Grass. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, 

introduced from Africa. [= F, HC, K, S] {not keyed} 

* Chloris pectinata Bentham. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, introduced from {}. 

[= K] {not keyed} 

* Chloris truncata R. Brown, Stargrass. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, 

introduced from Australia. [= HC, K] {not keyed} 

* Chloris ventricosa R. Brown. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, introduced from 

Australia. Also reported for VA (Hitchcock & Chase 1950; Kartesz 1999). [= HC, K] {not keyed} 

Chrysopogon Trinius (Goldbeard) 

A genus of about 26 species, tropical and subtropical, all species except Ch. pauciflorus native to the Old World. References: 

Hall & Thieret in FNA (2003a); Veldkamp (1999). 

Chrysopogon pauciflorus (Chapman) Bentham ex Vasey, Florida Goldbeard, Florida Rhaphis. Cp (NC): sandhill; rare 

(NC Watch List), perhaps only introduced. FL and Cuba; its occurrence in se. NC (at Carolina Beach State Park) is plausible 

either as a native, disjunct occurrence or as an introduction. [= FNA, HC, K; = Rhaphis pauciflora (Chapman) Nash – S] 

Cinna Linnaeus 1753 (Woodreed) 

(also see Limnodea) 

A genus of about 4 species, of temperate Eurasia, North America, and South America. References: Brandenburg, Blackwell, & 

Thieret (1991); Tucker (1996)=Z; Brandenburg & Thieret (2000).

POACEAE 858 

1 Spikelets (3.5-) 4-6 (-7.5) mm long; glumes firm, subherbaceous, rather dull, hyaline only narrowly and marginally, the 

second glume prominently 3-nerved.................................................................................................................. C. arundinacea 

1 Spikelets (2-) 2.5-4 (-5) mm long; glumes (at least the first and sometimes the second as well) glistening, hyaline except the 

midrib, the second glume 1-nerved (very rarely 3-nerved) ........................................................................................ C. latifolia 

Cinna arundinacea Linnaeus, Common Woodreed, Sweet Woodreed. Cp, Pd, Mt (GA, NC, SC, VA): bottomland forests, 

rocky bars in rivers, other low, wet habitats; common. August-October. New Brunswick and MN south to south to GA and TX. 

[= RAB, C, G, GW, K, S, W, Z; > C. arundinacea var. inexpansa Fernald & Griscom – F, HC] 

Cinna latifolia (Treviranus ex Goepp.) Grisebach, Drooping Woodreed, Slender Woodreed. Mt (NC, VA): moist forests at 

high elevations; rare (NC Watch List, VA Rare). June-August. Circumboreal, occurring in n. Eurasia and n. North America, 

south in North America to NC, TN, n. IL, MN, UT, NM, and CA. [= RAB, C, F, G, HC, K, W, Z] 

Coelorachis Brongniart (Jointgrass) 

A genus of about 20 species, widespread in the Old World and New World tropics and subtropics. Generic circumscription has 

been controversial and uncertain. References: Allen in FNA (2003a); Veldkamp, Koning, & Sosef (1986)=Z. 

1 Culms round in cross-section................................................................................................................................. C. cylindrica 

1 Culms compressed-keeled in cross-section. 

2 Lower glume with rectangular pits....................................................................................................................C. tesselata 

2 Lower glume smooth or with transverse ridges. 

3 Lower glume with transverse ridges ............................................................................................................C. rugosa 

3 Lower glume smooth ...........................................................................................................................C. tuberculosa 

Coelorachis cylindrica (Michaux) Nash, Carolina Jointgrass. Pd (GA, NC, SC), Cp (GA): open woodlands and roadsides, 

probably in areas formerly prairie-like and fire-maintained, perhaps now extirpated in our area; rare (NC Rare). June-August. 

Fairly widespread in se. North America, north to NC and SC (at least formerly), MS, MO, and TX. [= C, FNA, K; = Manisuris 

cylindrica (Michaux) Kuntze – RAB, F, G, GW, HC; = Manisuris campestris (Nuttall) A.S. Hitchcock – S; = Mnesithea 

cylindrica (Michaux) Koning & Sosef – Z] 

Coelorachis rugosa (Nuttall) Nash, Wrinkled Jointgrass. Cp (GA, NC, SC, VA), Pd (GA): limesink ponds (dolines), 

depression meadows, clay-based Carolina bays, wet savannas, always in places with a seasonally high water-table; rare (NC 

Watch List, VA Rare). June-October. A Southeastern Coastal Plain endemic: s. NJ south to FL and west to TX. [= C, FNA, K; 

= Manisuris rugosa (Nuttall) Kuntze – RAB, F, G, GW, HC, S; = Mnesithea rugosa (Nuttall) Koning & Sosef – Z] 

Coelorachis tesselata (Steudel) Nash, Pitted Jointgrass. Cp (GA): wet savannas and bogs; rare. Southeasatern Coastal 

Plain endemic: sw. GA and FL west to e. LA. [= FNA, K; = Manisuris tesselata (Steudel) Scribner – GW, HC, S; = Mnesithea 

tesselata (Steudel) Koning & Sosef – Z] 

Coelorachis tuberculosa (Nash) Nash, Smooth Jointgrass. Cp (GA): pond margins; rare. Southeastern Coastal Plain 

endemic: sw. GA (Mitchell County) (Sorrie 1998b) west to s. AL, and in the Florida peninsula. [= FNA, K; = Manisuris 

tuberculosa Nash – GW, HC, S; = Mnesithea tuberculosa (Nash) Koning & Sosef – Z] 

Coix Linnaeus 

A genus of about 5 species, native to tropical Asia. References: Thieret in FNA (2003a). 

* Coix lacryma-jobi Linnaeus, Job's-tears, reported for se. PA by Rhoads & Klein (1993), TN (Thieret in FNA 2003a), and NJ 

(Kartesz 1999). [= FNA, K] 

Cortaderia Stapf (Pampasgrass) 

A genus of ca. 20 species, native to South America. References: Allred in FNA (2003a). 

* Cortaderia selloana (J.A. & J.H. Schultes) Ascherson & Graebner, Pampasgrass. Cp (GA, SC), Pd (GA): disturbed areas; 

rare, introduced from South America. This grass is a popular ornamental, rarely escaping. [= RAB, FNA, HC, K] 

Ctenium Panzer (Toothache Grass) 

A genus of about 20-22 species, of tropical and subtropical Africa and the Americas. References: Barkworth in FNA (2003a); 

Longhi-Wagner & Renvoize (2004).

POACEAE 859 

1 Spikelets with numerous glands in rows on the back of the second glume; plant short-rhizomatous (nearly cespitose); 

[widespread in the Coastal Plain of our area].....................................................................................................Ct. aromaticum 

1 Spikelets with very few or no glands on the back of the second glume; plant rhizomatous (the rhizomes slender and scaly); 

[restricted to se. GA southwards]........................................................................................................................ Ct. floridanum 

Ctenium aromaticum (Walter) Wood, Toothache Grass, Orange Grass. Cp (GA, NC, SC, VA): wet savannas, pocosinsavanna 

ecotones, seepage bogs, sandhill-pocosin ecotones, sandhill seeps; common (VA Rare). June-August (or later in 

response to late summer fires). Southeastern Coastal Plain endemic: se. VA south to FL and west to LA and e. TX (Singhurst, 

Keith, & Holmes 2005). The entire plant is aromatic and numbs the mouth, tongue, and lips when chewed, hence the specific 

epithet and common names. Like many species of the longleaf pine ecosystem, toothache grass generally flowers only following 

fire (MacRoberts & MacRoberts 1992). Sterile clumps can be recognized by the rather broad, bicolored leaves (bluish on the 

upper surface, bright green on the lower surface). [= RAB, C, F, FNA, G, GW, HC, K; = Campulosus aromaticus (Walter) 

Trinius – S] 

Ctenium floridanum (A.S. Hitchcock) A.S. Hitchcock, Florida Toothache Grass. Cp (GA): dry pinelands, sandhills, upper 

ecotones of pineland pools; rare (GA Special Concern). June-September. Southeastern Coastal Plain endemic: se. GA to ne. FL. 

Like Ct. aromaticum, generally flowering only following fire. [= FNA, GW, HC, K; = Campulosus floridanus A.S. Hitchcock – 

S] 

Cynodon L.C. Richard (Bermuda Grass) 

A genus of ca. 9 species, native to the tropical Old World. References: Barkworth in FNA (2003a). 

* Cynodon dactylon (Linnaeus) Persoon var. dactylon, Bermuda Grass, Scutch Grass. Cp, Pd, Mt (GA, NC, SC, VA): lawns, 

gardens, roadsides, pastures, fields, disturbed areas; common, introduced from Eurasia. May-October. [= FNA; < C. dactylon – 

RAB, C, F, G, HC, K, W; < Capriola dactylon (Linnaeus) Kuntze – S] 


Cynosurus Linnaeus 1753 (Dog's-tail, Dogtail Grass) 

1 Panicle linear-oblong, 1-10 (-14) cm long, 0.4-1 cm wide; leaves 1-3 (-4) mm wide; perennial; fertile lemma 3-4 mm long, 

plus a 0-1 mm long mucro; [section Cynosurus] ......................................................................................................C. cristatus 

1 Panicle ovoid, 1-4 (-8) cm long, 0.7-2 cm wide; leaves (2-) 3-10 mm wide; annual; fertile lemma 4.5-7 mm long, plus a 6-16 

mm long awn; [section Falona] ..............................................................................................................................C. echinatus 

* Cynosurus cristatus Linnaeus, Crested Dog's-tail. Mt (NC), {VA}: lawns, roadsides; rare, introduced from Eurasia. June- 

July. [= RAB, C, F, G, HC, K, Z] 

* Cynosurus echinatus Linnaeus, Rough Dog's-tail. Cp (NC, SC, VA), Pd (GA, NC, SC, VA), Mt (GA): lawns, roadsides; 

rare, introduced from Eurasia. May-June. [= RAB, C, F, HC, K, Z] 


Dactylis Linnaeus 1753 (Orchard Grass) 

* Dactylis glomerata Linnaeus, Orchard Grass, Cock's-foot. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): pastures, fields, 

woodland edges, roadsides; common (less common in Coastal Plain, especially in NC and SC), introduced from Europe. May- 

October. In Europe there are various chromosome races, often accorded subspecies or species status. Their status in North 

America has been little investigated. See various references cited in Tucker (1996) for further information about these taxa in 

Europe. [= RAB, C, G, HC, S, W; > D. glomerata var. glomerata – F; > D. glomerata var. detonsa Fries – F; > D. glomerata 

var. ciliata Petermann – F; > D. glomerata ssp. glomerata – K, Z; > D. glomerata ssp. aschersoniana (Graebner) Thellung – K; 

> D. aschersoniana Graebner] 

Dactyloctenium Willdenow (Crowfoot Grass) 

* Dactyloctenium aegyptium (Linnaeus) Willdenow, Crowfoot Grass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): lawns, 

roadsides, disturbed areas; common, introduced from Old World tropics. June-November. [= RAB, C, F, G, HC, K, S] 

* Dactyloctenium radulans (R. Brown) Palisot de Beauvois is introduced in SC (Kartesz 1999). {investigate} [= K] {not 

keyed at this time}

POACEAE 860 

Danthonia Augustin de Candolle (Oat-grass) 

A genus of about 20 species, of North America, Europe, and the Americas, but the generic limits are unclear. References: 

Darbyshire in FNA (2003a). 

1 Lemma teeth (flanking the awn) 0.8-1.8 mm long, triangular, acuminate; glumes 8-13 mm long.............................. D. spicata 

1 Lemma teeth (flanking the awn) (1.8-) 2.0-4.5 mm long, setaceous; glumes 9-19 mm long. 

2 Lemma awn 4-10 mm long; glumes 9-13 mm long ...................................................................................... D. compressa 

2 Lemma awn 11-18 mm long; glumes 11-19 mm long. 

3 Sheaths villous; lemmas herbaceous in texture, villous on the back and sides; awn twisted at base several times, 

forming an awn column 2.5-3 mm long ...................................................................................................... D. sericea 

3 Sheaths glabrous; lemmas membranaceous in texture, villous only on the margins and towards the base; awn 

twisted at base a single time, forming a loose awn column 0.5-1.5 mm long ................................................D. epilis 

Danthonia compressa Austin ex Peck, Mountain Oat-grass. Mt, Pd (GA, NC, SC, VA), Cp (VA): grassy balds, thin soils 

around rock outcrops, woodlands; common (uncommon in upper Piedmont only). June-August. Fairly widespread in e. North 

America, primarily Appalachian, from s. Canada to SC and TN. [= RAB, C, F, FNA, G, HC, K, S, W] 

Danthonia epilis Scribner, Bog Oat-grass. Cp (NC, SC, VA), Mt (GA, NC, VA?), Pd (NC): peaty bogs in the Coastal Plain 

and Mountains, seeps around rock outcrops in the Piedmont and Mountains, granitic domes; rare (GA Special Concern, NC 

Watch List, VA Watch List). April-June. The range is apparently bogs in pinelands from NJ to SC, in mountain bogs in NC, VA 

(?), and GA, in seepage in the Cumberland Plateau and Blue Ridge of TN and AL. This taxon appears to be valid, with a distinct 

range, habitat, and variety of morphologic characters separating it from D. sericea, but further study is needed. Material from the 

mountains seems to differ from Coastal Plain material. RAB's description of the habitat as "dry woods, rare; pied. of N.C." 

appears to be in error. Blomquist listed the taxon (as a variety) for bogs in the mountains of sw. NC. It has since been found in 

bogs in the Sandhills region of NC and in seepage bogs in the adjacent Piedmont. [= F, HC, K, S; = D. sericea var. epilis 

(Scribner) Blomquist – RAB, C; < D. sericea Nuttall – FNA] 

Danthonia sericea Nuttall, Silky Oat-grass. Cp, Pd, Mt (GA, NC, SC, VA): dry woodlands, especially common in sandy 

soils in the Coastal Plain, dry oak, oak-pine, and pine forests in the Piedmont and low Mountains; common (uncommon in the 

Mountains). April-June. Primarily a Coastal Plain species northwards, ranging from e. MA south to FL and west to LA. [= F, 

HC, K, S, W; = D. sericea var. sericea – RAB, C, G; < D. sericea – FNA] 

Danthonia spicata (Linnaeus) Palisot de Beauvois ex Roemer & J.A. Schultes, Poverty Oat-grass. Mt, Pd (GA, NC, SC, 

VA), Cp (NC, SC, VA): dry woodlands, rock outcrops, shale barrens; common. May-July. Newfoundland and British Columbia 

south to FL and NM. [= RAB, C, FNA, G, HC, K, S, W; > D. spicata var. longipila Lamson-Scribner & Merrill – F; > D. 

spicata var. spicata – F; < D. allenii Austin – F] 

Deschampsia Palisot de Beauvois (Hairgrass) 

A genus of about 40 species, north and south temperate. References: Tucker (1996)=Z. 

1 Awn 2-3 mm long, straight or nearly so, scarcely (or not at all) exserted beyond the tips of the glumes; lemmas smooth; leaf 

blades flat or folded at the midvein (V-shaped in cross-section); ligule 3-10 (-17) mm long; [section Deschampsia]............... 

..............................................................................................................................................................D. cespitosa ssp. glauca 

1 Awn 4-8 mm long, geniculate, exserted beyond the tips of the glumes; lemmas minutely scabrous; leaf blades involute, 

appearing filiform (rounded in cross-section); ligule 0.5-3 (-5) mm long; [section Avenaria] ...........D. flexuosa var. flexuosa 

Deschampsia cespitosa (Linnaeus) Palisot de Beauvois ssp. glauca (Hartman) Hartman, Tufted Hairgrass. Mt (NC, VA): 

thin soil of rock outcrops or barrens over calcareous, mafic, and ultramafic rocks (such as serpentinized olivine, amphibolite, 

limestone, and dolostone); rare (NC Rare, VA Rare). June-July. D. cespitosa is a complex species, with a complicated polyploid 

and aneuploid series, variously subdivided (or not) by various taxonomists. As a whole, D. cespitosa is circumboreal, ranging 

south in North America to NJ, sw. NC, WV, IL, MN, and AZ. Ssp. glauca is the most widespread American subspecies, and 

extends the farthest south. Other subspecies occur farther north and in Eurasia. In our area, D. cespitosa is at its southern limit 

and is a rare species limited to barrens and outcrops over mafic or ultramafic rocks. [= K; = D. caespitosa var. glauca (Hartman) 

Lindman f. – RAB, F; < D. cespitosa – C, Z; < D. cespitosa var. cespitosa – G; < D. caespitosa var. caespitosa – HC; = D. 

cespitosa ssp. cespitosa var. glauca (Hartman) Lindman f.; < Aira caespitosa Linnaeus – S; < D. caespitosa – W] 

Deschampsia flexuosa (Linnaeus) Trinius var. flexuosa, Common Hairgrass, Wavy Hairgrass. Mt (GA, NC, SC, VA), Pd, 

Cp (NC, VA): grassy balds, high elevation rocky summits, rocky or sandy woodlands; common (uncommon in Piedmont and 

Coastal Plain) (SC Rare). April-August. Circumboreal, ranging south in North America to n. GA, OH, WI, and MN; disjunct in 

AR and OK, and in Mexico. [= F, K; < D. flexuosa – RAB, C, G, HC, W, Z; < Aira flexuosa Linnaeus – S] 

* Deschampsia elongata (Hooker) Munro, Slender Hairgrass. Cp (SC): waste areas near wool-combing mills; rare, perhaps 

only a waif, native of w. North America. [= HC, K] {not keyed}

POACEAE 861 

Desmazeria Dumortier 

(see Catapodium) 

Diarrhena Palisot de Beauvois (Beakgrain) 

A genus of about 4 species of perennial grasses of e. North America and e. Asia. References: Brandenburg, Estes, & Collins 

(1991)=Z. Key from Z. 

1 Callous pubescent on all mature lemmas except the first; lemmas widest below the middle and gradually tapering into a cusp 

at the apex, those of the first floret 7.1-10.8 mm long; mature fruit 1.3-1.8 mm broad, gradually tapering into a broad, blunt 

beak....................................................................................................................................................................... D. americana 

1 Callous glabrous on all mature lemmas; lemmas widest near or above the middle and more-or-less abruptly contracted into a 

cusp at the apex, those of the first floret 4.6-7.5 mm long; mature fruit 1.8-2.5 mm broad, abruptly contracted into a 

bottlenose-shaped beak ..............................................................................................................................................D. obovata 

Diarrhena americana Palisot de Beauvois, Eastern Beakgrain. Mt (GA, NC, VA): rich moist forests, usually over 

calcareous rocks; rare (NC Rare, VA Watch List). July-August; August-October. W. VA and WV west to IN, south to TN, sw. 

NC, and nw. GA (Jones & Coile 1988); disjunct in MO. [= K, Z; = Diarrhena americana var. americana – C, G; < Diarrhena 

americana – F, HC, W (also see Diarrhena obovata); < Diarina festucoides Rafinesque – S (also see Diarrhena obovata] 

Diarrhena obovata (Gleason) Brandenburg, Western Beakgrain. Pd (VA): alluvial forests; rare (VA Rare). July-August; 

August-October. Sw. PA and IN west to SD, KA, south to w. VA, nc. TN, and ne. TX. First reported for our area by Fleming & 

Ludwig (1996). The floodplain of the Potomac River (in Fairfax County, VA) has a number of disjuncts of species with more 

midwestern affinities, including Diarrhena obovata, Erigenia bulbosa, Valeriana pauciflora, and Erythronium albidum (Fleming 

& Ludwig 1996). [= K, Z; = Diarrhena americana var. obovata Gleason – C, G; < Diarrhena americana – F, HC, W; < 

Diarina festucoides Rafinesque – S] 

Dichanthelium (A.S. Hitchcock & Chase) Gould 1974 (Witch-grass) 

(by Richard J. LeBlond) 

A genus of 70-100 species, perennials, of temperate and tropical America. 

"We admit that our failure to distinguish the several named taxa ... was born of despair!" – Godfrey & Wooten (1979). 

"The recognition of only four species and six varieties in this complex [sabulorum] to which almost 50 species names have been 

applied admittedly is somewhat arbitrary and certainly not entirely satisfactory." – Gould & Clark (1978). 

Dichanthelium has often been treated as subgenus Dichanthelium of Panicum. It is most readily (though not consistently) 

separated from Panicum by the following combination of features: plants producing over-wintering rosettes of leaves often 

shorter and broader than the culm leaves; plants producing simple culms with terminal panicles in spring, the culms branching 

and producing panicles only on branches in the summer and autumn. 

Perhaps the most complex and confusing genus in our region, Dichanthelium requires careful collection and close observation of 

several characters to determine to which taxon a specimen belongs, or at least to which taxa it seems most closely aligned. A 

taxon that is distinct in one part of its range may be indistinguishable from another taxon elsewhere. This is particularly true of 

Coastal Plain species adapted to natural (and now human) disturbances. Although hybridization is frequently suspected in 

Dichanthelium, documentation of natural hybrids is rare. 

When collecting specimens in the field, mature spikelets are essential. This is determined by examining the usually whitish 

fertile lemma, which is firm and plump at maturity. Immature spikelets often are longer than mature ones (they shorten as they 

fatten); only mature spikelet length is used in the various manuals and keys. It is also important to note whether a plant is in its 

"vernal" or "autumnal" fruiting phase before collecting. "Vernal" plants produce panicles only at the summits of the culms 

(typically April-June). "Autumnal" plants produce panicles from leafy axillary branches below the summit (typically July- 

September). The autumnal panicles in most species are much smaller than the vernal panicles (and often hidden by fascicled 

leaves), but the spikelets are the same. When collecting autumnal plants, it is important to select specimens still possessing their 

vernal leaf blades and panicles, even though these will likely be senescent. It is also important to collect the whole plant, with the 

basal rosette intact (whether senescent or of current year's growth). When several plants are growing together, compare the culm, 

leaf, and spikelet features for differences; Dichanthelium taxa are gregarious. 

When analyzing the character of the culm internodes and nodes, look at the first elongate internode above the base (the lowest 

internode is often very short and uncharacteristic). Determining whether a node is bearded is often difficult. A bearded node 

usually is characterized by pubescence that is longer and of a different orientation or structure than that of the internodes and

POACEAE 862 

sheaths. Nodes with short pubescence generally are not regarded as bearded. Lower nodes are more likely to be bearded than 

upper nodes. When analyzing sheaths, look at those on the lower half of the culm. Senescent vernal sheaths often lose their 

pubescence (though in some species hair papillae are evident). All references in the key to sheath glabrousness or pubescence is 

without regard to the presence or absence of marginal hairs (cilia). A sheath that is glabrous except for marginal cilia is called 

glabrous. All culm leaves should be analyzed for blade characters; in general, the key relies on the size and character of the 

vernal blades. A "cordate" blade is one where the basal lobes of the blade extend outward and partially surround the culm when 

the culm is enclosed by the sheath. As with sheaths, references in the key to blade glabrousness or pubescence is without regard 

to marginal cilia. The ligule is an important diagnostic character for many Dichanthelium taxa; at least three ligules per specimen 

should be examined before making conclusions about its structure and length. Ligules form a distinct ring from a cartilaginous 

base at the inner summit of the sheath; in some species the ligule is membranous, but in most it is pubescent. Care must be taken 

to distinguish the pubescence of the ligule from any pubescence emanating from the inner surface of the blade base, and from 

marginal cilia. Ligules of senescent vernal leaves frequently lose their integrity. Spikelet shape as well as length should be 

determined only from mature spikelets. Measure the length from the base of the first glume (usually at an articulation) to the 

apex of the second glume or sterile lemma (whichever is longer). A micrometer is essential for determining the length of 

spikelets, first glumes, ligules, and various pilosity features. Sometimes one-tenth of a millimeter is all that separates two 

Dichanthelium taxa. 

Certain characters, particularly node bearding, cordate/non-cordate blade bases, and ligule length, can be quite variable, and an 

effort has been made to account for this variability in the key. Nonetheless, some specimens just won't "fit," and the road not 

taken may have to be reconsidered. 

References: Gould and Clark (1978)=Z; Freckmann (1981)=Y; Lelong (1984)=X; LeBlond (2001)=Q; Davidse and Polh (1992); 

Hansen & Wunderlin (1988); Hitchcock & Chase (1910); Freckmann & Lelong (2002). The treatment of Dichanthelium sect. 

Lanuginosa (=D. acuminatum group) is based closely on Y. The contributor must take responsibility for the treatment of sect. 

Angustifolia (including D. hirstii), sections Dichotoma and Ensifolia (the D. dichotomum group), and for sect. Lancearia. Other 

treatments are based largely on Z. 

1 Plants densely tufted, often cushion-forming; leaves basally disposed, the blades ascending or spreading-ascending, not 

forming a distinct rosette of basal leaves shorter than the culm leaves; autumnal culms branching basally or from the lower 

nodes.................................................................................................................................................................................Key A 

1 Plants less densely or sparsely tufted, not cushion-forming; leaves well-distributed on the culm, usually much longer than 

the short, often broad and spreading basal rosette leaves; autumnal culms usually branching from the mid and upper nodes. 

2 Spikelets 3.3-5.2 mm long ........................................................................................................................................Key B 



4 Larger culm blades 13-25 mm wide..........................................................................................................Key C 

4 Larger culm blades < 13 mm wide. 

5 Culm nodes (at least the lower) bearded............................................................................................Key D 

5 Culm nodes not bearded, the lowermost sometimes puberulent or sparsely hairy.............................Key E 


6 Lower culm internodes variously hairy..................................................................................................... Key F 

6 Lower culm internodes glabrous .............................................................................................................. Key G 

Key A - Plants densely tufted, often cushion-forming; leaves basally disposed, 

the blades ascending or spreading-ascending, 

not forming a distinct rosette of basal leaves 

shorter than the culm leaves; 

autumnal culms branching basally or from the lower nodes 


2 Nodes, internodes, and sheaths glabrous; blades 4-13 cm, 5-8 mm, the surfaces smooth, glabrous; spikelets 2.4-2.9 mm 

long, glabrous; not known to produce axillary (autumnal) inflorescences ...................................................[D. nudicaule] 

2 Nodes bearded or otherwise pubescent; internodes and sheaths variously pubescent to glabrate; blades 6-35 cm, 2-5 

mm, one or both surfaces scabrous and often pubescent; spikelets 1.7-4.5 mm long, glabrous or pubescent; plants 

produce axillary (autumnal) inflorescences. 

3 Spikelets 2.8-3.8 (-4.5) mm long, the second glume and sterile lemma pointed or beaked and extended beyond the 

summit of the fertile lemma; first glume 1.2-2 mm long................................................................. D. depauperatum 

3 Spikelets 1.7-2.8 mm long, the second glume and sterile lemma blunt or broadly pointed, not extending beyond 

the summit of the fertile lemma; first glume 0.7-1.2 mm long.......................................................... D. linearifolium 


4 Longer blades > 6 cm; if only 6 cm, then sheaths retrorsely long-pilose (D. laxiflorum). 

5 Spikelets 1.2-1.5 mm long, glabrous...................................................................[D. dichotomum var. glabrifolium] 

5 Spikelets 1.7-2.3 (-2.8) mm long, pubescent.

POACEAE 863 

6 Longer blades 10-35 cm long, 2-4 mm wide; sheaths glabrous to variously pilose, but not conspicuously 

retrorsely long-pilose; nodes variously pubescent to glabrate; spikelets 1.7-2.3 (-2.8) mm long........................ 

.................................................................................................................................................. D. linearifolium 

6 Longer blades 6-18 cm long, 7-12 mm wide; sheaths conspicuously retrorsely long-pilose; nodes bearded 

with retrorse or spreading hairs; spikelets 1.9-2.3 mm long......................................................... D. laxiflorum 

4 Longer blades 1.5-6 cm; sheaths glabrous or pubescent, but not retrorsely long-pilose. 

7 Blades 1-3 mm wide, glabrous, eciliate or basally ciliate; spikelets 0.9-1.4 mm long. 

8 Spikelets pubescent, 1.2-1.4 mm long; blades involute, often falcate, 2.5-6 cm long ......................................... 

............................................................................................................................ [D. chamaelonche ssp. breve] 

8 Spikelets glabrous, 0.9-1.2 mm long; blades flat, not falcate, 1.5-4 (-5) cm long ............................................... 

................................................................................................................ D. chamaelonche ssp. chamaelonche 

7 Blades 3-8 mm wide; spikelets 1.1-2.1 mm long (if < 1.5 mm, then blades either pubescent on one or both surfaces 

or ciliate to the apex). 

9 Spikelets pubescent, 1.5-2.1 mm long; blade surfaces glabrous..................... D. strigosum var. leucoblepharis 

9 Spikelets glabrous, 1.1-1.8 mm long; blade surfaces pubescent or glabrous. 

10 Blades pilose; spikelets 1.1-1.6 mm long ....................................................... D. strigosum var. strigosum 

10 Blades glabrous, or sparsely pilose only near the adaxial base; spikelets 1.4-1.8 mm long ........................ 

.................................................................................................................... D. strigosum var. glabrescens 

Key B - Spikelets 3.3-5.2 mm long 

1 Nodes (at least lower) densely bearded with retrorse hairs; spikelets 3.7-5.2 mm long. 

2 Ligule 2.5-4 mm long; internodes pubescent with long ascending or spreading hairs; blades 8-15 cm long, 10-25 mm 

wide; first glume 1.8-2.5 mm long................................................................................................................... D. ravenelii 

2 Ligule 0.4-0.9 (-1.3) mm long; internodes glabrous to puberulent; blades 7-12 cm long 12-40 mm wide; first glume 1.5- 

2.2 mm long .......................................................................................................................................................... D. boscii 

1 Nodes glabrous, pubescent, or sparsely pilose; spikelets (2.4-) 3.3-4.2 mm long. 

3 Ligule 1.6-3 mm long; blades 4-9 mm wide, > 10× as long as wide..............................D. oligosanthes var. oligosanthes 

3 Ligule 0.3-1.5 mm long; if larger blades < 9 mm wide and mostly 15× or more as long, then ligule 0.5-1 mm long (D. 

fusiforme). 

4 Larger blades 2-6 (-8) mm wide, mostly 15× or more as long as wide; spikelets fusiform to elliptic, acute, basally 

constricted...............................................................................................................................................D. fusiforme 

4 Larger blades 6-35 mm wide, mostly 10× or less as long as wide; spikelets broadly elliptic to obovate, rounded to 

sub-acute, not basally constricted. 

5 Spikelets strongly papillose-hispid with spreading hairs 0.5-1 mm long; blades papillose-hispid...................... 

........................................................................................................................................................[D. leibergii] 

5 Spikelets glabrous to pubescent with hairs < 0.5 mm long; blades glabrous, scabrous, or pubsecent. 

6 Ligules 1-1.5 mm long; blades 5-10 cm long by 6-15 mm wide, glabrous or pubescent, basally rounded; 

spikelets glabrous to pubescent .......................................................... D. oligosanthes var. scribnerianum 

6 Ligules 0.3-1 mm long; blades 7-35 cm long by 8-35 mm wide, glabrous or scabrous, basally cordate or 

rounded; spikelets pubescent to glabrate. 

7 Sheaths (at least lower) papillose-hispid with spreading hairs; blades 10-28 cm long; spikelets 2.4- 

3.6 mm long; first glume 1.2-1.8 mm long................................................................D. clandestinum 

7 Sheaths glabrous, pubescent, or sparsely pilose; blades 7-18 cm long; spikelets 2.9-4.1 mm long; 

first glume 1.5-2.2 mm long. 

8 Larger blades 15-32 mm wide, 4-7× as long as wide, basally cordate; sheaths glabrous to 

sprasely pilose ........................................................................................................ D. latifolium 

8 Larger blades 10-20 mm wide, 6-9× as long as wide, basally rounded; sheaths pubescent......... 

......................................................................................................................[D. xanthophysum] 

Key C - Spikelets 2.1-3.2 mm long, larger leaves 13-25 mm wide 

1 Culm nodes, at least the lower, bearded (often retrorsely). 

2 Ligule a stramineous to light brown membrane (with or without ciliate or lacerate extensions); peduncle and often 

internodes scabrous.................................................................................................................................D. scabriusculum 

2 Ligule entirely of white hairs; peduncle and internodes either smooth or densely hairy (velvety). 

3 Lower internodes glabrous, without a viscid band below the nodes; larger blades 7-14 mm wide ............................. 

.................................................................................................................................................D. dichotomum group 

3 Lower internodes densely hairy except for a viscid band below the nodes; larger blades 10-20 mm wide................. 

.............................................................................................................................................................. D. scoparium 

1 Culm nodes glabrous or slightly hairy, but not bearded.

POACEAE 864 

4 Second glume and sterile lemma acute to short-acuminate, conspicuously longer than the fertile lemma; spikelets 

glabrous (occasionally sparsely pubescent in D. scabriusculum). 

5 Panicle rachis pellucid-punctate; ligule a stramineous to light brown membrane, with or without terminal 

ciliations; peduncle and often internodes scabrous; first glume 0.3-0.6 (-0.8) mm long, reniform to suborbicular .... 

........................................................................................................................................................D. scabriusculum 

5 Panicle rachis not pellucid-punctate; ligule entirely of white hairs; peduncle and internodes smooth; first glume 

0.7-1.2 mm long, ovate to lanceolate ................................................................................................... D. yadkinense 

4 Second glume and sterile lemma blunt to subacute, shorter than, equaling, or barely exceeding the fertile lemma; 

spikelets pubescent (sometimes sparsely so in D. clandestinum). 

6 Sheaths, at least the lower, papillose-hispid with spreading hairs; blades 10-28 cm long.................D. clandestinum 

6 Sheaths glabrous, puberulent, finely pubescent, or sparsely pilose; blades 5-18 cm long. 

7 Ligule 0-0.3 mm long; spikelets 2.2-3.2 mm long, 1.1-1.3 mm wide; first glume 0.7-1.4 (-1.8) mm long ......... 

.................................................................................................................... D. commutatum var. commutatum 

7 Ligule 0.4-0.7 mm long; spikelets 2.9-3.9 mm long, 1.6-2.0 mm wide; first glume 1.5-2.2 mm long................ 

....................................................................................................................................................... D. latifolium 

Key D - Spikelets 2.1-3.2 mm long, larger culm blades < 13 mm wide, 

at least the lower culm nodes bearded with a usually spreading-ascending collar 

of dense and/or longish hairs 

1 Ligule with a dense ring of short hairs 0.5-1 mm long in front of a usually less dense ring of longer hairs (pseudoligule) 1-5 

mm long. 

2 Nodes retrorsely bearded; internode and sheath hairs spreading to restrorse; blade surfaces velvety-pubescent or longpilose. 

3 Spikelets 2.5-3.2 mm long; longer hairs of pseudoligule 1-3 mm long; blade surfaces velvety-pubescent; panicle 

rachis densely pubescent; [of cedar glades and dry limestone soils]...........................................[D. malacophyllum] 

3 Spikelets 1.8-2.5 mm long; longer hairs of pseudoligule 3-5 mm long; blade surfaces long-pilose; panicle rachis 

sparsely pilose; [of dry sandy soil of pine and oak woodlands] ......................... D. villosissimum var. villosissimum 

2 Node beard hairs spreading to ascending; internode and sheath hairs ascending to appressed; blade surfaces glabrate to 

appressed-pubescent. 

4 Spikelets 2.5-3.1 mm long; lower culm blades usually glabrous adaxially except for long hairs at or near the 

margin (appearing ciliate), appressed-pubescent abaxially ........................................................... D. ovale var. ovale 

4 Spikelets 2.1-2.6 mm long; lower culm blades usually sparsely appressed-pubescent on both surfaces, eciliate or 

ciliate at the base only.............................................................................................................D. ovale var. addisonii 

1 Ligule a single structure, without a pseudoligule. 

5 Ligule 2-5 mm long, ciliate.............................................................................................................D. acuminatum group 

5 Ligule < 2 mm long, ciliate or membranous. 

6 Ligule a stramineous to light brown membrane, with or without terminal ciliations; peduncle scabrous but not 

hairy. 

7 Panicle rachis smooth, pellucid-punctate; first glume 0.3-0.6 (-0.8) mm; larger leaves 10-25 cm long, 8-15 

mm wide; ligule 0.5-1.3 mm long; lowest elongate culm internode > 2 mm in diameter; lowest nodes usually 

glabrous or pubescent..............................................................................................................D. scabriusculum 

7 Panicle rachis scabrous, not pellucid-punctate; first glume 0.7-1.1 mm long; larger leaves 7-12 cm long, 6-9 

mm wide; ligule 0.3-0.6 mm long; lowest elongate culm internode < 1.7 mm in diameter; lowest nodes 

usually retrorsely bearded .................................................................................... D. species 9 (= cryptanthum) 

6 Ligule entirely of white hairs; peduncle variously hairy or glabrous, but not scabrous. 

8 Culms to 1.5 m tall, with a broad, glabrous, viscid band below the nodes; blades of the lower leaves typically 

villous or velvety-pubescent.......................................................................................................... D. scoparium 

8 Culms rarely exceeding 1 m, without a viscid band below the nodes; blades various. 

9 Sheaths retrorsely pilose with hairs 2-3 mm long; basal leaves usually numerous, ascending, similar in 

size and shape to the culm leaves; culms branching only at the base .................................... D. laxiflorum 

9 Sheaths glabrous or pilose (if pilose, then hairs not both retrorse and 2-3 mm long); basal leaves rosetteforming, 

usually much smaller than the culm leaves; culms branching at the nodes in age. 

10 Culm internodes glabrous to sparsely pilose; culm nodes bearded with long retrorse hairs; blade 

surfaces glabrous to velvety-pubescent ........................................................... D. dichotomum group 

10 Culm internodes, at least the lower, strigose, pilose, or villous; culm nodes bearded with ascending 

or spreading hairs; blade surfaces glabrous or variously hairy. 

11 Lower nodes bearded with erect-ascending, soft, and long hairs; mid-culm blades usually 20× 

or more as long as wide. 

12 Spikelets 2.9-4.0 mm long, fusiform to elliptic, acute, basally constricted; first glume 1.4- 

2.6 mm long.....................................................................................................D. fusiforme 

12 Spikelets 1.5-3.1 mm long, obovate to elliptic-obovate, obtuse to sub-acute, not basally 

constricted; first glume 0.6-1.5 mm long.

POACEAE 865 

13 Spikelets 1.5-2.2 mm long; first glume 0.6-0.8 mm long; blades to 8 cm long, 

usually involute........................................................................................ D. aciculare 

13 Spikelets 2.1-3.1 mm long; first glume 0.8-1.5 mm long; blades to 12 cm long, 

usually flat except at tip....................................................................D. angustifolium 

11 Lower and often mid-culm nodes bearded with spreading, stiffish, and short-to-long hairs; 

mid-culm blades usually 15× or less as long as wide. 

14 Blades stiff, often longitudinally ribbed, at least the lower villous or strongly pilose on 

the abaxial surface, and usually strongly pilose at least basally on the adaxial surface ....... 

.............................................................................................................. D. consanguineum 

14 Blades not noticeably stiff nor longitudinally ribbed, pubescent or strigose underneath, 

glabrous above or with a few long hairs near the base. 

15 Spikelets 2.5-3.1 mm long; lower culm blades usually glabrous adaxially except for 

long hairs at or near the margin (appearing ciliate), appressed-pubescent abaxially ... 

....................................................................................................... D. ovale var. ovale 

15 Spikelets 2.1-2.6 mm long; lower culm blades usually sparsely appressed-pubescent 

on both surfaces, eciliate or ciliate at the base only .................D. ovale var. addisonii 

Key E - Spikelets 2.1-3.2 mm long, 

larger culm blades < 13 mm wide, culm nodes not bearded, 

the lowermost sometimes puberulent or sparsely pilose 

1 Ligule 1.6-4 mm long ............................................................................................................D. oligosanthes var. oligosanthes 

1 Ligule < 1.5 mm long. 

2 Blades, at least the lower, cordate or subcordate at the base, mostly 6-12 mm wide. 

3 Spikelets obpyriform when viewed dorsally, strongly plano-convex when viewed laterally, usually markedly 

reddish-purple basally; fertile lemma papillose............................................................ D. species 2 (=webberianum) 

3 Spikelets elliptic to elliptic-obovoid when viewed dorsally or laterally, greenish to faintly purple-tinged basally; 

fertile lemma not papillose. 

4 Lowermost internodes crisp-puberulent; larger culm blades 4-8 (-11) cm long, 5-10 (-12) mm wide, broadest 

near the base, spreading from the culm .................................................................... D. commutatum var. ashei 

4 Lowermost internodes glabrous to puberulent to sparsely pilose; larger culm blades 6-14 cm long, 6-13 mm 

wide, broadest at or just below the middle, erect or erect-spreading, narrowed below to a moderately 

subcordate base .................................................................................................................................. D. boreale 

2 Blades tapering to the base, 2-12 mm wide. 

5 Ligule a stramineous to light brown membrane, with or without terminal ciliations. 

6 Panicle rachis smooth, pellucid-punctate; first glume 0.3-0.6 (-0.8) mm long; larger leaves 10-25 cm long, 8- 

15 mm wide; ligule 0.5-1.3 mm long; lowest elongate culm internode > 2 mm in diameter; lowest nodes 

usually glabrous or pubescent .................................................................................................D. scabriusculum 

6 Panicle rachis scabrous, not pellucid-punctate; first glume 0.7-1.1 mm long; larger leaves 7-12 cm long, 6-9 

mm wide; ligule 0.3-0.6 mm long; lowest elongate culm internode < 1.7 mm in diameter; lowest nodes 

usually retrorsely bearded .................................................................................... D. species 9 (= cryptanthum) 

5 Ligule of short white hairs or absent. 

7 Leaves basally disposed, usually matted or cushion-forming, larger than the mid and upper culm leaves; 

blade margins uniformly papillose-ciliate; culms branching only at the base, 0.5-3.5 dm tall; internodes 

glabrous or sparsely pubescent....................................................................... D. strigosum var. leucoblepharis 

7 Basal leaves rosette-forming, usually much smaller than culm leaves; blade margins glabrous, or ciliate only 

below the middle (or papillose-ciliate throughout in =lancearium, which has densely puberulent internodes); 

culms branching at the nodes in age, 1.5-7.5 mm tall. 

8 Blades of mid-culm leaves typically long and stiff, acuminate, linear or narrowly lanceolate, usually > 

10× as long as wide, only 2-5 mm wide when < 8 cm long. 

9 Spikelets 2.9-4.0 mm long, fusiform to elliptic, acute, basally constricted; first glume 1.4-2.6 mm 

long..............................................................................................................D. species 8 (=fusiforme) 

9 Spikelets 1.5-3.1 mm long, obovate to elliptic-obovate, obtuse to sub-acute, not constricted basally; 

first glume 0.6-1.5 mm long. 

10 Spikelets 1.5-2.2 mm long; first glume 0.6-0.8 mm long; blades to 8 cm long, usually involute 

................................................................................................................................. D. aciculare 

10 Spikelets 2.1-3.1 mm long; first glume 0.8-1.5 mm long; blades to 12 cm long, usually flat 

except at tip ......................................................................................................D. angustifolium 

8 Blades of mid-culm leaves lanceolate, thin or firm but not stiff, usually < 10× as long as wide, usually 7 

mm or more wide when as much as 8 cm long. 

11 Spikelets 2.9-3.8 mm long, broadly elliptic, rounded at the summit, with broad and thick nerves...... 

.................................................................................................... D. oligosanthes var. scribnerianum

POACEAE 866 

11 Spikelets 2.1-2.9 mm long, elliptic or obovate, rounded or pointed at the summit, the nerves often 

raised, but not broad and thick. 

12 Culm internodes and sheaths glabrous or sparsely pilose. 

13 Spikelets obpyriform when viewed dorsally, strongly plano-convex when viewed 

laterally; first glume and base of second glume usually strongly reddish-purple ................ 

.............................................................................................. D. species 2 (=webberianum) 

13 Spikelets variously shaped but not obpyriform when viewed dorsally, biconvex to elliptic 

when viewed laterally; first and second glumes various. 

14 Culms tending to be stiffly erect; blades erect or erect-spreading, broad, usually but 

not always tapering from just below the middle to both ends, often yellowish green; 

plants not or only sparingly branched in age, not developing leafy fascicles of 

reduced leaves and inflorescences .............................................................. D. boreale 

14 Culms not stiffly erect; leaves usually spreading, broad or narrow, dark to bright 

green; plants often freely branched in age, becoming top-heavy with a mass of 

fascicled, reduced leaves and inflorescences ............................D. dichotomum group 

12 Culm internodes crisp-puberulent (sparsely so in D. species 2 (=webberianum); sheaths 

puberulent or glabrous. 

15 Spikelets elliptic, sub-acute to pointed, greenish or faintly purple-tinged basally ............... 

.................................................................................................. D. commutatum var. ashei 

15 Spikelets strongly plano-convex when viewed laterally, obpyriform when viewed 

dorsally, broadly rounded, usually markedly reddish-purple basally. 

16 Fertile lemma and palea papillose; spikelets 2.2-2.6 mm long; lower culm blades 6- 

12 mm wide, glabrous................................................... D. species 2 (=webberianum) 

16 Fertile lemma and palea smooth (minutely reticulate but not papillose); spikelets 

(1.8) 1.9-2.2 (-2.3) mm long; lower culm blades 4-8 mm wide, glabrous, glabrate, or 

puberulent (especially abaxially) ......................................D. species 3 (=lancearium) 

Key F - Spikelets 0.8-2.0 mm long, lower culm internodes variously hairy 

1 Longer hairs of ligule 2-5 mm long. 

2 Ligule without a distinct ring of short hairs in front of the long hairs.............................................D. acuminatum group 

2 Ligule with a distinct ring of short hairs in front of the long hairs. 

3 Peduncle, panicle axis, and sheaths puberulent with hairs 0.1 mm long; larger blades 3-6 cm long, 3-5 mm wide; 

spikelets 1.3-1.7 mm long ...................................................................................................................D. meridionale 

3 Peduncle panicle axis hairs > 0.1 mm long; sheaths and internodes densely clothed with straight retrorse 

(occasionally spreading to spreading-ascending) hairs often > 4 mm long................................................................. 

........................................................................................................................... D. villosissimum var. villosissimum 

1 Longer hairs of ligule < 2 mm long. 

4 Culm leaves basally crowded, ascending, usually matted or cushion-forming, larger than the mid and upper culm 

blades. 

5 Sheaths conspicuously retrorsely long-pilose; longer blades 6-18 cm long and 7-12 mm wide; spikelets 1.9-2.3 

mm long ................................................................................................................................................ D. laxiflorum 

5 Sheaths variously pubescent or glabrous, but not conspicuously retrorsely long-pilose; longer blades 2-6 cm long 

and 1-8 mm wide; spikelets 0.9-2.1 mm long. 

6 Blades 1-4 mm wide, glabrous, the margins eciliate or basally ciliate; spikelets 0.9-1.5 mm long, glabrous; 

autumnal form branched from lower and mid nodes as well as from basal nodes. 

7 Blades 1.5-4 (-5) cm long; spikelets 0.9-1.2 mm long.....................................................D. chamaelonche 

7 Blades 4-12 (-20) cm long, some at least 7 cm long; spikelets 1.2-1.5 mm long ........................................ 

............................................................................................................[D. dichotomum var. glabrifolium] 

6 Blades 2-10 mm wide, pubescent or glabrous, the margins coarsely papillose-ciliate throughout; spikelets 1.1- 

2.1 mm long, glabrous or pubescent; autumnal form branched from basal nodes only. 

8 Spikelets pubescent, 1.5-2.1 mm long; blade surfaces glabrous............. D. strigosum var. leucoblepharis 


9 Blades pilose; spikelets 1.1-1.6 mm long ............................................... D. strigosum var. strigosum 

9 Blades glabrous, or sparsely pilose only near the adaxial base; spikelets 1.4-1.8 mm long ................ 

............................................................................................................. D. strigosum var. glabrescens 

4 Culm leaves not basally crowded, the lowest leaves spreading and rosette-forming, usually smaller than the culm 

leaves. 

10 Blades of mid-culm leaves typically long and stiff, acuminate, linear or narrowly lanceolate, often involute, only 

2-5 mm wide when < 8 cm long.............................................................................................................. D. aciculare 

10 Blades of mid-culm leaves lanceolate, thin or firm but not stiff, usually > 5 mm when > 8 cm long. 

11 Internodes crisp-puberulent.

POACEAE 867 

12 Ligule 0.7-1.5 mm long; first glume acute; spikelets elliptic when viewed dorsally, biconvex or elliptic 

when viewed laterally, not strongly nerved......................................................................D. columbianum 

12 Ligule < 0.5 mm long; first glume obtuse to truncate; spikelets obovate when viewed dorsally, planoconvex 

when viewed laterally, strongly nerved. 

13 Spikelets 1.5-1.8 mm long; first glume 0.5-0.8 mm long; lower culm blades 2-5 mm wide ............... 

.................................................................................................................................... D. portoricense 

13 Spikelets (1.8-) 1.9-2.2 (-2.3) mm long; first glume 0.8-1.2 mm long; lower culm blades 4-8 mm 

wide..........................................................................................................D. species 3 (=lancearium) 

11 Internodes variously hairy but not crisp-puberulent. 

14 Internodes (sparsely-) moderately to densely pubescent to pilose; ligule 1-5 mm long; blade margins 

either weakly ciliate, papillose-ciliate basally only, or eciliate, lacking a white-beige cartilagionous edge 

0.2 mm wide. 

15 Larger mid-culm blades 4-7 cm long, 4-7 mm wide, glabrous to sparsely pubescent adaxially; ligule 

1-5 mm long; spikelets 1.1-1.5 mm long........................................................................D. leucothrix 

15 Larger mid-culm blades 3-6 cm long, 3-5 mm wide, long-pilose adaxially; ligule often with a ring 

of hairs < 1 mm and scattered longer hairs to 4 mm; spikelets 1.3-1.7 mm long.........D. meridionale 

14 Internodes sparsely pilose; ligule < 1 mm long; blade margins either coarsely papillose-ciliate 

throughout or glabrous with a white-beige cartilaginous edge about 0.2 mm wide. 

16 Blades with white-beige cartilaginous margins 0.2 mm wide; spikelets 1.4-1.7 mm long; autumnal 

form branching from middle and upper nodes........................................................................D. tenue 

16 Blade margins coarsely papillose-ciliate throughout; spikelets 1.1-2.1 mm long; autumnal form 

branching from the base. 

17 Spikelets pubescent, 1.5-2.1 mm long; blade surfaces glabrous.................................................. 

................................................................................................ D. strigosum var. leucoblepharis 


18 Blades pilose; spikelets 1.1-1.6 mm long ............................... D. strigosum var. strigosum 

18 Blades glabrous, or sparsely pilose only near the adaxial base; spikelets 1.4-1.8 mm long 

............................................................................................. D. strigosum var. glabrescens 

Key G - Spikelets 0.8-2.0 mm long, lower culm internodes glabrous 

1 Ligule 1-5 mm long. 

2 Ligule 1-2 mm long; sheaths sparsely to moderately spreading short-pilose; internodes glabrous; nodes retrorsely 

bearded; leaves 1-4 cm long, 2-5 mm wide; spikelets 1.2-1.4 mm long ................................D. species 10 (=curtifolium) 

2 Ligule (1.5-) 2-5 mm long; sheaths glabrous to variously pubescent; internodes glabrous or pubescent; nodes glabrous, 

or bearded with ascending, spreading, or tangled hairs; leaves 3-10 cm long, 3-10 mm wide; spikelets 0.8-1.9 mm long. 

........................................................................................................................................................D. acuminatum group 

1 Ligule < 1 mm long. 

3 Basal leaves rosette-forming, usually much smaller than the culm leaves, not matted or cushion-forming; culms 

branching at the mid and upper nodes in age. 

4 Blades of mid-culm leaves typically long and acuminate, linear or narrowly lanceolate, usually 10-20× as long as 

wide, only 2-5 mm wide when < 8 cm long. 

5 Spikelets papillose-pubescent; blades 1-2 (-3) mm wide; panicle 2-3 cm wide; first glume 0.8-1.0 mm long, 

acute; culms to 4 dm tall ........................................................................................D. species 5 (=neuranthum) 

5 Spikelets glabrous; blades 3-8 mm wide; first glume 0.3-1.1 mm long, truncate to acute; culms to 10 dm tall. 

6 Leaves 3-8 mm wide; panicle (8-) 20-40 mm wide; first glume 0.6-1.1 mm long, blunt to acute............... 

................................................................................................................................ D. dichotomum group 

6 Leaves 3-5.5 mm wide; panicle 2-5 mm wide; first glume 0.3-0.4 mm long, truncate to obtuse ................ 

......................................................................................................................................................D. hirstii 

4 Blades of mid-culm leaves lanceolate, mostly 10× or less as long as wide, usually 7 mm or more wide when as 

much as 8 cm long. 

7 Spikelets elliptic, oblong, or obovate; lower culm blades 3-12 (-15) mm wide, thin, tapered to the base; plants 

often freely branching in age, becoming top-heavy with a mass of fascicled, reduced leafy branches and 

inflorescences.................................................................................................................. D. dichotomum group 

7 Spikelets broadly elliptic to suborbicular; lower culm blades 6-30 mm wide, thickish, broad, and cordate to 

subcordate at the base; plants sparingly branched in age, not becoming top-heavy with fascicled, reduced 

leafy branches and inflorescences. 

8 Spikelets 0.9-1.2 mm long; longer blades 6-8 cm long, erect to erect-ascending................D. erectifolium 

8 Spikelets 1.2-1.9 mm long; longer blades 8-20 cm long, ascending or the uppermost erect. 

9 Mid-culm blades 6-11 (-14) mm wide, the uppermost 3-9 cm long .................................................... 

................................................................................................D. sphaerocarpon var. sphaerocarpon 

9 Mid-culm blades, at least some, 15-30 mm wide, the uppermost 10-15 cm or more long................... 

.......................................................................................................................................D. polyanthes

POACEAE 868 

3 Basal leaves similar to or larger than the mid and upper culm leaves, often matted or cushion-forming; culms branching 

at the base (also at mid and upper nodes in D. chamaelonche vars. and D. dichotomum var. glabrifolium). 

10 Longer blades > 6 cm; if only 6 cm, then sheaths retrorsely long-pilose (D. laxiflorum). 

11 Spikelets 1.2-1.5 mm long, glabrous ...........................................................[D. dichotomum var. glabrifolium] 

11 Spikelets 1.7-2.3 (-2.8) mm long, pubescent. 

12 Longer blades 6-18 cm long by 7-12 mm wide; sheaths conspicuously retrorsely long-pilose; nodes 

bearded with retrorse or spreading hairs; spikelets 1.9-2.3 mm long ................................... D. laxiflorum 

12 Longer blades 10-35 cm long by 2-4 mm wide; sheaths glabrous to variously pilose, but not 

conspicuously retrorsely long-pilose; nodes variously pubescent to glabrate; spikelets 1.7-2.3 (-2.8) mm 

long ................................................................................................................................... D. linearifolium 

10 Longer blades 1.5-6 cm; sheaths glabrous or pubescent, but not retrorsely long-pilose. 

13 Blades 1-3 mm wide, glabrous, eciliate or basally ciliate; spikelets 0.9-1.4 mm long. 

14 Spikelets pubescent, 1.2-1.4 mm long; blades involute, often falcate, 2.5-6 cm long ................................. 

.................................................................................................................... [D. chamaelonche ssp. breve] 

14 Spikelets glabrous, 0.9-1.2 mm long; blades flat, not falcate, 1.5-4 (-5) cm long ....................................... 

........................................................................................................ D. chamaelonche ssp. chamaelonche 

13 Blades 3-8 mm wide; spikelets 1.1-2.1 mm long (if < 1.5 mm, then blades either pubescent on one or both 

surfaces or ciliate to the apex). 

15 Spikelets pubescent, 1.5-2.1 mm long; blade surfaces glabrous............. D. strigosum var. leucoblepharis 


16 Blades pilose; spikelets 1.1-1.6 mm long ............................................... D. strigosum var. strigosum 

16 Blades glabrous, or sparsely pilose only near the adaxial base; spikelets 1.4-1.8 mm long ................ 

............................................................................................................. D. strigosum var. glabrescens 

Key to the Dichanthelium acuminatum group 

1 Internodes glabrous. 

2 Ligule 1-2 mm long; sheaths sparsely to moderately spreading short-pilose; nodes retrorsely bearded; leaves 1-4 cm 

long, 2-5 mm wide; spikelets 1.2-1.4 mm long......................................................................D. species 10 (=curtifolium) 

2 Ligule (1.5-) 2-5 mm long; sheaths glabrous to variously pubescent, but not spreading short-pilose; nodes glabrous or 

pubescent, but not bearded; leaves 4-11 cm long, 4-8 mm wide. 

3 Panicles 8-12 cm long, ¼-⅓ as wide, bearing 250 or more spikelets; spikelets 1.4-1.6 mm long; ligule (1.5-) 2-3 

mm long; larger blades 7-11 cm long, often tinged with purple................................................................ D. spretum 

3 Panicles 3-8 cm long, > ½ as wide, bearing < 200 spikelets; spikelets 1.1-1.6 mm long; ligule 2-5 mm long; larger 

blades 4-10 cm long. 

4 Longer hairs of ligule 2-3 mm long; spikelets 1.1-1.5 mm long; blades often tinged with purple, the larger 4-8 

cm long................................................................................................................................... D. longiligulatum 

4 Longer hairs of ligule 3-5 mm long; spikelets 1.4-1.6 mm long; blades often yellowish-green, the larger 5-10 

cm long............................................................................................................D. acuminatum var. lindheimeri 

1 Internodes variously pubescent. 

5 Peduncle, panicle axis, and/or sheaths of vernal culms puberulent with hairs 0.1 mm long, sometimes also pubescent 

with longer hairs, but never grayish-villous; larger blades 2-7 cm long, 2-7 mm wide. 

6 Spikelets 0.8-1.1 mm long; blades 2-4.5 cm long, 2-5 mm wide; sheaths sparsely puberulent, lacking papillosebased 

longer hairs.............................................................................................................................. D. wrightianum 

6 Spikelets 1.1-1.7 mm long; mid-culm blades generally 3-7 cm long and 3-7 mm wide; sheaths with some 

papillose-based hairs 2 mm or more long. 

7 Larger mid-culm blades 4-7 cm long, 4-7 mm wide, glabrous to sparsely pubescent adaxially; ligule 1-5 mm 

long; spikelets 1.1-1.5 mm long .....................................................................................................D. leucothrix 

7 Larger mid-culm blades 3-6 cm long, 3-5 mm wide, long-pilose adaxially; ligule often with a ring of hairs < 1 

mm and scattered longer hairs to 4 mm; spikelets 1.3-1.7 mm long ...........................................D. meridionale 

5 Peduncle, panicle axis, and sheaths of vernal culms glabrous, or pilose, or grayish-villous with some shorter hairs 0.2- 

0.5 mm long, but not puberulent with hairs 0.1 mm long; larger blades 4-12 cm long, 4-12 mm wide. 

8 Sheaths and internodes of vernal culms gray-villous with a dense, tangled, or matted mixture of slender hairs 2-4 

mm long, variously ascending, spreading, and retrorse, papillose or non-papillose, often with shorter hairs 

beneath; blades velvety-pubescent on abaxial surface. 

9 Culms 15-60 cm tall and < 1.5 mm thick; panicle broadly ovoid, 5-8 cm long and > 1/2 as wide...................... 

...................................................................................................................... D. acuminatum var. acuminatum 

9 Culms 40-70 (-80) cm tall, the larger usually > 60 cm long and > 2 mm thick; panicle contracted, 8-11 cm 

long and < ½ as wide............................................................................................D. acuminatum var. thurowii 

8 Sheaths and internodes of vernal culms glabrous, or papillose-pilose to hispid with ascending straight hairs 1-3 

mm long; blades appressed-pilose to puberulent abaxially, but not velvety. 

10 Peduncle, panicle axis, and often middle and upper internodes glabrous; sheaths, at least near mid-length, 

lacking hairs or papillae; larger blades 4-8 mm wide, glabrous abaxially; spikelets 1.3-1.6 mm long................

POACEAE 869 

........................................................................................................................D. acuminatum var. lindheimeri 

10 Peduncle, panicle axis, and internodes pubescent to pilose; sheaths papillose-pilose to hispid, the hairs 

tending to break off but leaving evident papillae; larger blades 6-10 mm wide, short-pilose to glabrate 

abaxially; spikelets 1.5-2.0 mm long.............................................................D. acuminatum var. fasciculatum 

Key to the Dichanthelium dichotomum Group 

1 Lower cauline nodes glabrous or puberulent, but not bearded. 

2 Spikelets glabrous. 

3 Cauline leaves mostly basally disposed, strongly ascending, much larger than the 2-3 remote middle and upper 

cauline leaves of fertile culms; spikelets 2.4-2.9 mm long; culms branch from basal and lower nodes, but are not 

known to produce autumnal inflorescences..........................................................................................[D. nudicaule] 

3 Cauline leaves well-distributed along the culm, > 3, gradually reduced upwards and often spreading; spikelets 0.9- 

2.6 mm long; culms produce autumnal inflorescences from lower, middle, and/or upper nodes, if from lower only, 

then spikelets only 0.9-1.2 mm long. 

4 Fertile lemma and palea densely papillose; culms weak, soon sprawling over other vegetation....... D. lucidum 

4 Fertile lemma and palea smooth, with few or no papillae; culms stiffer, erect to ascending. 

5 Spikelets 0.9-1.5 mm long; vernal blades 1-4 mm wide. 

6 Spikelets 0.9-1.2 mm long; blades 1.5-4 (-5) cm long, 1-2.5 (-3) mm wide, mostly 15-20 times as 

long as wide; autumnal plants cushion-forming ...................... D. chamaelonche ssp. chamaelonche 

6 Spikelets 1.2-1.5 mm long; blades 1-12 (-20) cm long; autumnal plants not cushion-forming. 

7 Blades 1-3 (-5) cm long, 1.5-3 (-4) mm wide, about 10 times as long as wide; autumnal plants 

sparsely tufted........................................................................................................ D. ensifolium 

7 Blades 4-12 (-20) cm long (the longer at least 7 cm), 2-4 mm wide, 20-30 (-50) times as long 

as wide.................................................................................[D. dichotomum var. glabrifolium] 

5 Spikelets 1.4-2.6 mm long; vernal blades 3-15 mm wide (if spikelets < 1.6 mm long and vernal blades < 

5 mm wide, then larger blades > 5 cm long in D. caerulescens). 

8 Widest vernal cauline blades 7-15 mm wide; upper sheaths often glutinous-warty; spikelets 2.1-2.6 

mm long, some or most acute to beaked, second glume and sterile lemma extending 0.3-0.5 mm 

beyond fertile lemma in at least some spikelets............................................................ D. yadkinense 

8 Widest vernal cauline blades 3-10 mm wide; upper sheaths not glutinous-warty; spikelets 1.4–2.3 

mm long, blunt to subacute, second glume and sterile lemma often equal to or shorter than fertile 

lemma, or extending < 0.3 mm beyond it. 

9 Spikelets 1.4-1.8 mm long; first glume 0.3–0.8 mm long; fertile lemma 1.3-1.5 mm long; 

mature vernal panicles usually short-exerted with ascending branches; fresh foliage bluishglaucous..............................................................................................................D. 

caerulescens 

9 Spikelets 1.7-2.3 mm long; first glume 0.6-1.1 mm long; fertile lemma 1.6-1.9 mm long; 

mature vernal panicles exerted with spreading branches; fresh foliage not bluish-glaucous. 

10 Vernal cauline blades spreading to deflexed, flexuous; [of wet-mesic to dry woods and 

thickets] ..........................................................................D. dichotomum var. dichotomum 

10 Vernal cauline blades stiffly erect; [of wet pine savannas and open swamps]..................... 

......................................................................................... D. dichotomum var. roanokense 

2 Spikelets pubescent. 

11 Spikelets 1.2-1.7 mm long; fertile lemma and palea smooth; culms erect. 

12 Blades involute and often falcate, 3-6 cm long, about 1.5 mm wide when flattened, 20-50× as long as wide; 

lower internodes often strigose; spikelets 1.2-1.4 mm long; culms 5-20 cm long............................................... 

............................................................................................................................ [D. chamaelonche ssp. breve] 

12 Blades neither involute (except apically) nor falcate, 1-7 cm long, 1.5-7 mm wide, about 10× as long as wide; 

lower internodes glabrous or sparsely pilose, but not strigose; spikelets 1.1-1.7 mm long; culms 15-60 cm 

long. 

13 Blades 1-3 (-5) cm long, 1.5-3 (-4) mm wide, the cartilaginous margins typically gray-green to whitebeige 

and about 0.1 mm wide; spikelets 1.2-1.5 mm long; culms to 40 cm long .................. D. ensifolium 

13 Blades 2-7 mm long, 3-6 mm wide, the cartilaginous margins typically white-beige and about 0.2 mm 

wide; spikelets (1.2-) 1.4-1.7 mm long; culms to 60 cm long ........................................................D. tenue 

11 Spikelets (1.5-) 1.7-2.7 mm long, if shorter than 1.8 mm then fertile lemma and palea densely papillose; culms 

soon sprawling. 

14 Spikelets (1.5-) 1.7-2.3 mm long, glabrous (rarely pubescent); first glume 0.7-1.1 mm long; fertile lemma and 

palea densely papillose at 20×........................................................................................................... D. lucidum 

14 Spikelets 2.2-2.7 mm long, pubescent; first glume 1.0–1.4 mm long; fertile lemma and palea smooth or with a 

few weak papillae at 20×.............................................................................................................D. sphagnicola 

1 Lower cauline nodes bearded, the hairs usually retrorse. 

13 Spikelets glabrous.

POACEAE 870 

14 Spikelets 0.9-1.4 mm long; vernal cauline blades 1.5-4 (-5) cm long and 1-5 mm wide; internodes or sheaths 

glabrous or pubescent. 

15 Spikelets 1.2-1.4 mm long; sheaths spreading-pilose; vernal cauline blades 2-5 mm wide; ligule 1-2 mm long; 

node beard hairs usually spreading or reflexed; internodes glabrous ............................D. sp. 10 (=curtifolium) 

15 Spikelets 0.9-1.2 mm long; sheaths glabrous; vernal cauline blades 1-2 (-3) mm wide; ligule < 1 mm long; 

node beard hairs erect and often only partially encircling the node; internodes glabrous or puberulent ............. 

................................................................................................................................................D. chamaelonche 

14 Spikelets 1.4-2.3 mm long; vernal cauline blades 5-12 cm long and 3-15 mm wide; internodes and sheaths 

glabrous. 

16 Spikelets 1.8-2.3 mm long; first glume 0.6-1.1 mm long; fertile lemma 0.8–1.0 mm wide; widest vernal 

blades 3-8 (-10) mm wide ..............................................................................D. dichotomum var. dichotomum 

16 Spikelets 1.4-1.9 mm long; first glume 0.3-0.6 (-0.7) mm long; fertile lemma 0.6-0.8 mm wide; widest vernal 

blades 7-15 mm wide ...................................................................................... D. dichotomum var. ramulosum 

13 Spikelets pubescent. 

17 Spikelets 1.2-1.4 mm long; sheaths spreading-pilose; vernal cauline blades 1-4 cm long and 2-5 mm wide; ligule 

1-2 mm long..........................................................................................................................D. sp. 10 (=curtifolium) 

17 Spikelets 1.4-2.8 mm long; sheaths glabrous to appressed-pilose; vernal cauline blades 5-12 cm long and 5-15 mm 

wide; ligule < 1 mm long. 

18 Usually all culm nodes bearded; internodes glabrous, or middle and upper internodes and peduncle sparsely 

to moderately spreading short-hairy, sometimes also glandular; upper as well as lower vernal sheaths and 

both surfaces of cauline blades pubescent, often densely so; spikelets (1.5-) 1.8-2.1 mm long; [of dry rocky or 

sandy basic soil and barrens]............................................................................................................D. annulum 

18 Often only lower culm nodes bearded; internodes glabrous; at least middle and upper cauline blades glabrous; 

spikelets 1.4-2.8 mm long; [mostly of wet acid soils and mesic to dry woodlands]. 

19 Spikelets (2.0-) 2.2-2.8 mm long; first glume 0.5-1.3 mm long; fertile lemma 1.8-2.3 mm long; lowest 

vernal cauline blades pubescent at least abaxially.....................................................D. mattamuskeetense 

19 Spikelets 1.4-2.2 mm long; first glume 0.3-0.9 mm long; fertile lemma 1.4-1.7 mm long; lowest vernal 

cauline blades glabrous. 

20 Spikelets 1.7-2.2 mm long; first glume 0.6-0.9 mm long; fertile lemma 0.7-1.0 mm wide................. 

............................................................................................................... D. dichotomum var. nitidum 

20 Spikelets 1.4-1.9 mm long; first glume 0.3-0.6 (-0.8) mm long; fertile lemma 0.6-0.8 mm wide ....... 

......................................................................................................... D. dichotomum var. ramulosum 

Dichanthelium aciculare (Desvaux ex Poiret) Gould & Clark, Needle Witch Grass. Cp, Pd (GA, NC, SC, VA): sandy 

woods and fields; common in Coastal Plain, uncommon in Piedmont. May-October. NJ south to n. Fl, west to TX and OK, also 

in West Indies and n. South America. Blades typically are strongly involute. Can be confused with autumnal forms of D. ovale 

var. addisonii, which has vernal blades 5-10 mm wide. See note at end of descriptions regarding Panicum chrysopsidifolium. [= 

Panicum aciculare Desvaux ex Poiret – RAB, F, G; < P. aciculare – C; = D. aciculare ssp. aciculare – FNA; > P. aciculare – 

HC, S; > P. bennettense M.V. Brown – HC, S; < D. aciculare – K, Z] 

Dichanthelium acuminatum (Swartz) Gould & Clark. var. acuminatum, Woolly Witch Grass. Cp, Pd (GA, NC, SC, VA): 

on dryish sandy or clayey soils of open woods and disturbed areas; common. May-October. MA south to FL, west to TX, also in 

West Indies, Mexico, Central America, and n. South America. Internodes and sheaths gray-villous with usually non-papillate 

hairs. Plants tend to be low and "bushy" with several spreading-ascending culms and dense autumnal branching. See note at end 

of descriptions regarding Panicum chrysopsidifolium. [= Y; < Panicum lanuginosum Elliott – RAB; > P. lanuginosum var. 

lanuginosum – C, F, G; < P. leucothrix Nash – C; > P. auburne Ashe – F, G, HC, S; < D. acuminatum ssp. acuminatum – FNA; 

> P. lanuginosum – HC, S; >< D. acuminatum var. acuminatum – K, Z; >< D. acuminatum var. implicatum (Scribner) Gould & 

Clark – K, Z; < P. acuminatum Swartz var. acuminatum – X] 

Dichanthelium acuminatum (Swartz) Gould & Clark var. fasciculatum (Torrey) Freckmann, Slender-stemmed Witch 

Grass. Mt, Pd, Cp (GA, NC, SC, VA): open or cut-over woods, thickets, fields, meadows, and shores, frequently on disturbed 

soils; frequent (less common in the Coastal Plain). May-August. Newfoundland south to FL, west to CA, north to s. British 

Columbia. Typically much less pilose than var. acuminatum, the hairs usually papillate. See note at end of descriptions 

regarding Panicum glutinoscabrum. [= Y; < Panicum lanuginosum Elliott – RAB; > P. lanuginosum var. fasciculatum (Torrey) 

Fernald – C, F, G; > P. lanuginosum var. tennesseense (Ashe) Gleason – C, G; > P. lanuginosum var. implicatum (Scribner) 

Fernald – C, F, G; = D. acuminatum ssp. fasciculatum (Torrey) Freckmann & Lelong – FNA; > P. implicatum Scribner – HC; > 

P. huachucae Ashe var. huachucae – HC, S; > P. huachucae var. fasciculatum (Torrey) Hubb. – HC; > P. tennesseense Ashe – 

HC, S; > P. huachucae var. silvicola Hitchcock & Chase – S; >< D. acuminatum var. acuminatum – K, Z; >< D. acuminatum 

var. implicatum (Scribner) Gould & Clark – K, Z; > P. acuminatum Swartz var. fasciculatum (Torrey) Lelong – X; > P. 

acuminatum var. unciphyllum (Trinius) Lelong – X] 

Dichanthelium acuminatum (Swartz) Gould & Clark var. lindheimeri (Nash) Gould & Clark, Lindheimer's Witch Grass. 

Pd, Cp, Mt (GA, NC, SC, VA): open or cut-over woods, thickets, fields, meadows, and shores, often on wet soils; uncommon in 

Piedmont, rare in Coastal Plain and Mountains. May-September. Nova Scotia west to Manitoba, south to FL and MO, west to s. 

CA. Internodes as well as sheaths often nearly glabrous. Panicle axis sometimes sparsely pilose at branch nodes, but otherwise 

glabrous. [= Y; < Panicum lanuginosum Elliott – RAB; >< P. lanuginosum var. lindheimeri (Nash) Fernald – C, G; > P. 

lanuginosum var. septentrionale Fernald – C, F, G; > P. lanuginosum var. lindheimeri – F; = D. acuminatum ssp. lindheimeri

POACEAE 871 

(Nash) Freckmann & Lelong – FNA; < P. spretum Schultes – GW; > P. lindheimeri Nash – HC, S; < D. acuminatum var. 

acuminatum – K, Z; > D. acuminatum var. lindheimeri – K, Z; > P. acuminatum Swartz var. lindheimeri (Nash) Lelong – X; ? 

D. lanuginosum (Elliott) Gould var. lindheimeri (Nash) Harvill] 

Dichanthelium acuminatum (Swartz) Gould & Clark var. thurowii (Scribner & J.G. Smith) Gould & Clark, Thurow's 

Witch Grass. Cp (GA): in dry open woods, woodland edges, dry prairies, brushy pastures; occasional (possibly rare in GA). 

May?-October? Occasional from GA to AR and e. TX. [= K, Y, Z; < D. acuminatum (Swartz) Gould & Clark. ssp. acuminatum 

– FNA; = Panicum thurowii Scribner & J.G. Smith – HC, S] 

Dichanthelium angustifolium (Elliott) Gould, Narrow-leaved Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): sandy 

pinelands and fields; common in Coastal Plain and Piedmont, rare in mountains. May-October. NJ south to FL, west to AR and 

e. TX. Vernal blades typically are flat (often involute distally). Can be confused with D. consanguineum, which has spreadingpilose 

nodes and blades 10-15× as long as wide; D. angustifolium blades typically are 20× or more as long as wide. Plants with 

involute blades to 8 cm long, spikelets 2.1-2.5 mm long, and first glumes 0.7-1.1 mm long are referable to Panicum 

arenicoloides, here included in D. angustifolium. They are transitional to D. aciculare. [= Panicum angustifolium Elliott – RAB, 

F, G; < P. aciculare Desvaux ex Poiret – C; = D. aciculare ssp. angustifolium (Elliott) Freckmann & Lelong – FNA; > P. 

angustifolium – HC, S; > P. arenicoloides Ashe – HC, S; < D. aciculare – K, Z] 

Dichanthelium annulum (Ashe) LeBlond, Ringed Witch Grass. Pd (GA, NC, VA), Mt (VA): dry sandy or rocky soil of 

open woods, dry grasslands, and barrens, and glades over serpentine, limestone, calcareous shales, and other high pH dry soils; 

rare (NC Significantly Rare, VA Rare List). May-October. NJ, IN, and MO south to AL and MS, primarily in the Appalachian 

Province with very few occurrences in the Coastal Plain. One of the more distinctive taxa within the D. dichotomum group by 

morphology, habitat, and range. Plants from se. MA with all leaves pubescent, glabrous internodes, and spikelets 2.2-2.5 mm 

long were described as Panicum annulum var. glabrescens, but belong to D. mattamuskeetense. [= Q; < P. dichotomum 

Linnaeus – RAB, C, GW; = Panicum annulum Ashe – F, HC, S; = P. annulum var. annulum – G; < D. dichotomum ssp. 

mattamuskeetense (Ashe) Freckmann & Lelong – FNA; < D. dichotomum (Linnaeus) Gould – K, Z; < P. dichotomum var. 

mattamuskeetense (Ashe) Lelong - X] 

Dichanthelium boreale (Nash) Freckmann, Northern Witch Grass. Pd, Mt (GA, NC, VA): open woods and grassy slopes, 

usually in moist soil; rare (NC Watch List, VA Watch List). April-September. Newfoundland and Ontario south to NC, GA, and 

AR. Our plants are =Panicum bicknellii, regarded as a "putative hybrid" (along with =P. calliphyllum) by FNA, which cites WV 

as the southern limit of D. boreale. [= K, Z; > Panicum bicknellii Nash – RAB, F, HC, S; > P. boreale Nash – C, F, G, HC; > P. 

calliphyllum Ashe – F, HC; > D. boreale – FNA; > P. bicknellii var. bicknellii – G; > P. bicknellii var. calliphyllum (Ashe) 

Gleason – G] 

Dichanthelium boscii (Poiret) Gould & Clark, Bosc's Witch Grass. Pd, Mt, Cp (GA, NC, SC, VA): shaded mesic to dry 

woodlands; common. April-September. MA and IL south to n. FL and e. TX. [= FNA, K, Z; = Panicum boscii Poiret – RAB, 

C, G; > P. boscii var. boscii – F, HC, S; > P. boscii var. molle (Vasey) Hitchcock & Chase – F, HC, S] 

Dichanthelium caerulescens (Hackel ex Hitchcock) Correll, Blue Witch Grass. Cp (NC, VA): marshes, swamps, wet 

pinelands, maritime grasslands, damp sandy soil; rare (NC Watch List, VA Watch List). June-October. NJ to NC, and from FL 

to LA, also in the Bahamas and West Indies. Not treated by FNA, where it presumably would have been placed in synonymy 

with D. dichotomum ssp. roanokense. [= Q; < Panicum dichotomum Linnaeus – RAB, GW; = P. caerulescens Hackel ex 

Hitchcock – F, HC, S; < D. dichotomum ssp. roanokense – FNA; < P. roanokense Ashe – G; < D. dichotomum var. dichotomum 

– K, Z; < P. dichotomum var. roanokense (Ashe) Lelong – X] 

Dichanthelium chamaelonche (Trinius) Freckmann & Lelong ssp. chamaelonche, Carpet Witch Grass. Cp (GA, NC, SC, 

VA): moist pine savannas and flatwoods, pineland pondshores; occasional to rare (VA Watch List). April-September. Se. VA 

south to FL, west to LA, also in Cuba and Belize. Internodes can be glabrous or puberulent, and nodes glabrous, pubescent, or 

bearded, but the glabrous spikelets 0.9-1.2 mm long are diagnostic. The concept of this taxon in FNA (as ssp. chamaelonche) 

appears to include D. dichotomum var. glabrifolium (see descriptions of Floridian D. chamaelonche ssp. breve and D. 

dichotomum var. glabrifolium at end of this treatment). [= Panicum chamaelonche Trinius – RAB, G, GW, HC, S; < P. 

ensifolium Baldwin – C; < D. chamaelonche ssp. chamaelonche – FNA; < D. dichotomum (Linnaeus) Gould var. ensifolium 

(Baldwin) Gould & Clark – K, Z; = P. chamaelonche var. chamaelonche – X] 

Dichanthelium clandestinum (Linnaeus) Gould, Deer-tongue Witch Grass. Mt, Pd, Cp (GA, NC, SC, VA): shaded to 

filtered woodlands, ditches and low areas, and often in moist sandy soil; common (uncommon in Coastal Plain). May-October. 

Nova Scotia and Québec south to n. FL, west to IA, KA, and TX. [= FNA, K, Z; = Panicum clandestinum Linnaeus – RAB, C, 

F, G, HC, S, X] 

Dichanthelium columbianum (Scribner) Freckmann, American Witch Grass. Pd, Cp (GA, NC, SC, VA), Mt (NC, VA): 

dry to moist thin woods and open ground, usually in sandy soil; uncommon. June-October. S. ME, s. Ontario, and WI south to 

GA, TN, and IL. [= Panicum columbianum Scribner – RAB, C, G; > P. columbianum var. columbianum – F, HC; > P. 

columbianum var. oricola (Hitchcock & Chase) Fernald – F; = D. acuminatum ssp. columbianum (Scribner) Freckmann & 

Lelong – FNA; > P. columbianum var. thinium Hitchcock & Chase – HC; > P. oricola Hitchcock & Chase – HC; > P. 

tsugetorum Nash – HC, S; < D. sabulorum (Lamarck) Gould & Clark var. thinium (Hitchcock & Chase) Gould & Clark – K, Z; 

> P. columbianum – S; < P. acuminatum Swartz var. unciphyllum (Trinius) Lelong – X] 

Dichanthelium commutatum (Schultes) Gould var. ashei (Pearson ex Ashe) Mohlenbrock, Ashe's Witch Grass. Cp, Pd, Mt 

(GA, NC, SC, VA): dry rocky or sandy woods and openings; common. May-October. MA south to FL and MS, west to MI, 

MO, and OK. [< Panicum commutatum – RAB, C; = P. commutatum Schultes var. ashei (Pearson ex Ashe) Fernald – F, G; = D. 

commutatum ssp. ashei (Pearson ex Ashe) Freckman & Lelong – FNA; = P. ashei Pearson ex Ashe – HC, S; < D. commutatum – 

K]

POACEAE 872 

Dichanthelium commutatum (Schultes) Gould var. commutatum, Variable Witch Grass. Cp, Mt, Pd (GA, NC, SC, VA): 

low, shaded, moist woodlands and woodland edges, and dry, thin, often rocky woods and thickets; common. May-October. ME 

south to FL, west to MI, MO, OK, and TX, also in Mexico. [< Panicum commutatum Schultes – RAB, C; > P. commutatum var. 

commutatum – F, G; > P. commutatum – HC, S; > P. mutabile Scribner & Smith ex Nash – F, G, HC, S; > D. commutatum ssp. 

commutatum Freckmann & Lelong – FNA; > D. commutatum ssp. equilaterale (Scribner) Freckmann & Lelong – FNA; > D. 

commutatum ssp. joori (Vasey) Freckmann & Lelong – FNA; < D. commutatum – K; > P. joorii Vasey – HC, S; > P. 

equilaterale Scribner – HC, S] 

Dichanthelium consanguineum (Kunth) Gould & Clark, Kunth's Witch Grass. Cp, Pd (GA, NC, SC, VA): moist or dry 

sandy soils of pinelands; common in the Coastal Plain, uncommon in the Piedmont (VA Rare List). April-September. 

Occasional from se. VA south to FL, west to TX and IN. Often not easily separated from D. angustifolium and D. ovale. It is 

distinguished from D. angustifolium by spreading-hirsute nodes and leaves 10-15× as long as wide (D. angustifolium has 

beardless nodes, or nodes bearded with erect-ascending soft hairs, and longer leaves 20× or more as long as wide). D. 

consanguineum is distinguished from D. ovale by having strongly pilose upper blade surfaces (D. ovale upper blade surfaces are 

glabrous or with a few long hairs basally). The hairs of D. consanguineum frequently are strongly papillate. [= FNA, K, Z; = 

Panicum consanguineum Kunth – RAB, C, F, G, HC, S] 

Dichanthelium depauperatum (Muhlenberg) Gould, Starved Witch Grass. Pd, Mt, Cp (GA, NC, SC, VA): dry soils of 

grasslands and open woods, often on disturbed soils of roadsides and ditches; common (rare in Coastal Plain). May-September. 

Newfoundland and MN south to GA and TX. [= FNA, K, Z; = Panicum depauperatum Muhlenberg – RAB, C, HC, S; > P. 

depauperatum var. depauperatum – F, G; > P. depauperatum var. psilophyllum Fernald – F, G] 

Dichanthelium dichotomum (Linnaeus) Gould var. dichotomum, Forked Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): 

wet-mesic to dry woods, thickets, and woodland openings; common throughout. May-October. S. Canada and MI south to FL 

and TX. Plants with bearded nodes and larger leaves are referable to Panicum dichotomum var. barbulatum (here included) but 

intermediates abound. [= Q; < Panicum dichotomum Linnaeus – RAB, C, GW; > P. dichotomum var. dichotomum – F; P. 

dichotomum var. barbulatum (Michaux) Wood – F; = D. dichotomum ssp. dichotomum – FNA; = P. dichotomum – G; > P. 

dichotomum – HC, S; > P. barbulatum Michaux – HC, S; < D. dichotomum var. dichotomum – K, Z; = P. dichotomum var. 

dichotomum – X] 

Dichanthelium dichotomum (Linnaeus) Gould var. nitidum (Lamarck) LeBlond, Shining Witch Grass. Cp (GA, NC, SC, 

VA), Pd (NC), Mt (SC, VA): moist sandy or peaty soil of wet pine savannas and pocosin ecotones, wet meadows near the coast, 

swamps, and marshes; occasional to frequent in Coastal Plain, rare in Piedmont and Mountains. PA and NJ south to FL, west to 

MO and TX; also the Bahamas (Sorrie & LeBlond 1997) and West Indies, and Mexico to Venezuela. [= Q; < Panicum 

dichotomum Linnaeus – RAB, C, GW; = P. nitidum Lamarck – F, HC, S; = D. dichotomum ssp. nitidum (Lamarck) Freckmann 

& Lelong – FNA; = P. nitidum var. nitidum – G; < D. dichotomum var. dichotomum – K, Z; = P. dichotomum var. nitidum 

(Lamarck) Wood – X] 

Dichanthelium dichotomum (Linnaeus) Gould var. ramulosum (Torrey) LeBlond, Branched Witch Grass. Cp, Pd, Mt 

(GA, NC, SC, VA): floodplain forests, swamps, openings, and borders of streams and ponds, and occasionally in dry upland 

woods; widespread. May-October. MA and MI south to FL and TX. [= Q; < Panicum dichotomum Linnaeus – RAB, C, GW; = 

P. microcarpon Muhlenberg ex Elliott – F, HC, S; = D. dichotomum ssp. microcarpon (Muhlenberg ex Elliott) Freckmann & 

Lelong – FNA; = P. nitidum Lamarck var. ramulosum Torrey – G; < D. dichotomum var. dichotomum – K, Z; = P. dichotomum 

var. ramulosum (Torrey) Lelong – X] 

Dichanthelium dichotomum (Linnaeus) Gould var. roanokense (Ashe) LeBlond, Roanoke Witch Grass. Cp (GA, NC, SC, 

VA): wet pine savannas, swamp openings, and wet peaty meadows; uncommon (NC Watch List). May-September. DE south to 

FL, west to e. TX; also in Jamaica. See note under D. caerulescens regarding FNA treatment. [= Q; < Panicum dichotomum 

Linnaeus – RAB, C, GW; = P. roanokense Ashe – F, HC, S; < D. dichotomum ssp. roanokense (Ashe) Freckmann & Lelong – 

FNA; < P. roanokense – G; < D. dichotomum var. dichotomum – K, Z; < P. dichotomum var. roanokense (Ashe) Lelong – X] 

Dichanthelium ensifolium (Baldwin ex Elliott) Gould, Small-leaved Witch Grass. Cp, Pd (GA, NC, SC, VA): wet to mesic 

peaty, sandy, or mucky soils, often in open pinelands or with sphagnum; common in Coastal Plain, rare in Piedmont and 

Mountains. May-October. NJ south to FL, west to e. TX and AR. Plants with pubescent spikelets are frequent. [= D. ensifolium 

ssp. ensifolium – FNA; < Panicum ensifolium Baldwin ex Elliott – RAB, C, G, GW; = P. ensifolium – F; > P. ensifolium – HC, 

S; > P. flavovirens Nash – HC, S; > P. vernale Hitchcock & Chase – HC, S; < D. dichotomum (Linnaeus) Gould var. ensifolium 

(Baldwin ex Elliott) Gould & Clark – K, Z; < P. ensifolium var. ensifolium – X] 

Dichanthelium erectifolium (Nash) Gould & Clark, Erect-leaved Witch Grass. Cp (GA, NC, SC): limesink ponds, 

depression meadows, cypress savannas, pine savannas; rare (NC Watch List). May-August. Se. NC to FL, west to LA; Cuba. [= 

FNA, K, Z; = Panicum erectifolium Nash – RAB, GW, HC, S] 

Dichanthelium fusiforme (Hitchcock) Harvill, Spindle-fruited Witch Grass. Cp (GA, NC, SC, VA): dry to moist sand of 

open pine and pine/oak woods and clearings; rare (NC Significantly Rare, VA Watch List). May-November. Se. VA south to 

FL, west to MS, also in West Indies, Mexico, Central America, and Venezuela; perhaps most abundant in FL. Autumnal blades 

often flat. The autumnal form of D. oligosanthes var. oligosanthes can be very similar to D. fusiforme if the vernal blades of the 

former are missing. They are best separated by ligule length (0.5-1 mm in fusiforme, 1.5-3 mm in oligosanthes) and the more 

attenuated ends of the fusiforme spikelet. [= Panicum fusiforme Hitchcock – RAB, F, G, HC, S; < P. aciculare Desvaux ex Poiret 

– C; = D. aciculare ssp. fusiforme (Hitchcock) Freckmann & Lelong – FNA; < D. aciculare – K, Z] 

Dichanthelium hirstii (Swallen) Kartesz, Hirsts' Witch Grass. Cp (GA, NC): pond-cypress savannas and limesink 

depressions; rare (US Candidate, NC Endangered). June-September. This distinctive species is known from only seven sites: 

two in NC, one in DE, two in NJ (one not seen since 1992), and two historical populations in GA. Described in 1961 (Swallen 

1961), it is treated by some taxonomists as part of the D. aciculare group and by others as part of the D. dichotomum group; its

POACEAE 873 

affinities appear to lie with the former. See Schuyler (1996) for a discussion of the taxonomic distinctiveness of this species. 

The occurrence of this species in NC is documented in LeBlond & Sorrie (2001). [= K; < P. aciculare Desvaux ex Poiret – C; < 

D. dichotomum ssp. roanokense (Ashe) Freckmann & Lelong – FNA] 

Dichanthelium latifolium (Linnaeus) Harvill, Broad-leaved Witch Grass. Mt (GA, NC, SC, VA): open or shady welldrained 

forests; rare (NC Watch List). Late May-September. ME south to n. GA, west to WI and MS. [= FNA; = Panicum 

latifolium Linnaeus – RAB, C, F, G, HC, S; = D. latifolium (Linnaeus) Gould & Clark – K, Z, a later combination] 

Dichanthelium laxiflorum (Lamarck) Gould, Open-flower Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): open or shaded 

woodlands, often in moist soil; common. April-September. MD south to FL, west to TX, north to IN, also in Mexico, Central 

America, and West Indies. [= FNA, K, Z; = Panicum laxiflorum Lamarck – RAB, C, F, G; > P. laxiflorum – HC, S; > P. 

xalapense Humboldt, Bonpland, & Kunth var. xalapense – HC, S; > P. xalapense var. strictirameum Hitchcock & Chase – HC, 

S] 

Dichanthelium leucothrix (Nash) Freckmann, Roughish Witch Grass. Cp, Pd (GA, NC, SC, VA): wet sandy, peaty, or 

mucky soil of pinelands; occasional in Coastal Plain, rare in Piedmont (VA Watch List). May-October. S. NJ south to FL, west 

to TX, also in TN, West Indies and n. South America. A micrometer is needed to measure the very short puberulence (0.1 mm) 

that distinguishes this taxon, D. meridionale, and D. wrightianum from other members of the D. acuminatum group. [= K, Y; = 

Panicum leucothrix Nash – RAB, F, G, HC, S; < P. leucothrix – C; = D. acuminatum ssp. leucothrix (Nash) Freckmann & 

Lelong – FNA; < P. spretum Schultes – GW; = P. acuminatum Swartz var. leucothrix (Nash) Lelong – X; < D. acuminatum 

(Swartz) Gould & Clark var. implicatum (Scribner) Gould & Clark – Z] 

Dichanthelium linearifolium (Scribner) Gould, Low White-haired Witch Grass. Pd, Mt (GA, NC, SC?, VA): dry open 

woods; rare (NC Watch List). May-October. Se. Canada and MN south to GA and TX. [= FNA, K, Z; = Panicum linearifolium 

Scribner – RAB, C, S; > P. linearifolium var. linearifolium – F, G; > P. linearifolium var. werneri (Scribner) Fernald – F, G; > 

P. linearifolium – HC; > P. werneri Scribner – HC] 

Dichanthelium longiligulatum (Nash) Freckmann, Long-ligule Witch Grass. Cp, Pd (GA, NC, SC, VA): limesink ponds, 

depression meadows, cypress savannas, pine savannas, bogs, swamps; frequent in Coastal Plain, rare in Piedmont. May- 

September. NJ and PA south to FL, also in TN, e. TX, and Central America. Intermediate forms between this taxon and D. 

spretum occur. [= K, Y; = Panicum longiligulatum Nash – RAB, HC, S; < P. lanuginosum Elliott var. lindheimeri (Nash) 

Fernald – C, G; = D. acuminatum ssp. longiligulatum (Nash) Freckmann & Lelong – FNA; < P. spretum Schultes – GW; = P. 

acuminatum Swartz var. longiligulatum (Nash) Lelong – X; = D. acuminatum (Swartz) Gould & Clark var. longiligulatum 

(Nash) Gould & Clark – Z] 

Dichanthelium lucidum (Ashe) LeBlond, Bog Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): wet meadows, sphagnous 

swamps, bogs, wet woods, sphagnous streamhead pocosins, baygalls; frequent (uncommon in Mountains). May-October. MA 

and MI south to FL and TX. Vernal culms soon recline, producing a tangled mass. The papillose fertile lemma is diagnostic. [= 

Q; < Panicum dichotomum Linnaeus – RAB, C, GW; = P. lucidum Ashe – G, S; > P. lucidum var. lucidum – F, HC; > P. 

lucidum var. opacum Fernald – F, HC; < D. dichotomum ssp. lucidum (Ashe) Freckmann & Lelong – FNA; < D. dichotomum 

var. dichotomum – K, Z; < P. dichotomum var. lucidum (Ashe) Lelong – X] 

Dichanthelium mattamuskeetense (Ashe) Mohlenbrock, Mattamuskeet Witch Grass. Cp (NC, SC, VA): wet savannas, 

meadows, borders of pocosin shrub swamps, thickets; occasional in Coastal Plain (VA Watch List). May-October. Se. MA 

south to ne. SC. Typically a robust plant, often richly tinged with dark purple-maroon. [= Q; < Panicum dichotomum Linnaeus 

– RAB, C, GW; > P. mattamuskeetense var. mattamuskeetense – F; > P. mattamuskeetense var. clutei (Nash) Fernald – F; < D. 

dichotomum (Linnaeus) Gould ssp. mattamuskeetense (Ashe) Freckmann & Lelong – FNA; > P. mattamuskeetense Ashe – G, 

HC, S; > P. annulum Ashe var. glabrescens Gleason – G; > P. clutei Nash – HC, S; < D. dichotomum var. dichotomum – K, Z; 

< P. dichotomum var. mattamuskeetense (Ashe) Lelong – X] 

Dichanthelium meridionale (Ashe) Freckmann, Matting Witch Grass. Pd, Cp, Mt (GA, NC, SC, VA): dry to damp sand of 

shores and woods; occasional. May-October. Sw. Nova Scotia and MA to MN, south to e. NC, n. GA, and n. AL. A micrometer 

is needed to measure the very short puberulence (0.1 mm) that distinguishes this taxon, D. leucothrix, and D. wrightianum from 

other members of the D. acuminatum group. [= K, Y; < Panicum lanuginosum Elliott – RAB; < P. leucothrix Nash – C; > P. 

meridionale var. meridionale – F; > P. meridionale var. albemarlense (Ashe) Fernald – F; = D. acuminatum (Swartz) Gould & 

Clark ssp. implicatum (Scribner ex Nash) Freckmann & Lelong – FNA; = P. meridionale Ashe – G; > P. meridionale – HC, S; > 

P. albemarlense Ashe – HC, S; < P. acuminatum Swartz var. unciphyllum (Trinius) Lelong – X; < D. acuminatum var. 

implicatum (Scribner) Gould & Clark – Z] 

Dichanthelium oligosanthes (Schultes) Gould var. oligosanthes, Few-flowered Witch Grass. Cp, Pd (GA, NC, SC, VA): 

sandy fields and open woods; occasional in Coastal Plain, rare in Piedmont (VA Watch List). April-October. MA and MN south 

to FL and TX. See note under D. fusiforme. [= K, Z; = Panicum oligosanthes Schultes – RAB, HC, S; < P. oligosanthes – C, G; 

= P oligosanthes var. oligosanthes – F; = D. oligosanthes ssp. oligosanthes – FNA] 

Dichanthelium oligosanthes (Schultes) Gould var. scribnerianum (Nash) Gould, Scribner's Witch Grass. Mt, Pd (GA, 

VA), Cp (NC, VA): calcareous maritime forests, dry thin woods and openings, dry prairies, usually in basic soil; uncommon in 

Mountains, rare in Piedmont and Coastal Plain (NC Watch List). April-November. Sw. ME to s. British Columbia, south to se. 

NC, n. GA, and CA, also in n. Mexico. Throughout the U.S., but infrequent in the southeastern and western states. [= K, Z; < 

Panicum oligosanthes Schultes – RAB, C, G; = P. oligosanthes var. scribnerianum (Nash) Fernald – F; = D. oligosanthes ssp. 

scribnerianum (Nash) Freckmann & Lelong – FNA; = P. scribnerianum Nash – HC, S] 

Dichanthelium ovale (Elliott) Gould & Clark var. addisonii (Nash) Gould & Clark, Low Stiff Witch Grass. Cp (GA, NC, 

SC, VA): dry to damp sandy woods and fields; uncommon (VA Watch List). May-October. MA and MN south to FL and TX, 

also in n. Mexico. See note under D. ovale var. ovale. [= K, Z; = Panicum commonsianum Ashe – RAB, C; > P. 

commonsianum var. commonsianum – F, G; > P. commonsianum var. addisonii (Nash) Fernald – F, G; > P. mundum Fernald –

POACEAE 874 

F, G, HC; > P. villosissimum var. pseudopubescens (Nash) Fernald – F, G; = D. ovale ssp. pseudopubescens (Nash) Freckmann 

& Lelong – FNA; > P. commonsianum – HC, S; P. pseudopubescens Nash – HC, S; P. addisonii Nash – HC, S; P. 

wilmingtonense Ashe – HC, S; = P. ovale Elliott var. pseudopubescens (Nash) Lelong – X] 

Dichanthelium ovale (Elliott) Gould & Clark var. ovale, Oval-flowered Witch Grass. Cp (GA, NC, SC, VA): dry to damp 

sandy pinelands; rare (NC Watch List, VA Rare List). May-October. NY to WI, south to FL and e. TX. Infrequent over most of 

its range except FL. In our region, the D. ovale double ligule character is more evident in var. ovale, with var. addisonii often 

having only a single ligule about 1 mm long. Also see note under D. consanguineum, and note at end of descriptions regarding 

Panicum malacon (placed in synonymy here). [= K, Z; = Panicum ovale Elliott – RAB, X; = D. ovale ssp. ovale – FNA; > P. 

ovale – HC, S; > P. malacon Nash – HC, S; = P. ovale var. ovale – X] 

Dichanthelium polyanthes (Schultes) Mohlenbrock, Small-fruited Witch Grass. Pd, Mt, Cp (GA, NC, SC, VA): damp to 

dry soil of open woods and ditches; common (absent from the outer Coastal Plain south of the Neuse River). June-October. VA 

to s. IL, south to GA and e. TX. [= FNA; = Panicum polyanthes Schultes – RAB, C, F, G, GW, HC, S; = D. sphaerocarpon 

(Elliott) Gould var. isophyllum (Scribner) Gould & Clark – K, Z] 

Dichanthelium portoricense (Desvaux ex Hamilton) B.F. Hansen & Wunderlin, Puerto Rican Witch Grass. Cp (GA, NC, 

SC): moist pine savannas and flatwoods; frequent in North Carolina, uncommon in South Carolina. May-September. NC south 

to FL, west to TX, also in West Indies. [= Panicum portoricense Desvaux ex Hamilton – RAB, HC, S; = D. portoricense ssp. 

portoricense – FNA; < D. sabulorum (Lamarck) Gould & Clark var. thinium (Hitchcock & Chase) Gould & Clark – K, Z; = P. 

portoricense var. portoricense – X] 

Dichanthelium ravenelii (Scribner & Merrill) Gould, Ravenel's Witch Grass. Cp, Pd (GA, NC, SC, VA): dry sandy or 

rocky thin woods and openings, sometimes in moist soils; common (VA Watch List). May-October. DE south to FL, west to e. 

TX, north to IA. [= FNA, K, Z; = Panicum ravenelii Scribner & Merrill – RAB, C, F, G, HC, S] 

Dichanthelium scabriusculum (Elliott) Gould & Clark, Tall Swamp Witch Grass. Cp, Pd (GA, NC, SC, VA): moist, low, 

open or shaded woodlands, often along streams or ditches; common in Coastal Plain, uncommon in Piedmont (VA Watch List). 

May-October. Se. MA south to FL, west to e. TX and AR. [< Panicum scabriusculum – RAB, C, GW; > P. scabriusculum – F, 

HC, S; > P. aculeatum Hitchcock & Chase – F, G, HC, S; < D. scabriusculum – FNA, K, Z; > P. scabriusculum var. 

scabriusculum – G] 

Dichanthelium scoparium (Lamarck) Gould, Velvet Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): moist sandy soil of 

woodland openings and ditches; common (uncommon in Mountains). May-October. MA and MI south to FL and TX, also in 

Mexico and West Indies. The dense, velvety pubescence of the internodes, sheaths, and blades of this taxon, combined with the 

viscid band below the nodes, are diagnostic. [= FNA, K, Z; = Panicum scoparium Lamarck – RAB, C, F, G, GW, HC, S] 

Dichanthelium species 2 (=webberianum), Webber's Witch Grass. Cp (GA, NC, SC): moist pine savannas and flatwoods; 

uncommon. May-August. Disjunct in se. NC and SC from GA and FL. [= Panicum webberianum Nash – RAB, HC, S; < D. 

portoricense (Desvaux ex Hamilton) B.F. Hansen & Wunderlin ssp. patulum (Scribner & Merrill) Freckmann & Lelong – FNA; 

< D. sabulorum (Lamarck) Gould & Clark var. patulum (Scribner & Merrill) Gould & Clark – K, Z; < P. portoricense Desvaux 

ex Hamilton var. nashianum (Scribner) Lelong – X] 

Dichanthelium species 3 (=lancearium), Nash's Witch Grass. Cp, Pd (GA, NC, SC, VA): moist pine savannas and 

flatwoods, moist to dry openings in maritime forests, dry pine and oak sandhills; common in Coastal Plain, uncommon in 

Piedmont (VA Watch List). May-September. Se. VA to FL, west to e. TX, also in West Indies and Central America. This and 

D. portoricense appear to intergrade in our region, and =lancearium is treated as part of D. portoricense ssp. patulum in FNA, 

whose concept of ssp. patulum includes Panicum webberianum (D. species 2 here) and P. patentifolium, both of which appear to 

merit recognition (=webberianum as a species and =patentifolium as at least a variety). [= Panicum lancearium Trinius – RAB, 

C, G; > P. lancearium var. lancearium – F; > P. lancearium var. patulum (Scribner & Merrill) Fernald – F; < D. portoricense 

(Desvaux ex Hamilton) B.F. Hansen & Wunderlin ssp. patulum (Scribner & Merrill) Freckmann & Lelong – FNA; > P. 

lancearium – HC, S; > P. patulum (Scribner & Merrill) Hitchcock – HC, S; < D. sabulorum (Lamarck) Gould & Clark var. 

patulum (Scribner & Merrill) Gould & Clark – K, Z; < P. portoricense Desvaux ex Hamilton var. nashianum (Scribner) Lelong – 

X] 

Dichanthelium species 5 (=neuranthum), Nerved Witch Grass. Cp (GA, NC): maritime wet grasslands and wet savannas 

near the coast; rare (GA Special Concern, NC Significantly Rare). May-September. Disjunctly in se. NC, se. SC, GA, FL, s. 

MS, TX, Bahamas, Cuba, and Belize. Treated as D. aciculare ssp. neuranthum in FNA, but distinctive morphology and habitat 

argue for recognition as a species. Can occur with the similar-appearing D. caerulescens, from which it differs by having 

spikelets that are longer (1.8-2.0 mm vs. 1.4-1.8), rounded vs. obtuse to sub-acute, and pubescent vs. glabrous; longer first 

glumes (0.8-1.0 mm vs. 0.3-0.8); leaves 15× or more as long as wide vs. 10-15×; and a nearly strict panicle. FNA gives a 

spikelet length of 2-2.8 mm, well beyond the length of spikelets on specimens we have seen throughout the range. As FNA 

provides no synonymy, it is possible that its concept of "neuranthum" includes entities treated separately here, or entities outside 

the range of this flora. [= Panicum neuranthum Grisebach – RAB, HC, S; = D. aciculare (Desvaux ex Poiret) Gould & Clark 

ssp. neuranthum (Grisebach) Freckmann & Lelong – FNA; < D. aciculare – K, Z] 

Dichanthelium species 9 (=cryptanthum), Hidden-flowered Witch Grass. Cp (NC, SC): wet meadows and ditches, 

streamside openings (NC Significantly Rare). May-September. NC (or NJ?) to MS (or TX?) (previous concepts of this taxon 

and its range are unclear). In the field, this taxon can be mistaken for D. yadkinense; it is readily distinguished by its scabrous 

peduncle and membranous ligules. [= Panicum cryptanthum Ashe – F, HC, S; < P. scabriusculum Elliott – RAB, C, GW; < D. 

scabriusculum (Elliott) Gould & Clark – FNA, K, Z; = P. scabriusculum var. cryptanthum (Ashe) Gleason – G] 

Dichanthelium species 10 (=curtifolium), Short-leaved Witch Grass. Cp (SC), Mt (NC): bogs, sphagnous streamhead 

swamps, mountain streams; uncommon in Coastal Plain, rare in Piedmont and Mountains. April-September. Ranging disjunctly 

in w. NC and e. TN, e. SC, FL, and MS. The combination of characters is quite distinctive for the genus in our region. [=

POACEAE 875 

Panicum curtifolium Nash – RAB, HC, S; = D. ensifolium (Baldwin ex Elliott) Gould ssp. curtifolium (Nash) Freckmann & 

Lelong – FNA; < D. acuminatum (Swartz) Gould & C.A. Clark var. implicatum (Scribn.) Gould & C.A. Clark – K, Z; = 

Panicum ensifolium Baldwin ex Elliott var. curtifolium (Nash) Lelong – X] 

Dichanthelium sphaerocarpon (Elliott) Gould, Round-fruited Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): moist or dry 

thin woods, meadows, and ditches, often in dry sandy soil; common. May-October. MA, VT, OH, and KA south to FL and TX, 

also in Mexico. [= FNA, K, Z; = Panicum sphaerocarpon Elliott – RAB, C; > P. sphaerocarpon var. sphaerocarpon – F, G, 

HC, S; > P. sphaerocarpon var. inflatum (Scribner & J.G. Smith) Hitchcock & Chase – F, G, HC, S] 

Dichanthelium sphagnicola (Nash) LeBlond, Peaty Witchgrass. Cp (GA): edges of cypress swamps, in sphagnous bogs, 

moist shady places; poorly known and possibly very rare. May-October. GA (Chatham County) to FL; should be sought in se. 

SC. Treated in synonymy with Panicum dichotomum by RAB, but no specimen is known from the Carolinas. This species is 

similar to D. lucidum in appearance, and differs most readily by its larger pubescent spikelets with smooth fertile lemma and 

palea. [= Q; < Panicum dichotomum Linnaeus – RAB; < D. dichotomum (Linnaeus) Gould ssp. lucidum (Ashe) Freckmann & 

Lelong – FNA; = P. sphagnicola Nash – HC, S; < D. dichotomum var. dichotomum – K, Z; < P. dichotomum var. lucidum 

(Ashe) Lelong – X] 

Dichanthelium spretum (Schultes) Freckmann, Eaton's Witch Grass. Pd, Cp, Mt (GA, NC, SC, VA): wet sands and peats 

of bogs, savannas, meadows, and shores; rare (NC Watch List, VA Watch List). May-September. ME south to n. FL, LA and e. 

TX. Intermediate forms between this taxon and D. longiligulatum occur. [= K, Y; = Panicum spretum Schultes – RAB, C, F, G, 

HC, S; = D. acuminatum (Swartz) Gould & Clark ssp. spretum (Schultes) Freckmann & Lelong – FNA; < P. spretum – GW; = 

P. acuminatum Swartz var. densiflorum (Rand & Redfield) Lelong – X; = D. acuminatum var. densiflorum (Rand & Redfield) 

Gould & Clark – Z] 

Dichanthelium strigosum (Muhlenberg) Freckmann var. glabrescens (Grisebach) Freckmann, Hairless Witch Grass. Cp 

(GA): low, open sandy pinelands and hammocks. May-October. S. GA and FL west to LA; also in West Indies, Belize. 

Included in synonymy with Panicum strigosum by RAB, but no specimen from the Carolinas has been found. [= K; = D. 

strigosum ssp. glabrescens (Grisebach) Freckmann & Lelong – FNA; < Panicum strigosum Muhlenberg – GW; = P. polycaulon 

Nash – HC, S; = D. leucoblepharis (Trinius) Gould & Clark var. glabrescens (Grisebach) Gould & Clark – Z] 

Dichanthelium strigosum (Muhlenberg) Freckmann var. leucoblepharis (Trinius) Freckmann, Dwarf Witch Grass. Cp, Mt 

(GA, NC, SC): sandy, acidic soils of pinelands; occasional in Coastal Plain, rare in Mountains. May-October. NC south to FL, 

west to TX, also in Mexico. [= K; = Panicum ciliatum Elliott – RAB, HC, S; = D. strigosum ssp. leucoblepharis (Trinius) 

Freckmann & Lelong – FNA; = P. strigosum Muhlenberg var. leucoblepharis (Trinius) Lelong – X; = D. leucoblepharis 

(Trinius) Gould & Clark var. leucoblepharis – Z] 

Dichanthelium strigosum (Muhlenberg) Freckmann var. strigosum, Rough-hairy Witch Grass. Cp (GA, NC, SC, VA): in 

moist soils of pine flatwoods, savannas, and pocosins, also in boggy situations; uncommon (VA Rare List). May-September. Se. 

VA south to FL, west to TX, also in TN, e. Mexico, Mesoamerica, n. South America, and West Indies. [= K; = Panicum 

strigosum Muhlenberg – RAB, C, F, G, GW, HC, S; = D. strigosum ssp. strigosum – FNA; = P. strigosum var. strigosum – X; = 

D. leucoblepharis (Trinius) Gould & Clark var. pubescens (Vasey) Gould & Clark – Z] 

Dichanthelium tenue (Muhlenberg) Freckmann & Lelong, White-edged Witch Grass. Cp, Pd, Mt (GA, NC, SC, VA): wet 

peaty or sandy soil pineland savannas, flatwoods, bogs, and meadows; common in Coastal Plain, occasional in Piedmont, rare in 

Mountains. May-October. NJ south to FL, west to TX, also in Mesoamerica and Cuba. This treatment of D. tenue includes 

plants from northern Alabama formerly recognized as Panicum concinnius, with spikelets 1.2-1.4 mm long but otherwise 

possessing the characters of D. tenue. [= FNA; = Panicum tenue Muhlenberg – RAB, C; > P. tenue – F, HC, S; > P. 

albomarginatum Nash – F, HC, S; > P. trifolium Nash – F, G, HC, S; < P. ensifolium Baldwin – G; > P. concinnius Hitchcock & 

Chase – HC, S; < D. dichotomum (Linnaeus) Gould var. tenue (Muhlenberg) Gould & Clark – K, Z] 

Dichanthelium villosissimum (Nash) Freckmann var. villosissimum, White-haired Witch Grass. Cp, Pd, Mt (GA, NC, SC, 

VA): dry sandy soil of open woods and prairies; frequent (VA Watch List). April-September. MA south to FL, west to TX, also 

in Mexico and Mesoamerica. Appearing to be related to D. ovale based on such characters as the double ligule. [= K, Y; = 

Panicum villosissimum Nash – RAB, C, HC, S; = P. villosissimum var. villosissimum – F, G; = D. ovale (Elliott) Gould & Clark 

ssp. villosissimum (Nash) Freckmann & Lelong – FNA; = P. ovale Elliott var. villosum (A. Gray) Lelong – X; < D. acuminatum 

(Swartz) Gould & Clark var. villosum (A. Gray) Gould & Clark – Z] 

Dichanthelium wrightianum (Scribner) Freckmann, Wright's Witch Grass. Cp (GA, NC, SC, VA): limesink ponds and 

meadows, cypress savannas, pine savannas, bogs; uncommon (VA Watch List). May-September. MA south to FL, west to TX, 

also in Cuba and Mesoamerica. A micrometer is needed to measure the very short puberulence (0.1 mm) that distinguishes this 

taxon, D. meridionale, and D. leucothrix from other members of the D. acuminatum group. [= FNA, K, Y; = Panicum 

wrightianum Scribner – RAB, C, F, G, HC, S; < P. spretum Schultes – GW; = D. acuminatum (Swartz) Gould & Clark var. 

wrightianum (Scribner) Gould & Clark – Z] 

Dichanthelium yadkinense (Ashe) Mohlenbrock, Spotted-sheath Witch Grass. Pd, Cp, Mt (GA, NC, SC, VA): floodplain 

forests, thickets, bottomlands, and swamps, often on alluvial deposits; frequent in Piedmont, occasional in Coastal Plain, 

uncommon in Mountains. May-October. NJ and MI south to GA and TX, also in Mexico. Sheaths often with wart-like glands. 

This taxon resembles D. species 9 (=cryptanthum), from which it differs most readily by its hairy ligule (vs. membranous) and 

smooth peduncle (vs. antrorsely scabrous). [= Q; < Panicum dichotomum Linnaeus – RAB, GW; = P. yadkinense Ashe – C, F, 

G, HC, S; = D. dichotomum ssp. yadkinense (Ashe) Freckmann & Lelong – FNA; < D. dichotomum var. dichotomum – K, Z; = 

P. dichotomum var. yadkinense (Ashe) Lelong – X] 

Dichanthelium chamaelonche (Trinius) Freckmann & Lelong ssp. breve (Hitchcock & Chase) Freckmann & Lelong, Short 

Witch Grass, endemic to c. and s. FL, primarily near the east coast. [= FNA; = Panicum breve Hitchcock & Chase – HC, S; = D.

POACEAE 876 

dichotomum (Linnaeus) Gould var. breve (Hitchcock & Chase) Gould & Clark – K, Z; = P. chamaelonche Trinius var. breve 

(Hitchcock & Chase) Lelong – X] 

Dichanthelium dichotomum (Linnaeus) Gould var. glabrifolium (Nash) Gould & Clark, Smooth-leaved Witch Grass, 

endemic to peninsular FL, mostly near the west coast. Like D. chamaelonche ssp. breve, this taxon appears to be more closely 

related to D. chamaelonche than to D. dichotomum or D. ensifolium. [= K, Z; = Panicum glabrifolium Nash – HC, S; < P. 

chamaelonche Trinius var. chamaelonche – X] 

Dichanthelium leibergii (Vasey) Freckmann, Leiberg's Witch Grass, NY and PA west to Alberta, ND, and KS. [= FNA, K, 

Z; = Panicum leibergii (Vasey) Scribner – C, F, G, HC] 

Dichanthelium malacophyllum (Nash) Gould, Soft-leaf Witch Grass, KY and TN west to KS and TX. Primarily a plant of 

cedar glades and dry calcareous soils. Reported from SC by FNA, but source of record has not been identified. [= FNA, K, Z; = 

Panicum malacophyllum – F, G, HC, S] 

Dichanthelium nudicaule (Vasey) B.F. Hansen & Wunderlin, ranges from the FL Panhandle and s. AL west to MS. [= Q; = 

Panicum nudicaule Vasey] 

Dichanthelium wilcoxianum (Vasey) Freckmann is shown as occurring in SC and MS on the range map in FNA, but the 

source of these records is not known for this plant primarily of dry prairies in the Upper Midwest. It is not treated here. 

Dichanthelium xanthophysum (A. Gray) Freckmann, Slender Witch Grass, ME south to PA, west to SD; Nova Scotia to 

Saskatchewan. [= FNA, K, Z; = Panicum xanthophysum A. Gray – C, F, G, HC] 

Panicum chrysopsidifolium Nash is treated variously by the cited sources. According to Z, who examined an isotype 

collection and found the ligule to be 2.5 mm long, it belongs to D. acuminatum var. acuminatum. Plants referred to by HC all 

have ligules < 1 mm long, and apparently belong to the D. aciculare complex. These plants are described as having densely 

villous nodes, internodes, sheaths, and blades; the blades 5-10 cm long and 3-5 mm wide; and spikelets 1.9-2.2 mm long, 

obovate, and villous. HC gives a range of se. VA to FL, west to TX, along the Coastal Plain in sandy oak or pine woods. This 

entity needs further scrutiny. 

Panicum glutinoscabrum Fernald is an entity known only locally from "boggy spots" in southeastern Virginia. It is treated 

as a full species by F, but is placed in synonymy with Dichanthelium scoparium (Lamarck) Gould by FNA; with D. acuminatum 

(Swartz) Gould & Clark var. acuminatum by Z; with P. lanuginosum Elliott var. fasciculatum (Torrey) Fernald by C; and with P. 

huachucae Ashe var. fasciculatum (Torrey) Hubb. by HC. Panicum glutinoscabrum is described as having culms 7-9 dm high; 

elongate internodes with cinereous puberulence and black, warty, viscid glands; villous nodes; glutinous-warty and scabrous 

sheaths and blades; ligule 4-5 mm long; minutely puberulent panicle axis; spikelets ellipsoid, subacute, 1.7-1.8 mm long, 

pubescent; first glume subacute, 0.6-0.7 mm long. Y concedes that "I am not able to render a decision on the poorly known P. 

glutinoscabrum Fernald, but I suggest that it is a hybrid between a member of the D. acuminatum complex and D. scoparium 

(Lam.) Gould - the latter contributing the genes for height and viscid sheaths." 

Panicum malacon Nash, here placed in synonymy with Dichanthelium ovale var. ovale, needs additional study. It is 

distinguished by HC and S as having spikelets 3-3.2 mm long with a first glume situated conspicuously below the second glume 

and sterile lemma, half or more as long as the spikelet; and leaves 3-5 mm wide, puberulent beneath, and puberulent to glabrous 

above. The leaf width and puberulence characters are not consistent with descriptions of D. ovale, and the placement of P. 

malacon within D. ovale by current treatments (including this one) may be in error. The description of P. malacon in RAB 

differs greatly from that of HC and S, and falls well outside the range of D. ovale characters. In RAB, P. malacon is described as 

having spikelets 3.5-4 mm long, and leaves 5-14 mm wide. This description may be based on specimens from the Sandhills of 

SC at NCU identified as P. malacon and matching the RAB description. These specimens appear to be misidentified collections 

of D. oligosanthes var. oligosanthes. 

Digitaria Haller (Crab Grass) 

A genus of about 200 species, primarily in the tropics and subtropics. Most of our species occur primarily in disturbed situations; 

their original distributions and habitats are now obscure. References: Wipff in FNA (2003a); Webster (1987)=Z; Wipff & Hatch 

(1994)=Y; Wipff (1996b)=X; Webster (1980). 

1 Inflorescence an open panicle; spikelets long-pedicellate, borne singly at the ends of long panicle branches; [section 

Pennatae] ............................................................................................................................................ D. cognata var. cognata 

1 Inflorescence of 2-several spikelike racemes borne digitately or in close proximity near the summit of the culm; spikelets 

sessile or short-pedicellate, borne more-or-less closely spaced along the racemes. 

2 Rachis of each raceme narrow, trigonous, only slightly (if at all) winged. 

3 Spikelets 1.7-2.2 mm long; plants 3-10 dm tall; racemes to 10 cm long; upper sheaths glabrous, lower sheaths 

glabrous to sparsely pilose ............................................................................................... D. filiformis var. filiformis 

3 Spikelets 2.0-2.8 mm long; plants 8-15 dm tall; racemes to 25 cm long; upper sheaths glabrous or pilose, lower 

sheaths densely pilose ..................................................................................................................................D. villosa 

2 Rachis of each raceme broad (0.5-1 mm wide), winged, the wings as wide as or wider than the rachis proper. 

4 Lower sheaths glabrous; second glume 0.75-1× as long as the first glume; fertile lemma dark brown or black at 

maturity. 

5 Hairs of the spikelet not minutely capitate; second glume ca. 0.75 as long as the first glume; spikelets 

averaging 1.5 mm long...................................................................................................................D. violascens

POACEAE 877 

5 Hairs of the spikelet minutely capitate; second glume ca. 1× as long as the first glume; spikelets averaging 2.0 

mm long .......................................................................................................................................D. ischaemum 

4 Lower sheaths pilose; second glume 1/3-3/5 (to 4/5 in D. ciliaris) as long as the first glume; fertile lemma white, 

tan, or grayish-brown at maturity. 

6 Spikelets 1.5-1.8 mm long, villous with crinkled hairs; pedicels glabrous, terete in cross-section ..................... 

.......................................................................................................................................................... D. serotina 

6 Spikelets (1.7-) 2.4-4.1 mm long, glabrous, scabrous, or pubescent with straight hairs; pedicels scabrous, 3- 

angled in cross-section; [section Digitaria]. 

7 Spikelets (1.7-) 2.5-3.4 mm long, averaging 3.0 mm long or shorter; leaf blades pilose over the upper 

surface ..................................................................................................................................D. sanguinalis 

7 Spikelets 2.6-4.1 mm long, averaging 3.1 mm long or longer; leaf blades glabrous except for a few hairs 

on the upper surface at the base. 

8 Lower lemma of the sessile spikelet with 5 equidistant nerves; lowermost inflorescence node 

glabrous or pubescent with hairs < 0.4 mm long; apex of the first glume rounded to truncate ........... 

........................................................................................................................................... D. bicornis 

8 Lower lemma of sessile spikelet with the lateral nerves crowded to the margins; lowermost 

inflorescence node pubescent with hairs > 0.4 mm long; apex of the first glume acute ...................... 

............................................................................................................................................. D. ciliaris 

Digitaria bicornis (Lamarck) Roemer & J.A. Schultes. Cp (GA, NC, SC): sandy fields, lawns, roadsides, disturbed places; 

common. Webster (1980) believed that this species is likely to occur in VA and MD, as well. Whether or not it is introduced is 

unclear; it is now widely distributed in the tropics and subtropics worldwide. [= FNA, K, Z] 

Digitaria ciliaris (Retzius) Köler, Southern Crab Grass. Cp (GA, NC, SC, VA), Pd, Mt (GA, NC, SC): sandy fields, 

roadsides, and disturbed areas; common? August-October. [= C, FNA, K, Z; = D. sanguinalis var. ciliaris (Retzius) Parlatore – 

F, HC] 

Digitaria cognata (J.A. Schultes) Pilger, Fall Witch Grass. Cp, Pd (GA, NC, SC, VA): sandy fields and roadsides; 

common (VA Rare). July-October. Wipff & Hatch (1994) discuss the reasons for including Leptoloma in Digitaria. [= FNA; = 

Digitaria cognata var. cognata – K; = Leptoloma cognatum (J.A. Schultes) Chase – RAB, C, F, G, HC, S; = D. cognatum ssp. 

cognatum – Y] 

Digitaria filiformis (Linnaeus) Köler var. filiformis. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disturbed areas; 

common. September-October. Var. filiformis, with pubescent spikelets, is widespread in e. North America. Var. laeviglumis 

(Fernald) J. Wipff, with glabrous spikelets, occurs in New England. Var. dolichophylla (Henrard) J. Wipff occurs in s. FL, Cuba, 

and PR. See Wipff (1996) for additional discusion. [= RAB, C, F, FNA, G, X; = D. filiformis – HC, K; = Syntherisma filiformis 

(Linnaeus) Nash – S; < D. filiformis – W] 

* Digitaria ischaemum (Schreber) Muhlenberg, Smooth Crab Grass. Cp, Pd, Mt (GA, NC, SC, VA): fields, lawns, disturbed 

areas; common, introduced from Eurasia. July-October. Two varieties have sometimes been recognized. Var. ischaemum has 

racemes (1-) 2-6, 1-9 (-10) cm long, mostly curved and plants mostly to 4 dm tall. Var. mississippiensis (Gattinger) Fernald has 

racemes 5-7, 6-15 cm long, mostly stiff and straight and plants to 10 dm tall. [= C, FNA, K, W; > D. ischaemum var. ischaemum 

– F, G, HC; > D. ischaemum (Schreber) Muhlenberg var. mississippiensis (Gattinger) Fernald – F, G, HC; = D. ischaemum var. 

ischaemum – RAB; = Syntherisma ischaemum (Schreber) Nash – S] 

* Digitaria sanguinalis (Linnaeus) Scopoli, Northern Crab Grass. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, 

disturbed areas; common. July-October. [= RAB, FNA, C, G, K, W, Z; = D. sanguinalis var. sanguinalis – F, HC; Syntherisma 

sanguinalis (Linnaeus) Dulac – S] 

Digitaria serotina (Walter) Michaux, Dwarf Crab Grass. Cp (GA, SC, VA): sandy woodlands; rare (VA Rare). October. 

[= RAB, C, F, FNA, G, GW, HC, K; = Syntherisma serotina Walter – S] 

Digitaria villosa (Walter) Persoon. Cp (GA, NC, SC, VA), Pd, Mt (GA, NC, SC): sandy fields, roadsides; common (VA 

Watch List). September-October. [= HC, K; = D. filiformis var. villosa (Walter) Fernald – RAB, C, F, FNA, G, X; = 

Syntherisma villosa Walter – S] 

* Digitaria violascens Link. Cp (GA, NC, SC), Pd (GA, SC), Mt (GA): sandy fields, roadsides, and woodland borders; 

common. September-October. [= C, FNA, G, HC, K; = D. ischaemum var. violascens (Link) Radford – RAB; ? Syntherisma 

floridana (A.S. Hitchcock) A.S. Hitchcock – S] 

* Digitaria horizontalis Willdenow, Jamaican Crabgrass, is reported for SC on the basis of a specimen at NCU (Kartesz 

1999). {check specimen} [= FNA, K] {not keyed at this time} 

Digitaria insularis (Linnaeus) Mez ex Ekman. AL, FL, MS. [= K] {not keyed at this time} 

References: Barkworth in FNA (2003a). 

Dinebra Jacquin (Viper Grass) 

* Dinebra retroflexa (Vahl) Panzer, Viper Grass, native of Africa and s. Asia, has been collected as a waif in Mecklenburg 

County, NC (Mellichamp, Matthews, & Smithka 1987). [= FNA, K] {not keyed}

POACEAE 878 

Distichlis Rafinesque (Saltgrass) 

A genus of about 5 species, of North, Central, and South America, and Australia. References: Barkworth in FNA (2003a). 

Identification notes: When sterile, Distichlis spicata is easily confused with Sporobolus virginicus, with which it sometimes 

occurs. Distichlis spicata is generally a coarser plant, and lacks long hairs around the collar of the sheath; Sporobolus virginicus 

is more delicate, and typically has long hairs on either side of the collar. 

Distichlis spicata (Linnaeus) Greene, Saltgrass, Spike Grass. Cp (GA, NC, SC, VA): coastal marshes and shores, 

especially common in hypersaline flats (where infrequent tidal inundation is followed by evaporation); common. June-October. 

Two varieties (or subspecies or species) have often been recognized: var. spicata ranging along the Atlantic coast from Nova 

Scotia and Prince Edward Island south to tropical America, and on the Pacific coast of North America, and var. stricta (Torrey) 

Scribner widespread in saline situations in western North America. These do not appear to warrant taxonomic recognition 

(Barkworth in FNA 2003a). [= RAB, FNA, GW, K, S; > D. spicata var. spicata – C; > D. spicata – F, G, HC; > D. spicata ssp. 

spicata] 

Echinochloa Palisot de Beauvois (Barnyard-grass, Jungle-rice) 

A genus of 4-5- species of the tropics and warm temperate regions. References: Michael in FNA (2003a). Key based in part on 

C. 

1 Panicle elongate, the branches few, distant, unbranched, and short, to 2 (-3) cm long; spikelets awnless; leaves 3-6 (-9) mm 

wide ......................................................................................................................................................................... E. colonum 

1 Panicle broader, the branches numerous, approximate, often further branched, short to long, some (at least) exceeding 2 cm 

long; spikelets awnless or awned; leaves 5-30 mm wide. 

2 Lower sheaths usually papillate-pubescent; fertile lemma 2.5-4× as long as wide ..............................................E. walteri 

2 Lower sheaths glabrous; fertile lemma 1.5-2.5× as long as wide. 

3 Inflorescence nodding; awns 4-29 mm long............................................................ E. cruspavonis var. cruspavonis 

3 Inflorescence erect, stiff; awns 0-25 mm long. 

4 Second glume and sterile lemma hairy or scabrous to nearly glabrous, the hairs usually not papillose-based; 

fertile lemma obtuse or broadly acute, with a thin, membranous (later withering) tip set off from the body by 

a line of minute hairs. 

5 Panicle fairly open, the branches erect, appressed, or spreading; spikelets green or purple-tinged, 

awnless or with a well-developed awn (to 25 mm long); leaves 5-15 mm wide; plants mostly 3-7 dm tall 

...........................................................................................................................E. crusgalli var. crusgalli 

5 Panicle very crowded, the branches appressed to slightly spreading, the tips often incurved; spikelets 

purplish-brown, awnless (or with awn to 2 mm long); leaves mostly 15-30 mm wide; plants mostly 7-15 

dm tall ................................................................................................................................ E. frumentacea 

4 Second glume and sterile lemma usually with stout, papillose-based hairs on the veins; fertile lemma 

acuminate, abruptly narrowed to a firm, persistent tip. 

6 Spikelets < 3.5 mm long, not including the awn (if present); sterile lemma awnless or with an awn to 6 (- 

10) mm long ............................................................................................... E. muricata var. microstachya 

6 Spikelets > 3.5 mm long, not including the awn (if present); sterile lemma usually awned (rarely 

awnless), the awn 6-25 mm long....................................................................... E. muricata var. muricata 

* Echinochloa colonum (Linnaeus) Link, Jungle-rice. Cp, Pd (GA, NC, SC, VA), Mt (GA, SC, NC): fields, ditches, 

disturbed wet areas; uncommon, introduced from the Old World tropics. July-October. The debate over the appropriate 

grammatical treatment and therefore spelling of the epithet is discussed in detail in Ward (2005b). [= E. colonum – RAB, C, F, 

G, GW, HC; = E. colona -- FNA, K, S, orthographic variant] 

* Echinochloa crusgalli (Linnaeus) Palisot de Beauvois var. crusgalli, Barnyard-grass. {GA, NC, SC, VA} July-October. 

[= C, G; < E. crusgalli – RAB, GW (also see E. muricatum); = E. crus-galli – K, orthographic variant; < E. crusgalli – F, FNA; 

< E. crus-galli ssp. crus-galli – S (also see E. muricata)] 

Echinochloa cruspavonis (Kunth) J.A. Schultes var. cruspavonis. {AL, MS, FL} July-October. [< E. crus-pavonis – HC; 

= E. crus-pavonis var. crus-pavonis – FNA, K, orthographic variant] 

* Echinochloa frumentacea Link, Japanese Millet, Billion-dollar Grass, White Panic. Cp (NC), {VA}: disturbed areas; rare, 

introduced from Asia. July-October. [= F, FNA, K; < E. crusgalli – RAB, GW; = E. crusgalli (Linnaeus) Palisot de Beauvois 

var. frumentacea (Link) W. Wight – C, G; = E. crus-galli ssp. edulis A.S. Hitchcock – S] 

Echinochloa muricata (Palisot de Beauvois) Fernald var. microstachya Wiegand, Barnyard-grass. {GA, NC, SC, VA} 

July-October. [= C, FNA, K; < E. crusgalli – RAB, GW; = E. pungens (Poiret) Rydberg var. microstachya (Wiegand) Fernald 

& Griscom – F; = E. microstachya (Wiegand) Rydberg – G; < E. crus-galli ssp. crus-galli – S]

POACEAE 879 

Echinochloa muricata (Palisot de Beauvois) Fernald var. muricata, Barnyard-grass. Cp (NC): interdune wetlands. July- 

October. [= C, FNA, K; < E. crusgalli – RAB, GW; > E. pungens (Poiret) Rydberg var. pungens – F; > E. pungens var. 

ludoviciana (Wiegand) Fernald & Griscom – F; = E. muricata – G; < E. crus-galli ssp. crus-galli – S] 

Echinochloa walteri (Pursh) Heller. Cp (GA, NC, SC, VA), Pd (GA): marshes; common. July-October. MA south to FL, 

west to TX on the outer Coastal Plain; also inland from OH west to WI, south to MO and AR. [= RAB, C, F, FNA, GW, HC, K, 

S, W] 

References: Hilu in FNA (2003a). Key based on FNA. 

Eleusine Gaertner (Yard Grass) 

1 Ligule pubescent................................................................................................................................E. coracana ssp. africana 

1 Ligule membranaceous, ciliate to erose ........................................................................................................................E. indica 

* Eleusine coracana (Linnaeus) Gaertner ssp. africana (Kennedy & O'Byrne) Hilu & de Wet, Finger Millet. Cp (SC): 

disturbed areas; rare, introduced from Africa. There remains some doubt about the identity of the population discovered. 

Reported by Werth, Zeng, & Baird (1997). [= FNA, K; = E. africana Kennedy & O'Byrne] 

* Eleusine indica (Linnaeus) Gaertner, Yard Grass, Goose Grass. Cp, Pd, Mt (GA, NC, SC, VA): lawns, roadsides, gardens, 

disturbed areas; common, introduced from Old World. [= RAB, C, F, FNA, G, HC, K, S, W] 

* Eleusine tristachya (Lamarck) Lamarck, is introduced from South America in scattered states in e. United States, including 

VA, NJ (Hilu 1980) and AL (Small 1933). [= FNA, K, S] {not keyed at this time} 

Elionurus Humboldt & Bonpland ex Willdenow (Balsamscale) 

A genus of about 15 species, native to tropical and subtropical parts of Africa and the Americas. References: Barkworth in FNA 

(2003a). 

Elionurus tripsacoides Humboldt & Bonpland ex Willdenow, Pan-American Balsamscale. Cp (GA): wet savannas; rare 

(GA Special Concern). S. GA south to s. FL, west to s. and w. TX, and south through Central America to s. South America. 

Reported for sw. GA by Jones & Coile (1988), for s. MS and FL (Sorrie & Leonard 1999). [= FNA, K; = Elyonurus tripsacoides 

– GW, HC, S, orthographic variant] 

Elymus Linnaeus 1753 (Wild-rye, Rye Grass) 

(also see Thinopyrum) 

A genus of about 150 species, semicosmopolitan in temperate regions. The genus, as now circumscribed, includes all 

allopolyploid taxa with at least one chromosome complement contributed from Pseudoroegneria. North American Elymus are 

allopolyploids of Pseudoroegneria and Hordeum (Helfgott & Mason-Gamer 2004). Reference: Barkworth & Campbell in FNA 

(in prep.); Campbell (2000); Church (1967); Tucker (1996)=Z; Barkworth (1997)=X. This treatment largely follows Barkworth 

& Campbell in FNA (in prep.). 

Identification notes: Measurements of the spike include the awns, but measurements of spikelets and its components do not. 

Rachis internodes should be measured near the middle of the spike. Glume widths are measured at the widest point, or if the 

widest point is not apparent, at about 5 mm above the glume base. 

1 Spikelets solitary at each node (occasionally paired at the lowest nodes); glumes and lemmas awned or unawned; plants 

cespitose to strongly rhizomatous. 

2 Plants strongly rhizomatous; [common and weedy introduced species]; [section Elytrigia] ...............................E. repens 

2 Plants cespitose; [rare natives and introductions]; [section Goulardia]. 

3 Spikelets 20-30 mm long; anthers 3-6 mm long; rachis internodes hirtellous below the spikelets; [very rare 

introduction, reported for c. GA]......................................................................................................[E. semicostatus] 

3 Spikelets 8-25 mm long; anthers 0.8-3 mm long; rachis internodes glabrous below the spikelets; [rare natives of 

glades and barrens]. 

4 Lemma awns 15-40 mm long, longer than the body of the lemma............. [E. trachycaulus ssp. subsecundus] 

4 Lemma awns 1-13 mm long, shorter than the body of the lemma.................E. trachycaulus ssp. trachycaulus 

1 Spikelets 2-3 (-5) at each node; glumes and lemmas usually awned; plants usually cespitose, occasionally short-rhizomatous. 

5 Both glumes (including their awn) either 0-3 mm long and subulate or 1-20 mm long and differing in length by > 5 

mm, 0.1-0.6 mm wide, tapering from the base, with 0-1 distinct veins, persistent; rachis internodes 4-12 mm long, ca. 

0.5 mm thick at the narrowest section.

POACEAE 880 

6 Spikelets appressed; lemma awns straight or curving; glumes sometimes absent, but usually 1-20 mm long, 0.1-0.6 

mm wide, with a distinct vein; spikes erect or nodding.........................................................................[E. svensonii] 

6 Spikelets widely spreading to horizontal; lemma awns straight (rarely slightly curving); glumes 0-3 mm long, with 

no distinct veins (rarely 1 glume to 20 mm long, 0.2 mm wide); spikes usually erect. 

7 Lemmas pubescent..................................................................................................E. hystrix var. bigelovianus 

7 Lemmas glabrous to scabrous .......................................................................................... E. hystrix var. hystrix 

5 Both glumes (including the awns) 10-40 mm long, usually differing in length by < 5 mm, 0.2-2.3 mm wide, lanceolate 

to setaceous, usually widest above the base, with 2-8 veins, persistent or disarticulating; rachis internodes slender (as 

above) or stout (2-5 mm long and ca. 1 mm thick at the narrowest section). 

8 Glume bases flat, thin, and evidently veined, or indurate for < 1 mm, the bodies not exceeding the adjacent 

(usually 8-15 mm long) lemmas; lemma awns usually curving outward; spikes usually nodding to pendent; 

internodes (2-) 4-12 mm long. 

9 Glumes 0.5-1.6 mm wide; lemma awns 15-40 (-50) mm long; paleas acute; rachis internodes 2-5 (-7) mm 

long; blades (3-) 4-15 (-20) mm wide, pale green, usually glabrous or scabridulous above................................ 

............................................................................................................................E. canadensis var. canadensis 

9 Glumes 0.3-0.8 mm wide; lemma awns 15-25 (-35) mm long; paleas narrowly truncate; rachis internodes 5-8 

(-12) mm long; blades 8-24 mm wide, dark green, usually thinly pilose above........................... [E. wiegandii] 

8 Glume bases terete, indurate, and lacking evident veins for 0.5-4 mm, the bodies (unless indistinct from the awns) 

exceeding the adjacent (usually 6-12 mm long) lemmas; lemma awns straight; spikes erect or nodding; internodes 

2-5 mm long (to 7 mm in E. sp. 1). 

10 Glumes persistent, 0.2-1 mm wide, with 2-4 veins, the basal 0.5-2 mm essentially straight; lemmas rarely 

glabrous; spikelets with 1-3 (-4) florets; spikes nodding, exserted. 

11 Blades glabrous to scabrous, pale dull green; spikes 7-25 cm long; internodes usually 3-5 mm long; 

spikelets with 2-3 (-4) florets; lemmas usually scabrous, 7-14 mm long, 1-5 mm longer than the acute 

paleas; flowering usually late June to late July...........................................................................E. riparius 

11 Blades villous to pilose, dark glossy green; spikes 4-12 cm long; internodes usually 2-3 mm long; 

spikelets with 1-2 (-3) florets; lemmas usually villous, 5.5-9 mm long, 0-1.5 mm longer than the obtuse 

paleas; flowering usually early June to early July ...................................................................... E. villosus 

10 Glumes disarticulating with the lowest floret, 0.7-2.3 mm wide, with (2-) 3-5 (-8) veins, the basal 1-4 mm 

clearly bowed-out; lemmas often glabrous; spikelets with (2-) 3-5 (-6) florets; [Elymus virginicus complex]. 

12 Spikes 2.5-6 cm wide, exserted; lemma awns 15-40 mm long; blades glabrous or villous. 

13 Spikes with 9-18 nodes; internodes 4-7 mm long; blades usually lax, dark glossy green under the 

glaucous bloom; auricles 2-3 mm long, blackish at maturity; flowering usually in mid-May to mid- 

June .............................................................................................................................E. macgregorii 

13 Spikes with 15-30 nodes; internodes 3-5 mm long; blades lax, or often ascending and involute, pale 

dull green; auricles 0-2 mm long, brownish at maturity; flowering usually in mid-June to late July. 

14 Spikelets (and usually also the foliage) pubescent; spikes usually 6-12 cm long; lemmas 6-10 

mm long......................................................................................... E. glabriflorus var. australis 

14 Spikelets (and usually also the foliage) glabrous to scabrous); spikes usually 9-16 cm long; 

lemmas 7-13 mm long ..............................................................E. glabriflorus var. glabriflorus 

12 Spikes 0.7-2 cm wide (including the awns), exserted or sheathed; lemma awns 1-15(20) mm long; 

spikelets appressed to slightly spreading; blades usually glabrous to scabridulous. 

15 Lemma awns 1-3(5) mm long; blades often ascending, somewhat involute, those higher on the 

stiffly erect culms broader and more persistent; flowering usually in early July to mid-August......... 

........................................................................................................................................ [E. curvatus] 

15 Lemma awns 5-15(20) mm long; blades usually spreading or lax, not markedly broader or more 

persistent towards the culm summit; flowering usually in mid-June to late July. 

16 Spikes glaucous, hispidulous to villous-hirsute, often intermediate in exsertion; glumes 

indurate in the lowest 1-2 mm; ligules and auricles usually absent; flowering usually early July 

to mid-August..............................................................................E. virginicus var. intermedius 

16 Spikes green to glaucous, usually glabrous to scabrous, partly included in the sheath to fully 

exserted; ligules and auricles often present; flowering usually mid-June to mid-July. 

17 Spikes partly sheathed; glumes 1-2.3 mm wide, strongly indurate and bowed-out in the 

lowest 2-4 mm; plants usually green to yellowish-brown; nodes mostly covered ............... 

................................................................................................ E. virginicus var. virginicus 

17 Spikes usually exserted; glumes (0.5-) 0.7-1.5 (-1.8) mm wide, moderately indurate and 

bowed out in the lowest 1-2 mm; plants usually glaucous, sometimes reddish-brown at 

maturity; nodes often exposed. 

18 Culms usually 3-8 dm tall, with 4-6 nodes; blades 2-9 mm wide, becoming involute; 

spikes 3.5-11 cm long, strongly glaucous; glumes usually indurate in the lowest 1-2 

mm.................................................................................E. virginicus var. halophilus 

18 Culms usually 7-10 dm tall, with 6-8 nodes; blades 3-15 mm wide, flat; spikes 4-20 

cm long, pale green or glaucous; glumes indurate only in the lowest 1 mm................ 

............................................................................................ E. virginicus var. jejunus

POACEAE 881 

Elymus canadensis Linnaeus var. canadensis, Great Plains Wild-rye, Nodding Wild-rye. Mt, Pd (NC, VA), Cp? (SC?): 

moist forests; rare (NC Watch List, VA Rare). [= FNA; < E. canadensis – RAB, C, F, G, GW, K, W] 

Elymus glabriflorus (Vasey) Scribner & Ball var. australis (Scribner & C.R. Ball) J.J.N. Campbell, Southeastern Wild-rye. 

{Cp, Pd, Mt (GA, NC, SC, VA): } [< E. glabriflorus – FNA; < E. virginicus – RAB, C, GW, W; < E. virginicus var. 

glabriflorus (Vasey) Bush – F, "forma australis"; < E. virginicus var. virginicus – G, K; = E. virginicus var. australis – S] 

Elymus glabriflorus (Vasey) Scribner & Ball var. glabriflorus, Southeastern Wild-rye. {Cp, Pd, Mt (GA, NC, SC, VA): } 

[< E. glabriflorus – FNA; < E. virginicus – RAB, C, GW, W; < E. virginicus var. glabriflorus (Vasey) Bush – F, "forma 

glabriflorus"; < E. virginicus var. virginicus – G, K; = E. virginicus var. glabriflorus – S] 

Elymus hystrix Linnaeus var. bigelovianus (Fernald) Bowden, Northern Bottlebrush Grass. Mt (NC): high elevation 

forests; rare. [< Hystrix patula Moench – RAB, G; < Elymus hystrix – C, FNA; = Hystrix patula var. bigeloviana (Fernald) 

Deam – F; = E. hystrix var. bigeloviana – K, orthographic variant; < Hystrix hystrix (Linnaeus) Millspaugh – S] 

Elymus hystrix Linnaeus var. hystrix, Common Bottlebrush Grass. Mt, Pd (GA, NC, SC, VA): moist forests, dry forests 

especially over more fertile soils; common. [= K; < Hystrix patula Moench – RAB, G; < Elymus hystrix – C, FNA; = Hystrix 

patula var. patula – F; < Hystrix hystrix (Linnaeus) Millspaugh – S] 

Elymus macgregorii R. Brooks & J.J.N. Campb., Early Wild-rye. Pd (GA, NC, VA), Mt, Cp (NC, VA): rich mesic forests, 

especially bottomlands; uncommon. ME west to SD, south to Panhandle FL and s. TX. See Campbell (2000). [= FNA; < E. 

virginicus – RAB, C, GW, W; < E. virginicus var. virginicus – F, G, K, S] 

* Elymus repens (Linnaeus) Gould, Quackgrass, Dog-grass, Witchgrass. Mt, Pd, Cp (NC, VA): roadsides, disturbed areas, 

pastures; uncommon, probably introduced from Europe (sometimes considered to be partially native along the coast). June- 

August. [= K, X; = Elytrigia repens (Linnaeus) Nevski – C, Z; = Agropyron repens (Linnaeus) Palisot de Beauvois – RAB, G, 

HC, S, W; > Agropyron repens var. repens – F; > Agropyron repens var. subulatum (Schreber) Roemer & J.A. Schultes – F] 

Elymus riparius Wiegand, Eastern Riverbank Wild-rye. Mt (GA, NC, SC, VA), Pd (GA, NC, SC, VA), Cp (SC?, VA): 

moist forests; uncommon (NC Watch List). [= RAB, C, F, FNA, G, GW, K, S, W] 

Elymus trachycaulus (Link) Gould ex Shinners ssp. trachycaulus, Slender Wheatgrass. Mt (NC, VA): glades and barrens, 

over serpentine, etc.; rare (VA Rare). [= K; < Agropyron trachycaulum (Link) Malte ex H.F. Lewis – RAB, W; < Elymus 

trachycaulus – C; > Agropyron trachycaulum var. novae-angliae (Scribner) Fernald – F; > Agropyron trachycaulum var. 

ciliatum (Scribner & J.G. Smith) Gleason – G; = Agropyron trachycaulum – HC] 

Elymus villosus Muhlenberg ex Willdenow, Downy Wild-rye. Mt, Pd, Cp (NC, SC, VA), {GA}: moist forests; 

uncommon. And reported for PA by Rhoads & Klein (1993). [= RAB, C, F, FNA, G, GW, K, W; = E. striatus Willdenow – S] 

Elymus virginicus Linnaeus var. halophilus (Bicknell) Wiegand, Salt-marsh Wild-rye. Cp (NC, VA): brackish marshes, 

maritime forests and hammocks; uncommon (VA Watch List). [= F, FNA, G, K; < E. virginicus – RAB, C, GW; < E. virginicus 

var. virginicus – S] 

Elymus virginicus Linnaeus var. intermedius (Vasey) Bush. [= FNA, G; < E. virginicus – RAB, C, GW, W; < E. 

virginicus var. virginicus – F, K; < E. virginicus var. hirsutiglumis (Scribner) A.S. Hitchcock – S] 

Elymus virginicus Linnaeus var. jejunus (Ramaley) Bush. [= F, FNA, G; < E. virginicus – RAB, C, GW, W; < E. 

virginicus var. virginicus – K; < E. virginicus var. virginicus – S] 

Elymus virginicus Linnaeus var. virginicus, Common Eastern Wild-rye, Terrell Grass. Mt, Pd, Cp (NC, SC, VA): moist 

forests; common. [= FNA; < E. virginicus – RAB, C, GW, W; < E. virginicus var. virginicus – F, G, K; < E. virginicus var. 

virginicus – S; ? E. striatus Willdenow – S] 

Elymus curvatus Piper, Awnless Wild-rye, east to c. TN and KY. [= FNA; < E. virginicus Linnaeus – C; = E. submuticus 

(Hooker) Smyth & Smyth – K; = E. virginicus Linnaeus var. submuticus Hooker – F, G; < E. virginicus var. virginicus – S] 

* Elymus semicostatus (Nees ex Steudel) Melderis. Pd (GA): disturbed areas; rare, introduced. Reported for c. GA by Jones 

& Coile (1988), as Agropyron semicostatum Nees ex Steudel. [= FNA, K; = Agropyrum semicostatum Nees ex Steudel] 

{synonymy incomplete} 

Elymus svensonii G.L. Church, Svenson's Wild-rye. Nc. KY south to c. TN; disjunct westwards in e. MO, and c. AR. [= 

FNA, K] 

Elymus trachycaulus (Link) Gould ex Shinners ssp. subsecundus (Link) A. & D. Löve, Bearded Wheatgrass, in MD, WV, 

and KY (Kartesz 1999). [= K; ? Agropyron trachycaulum (Link) Malte ex H.F. Lewis var. glaucum (Pease & Moore) Malte – F, 

G; = Agropyron subsecundum (Link) A.S. Hitchcock var. subsecundum – HC] 

Elymus wiegandii Fernald, Northern Riverbank Wild-rye. South to sc. PA and NJ. [= C, F, FNA, K; < E. canadensis – G] 

Elytrigia Desvaux (Quackgrass) 

(see Elymus, Pascopyrum) 

Enteropogon Nees 

* Enteropogon prieurii (Kunth) W.D. Clayton. Cp (NC): on ballast at Wilmington, New Hanover County, NC; rare, native 

of Africa, probably only a waif. Also reported from Mobile, Baldwin County, AL (Hitchock & Chase 1950). [= K; = Chloris 

prieurii Kunth – S]

POACEAE 882 

Eragrostis Wolf 1776 (Lovegrass) 

A genus of about 350 species of temperate and tropical areas. References: Peterson in FNA (2003a); Koch (1978); Peterson & 

Harvey (in prep.)=Z. Key adapted from Peterson & Harvey (in prep.). 

1 Plants cespitose or rhizomatous perennials, with innovations near the base, and with or without buds in the basal sheaths. 

2 Plants with short, knotty, thick rhizomes; florets articulating whole .............................................................E. spectabilis 

2 Plants without short or thick rhizomes; florets usually disarticulating. 

3 Caryopsis with a deep to shallow groove along the adaxial surface. 

4 Caryopsis dorso-ventrally compressed, flattened parallel to the side of the embryo, translucent, light brownish 

........................................................................................................................................................... E. curvula 

4 Caryopsis laterally compressed, flattened on the side perpendicular to the embryo, or cylindric, opaque 

(rarely translucent), usually reddish brown. 

5 Lateral veins of the lemmas conspicuous, often greenish, the lemmas strongly keeled ...........E. trichodes 

5 Lateral veins of the lemmas inconspicuous and hardly evident, the lemmas sometimes weakly keeled. 

6 Lemmas 1.2-1.8 mm long; culms 30-70 cm tall ...................................................................E. lugens 

6 Lemmas 1.6-3.0 mm long; culms (30-) 40-110 (-120) cm tall. 

7 Spikelets 2-6-flowered, greenish with purple tinges; leaf blades 3-8 (-11) mm wide, 25-60 cm 

long; sheaths often densely papillose-hirsute............................................................... E. hirsuta 

7 Spikelets (3-) 5-12-flowered, olive green to lead gray; leaf blades 1-3.8 mm wide, (4-) 10-35 

cm long; sheaths never papillose-hirsute ............................................................... E. intermedia 

3 Caryopsis not grooved on the adaxial surface. 

8 Stamens 3. 

9 Spikelets 4-8.2 (-10) mm long.................................................................................................... E. curvula 

9 Spikelets 2-4.5 (-5) mm long. 

10 Leaf blades 25-60 cm long, 3-8 (-11) mm wide; lemmas 1.6-2.4 mm long; spikelets 1.0-1.7 mm 

wide..................................................................................................................................... E. hirsuta 

10 Leaf blades (4-) 8-22 cm long, 1-3.5 mm wide; lemmas 1.2-1.8 mm long; spikelets 0.5-1.0 (-1.3) 

mm wide ..............................................................................................................................E. lugens 

8 Stamens 2. 

11 Panicle 15-45 cm wide, open, diffuse, broadly ovate to obovate in outline, the panicle branches 

capillary; pedicels 0.5-35 (-50) mm long, longer than or shorter than the spikelets. 

12 Spikelets with widely spreading pedicels, the lower pedicels all generally longer than the spikelets; 

disarticulation of the lemmas only, the paleas persistent..................................................... E. elliottii 

12 Spikelets with appressed pedicels, lower pedicels of each branch shorter than the spikelets; 

disarticulation usually of the whole floret ..........................................................................E. refracta 

11 Panicle (1-) 2-17 (-20) cm wide, contracted to open, narrowly ovate to oblong in outline;, the panicle 

branches stiffly spreading; pedicels (0-) 0.3-6 mm long, always shorter than the spikelets. 

13 Spikelets 0.7-2.4 mm wide; glumes 0.3-2.2 mm long; lemma 1.5-2.5 mm long, the apex acute 

(sometimes acuminate).................................................................................................... E. bahiensis 

13 Spikelets 2.4-5 mm wide; glumes 1.4-4 mm long; lemma 2-6 mm long, the apex acuminate to 

attenuate ................................................................................................E. secundiflora var. oxylepis 

1 Plants cespitose, geniculate or mat-forming annuals, lacking innovations or buds in the lower sheaths. 

14 Paleas prominently ciliate-pectinate on the keels, the hairs 0.1-0.8 mm long. 

15 Panicles contracted, narrow, spike-like, usually < 1.5 cm wide ............................................... E. ciliaris var. ciliaris 

15 Panicles open, cylindrical to narrowly ovate, usually 1-8 cm wide. 

16 Spikelets (1.0-) 1.5-3.5 mm long, 0.9-1.4 mm wide, 4-12-flowered; lemmas 0.7-1.1 mm long, membranous, 

the apex truncate to obtuse ................................................................................................................E. amabilis 

16 Spikelets 5-12 (-18) mm long, 1.4-2.4 mm wide, 12-42-flowered; lemmas (1.3-) 1.5-2.0 mm long, 

chartaceous, the apex acute .............................................................................................................. E. cumingii 

14 Paleas smooth to scaberulous on the keels, the hairs (if present) < 0.1 mm long. 

17 Plants extensively stoloniferous, creeping and forming flat mats; inflorescences 1-3.5 cm long; culms (2-) 5-12 (- 

20) cm tall on the erect portions............................................................................................................. E. hypnoides 

17 Plants not stoloniferous (sometimes creeping and forming flat mats); inflorescences 3-55 cm long; culms (2-) 6- 

130 cm tall. 

18 Ligules membranous, glabrous..........................................................................................................E. japonica 

18 Ligules ciliate, with a row of tiny white hairs. 

19 Caryopsis with a deep to shallow groove along the adaxial surface. 

20 Spikelets (4-) 5-10 (-11) mm long, 5-11 (-15)-flowered; pedicels ascending, somewhat appressed 

along the branches. 

21 Spikelets ovate to oblong in outline, >1.4 mm wide; lower glume 1.2-2.3 mm long................... 

......................................................................................................... E. mexicana ssp. mexicana 

21 Spikelets linear to linear-lanceolate,

POACEAE 883 

....................................................................................................... [E. mexicana ssp. virescens] 

20 Spikelets (1.4-) 2-5 mm long, 2-6 (-7)-flowered; pedicels erect, spreading along the branches. 

22 Panicle 10-45 (-55) cm long, 2/3 or more the height of the plant; pedicels (4-) 5-25 mm long; 

glandular pits absent below the nodes, branches, and rachis ....................................E. capillaris 

22 Panicle 4-20 cm long, < ½ the height of the plant; pedicels 1.5-5 mm long; glandular pits often 

present below the nodes, branches, and rachis..............................................................E. frankii 

19 Caryopsis not grooved on the adaxial surface. 

23 Plants with glandular pits or bands on the culm below the nodes, on the veins of the sheath, on the 

margins and veins of the blade, on the rachis, on the inflorescence branches and pedicels, and/or on 

the midveins of the lemma and palea. 

24 Spikelets (1.7-) 2-4 mm long, 3-6-flowered .................................................................E. frankii 

24 Spikelets (2-) 3.5-20 mm long, (3-) 5-40-flowered. 

25 Spikelets 0.6-1.3 mm wide; pedicels 1-10 mm long, flexuous and delicate, appressed or 

spreading ............................................................................................................... E. pilosa 

25 Spikelets 1.1-4 mm wide; pedicels 0.2-4 mm long, straight and rigid, mostly spreading. 

26 Spikelets 6-20 mm long, 2-4 mm wide, 10-40-flowered; lemmas 2-2.8 mm long, 

with 1-3 crateriform glands along the keel; disarticulation of the entire florets from 

the persistent rachilla; anthers yellow.................................................... E. cilianensis 

26 Spikelets 4-7 (-11) mm long, 1.1-2.2 mm wide, 7-12 (-20)-flowered; lemmas 1.4-1.8 

mm long, rarely with 1-2 crateriform glands along the keel; disarticulation of the 

lemmas only, the palea and rachilla usually persistent; anthers reddish-brown. 

27 Inflorescence with glandular areas of spots or rings on the rachis below the 

panicle branch bases, the glands often shiny or yellowish; stamens 3; blade 

margins lacking crateriform glands.................................................. E. barrelieri 

27 Inflorescence sometimes with glandular areas of spots or crateriform pits on the 

rachis below the panicle branch bases, the glands usually dull and greenishgray 

to straw-colored; stamens 2; blade margins sometimes with crateriform 

glands.....................................................................................................E. minor 

23 Plants lacking glandular pits or bands on the culm below the nodes, on the veins of the sheath, on 

the margins and veins of the blade, on the rachis, on the inflorescence branches and pedicels, and/or 

on the midveins of the lemma and palea. 

28 Spikelets (1.6-) 2-4 mm wide, 12-42-flowered; disarticulation of entire florets from a 

persistent rachilla................................................................................................. [E. unioloides] 

28 Spikelets 0.6-2.5 mm wide, 3-22-flowered; disarticulation of the lemmas only, the paleas 

usually persistent (or deciduous), the rachilla persistent. 

29 Spikelets 3-6-flowered..........................................................................................E. frankii 

29 Spikelets (3-) 5-22-flowered. 

30 First glume 0.3-0.6 (-0.8) mm long, 0.5× as long as the lowest lemma; spikelets 1.2-2.5 

mm wide; panicle branches solitary or paired at the 2 lowest nodes. 

31 Pedicels widely spreading...................................[E. pectinacea var. miserrima] 

31 Pedicels appressed or rarely diverging up to 20 degrees from the branches ........ 

..............................................................................E. pectinacea var. pectinacea 

* Eragrostis amabilis (Linnaeus) Wright & Arnott ex Nees, Japanese Lovegrass. Cp (GA, SC), Pd? (GA?): disturbed areas; 

rare, introduced the Old World. June. [= RAB, FNA, HC, S, Z; ? E. tenella (Linnaeus) Palisot de Beauvois ex Roemer & J.A. 

Schultes – K] 

* Eragrostis bahiensis (Schrader ex J.A. Schultes) J.A. Schultes, Bahia Lovegrass. Cp (GA, SC): disturbed areas; rare. 

Reported for SC (Kartesz 1999) and sw. GA (Jones & Coile 1988, GW, Kartesz 1999). [= FNA, GW, HC, K, S, Z] 

* Eragrostis barrelieri Daveau, Mediterranean Lovegrass. Cp (SC): waste areas near wool-combing mills; rare, introduced 

from Mediterranean Europe. Also reported for e. TN (Chester et al. 1993). [= FNA, HC, K, Z] 

Eragrostis capillaris (Linnaeus) Nees, Lacegrass. Mt, Pd, Cp (GA, NC, SC, VA): fields, roadsides, disturbed areas; 

common (uncommon in Piedmont, rare in Coastal Plain). July-October. ME and WI south to GA and TX. [= RAB, C, F, FNA, 

G, HC, K, S, W, Z] 

* Eragrostis cilianensis (Allioni) Vignolo ex Janchen, Stinkgrass. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): fields, 

disturbed areas; common, introduced from Europe. July-October. [= RAB, C, FNA, G, HC, K, S, W, Z; ? E. megastachya 

(Koel.) Link – F] 

Eragrostis ciliaris (Linnaeus) R. Brown var. ciliaris. Cp (GA, SC): sandy shores; rare. S. SC south to TX, Central 

America, West Indies, South America, Africa, and Asia. [= FNA, HC; < E. ciliaris – RAB, G, K, S, Z] 

* Eragrostis cumingii Steudel, Fortyflower Lovegrass, Cuming's Lovegrass. Cp (GA), {NC}. Reported for NC (Kartesz 

1999) and sw. GA (Jones & Coile 1988, HC). [= FNA, K, Z; ? E. simplex Scribner – HC]

POACEAE 884 

* Eragrostis curvula (Schrader) Nees, Weeping Lovegrass. Cp, Pd, Mt (GA, NC, SC, VA): roadsides; common, introduced 

from s. Africa. May-June. Very commonly planted as a roadbank stabilizer, E. curvula is fire resistant and shows some 

capability to spread into adjacent natural habitats. [= RAB, C, FNA, HC, K, Z; > E. curvula var. conferta Stapf] 

Eragrostis elliottii S. Watson, Elliott's Lovegrass. Cp (GA, NC, SC): ultisol wet pine savannas, maritime wet grasslands, 

inland edges of brackish marshes, inland edges of freshwater tidal marshes, calcareously-influenced wet pine savannas; rare. 

September-October. NC south to FL, west to TX. [= RAB, FNA, GW, HC, K, S, Z] 

Eragrostis frankii C.A. Meyer ex Steudel, Lacegrass. Mt (VA), Pd (NC, VA), Cp (VA), {GA}: disturbed areas; 

uncommon (NC Watch List). September. MA and MN south to FL and AR. [= RAB, C, FNA, G, GW, K, S, W, Z; > E. frankii 

var. frankii – F, HC] 

Eragrostis hirsuta (Michaux) Nees, Bigtop Lovegrass. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disturbed areas; 

common (uncommon in Mountains). July-October. MD south to FL, west to TX, north in the interior to TN, AR, and MO; 

Central America. [= RAB, C, FNA, K, S, W, Z; > E. hirsuta var. hirsuta – F, G, HC; > E. hirsuta var. laevivaginata Fernald – F, 

G, HC] 

Eragrostis hypnoides (Lamarck) Britton, Sterns, & Poggenburg, Creeping Lovegrass, Teal Lovegrass. Cp, Pd (GA, NC, 

SC, VA), Mt (NC, VA): marshes, shores; uncommon. September. Throughout most of North America, south to South America. 

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

Eragrostis intermedia A.S. Hitchcock, Plains Lovegrass. Cp, Pd (GA), {NC, SC, VA}. Reported for scattered locations as 

far east as NC, SC, VA (Kartesz 1999), e. GA (Jones & Coile 1988), e. TN (Chester et al. 1993). [= C, F, FNA, G, HC, K, Z] 

*? Eragrostis japonica (Thunberg) Trinius, Pond Lovegrass. Cp (GA, SC): moist or wet sandy areas; rare. SC and TN south 

to Central America, South America, and West Indies; old World tropics. Perhaps introduced from the Old World. Reported for 

SC by HC, G, and Small (1933), sw. GA by Jones & Coile (1988), and for w. TN by Chester et al. (1993). [= FNA, K, Z; ? E. 

glomerata (Walter) L.H. Dewey – G, GW, HC, S] 

*? Eragrostis lugens Nees, Mourning Lovegrass. Cp (NC, SC), Pd (GA, NC, SC): marshes, roadsides, low fields; rare, 

introduced (NC Watch List). June-October. Perhaps only introduced from further south and west. [= RAB, FNA, HC, K, S, W, 

Z] 

* Eragrostis mexicana (Hornemann) Link ssp. mexicana, Mexican Lovegrass. Cp (SC): waste areas near wool-combing 

mills; rare, introduced from . Reported to be naturalized as far east and north as SC, DE, and MD (Kartesz 1999). [= FNA, K; > 

E. neomexicana Vasey – C, F, G, HC; > E. mexicana – C, F, G, HC] 

* Eragrostis minor Host, Little Lovegrass. Pd (GA, NC, SC, VA), Mt (NC, SC, VA), Cp (VA): disturbed areas; uncommon, 

introduced from Europe. July-September. [= C, FNA, K, Z; ? E. poaeoides Palisot de Beauvois ex Roemer & J.A. Schultes – 

RAB, F, G, HC, W; ? E. eragrostis (Linnaeus) Palisot de Beauvois – S] 

Eragrostis pectinacea (Michaux) Nees ex Steudel var. pectinacea, Carolina Lovegrass. Cp, Pd, Mt (GA, NC, SC, VA): 

fields, roadsides, disturbed areas; common (VA Watch List). ME and WA south to Central America and West Indies. [= FNA, 

K, Z; < E. pectinacea – C, GW, W; = E. pectinacea – F, HC, S; > E. pectinacea – G; > E. diffusa Buckley – G] 

* Eragrostis pilosa (Linnaeus) Palisot de Beauvois var. pilosa. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, disturbed 

areas; common, introduced from tropical regions of the Old and New World. July-October. Var. perplexa (L.H. Harvey) S.D. 

Koch is also introduced but is not known from our area. [= FNA; = E. pilosa – RAB, S, W; > E. multicaulis Steudel – F, G, HC; 

> E. pilosa – F, G, HC; < E. pilosa – K, Z] 

Eragrostis refracta (Muhlenberg) Scribner, Coastal Lovegrass. Cp (GA, NC, SC, VA), Pd (NC, SC, VA): pinelands, 

savannas, woodlands, marshes; common (uncommon in Piedmont). July-October. DE south to FL, west to TX. [= RAB, C, F, 

FNA, G, GW, HC, K, S, Z; ? E. virginica (Zuccarini ex Roemer) Steudel] 

* Eragrostis secundiflora J. Presl var. oxylepis (Torrey) S.D. Koch, Red Lovegrass. Cp (GA, NC, SC, VA): sandy 

roadsides, coastal dunes, and disturbed areas; rare, introduced from sw. United States. First reported for SC by Nelson & Kelly 

(1997). [= E. oxylepis (Torrey) Torrey – GW, HC; = E. secundiflora ssp. oxylepis S.D. Koch – FNA, K, Z; < E. secundiflora – 

S] 

Eragrostis spectabilis (Pursh) Steudel, Purple Lovegrass, Tumblegrass. Cp, Pd, Mt (GA, NC, SC, VA): sandy fields, 

roadsides, woodlands; common. August-October. ME west to ND, south to FL and TX. [= RAB, C, FNA, G, GW, HC, K, S, 

W, Z; > E. spectabilis var. spectabilis – F; > E. spectabilis var. sparsihirsuta Farwell – F; E. pectinacea, misapplied] 

* Eragrostis trichodes (Nuttall) Wood. Pd (VA): disturbed areas; rare, introduced from w. North America. [= C, FNA, K, Z; 

> E. trichodes var. trichodes – F, HC] 

* Eragrostis elongata (Willdenow) Jacquin f., Long Lovegrass. Cp (SC): waste areas near wool-combing mills; rare, 

introduced from se. Asia and Australia. [= FNA, K] {not keyed at this time} 

* Eragrostis leptostachya (R. Brown) Steudel, Australian Lovegrass, is reported for NC (Kartesz 1999). {investigate} [= 

FNA, K] {not keyed at this time} 

* Eragrostis mexicana (Hornemann) Link ssp. virescens (J. Presl) S.D. Koch & Sánchez, reported as an introduction on 

ballast in MD and FL. [= FNA, K; = E. virescens J. Presl – HC] 

Eragrostis pectinacea (Michaux) Nees ex Steudel var. miserrima (Fournier) J. Reeder, from FL and westwards and 

southwards, may be in our area. [= FNA, K, Z; = E. tephrosanthos J.A. Schultes – HC, S; < E. pectinacea – GW] 

* Eragrostis plana Nees, South African Lovegrass. Cp (SC): waste areas near wool-combing mills; rare, introduced from 

South Africa. [= FNA, K] {not keyed at this time} 

* Eragrostis setifolia Nees, Neverfail. Cp (SC): waste areas near wool-combing mills; rare, introduced from Australia. [= 

FNA, K] {not keyed at this time}

POACEAE 885 

* Eragrostis tef (Zuccagni) Trotter, Teff. Cp (SC): waste areas near wool-combing mills; rare, introduced from Africa. This 

is the grain used in making Ethiopian bread. [= FNA, HC, K] {not keyed at this time} 

* Eragrostis unioloides (Retzius) Nees ex Steudel, Chinese Lovegrass. Cp (GA): rare, introduced from Asia. Reported for s. 

GA (Jones & Coile 1988, FNA, GW, HC). [= FNA, GW, HC, K, S, Z] 

Eremochloa Büse (Centipede Grass) 

A genus of about 11 species, native of Asia and Australia. References: Thieret in FNA (2003a). 

Identification notes: In the autumn, the inflorescences make this grass readily recognizable at a distance: a short, tight lawn 

grass with a reddish aspect. 

* Eremochloa ophiuroides (Munro) Hackel, Centipede Grass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): lawns, roadsides, 

sometimes weedy in more natural sites; common, introduced from se. Asia. Now very commonly planted as a lawn and roadside 

grass in the Coastal Plain from se. NC southward. Stalter & Lamont (1996) report the VA occurrence of this species. [= RAB, 

FNA, HC, K] 

Erianthus 

(see Saccharum) 

Eriochloa Kunth (Cup Grass) 

A genus of 320-30 species, of the tropical, subtropical, and warm temperate Old World and New World. References: Crins 

(1991)=Z; Shaw, Webster, & Bern in FNA (2003a); Shaw & Webster (1987)=Y. 

1 Lemma of fertile floret with an awn >0.2 mm long; second glume awned; panicle compact, the raceme-like lateral branches 

close together and ascending-appressed, of irregular lengths; spikelets 8-16 on a typical, primary branch............ E. contracta 

1 Lemma of fertile floret lacking an awn; second glume not awned; panicle open, the raceme-like lateral branches remote and 

divergent, the lowermost longest, the upper gradually reduced in length to the apex (E. acuminata var. acuminata, E. 

michauxii var. michauxii) or the panicle compact (E. villosa); spikelets 12-40 on a typical, primary branch. 

2 Spikelets 2.0-2.5 mm wide................................................................................................................................... E. villosa 

2 Spikelets 1.1-1.8 mm wide. 

3 Annual, 3-12 dm tall; spikelets 1.1-1.4 mm wide.........................................................E. acuminata var. acuminata 

3 Perennial, 5-25 dm tall; spikelets 1.3-1.8 mm wide .......................................................E. michauxii var. michauxii 

* Eriochloa acuminata (J. Presl) Kunth var. acuminata. Cp (GA, SC), Pd (GA): disturbed areas, waste areas near woolcombing 

mills; rare, presumably introduced from further south. Reported for scattered locations in GA (Jones & Coile 1988, as 

E. gracilis). Reported for NC (Kartesz 1999), but the specimen basis is of cultivated material. [= FNA, K, Y, Z; < E. acuminata 

– C; = E. gracilis (Fournier) A.S. Hitchcock var. gracilis – HC] 

* Eriochloa contracta A.S. Hitchcock, Prairie Cupgrass. Pd, Mt (VA), Cp (SC): disturbed areas, waste areas around woolcombing 

mills; rare, introduced from midwestern United States. [= C, F, FNA, G, GW, HC, K, Y, Z] 

Eriochloa michauxii (Poiret) A.S. Hitchcock var. michauxii, Longleaf Cupgrass. Cp (GA, SC): coastal freshwater and 

slightly brackish marshes, flatwoods, disturbed areas; rare (GA Special Concern). Se. SC south to FL, west to AL, or possibly 

LA. Var. simpsonii A.S. Hitchcock is endemic to sw. FL. [= FNA, HC, K, Y, Z; < E. michauxii – GW, S] 

* Eriochloa villosa (Thunberg) Kunth, Chinese Cupgrass. Mt (VA): disturbed area (open edge of railroad bed); rare, 

introduced from e. Asia. See Belden et al. (2004) for additional information about the first occurrence in Virginia. [= C, FNA, 

HC, K, Y] 

Eriochloa punctata (Linnaeus) Desvaux ex Hamilton, Louisiana Cupgrass. Cp (GA): marshes, creek banks; rare. MS west 

to TX, and south into the New World Tropics; reported for e. GA (FNA). [= FNA, HC, K] {not keyed at this time; synonymy 

incomplete} 

Eustachys Desvaux (Finger-grass) 

A genus of ca. 12 species, of tropical and warm temperate regions. References: Aulbach in FNA (2003a). McKenzie, Urbatsch, 

& Aulbach-Smith (1987)=Z. Key based on Z. 

1 Lateral nerves of the fertile lemma glabrous; culms stout, 7-15 dm tall; spikes 8-16 (-20), 7-12 cm long ..................E. glauca 

1 Lateral nerves of the fertile lemma pubescent; culms slender, 3-10 dm tall; spikes 1-20, 2.5-9 cm long. 

2 Keel of the fertile lemma appressed brownish-ciliate; spikes 1-6; [native].

POACEAE 886 

3 Spikes 5-10 cm long; spikelets >3 mm long ............................................................................................E. floridana 

3 Spikes 2.5-6 cm long; spikelets 2.4 mm long; sterile floret oblanceolate, acute...............................................................E. distichophylla 

4 Spikelets F. rubra var. rubra – F; > 

F. rubra var. commutata Gaudin – F; ? F. rubra ssp. rubra – K] 

Festuca subverticillata (Persoon) Alexeev, Nodding Fescue. Mt, Pd, Cp (GA, NC, SC, VA): moist to wet forests, 

woodlands, and disturbed areas; common. May-June. ME, Québec, and Manitoba south to FL and e. TX. [= C, K, Y, Z; ? F. 

obtusa Biehler – RAB, F, G, GW, HC, S, W] 

* Festuca trachyphylla (Hackel) Krajina, Hard Fescue. Pd (GA, NC, VA), Mt (NC, VA), Cp (NC, SC, VA): meadows, 

pastures, disturbed areas; uncommon, introduced from Eurasia. May-June. The nomenclatural debate about the application of 

the name F. trachyphylla is summarized in Darbyshire & Pavlick (1997). [= C, K, Y, Z; < F. ovina – RAB, S, W, in the broad

POACEAE 887 

sense (misapplied as to our material); < F. ovina var. ovina – F, G, HC; < F. ovina var. duriuscula (Linnaeus) W.D.J. Kock – F, 

G, HC, misapplied as to our material] 

* Festuca thurberi Vasey. Introduced in SC. [= K] {not keyed; investigate} 

Festuca versuta Beal, Texas Fescue. Native, east to TN. [= K] {not keyed; investigate} 

Glyceria R. Brown (Mannagrass) 

(also see Torreyochloa) 

A genus of about 40 species, nearly cosmopolitan. References: Tucker (1996)=Z. 

1 Spikelets (10-) 15-40 mm long, linear, subterete, 5-15× as long as wide; [section Glyceria]. 

2 Lemma (6-) 7-8.5 (-10) mm long, acute to acuminate; palea longer than the lemma, extending 1.5-3 mm beyond the 

lemma apex.....................................................................................................................................................G. acutiflora 

2 Lemma 2.5-5.3 mm long, obtuse to notched; palea about as long as the lemma (ranging from shorter than the lemma 

and included, to projecting up to 1 mm beyond the lemma apex). 

3 Anthers 0.5-0.8 mm long; lemma mostly 2.5-3.5 mm long................................................................... G. arkansana 

3 Anthers 1.0-2.0 mm long; lemma mostly 3.6-5.5 mm long.............................................................G. septentrionalis 

1 Spikelets 2.5-8 mm long, ovate to oblong, 1.5-3× as long as wide; [section Hydropoa]. 

4 Inflorescence compact (at maturity), the branches stiffly ascending to appressed, the tips never nodding; ligule < 1 mm 

long. 

5 Inflorescence branches elongate, appressed; lower internodes of the inflorescence 2-8 cm long; spikelets with 3-4 

flowers, 3.5-4 mm long; lemma 1.9-2.8 mm long; leaves 2-5 mm wide; [of the Mountains, rarely elsewhere] ......... 

................................................................................................................................................................ G. melicaria 

5 Inflorescence branches short, stiffly ascending; lower internodes of the inflorescence 0.8-2.0 (-2.5) cm long; 

spikelets with 4-7 flowers, 4-8 mm long; lemma 3.0-3.7 mm long; leaves 3-10 mm wide; [of the Coastal Plain, 

rarely disjunct inland to the Mountains of VA]............................................................................................G. obtusa 

4 Inflorescence lax and diffuse (at maturity), the branches spreading to somewhat ascending, the tips often nodding or 

drooping; ligule 1-6 mm long. 

6 Veins of the lemma visible, but not raised; lemma 2.3-4.0 mm long; ligule 2-6 mm long. 

7 Lemma 2.9-4.0 mm long, projecting conspicuously beyond the palea; spikelets 4-8 mm long, with 5-10 

flowers...........................................................................................................................................G. canadensis 

7 Lemma 2.3-2.9 mm long, more-or-less equal to the palea; spikelets 3-5 mm long, with (2-) 3-5 (-6) flowers ... 

................................................................................................................................................................ G. laxa 

6 Veins of the lemma prominently raised; lemma 1.4-3.0 mm long; ligule 1-4 mm long. 

8 Lemma 1.4-2.1 mm long, usually green; first glume 0.5-1.0 mm long; second glume 0.8-1.3 mm long; culms 

mostly 5-12 dm tall; leaves 2-5 (-8) mm wide ...................................................................G. striata var. striata 

8 Lemma 2.5-3.0 mm long, purple or green; first glume 1.2-1.9 mm long; second glume 1.5-2.4 mm long; 

culms mostly 10-20 dm tall; leaves 5-12 mm wide. 

9 Glumes obtuse, subequal (the first 1.2-1.9 mm long, the second 1.5-2.4 mm long); spikelets 4-6.5 mm 

long, (4-) 5-9-flowered; lemmas purplish, contrasting with the pale glumes; ligule 2-4 (-5) mm long; [of 

various montane wetlands of VA and possibly NC]...............................................G. grandis var. grandis 

9 Glumes acute, unequal (the first 1.4-1.6 mm long, the second 1.8-2.1 mm long); spikelets mostly 3-4 

mm long, 2-4-flowered; lemmas dark green (or purplish?); ligule 1.5-3 mm long; [of high elevation 

seepages in the Great Smoky Mountains of NC and TN]........................................................ G. nubigena 

Glyceria acutiflora Torrey. Mt (GA, VA): shallow water and wet mucky soils in mountain ponds, wet pastures; rare (GA 

Special Concern). June-July. ME west to MI, south to DE, VA, nw. GA (Jones & Coile 1988), e. TN, and MO; also in e. Asia. 

[= C, F, G, GW, HC, K, W, Z; = Panicularia acutiflora (Torrey) Kuntze – S] 

Glyceria arkansana Fernald, Arkansas Mannagrass. Cp (VA): swamps; rare. May-June. IL south to LA and AR; disjunct 

in se. VA and w. NY. The appropriate treatment of this taxon needs further investigation. [= F, HC, K, Z; < G. septentrionalis – 

C, G; = G. septentrionalis A.S. Hitchcock var. arkansana (Fernald) Steyermark & Kučera] 

Glyceria canadensis (Michaux) Trinius, Rattlesnake Mannagrass. Mt (NC, VA), Cp (VA): bogs, seepages, and wet 

meadows; rare. June-July. Newfoundland west to MN, south to NJ, VA, nw. NC, and IL. [= C, F, G, K, Z; = G. canadensis var. 

canadensis – HC, W] 

Glyceria grandis S. Watson var. grandis, American Mannagrass. Mt (NC?, VA), Cp (VA): wet, mucky soils of open 

wetlands; rare. Nova Scotia west to AK, south to VA, IA, NM, and OR. Attributed to w. NC by Tucker (1996). [= K; < G. 

grandis – C, F, G, GW, HC, Z; = Panicularia grandis (S. Watson) Nash – S; ? G. maxima (Hartman) Holmberg ssp. grandis (S. 

Watson) Hultén] 

Glyceria laxa (Scribner) Scribner, Lax Mannagrass. Mt (NC, VA): bogs; rare. June-July. Prince Edward Island south to 

NC, mostly Appalachian. Though often described as a hybrid of G. canadensis and either G. striata var. striata and/or G. 

grandis var. grandis, G. laxa ranges south of the distribution of both G. canadensis and G. grandis var. grandis. It is best

POACEAE 888 

considered as a species, perhaps of hybrid origin. [= F, G, K; = G. canadensis (Michaux) Trinius var. laxa (Scribner) A.S. 

Hitchcock – RAB, HC; = G. ×laxa – C; < G. canadensis – GW] 

Glyceria melicaria (Michaux) F.T. Hubbard, Northeastern Mannagrass. Mt (GA, NC, VA), Pd (NC): mountain swamp 

forests and seepages; uncommon (GA Special Concern). June-August. Nova Scotia west to Québec, south to n. GA (Jones & 

Coile 1988) and KY. [= RAB, C, F, G, GW, HC, K, W, Z; = Panicularia melicaria (Michaux) A.S. Hitchcock – S] 

Glyceria nubigena W.A. Anderson, Smoky Mountain Mannagrass. Mt (NC): moderate to high elevation seepages in the 

Great Smoky Mountains, sometimes in areas appearing dry (such as heath balds), nearly endemic to Great Smoky Mountains 

National Park; rare (US Species of Concern, NC Rare). June-July. Endemic to the Great Smoky Mountains of w. NC and e. TN. 

G. nubigena has nearly the same range as Rugelia nudicaulis, but is more restricted to seepage. The distinctions and relationship 

between this taxon and G. grandis need further investigation. [= RAB, HC, K, W, Z] 

Glyceria obtusa (Muhlenberg) Trinius, Coastal Mannagrass. Cp (NC, SC, VA), Mt (VA): blackwater swamp forests, wet 

meadows, freshwater marshes; uncommon. June-September. Nova Scotia south to SC, on or near the Coastal Plain. [= RAB, C, 

F, G, GW, HC, K, W, Z; = Panicularia obtusa (Muhlenberg) Kuntze – S] 

Glyceria septentrionalis A.S. Hitchcock, Floating Mannagrass, Eastern Mannagrass. Cp, Pd (GA, NC, SC, VA), Mt (NC, 

VA): shallow water, wet mucky soils, floodplain sloughs, cypress ponds; uncommon (GA Special Concern). May-June. MA 

west to MN, south to SC, ne. GA, and TX. [= RAB, F, GW, HC, K, W, Z; < G. septentrionalis – C, G (also see G. arkansana); 

= Panicularia septentrionalis (A.S. Hitchcock) Bicknell – S] 

Glyceria striata (Lamarck) A.S. Hitchcock var. striata, Fowl Mannagrass. Mt, Pd, Cp (GA, NC, SC, VA): wet meadows, 

seepages, bogs, marshes, swamp forests; common. April-June. Newfoundland west to British Columbia, south to FL and CA. 

Var. stricta (Scribner) Fernald is more northern. [= C, F, G, HC, Z; < G. striata – RAB, GW, K, W; = Panicularia striata 

(Lamarck) A.S. Hitchcock – S; = G. striata ssp. striata] 

Gymnopogon Palisot de Beauvois (Beard Grass, Skeleton Grass) 

A genus of about 15 species, in temperate and tropical areas of the Americas. References: Smith (1971)=Z. 

Identification notes: When sterile, Gymnopogon is sometimes confused with Dichanthelium. Gymnopogon differs in having 

the sheaths conspicuously overlapping (vs. not overlapping in Dichanthelium) and leaves that are definitely cordate-clasping and 

of stiff texture (only a few Dichanthelium have this combination). 

1 Awn of the lemma 4.5-12 mm long; inflorescence branches with spikelets distributed from the tip nearly to the base; leaves 

5-15 mm wide; [of the Coastal Plain, Piedmont, and Mountains]...........................................................................G. ambiguus 

1 Awn of the lemma 0.8-1.6 (-3.5) mm long; inflorescence branches with spikelets distributed from the tip nearly to the base 

(G. chapmanianus) or to roughly the midpoint, the basal portion naked (or some branches rarely with a few spikelets) (G. 

brevifolius); leaves 2-8 mm wide; [of the Coastal Plain and lower Piedmont]. 

2 Spikelets 1-flowered; first glume 2.3-3.7 mm long....................................................................................... G. brevifolius 

2 Spikelets 2-4-flowered; first glume 3.8-5 mm long ............................................................................... G. chapmanianus 

Gymnopogon ambiguus (Michaux) Britton, Sterns, & Poggenburg, Eastern Beard Grass. Cp, Pd (GA, NC, SC, VA), Mt 

(NC, SC, VA): prairies, glades, barrens, dry pinelands and woodlands, dry fields; common (rare in Mountains). August-October. 

S. NJ west to KY, OH, and MO, south to FL and TX. [= RAB, C, F, G, HC, K, S, W, Z] 

Gymnopogon brevifolius Trinius, Pineland Beard Grass. Cp, Pd (GA, NC, SC, VA), Mt (GA): pine savannas, sandhills, 

dry woodlands, prairies, calcareous glades; common (rare in lower Piedmont) (GA Special Concern, VA Watch List). August- 

October. S. NJ south to FL, west to LA and AR; disjunct in the Highland Rim of KY and TN. [= RAB, C, F, G, HC, K, S, Z] 

Gymnopogon chapmanianus A.S.Hitchcock, Chapman's Beard Grass. Cp (GA): sandhills and other xeric, sandy habitats; 

rare (GA Special Concern). Se. GA south to FL. [= HC, K, S; > G. chapmanianus – Z; G. floridanus Swallen – Z] 

Hackelochloa Kuntze (Pitscale Grass 

* Hackelochloa granularis (Linnaeus) Kuntze, Pitscale Grass. Cp (GA): disturbed areas; rare, introduced from the Old 

World. Reported for sw. GA and other Gulf Coast states (Thieret in FNA 2003a, Jones & Coile 1988, Kartesz 1999). [= FNA, 

HC, K; = Rytilix granularis (Linnaeus) Skeels – S; = Mnesithea granularis (Linnaeus) Koning & Sosef] 


Hainardia W. Greuter (Thintail) 

* Hainardia cylindrica (Willdenow) W. Greuter, Thintail. Cp (SC): waste areas around wool-combing mills; rare, 

introduced from the Old World. April-June. [= K, Z; = Lepturus cylindricus (Willdenow) Trinius – RAB; = Monerma 

cylindrica (Willdenow) Cosson & Durieu – HC]

POACEAE 889 

Heteropogon Persoon (Tanglehead) 

A genus of about 10 species, pantropical and extending into subtropical and warm temperate areas. References: Barkworth in 

FNA (2003a). 

* Heteropogon melanocarpus (Elliott) Elliott ex Bentham, Sweet Tanglehead. Cp (GA, NC, SC): sandy roadsides, disturbed 

areas; rare, probably naturalized from further south (or even from the Old World). September-October. The species is 

widespread in the Old World and New World tropics, north in North America to se. NC. [= RAB, FNA, HC, K, S] 

Hierochloe R. Brown (Holy Grass, Sweet Grass, Vanilla Grass) 

A genus of about 30 species, temperate and boreal in the northern and southern hemispheres. Tucker (1996) and Soreng et al. 

(2003) propose the inclusion of Hierochloe into a more broadly circumscribed Anthoxanthum. References: Tucker (1996)=Z; 

Soreng et al. (2003)=Y. 

Hierochloe odorata (Linnaeus) Palisot de Beauvois, Holy Grass, Sweet Grass, Vanilla Grass. Mt (NC, VA): fens, wet 

calcareous medaows, high elevation pastures and openings; rare (NC Rare). April-May. A circumboreal species and subspecies, 

widespread in n. Eurasia and n. North America, ranging south in North America to NJ, MD, PA, OH, IN, IL, IA, SD, CO, UT, 

NM, and CA, with several disjunct occurrences in North Carolina, in Long Hope Valley, Ashe County, the Nantahala River 

Bogs, Macon County, and Pond Mountain, Ashe County. The report by S ("recorded by Chapman from Statesville, N.C.") can 

be discounted; the record reflects a collection made in the mountains by Mordecai E. Hyams, a botanist based in Statesville. 

Belden et al. (2004) document the first occurrence in Virginia. The sweet, vanilla-like odor of this grass is responsible for 

various folk uses – by Native Americans for making fragrant baskets, in Scandinavia strewn on church floors on festival days. 

Kartesz (1999) maps the NC occurrence of Hierochloe as H. hirta ssp. arctica; the reasons for this are unknown. {investigate} 

[= C, F, G, HC; ? H. odorata ssp. odorata – K; ? H. hirta (Schrank) ssp Borbás ssp. arctica (J. Presl) G. Weimarck – K; Torresia 

odorata (Linnaeus) A.S. Hitchcock – S; = Anthoxanthum nitens (Weber) Y. Schouten & Veldkamp – Z; ? Anthoxanthum nitens 

(Weber) Y. Schouten & Veldkamp spp. nitens – Y; > H. odorata var. fragrans (Willdenow) Richter (the North American plants)] 


Holcus Linnaeus (Velvet Grass, Soft Grass) 

1 Plant not rhizomatous; upper culm internodes velvety-villous; lemma awn recurved ............................................... H. lanatus 

1 Plant strongly rhizomatous; upper culm internodes glabrous; lemma awn straight ..................................................... H. mollis 

* Holcus lanatus Linnaeus, Velvet Grass, Soft Grass, Yorkshire-fog. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): 

pastures, disturbed areas, roadsides, hedge-rows; common (rare in SC), introduced from Europe. May-October. [= RAB, C, F, 

G, HC, K, W, Z; = Notholcus lanatus (Linnaeus) Nash – S] 

* Holcus mollis Linnaeus, Creeping Soft Grass. Mt (NC): lawns; rare, introduced from Europe. September. This European 

species is known from scattered sites in e. North America. The species was documented for our area by Clay (1995). [= C, F, G, 

HC, K, Z] 

Hordeum Linnaeus 1753 (Barley) 

A genus of about 40 species, north temperate and in South America. Many recent authors place most of our species (other than 

H. vulgare) in Critesion Rafinesque. References: Tucker (1996)=Z; Petersen & Seberg (2003); Blattner (2004). 

1 Rachis remaining intact at maturity; leaves 5-12 mm wide, with well-developed auricles; [section Hordeum] ........ H. vulgare 

1 Rachis disarticulating at maturity; leaves 1-5 mm wide, not auriculate (except in H. murinum ssp. leporinum). 

2 Perennial; glumes 25-150 mm long; [intersectional hybrid derivative of section Sibirica and section Critesion].............. 

..........................................................................................................................................................................H. jubatum 

2 Annual; glumes 7-22 (-28) mm long. 

3 Leaves auriculate; glumes of the central spikelet (in the triad) with ciliate margins; [section Hordeum] ................... 

....................................................................................................................................... H. murinum ssp. leporinum 

3 Leaves not auriculate; glumes of the central spikelet (in the triad) with scabrous margins; [section Critesion] ......... 

................................................................................................................................................................. H. pusillum 

* Hordeum jubatum Linnaeus, Foxtail Barley, Squirreltail Barley. Mt, Pd (VA), Cp (NC, SC): disturbed areas; rare, 

apparently introduced in our area, introduced from w. United States. May-August. A tetraploid taxon. [= RAB, C, F, G, HC, W, 

Z; ? H. jubatum ssp. jubatum – K; = Critesion jubatum (Linnaeus) Nevski]

POACEAE 890 

* Hordeum murinum Linnaeus ssp. leporinum (Link) Arcangeli. Pd (GA, NC, VA), Cp (SC, VA): disturbed areas; rare, 

introduced from Mediterranean Europe. May. A tetraploid taxon. [= K, Z; = H. leporinum Link – RAB, C, HC; < Hordeum 

murinum Linnaeus – G, S; = Critesion murinum (Linnaeus) Á. Löve ssp. leporinum (Link) Á. Löve] 

Hordeum pusillum Nuttall, Little Barley. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, ditches, disturbed areas; common. 

April-June. Se. NY west to MN, south to n. FL, s. TX, and s. AZ. A diploid taxon. [= RAB, C, F, G, HC, K, S, W, Z; = 

Critesion pusillum (Nuttall) Á. Löve] 

* Hordeum vulgare Linnaeus, Barley. Cp, Pd (NC, SC, VA), Mt (VA): cultivated fields, occasionally persistent as a waif; 

commonly cultivated, rare as a waif, introduced from Eurasia. May-June. A diploid taxon. The original wild form is often 

treated as H. vulgare ssp. spontaneum and the cultivated, non-shattering derivative as ssp. vulgare (Hancock 2004). The wild 

form was used as a food source since at least 19,000 years ago, and ssp. vulgare developed by 8500 years ago. [= RAB, C, F, K, 

Z; > H. aegiceras Nees ex Royle – G; > H. vulgare var. vulgare – G, HC; > H. vulgare var. trifurcatum (Schlechtendahl) Alefeld 

– G, HC; > H. vulgare ssp. vulgare; > H. vulgare ssp. spontaneum (K. Koch) Thellung] 

*? Hordeum brachyantherum Nevski ssp. brachyantherum is reported for se. PA (Rhoads & Klein 1993) and also is 

apparently known from specimens from GA (Sorrie, pers. comm.). A tetraploid taxon. [= K; Critesion brachyantherum (Nevski) 

Barkworth & D.R. Dewey] {not keyed at this time} 

* Hordeum depressum (Scribner & J.G. Smith) Rydberg. Cp (SC): waste areas around wool-combing mills; rare, introduced 

from w. North America. A tetraploid taxon. [= HC, K; = Critesion depressum (Scribner & J.G. Smith) Á. Löve] {not keyed at 

this time} 

Hystrix Moench 

(see Elymus) 

Imperata Cirillo (Cogongrass, Satintail) 

A genus of about 8-9 species, of tropical, subtropical, and warm temperate areas of both hemispheres. References: Gabel in 

FNA (2003a); Ward (2004c)=Z; Hall (1998)=Y. 

* Imperata cylindrica (Linnaeus) Palisot de Beauvois, Cogongrass, Brazilian Satintail. Cp (GA, SC): grassy roadside; rare, 

introduced fom the tropics. See Nelson (1993) for first report from SC. An extremely aggressive and dangerous weed, now wellestablished 

and rapidly invading fire-maintained Coastal Plain areas (such as longleaf pine and slash pine flatwoods and longleaf 

pine clayhills) on the Gulf Coastal Plain of FL, AL, and MS. Hall (1998) argues that I. cylindrica and I. brasiliensis are not 

distinct. The only character considered to separate them is that I. brasiliensis has 1 anther and I. cylindrica has 2. Ward (2004c) 

treats the 2 taxa at varietal level. Both putative taxa are present in the Gulf Coast area of FL, GA, AL, and LA. [= Y; > I. 

cylindrica – FNA, HC, K; > I. brasiliensis Trinius – FNA, HC, K, S; > I. cylindrica var. cylindrica – Z; > I. cylindrica var. 

mexicana (Ruprecht) D.B. Ward – Z] 

A genus of about 60 species, north and south temperate. 

Koeleria Persoon (Junegrass, Koeleria) 

Koeleria macrantha (Ledebour) J.A. Schultes, Junegrass, South to DE, MD, PA, KY, and AL (Kartesz 1999). [= K; K. 

pyramidata (Lamarck) Palisot de Beavois – C] 

Lachnagrostis Trinius 1820 

A genus of about 20 species, of the Southern Hemisphere. References: Soreng et al. (2003); FNA in prep. 

* Lachnagrostis filiformis (G. Forst.) Trinius, Pacific Bentgrass. Cp (SC): waste areas around wool-combing mill; rare, 

perhaps only a waif, native of Australia. [= FNA; = Agrostis avenacea J.F. Gmelin – K] 

Lagurus Linnaeus (Hare's-tail Grass) 

A monotypic genus, of the Mediterranean region. References: Tucker (1996)=Z. 

* Lagurus ovatus Linnaeus, Hare's-tail Grass. Cp (NC): on ballast; rare, introduced from Mediterranean Europe. April-June. 

[= RAB, HC, K, Z]

POACEAE 891 

Leersia Swartz (Cutgrass) 

A genus of about 18 species, tropical and warm temperate. References: Tucker (1988)=Z. 

1 Lower panicle branches whorled or closely approximate; spikelets 4.0-5.5 mm long, 1.5-2.0 mm broad; stamens 3 ............... 

.................................................................................................................................................................................L. oryzoides 

1 Lower panicle branches alternate (rarely opposite); spikelets 2.2-5.0 mm long, 0.8-4.0 mm broad; stamens 2 or 6. 

2 Spikelets suborbicular-falcate, 3.0-4.0 mm broad, < 2× as long as broad; principal leaf-blades 10-15 mm wide; stamens 

2 ....................................................................................................................................................................L. lenticularis 

2 Spikelets narrowly elliptic-falcate, 1.0-2.0 mm broad, > 2× as long as wide; principal leaf-blades usually < 7 mm wide; 

stamens 2 or 6. 

3 Spikelets 3.8-4.7 mm long, 1.5-2.0 mm broad; panicle branches short, bearing spikelets nearly to their bases; 

stamens 6................................................................................................................................................. L. hexandra 

3 Spikelets 2.2-3.5 mm long, 0.8-1.2 mm broad; panicle branches long, filiform, the longer ones bearing spikelets 

only in their upper half; stamens 2 ........................................................................................................... L. virginica 

Leersia hexandra Swartz, Southern Cutgrass. Cp (GA, NC, SC, VA): clay-based Carolina bays, limesink ponds, lakes, 

pools, usually in places where periodically or seasonally inundated; uncommon (VA Rare). June-August. Pantropical, ranging 

north in North America to MD, TN, and TX. This species is considered a serious weed in the Old World and New World tropics; 

in our area, however, it is uncommon and not weedy. [= RAB, C, F, G, GW, HC, K, Z; = Homalocenchrus hexandrus (Swartz) 

Kuntze – S] 

Leersia lenticularis Michaux, Catchfly Cutgrass. Cp (GA, NC, SC, VA): floodplain forests and swamps; uncommon. 

September-October. Se. VA south to n. FL, west to e. TX, north in the interior to IN and MN. [= RAB, C, F, G, GW, HC, K, Z; 

= Homalocenchrus lenticularis (Michaux) Kuntze – S] 

Leersia oryzoides (Linnaeus) Swartz, Rice Cutgrass. Cp, Pd, Mt (GA, NC, SC, VA): marshes, riverbanks, pond-shores; 

common. August-October. Nova Scotia west to British Columbia, south to FL and CA; also in Europe and e. Asia. [= RAB, C, 

F, G, GW, HC, K, Z; = Homalocenchrus oryzoides (Linnaeus) Pollich – S] 

Leersia virginica Willdenow, White Grass, White Cutgrass. Cp, Pd, Mt (GA, NC, SC, VA): floodplain forests, swamps, 

streambanks; common. August-October. Québec west to MN and SD, south to FL and TX. [= RAB, C, G, GW, HC, K, Z; > L. 

virginica var. virginica – F; > L. virginica var. ovata (Poiret) Fernald – F; = Homalocenchrus virginicus (Willdenow) Britton – 

S] 

Leptochloa Palisot de Beauvois (Sprangletop, Feathergrass) 

A genus of about 30 species, pantropical and extending into warm temperate areas. The circumscription of Leptochloa has been 

controversial; many earlier authors have preferred to separate Diplachne as a separate genus. References: Snow in FNA 

(2003a); Snow (1998); Cronquist (1991). 

1 Spikelets 1-2.5 mm long, with 2-4 flowers; sheaths sparsely pilose with long, pustular-based hairs ......................................... 

............................................................................................................................................................L. panicea ssp. brachiata 

1 Spikelets 3.5-10 mm long, with 5-12 flowers; sheaths glabrous (rarely slightly scabrous). 

2 Lemmas 2-3 mm long, the apex obtuse to truncate, with the midrib often extended as a mucro .................... L. uninervia 

2 Lemmas 3-5 mm long, the apex acuminate or awned. 

3 Lemmas acuminate; leaf blades 5-10 mm wide .....................................................................................L. panicoides 

3 Lemmas awned; leaf blades 1-3 mm wide. 

4 Low sprawling grasses, < 5 dm tall; lemma awns (1-) 2.5-5 mm long; first glume 2.5-3.5 mm long; second 

glume 4-7 mm long ...............................................................................................L. fascicularis var. maritima 

4 Taller grasses, usually 5-10 dm tall; lemma awns 0.5-2.5 mm long; first glume 1.3-3.4 mm long; second 

glume 2.2-5 mm long. 

5 First glume 2.3-3.4 mm long; second glume 3.4-5.0; lemmas 4-5 mm long, with an awn 0.5-2.5 mm long 

.................................................................................................................[L. fascicularis var. acuminata] 

5 First glume 1.3-2 mm long; second glume 2.2-3.5; lemmas 3-4 mm long, with an awn 0.5-1 mm long..... 

.................................................................................................................. L. fascicularis var. fascicularis 

* Leptochloa fascicularis (Lamarck) A. Gray var. fascicularis, Bearded Sprangletop. Pd (NC): bed of artificial 

impoundment; rare, adventive from further west. September. Widespread in e. North America, primarily west of the 

Appalachians (adventive further east), and extending into South America. [= C, G; < L. fascicularis – RAB, GW, HC, S; < L. 

fusca (Linnaeus) Kunth ssp. fascicularis (Lamarck) N. Snow – FNA, K; Diplachne fascicularis (Lamarck) Palisot de Beauvois – 

F] 

Leptochloa fascicularis (Lamarck) A. Gray var. maritima (Bicknell) Gleason, Salt-meadow Grass. Cp (NC, SC, VA): 

fresh to brackish marshes, overwash flats, other disturbed brackish habitats; rare (NC Rare, VA Watch List). August-October. 

Along the coast from s. NH south to SC. This taxon appears to warrant status as a species separate from L. fascicularis.

POACEAE 892 

Reported (as L. fascicularis) for SC by Nelson & Kelly (1997). [= C, G; < L. fascicularis – RAB, GW, HC, S; = Diplachne 

maritima Bicknell – F; < L. fusca (Linnaeus) Kunth ssp. fascicularis (Lamarck) N. Snow – FNA, K] 

Leptochloa panicea (Retzius) Ohwi ssp. brachiata (Steudel) N. Snow, Red Sprangletop. Pd (GA, NC, SC, VA), Cp (NC, 

SC, VA): disturbed areas; uncommon (VA Watch List). June-October. Widespread in the Western Hemisphere. The more 

familiar name, L. filiformis, must be replaced for reasons of nomenclatural priority. [= FNA, K; < L. filiformis (Lamarck) Palisot 

de Beauvois – RAB, C, F, G, GW, HC, K, S, W] 

* Leptochloa panicoides (J. Presl) A. Hitchcock & Chase, Amazon Sprangletop. Pd (VA), Cp (GA): drawdown habitats on 

lake margins; rare, introduced from South America. Belden et al. (2004) discuss the Virginia occurrences along the banks of the 

Roanoke (Staunton) River at Kerr Reservoir. Also reported for e. GA in the Coastal Plain (Sorrie, pers. comm.). [= C, FNA, G, 

GW, HC, K; ? Diplachne halei Nash – F; ? Leptochloa floribunda Doell – S; = Diplachne panicoides (J. Presl) McNeill] 

* Leptochloa uninervia (J. Presl) A. Hitchcock & Chase. Cp (GA, SC, VA), Pd (NC): disturbed areas; rare, adventive from 

further west. July-August. Widespread in the Western Hemisphere, the native range obscure, but not likely native in our area. 

Reported for SC by Nelson & Kelly (1997). [= RAB, C, G, GW, HC, K, S; = L. fusca (Linnaeus) Kunth ssp. uninervia (J. Presl) 

N. Snow – FNA, K; = Diplachne uninervia (J. Presl) Parodi] 

* Leptochloa decipiens (R. Brown) Stapf ex Maiden ssp. peacockii (Maiden & Betche) N. Snow. Cp (SC): waif at woolcombing 

mill; rare, introduced, probably not established. [= K] {not keyed} 

* Leptochloa digitata (R. Brown) Domin. Cp (SC): waif at wool-combing mill; rare, introduced, probably not established. 


* Leptochloa divaricatissima S.T. Blake. Cp (SC): waif at wool-combing mill; rare, introduced, probably not established. [= 

K] {not keyed} 

* Leptochloa dubia (Kunth) Nees. Cp (SC): waif at wool-combing mill; rare, introduced, probably not established. Also 

reported for NC by Kartesz (1999), but the documentation indicates that it was cultivated at a Soil Conservation Service test 

nursery in Chapel Hill, Orange County. [= FNA, HC, K] {not keyed} 

* Leptochloa fascicularis (Lamarck) A. Gray var. acuminata (Nash) Gleason has been reported as adventive in PA and along 

highways in WV from halophytic habitats of w. United States (Cusick 1994). [= C, G; Diplachne acuminata Nash – F; < L. 

fascicularis – HC; < L. fusca (Linnaeus) Kunth ssp. fascicularis (Lamarck) N. Snow – FNA, K; L. acuminata (Nash) 

Mohlenbrock] {not keyed} 

* Leptochloa virgata (Linnaeus) Palisot de Beauvois, Tropical Sprangletop. Cp (SC): waif at wool-combing mill; rare, 

introduced, probably not established. [= FNA, HC, K] {not keyed} 

Leptoloma 

(see Digitaria) 

References: Brandenburg & Thieret (2000)=Z. 

Limnodea L.H. Dewey 1894 

* Limnodea arkansana (Nuttall) L.H. Dewey. Cp (SC): waste at wool-combing mill, probably not established; rare, 

introduced from sc. United States. [= HC, K, S, Z; = Cinna arkansana (Nuttall) G. Tucker] 

Lolium Linnaeus (Rye-grass, Darnel, Fescue) 

(also see Schedonorus) 

References: Darbyshire (1993)=Y; Aiken & Darbyshire (1990)=X; Tucker (1996)=Z. Key based in part on C and X. 

1 Inflorescence paniculate (spikelets borne on branches off the central axis).................................................. [see Schedonorus] 

1 Inflorescence spikelike (spikelets sessile on the central axis); [subgenus Lolium]. 

2 Glumes (12-) 15-25 mm long, subcoriaceous, equalling or surpassing the uppermost lemma (therefore the length of the 

spikelet); florets 4-9 per spikelet; annual .................................................................................................... L. temulentum 

2 Glumes 4-12 mm long, herbaceous, shorter than the lemmas (therefore shorter than the spikelet); florets (2-) 5-22 per 

spikelet; annual or perennial. 

3 Lemmas (at least the upper) awned, the awns to 15 mm long; florets 11-22 per spikelet; annual or perennial........... 

........................................................................................................................................... L. perenne var. aristatum 

3 Lemmas awnless; florets (2-) 5-10 per spikelet; perennial................................................... L. perenne var. perenne 

* Lolium perenne Linnaeus var. aristatum Willdenow, Italian Rye-grass. Cp (GA, NC, SC, VA), Pd, Mt (NC, SC, VA): 

fields, roadsides, pastures, disturbed areas; common, introduced from Eurasia. April-July. [= C, Z; = L. multiflorum Lamarck – 

RAB, F, G, HC, S; = L. perenne ssp. multiflorum (Lamarck) Husnot – K; < L. perenne – W]

POACEAE 893 

* Lolium perenne Linnaeus var. perenne, English Rye-grass, Perennial Rye-grass. Cp (GA, NC, SC, VA), Pd, Mt (GA, NC, 

VA): fields, roadsides, pastures, disturbed areas; uncommon, introduced from Eurasia. April-July. [= C, Z; = L. perenne – 

RAB, F, G, HC, S; = L. perenne ssp. perenne – K; < L. perenne – W] 

* Lolium temulentum Linnaeus, Darnel. Pd (GA, NC, VA), Cp (NC, VA), {SC}: fields, roadsides, pastures, disturbed areas; 

common, introduced from Eurasia. May-June. [= RAB, C, F, HC, S, Z; > L. temulentum var. leptochaetum A. Braun – G; > L. 

temulentum var. macrochaeton A. Braun – G; > L. temulentum ssp. temulentum – K] 

Luziola Antoine Laurent de Jussieu (Southern Water Grass) 

A genus of about 12 species, from s. North America south to tropical South America. References: Tucker (1988)=Z; Judziewicz 

et al. (2000)=Y. 

Luziola fluitans (Michaux) Terrell & H. Robinson var. fluitans, Southern Water Grass. Cp (GA, NC, SC), Pd? (GA?): 

aquatic in water of natural lakes, slow-moving blackwater rivers, and other stagnant waters; rare (NC Watch List). August- 

October. Var. fluitans ranges from ne. NC to c. FL and west to e. TX; var. oconnerii (Guzman M.) G. Tucker occurs in the 

highlands of w. Mexico (Tucker 1988). A very unusual grass, truly aquatic, with flexuous stems and unwettable, floating leaves. 

In addition to floating leaves (helpful in the field but not in the herbarium!), other useful characters include two secondary blade 

nerves on either side of the midnerve and virtually as prominent as the midnerve, and which extend onto the sheath where they 

occur with another 5 or so strong nerves; often with cilia 0.5-1 mm long at the summit of the ventral face of the sheath (an 

unusual place); and a hyaline ligule about 1 mm long on the same plane as the sheath (i.e., free from the base of diverging 

blades). [= Y, Z; < L. fluitans – K; < Hydrochloa carolinensis Palisot de Beauvois – RAB, GW, HC, S] 

Luziola bahiensis (Steudel) Hitchcock. Streams and riverbanks. Apparently native, in AL, FL, MS. See Anderson & Hall 

(1993). [= HC, K] {not keyed} 

* Luziola peruviana Gmelin, Peruvian Water Grass. Disturbed wet areas. Apparently an introduction, occurring in disturbed 

situations. See Anderson & Hall (1993). [= HC, K] {not keyed} 

Melica Linnaeus (Melic) 

A genus of about 80 species, north temperate, s. Africa and s. South America. References: Tucker (1996)=Z. 

1 First glume oblong, 6.5-10 mm long, 2-4× as long as wide, acute to obtuse at the apex, about the same length and width as 

the second glume; inflorescence with (0-) 1-5 branches from the lower nodes only; fertile lemmas 2; leaves 1-6 mm wide; 

[common species, widespread in our area] .................................................................................................................M. mutica 

1 First glume broadly ovate, 5-8 mm long, 1.5-2× as long as wide, obtuse to rounded at the apex, shorter and broader than the 

second glume; inflorescence with 2-10 (or more) branches from most nodes; fertile lemmas (2-) 3; leaves 3-12 mm wide; 

[rare species of the Mountains of NC and VA, northwards and westwards]................................................................M. nitens 

Melica mutica Walter, Two-flower Melic. Cp, Pd, Mt (GA, NC, SC, VA): forests and woodlands, including coastal fringe 

and maritime forests; common. April-May. MD west to IN and IL, south to FL and TX. [= RAB, C, F, G, HC, K, S, W, Z] 

Melica nitens (Scribner) Nuttall ex Piper, Three-flower Melic. Mt (GA, NC, VA): rocky upland woodlands, barrens, and 

glades, over calcareous rocks (such as limestone, calcareous shale); rare (NC Rare, VA Rare). May. PA west to s. MN and NE, 

south to nw. GA and TX. [= RAB, C, F, G, HC, K, W, Z] 

Melinis Palisot de Beauvois (Natalgrass) 

A genus of ca. 22 species, native to Africa and w. Asia. References: Wipff in FNA (2003a). 

* Melinis repens (Willdenow) Zizka ssp. repens, Rose Natalgrass. Cp, Pd, Mt (GA): disturbed areas, roadsides; rare, 

introduced from Africa. The report for NC by Kartesz (1999) is an error. [= FNA; < M. repens – K; ? Rhynchelytrum roseum 

(Nees) Stapf & C.E. Hubbard ex Bews – HC; < Rhynchelytum repens (Willdenow) C.E. Hubbard] 

Microstegium Nees in Lindley 1836 (Sasa-grass, Japanese-grass) 

A genus of about 15 species, of subtropical Asia and Africa. References: Barden (1987); Fairbrothers & Gray (1972); Winter, 

Schmitt, & Edwards (1982); Koyama (1987); Thieret in FNA (2003a). 

* Microstegium vimineum (Trinius) A. Camus, Flexible Sasa-grass, Japanese-grass. Pd, Mt, Cp (GA, NC, SC, VA): 

disturbed areas, colonizing moist, rich soil, especially in floodplains; common, native of tropical se. Asia. The following 

chronological synopsis of flora accounts of Microstegium is perhaps instructive: not treated by Small (1933), "local" (Fernald

POACEAE 894 

1950), "rarely introduced and possibly not established" (Gleason & Cronquist 1952), "sporadically naturalized" (Godfrey & 

Wooten 1979), "a rapidly spreading pernicious invader on moist ground, too common" (Wofford 1989). RAB report it from 

fewer than 1/3 of the counties of the Carolinas (in 1968); it is now undoubtedly in every county, an abundant weed in most of 

them. This species has become a very serious pest, now ranking as one of the most destructive introduced plants in our area, 

forming extensive and dense patches, sprawling over and eliminating nearly all other herbaceous plants. Eradication is very 

difficult, and considering its obvious colonizing abilities, only temporary. Hunt & Zaremba (1992) document the continuing 

northern expansion of Microstegium into NY and CT. Redman (1995) discusses its habitat preferences in MD and DC. Koyama 

(1987) reports it as "common as undergrowth of forests" in Japan, part of its native distribution. [= RAB, C, FNA, GW, K, W; = 

Eulalia viminea (Trinius) Kuntze – G; > Eulalia viminea var. viminea – F; > Eulalia viminea var. variabilis Kuntze – F; > M. 

vimineum var. vimineum – HC; > M. vimineum var. imberbe (Nees) Honda – HC] 

Milium Linnaeus (Wood-millet, Millet-grass) 

A genus of 3-4 species, north temperate. References: Tucker (1996)=Z; Fernald (1950b)=Y. 

Milium effusum Linnaeus var. cisatlanticum Fernald, American Wood-millet, Millet-grass. Mt (NC, VA): forests at high 

(or rarely moderate) elevations; rare (NC Rare, VA Watch List). June. A circumboreal species, ranging in North America south 

to w. NC (Swain County), e. TN (Sevier County), w. VA, WV, OH, IN, IL, and MN. The American plants are sometimes 

segregated as var. cisatlanticum Fernald (Fernald 1950b). Though considered "probably accidentally introduced and established" 

in NC by Radford, Ahles, & Bell (1968), the native occurrence of this northern species is more plausible; the only known 

occurrence in NC (not recently seen) is in the Great Smoky Mountains National Park. [= K, Y; < M. effusum – RAB, C, F, G, 

HC, W, Z] 

References: Barkworth in FNA (2003a). 

Miscanthus Andersson (Eulalia) 

* Miscanthus sinensis Andersson, Eulalia, Chinese Silver Grass. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): roadsides; 

common, introduced from e. Asia. September-November. This species is becoming aggressively weedy. Forms with leaves 

cross-variegated or linear-variegated with yellow are cultivated and sometimes escape or persist (in addition to the much more 

common green-leaved form). [= RAB, C, FNA, G, K, S, W; > M. sinensis var. variegatus Beal – F, HC; > M. sinensis var. 

zebrinus Beal – F, HC] 

Muhlenbergia Schreber 1789 (Muhly) 

A genus of about 160 species, of North America south to Andean South America, and e. and se. Asia. Muhlenbergia is a large 

and diverse genus; the various groups seem very different. References: Pohl (1969); Morden & Hatch (1989); Peterson in FNA 

(2003a). 

1 Panicle open and diffuse, > 4 cm broad, the spikelets borne on slender or capillary pedicels longer than the lemmas. 

2 Plant with rhizomes, the rhizomes prominent, creeping, and covered with imbricate scales; culms and sheaths strongly 

compressed at base, the leaves distichous; spikelets 1.5-2 mm long.............................................................. M. torreyana 

2 Plant without rhizomes, tufted with erect culms (a "bunchgrass"); culm and sheaths terete, the leaves not distichous; 

spikelets 1.5-5 mm long (excluding awns, if present); [subgenus Podosemum]. 

3 Spikelets 1.5-2 mm long, awnless..........................................................................................................[M. uniflora] 

3 Spikelets 2.5-5 mm long (excluding awns), awned or awnless. 

4 Lemma awn 0-1.5 (-4) mm long; glumes (1.1-) 2.0-3.3 (-3.6) mm long, acuminate, not awned (rarely the 

second with a short awn < 0.6 mm long); spikelets usually brown or bronze (when fresh); basal sheaths 

usually very fibrous.......................................................................................................................... M. expansa 

4 Lemma awn (2-) 3-33 mm long; glumes (0.3-) 0.7-1.7 (-2.4) mm long, one or both glumes sometimes awned; 

spikelets usually purple (when fresh); basal sheaths rarely strongly fibrous. 

5 Lemma awn (0-) 3-13 (-18) mm long, first glume awnless (or rarely with an awn to 3.2 mm long), 

second glume awnless (or rarely with an awn up to 5.0 mm long), palea awnless; lemma lacking 

setaceous teeth flanking the awn; flowering late August-October; [widespread in our area, particularly in 

rocky, clayey, or sandy glades, barrens, and woodlands with prairie affinities]......................M. capillaris 

5 Lemma awn (8-) 12-26 (-35) mm long, first glume awn (0.5-) 1-7 (-10) mm long, second glume awn (1- 

) 5-19 (-25) mm long, palea awn-tipped; lemma with two setaceous teeth flanking the awn, the teeth 0.5- 

2.5 (-4.7) mm long; flowering October-November; [of sandy maritime situations on barrier islands of the 

outer Coastal Plain] .................................................................................................................... M. sericea 

1 Panicle slender, dense, < 2.5 cm broad, the spikelets sessile or on non-capillary pedicels shorter than the lemmas; [subgenus 

Muhlenbergia].

POACEAE 895 

5 Glumes minute, 0-0.5 mm long; plant lacking rhizomes; culms weak, decumbent and cespitosely branching in their 

lower portions, rooting at the nodes, the upper portions erect and sparsely branched.....................................M. schreberi 

5 Glumes well-developed, 1-7 mm long; plant with scaly rhizomes (except for M. cuspidata); culms firm (rarely 

sprawling), few or solitary (rarely forming dense colonies). 

6 Glumes 5-7 mm long (tapered to arched or straight awns), about double the length of the lemma (excluding its 

awn); panicle dense and spike-like, 2-6 cm long and 3-10 mm broad................................................... M. glomerata 

6 Glumes 1.2-3 mm long, shorter than to barely exceeding the lemma; panicle usually slender, arching, generally 

less dense and not spike-like, often with some elongated (though appressed) branches, 4-50 cm long, 2-15 mm 

broad. 

7 Callus glabrous; plant lacking scaly rhizomes (with slender stolons and a hard, knotty crown); leaves 0.5-2 

mm wide; [of calcareous cliffs]...................................................................................................... M. cuspidata 

7 Callus bearded (sometimes only slightly so) (glabrous in M. glabriflora); plant with scaly rhizomes; leaves 

(1-) 2-14 mm wide; [collectively of various habitats]. 

8 Panicle linear, loosely flowered, much exceeding the leaves; culm erect, simple or sparingly branched; 

glumes relatively broad, the body ovate, 1.2-2.5 mm long, abruptly narrowed to the acuminate tip; ligule 

obsolete or shorter than the elongate cartilaginous summit of the leaf sheath. 

9 Lemmas awnless or awn < 0.5 mm long; spikelets 1.5-2.5 mm long; leaf blades usually (1-) 2-6 

mm wide.........................................................................................................................M. sobolifera 

9 Lemma awn 1-11 mm long (rarely awnless); spikelets 3-5 mm long; leaf blades (2) 6-10 (-13) mm 

wide (often > 8 mm wide) ..............................................................................................M. tenuiflora 

8 Panicle lanceolate, densely (rarely loosely) flowered, leaves often extending conspicuously into the 

inflorescence; culm geniculate, freely branched; glumes relatively narrow, the body lanceolate, 2-3 mm 

long, tapering from base to apex; ligule usually obvious above the short cartilaginous summit of the leaf 

sheath. 

10 Culm glabrous throughout (including below the nodes). 

11 Glumes 1.4-2.0 mm long; ligule 0.2-0.5 mm long........................................................M. bushii 

11 Glumes 2-4 (-5) mm long; ligule 0.8-1.5 mm long.................................................. M. frondosa 

10 Culm pubescent, at least below the nodes. 

12 Lemma awn 7-12 mm long; spikelets loosely clustered, on pedicels 2-4 mm long..................... 

..................................................................................................................................M. sylvatica 

12 Lemma awnless or with a short awn tip (rarely to 9 mm long); spikelets densely clustered, on 

pedicels < 1 mm long. 

13 Lemma glabrous below, or with short basal bearding; ligule 0.5-1.5 mm long ................... 

......................................................................................................................M. glabrifloris 

13 Lemma pilose basally; ligule 0.5-1 mm long.................................................. M. mexicana 

Muhlenbergia bushii Pohl, Bush's Muhly. Pd (VA), Mt (GA), {NC}: bottomlands and other moist forests; rare (VA Rare). 

IN west IA, south to NE and TX; apparently disjunct eastward in scattered localities, including in n. GA (Jones & Coile 1988) 

and VA. The habitat is variously given in floras as "dry woods" or "moist woods." [= C, FNA, K; = M. brachyphylla Bush – F, 

G, HC] 

Muhlenbergia capillaris (Lamarck) Trinius, Hairgrass. Pd, Cp (GA, NC, SC, VA), Mt (GA, NC, VA): in the Piedmont 

primarily in clayey or thin rocky soils (especially in areas which formerly burned and were prairie-like), open woodlands, in the 

Coastal Plain in savannas, dry woodlands, and coastal grasslands (where sometimes in close proximity with M. sericea), in the 

mountains around calcareous rock outcrops; uncommon. Late August-October. The species is widespread in e. North America. 

M. capillaris and its relatives, M. expansa and M. sericea, have been the subject of an herbarium morphological study by Morden 

& Hatch (1989), who conclude that the three taxa are not sharply separable and should be recognized only at the varietal level. If 

one considers behavior in the field, ecology, and geography in conjunction with morphologic characters, however, there is little 

doubt that the three taxa are biological species. Distribution and typical habitat are different for the three species, but M. 

capillaris can be found growing with or in proximity to each of the other two (I have not seen M. sericea and M. expansa 

together). In such situations, the two taxa present are readily distinguishable at a glance, and there is no evidence of 

intermediates or hybrids. [= F, FNA, G, W; < M. capillaris – RAB, GW (also see M. sericea); = M. capillaris var. capillaris – 

C, HC, K, S] 

Muhlenbergia cuspidata (Torrey ex Hooker) Rydberg, Plains Muhly. Mt (VA): dolomite and limestone palisade cliffs 

along the New, Roanoke, and Shenandoah rivers; rare (VA Rare). OH west to MT and Alberta, south to sw. VA, KY, MO, OK, 

and NM. [= C, F, FNA, G, HC, K] 

Muhlenbergia expansa (Poiret) Trinius, Savanna Hairgrass. Cp (GA, NC, SC, VA): pine savannas, pine flatwoods, mesic 

areas in sandhill-pocosin ecotones; common (VA Rare). September-October. An important part of the grassy component of 

many longleaf pine savannas, M. expansa is a Coastal Plain species, ranging from se. VA south to FL, and west to e. TX (nearly 

exactly the range of Pinus palustris). Its flowering is stimulated by fire, and, lacking fire, it may be found in large populations in 

solely vegetative condition. It can be distinguished in sterile condition from other savanna bunchgrasses (Sporobolus teretifolius, 

S. pinetorum, S. floridanus, S. curtissii, Aristida stricta, and A. beyrichiana) by the following characteristics: old leaf bases 

fibrous and curly (rather than hardened and cartilaginous), ligules 1-3 mm long (rather than 0.2 to 0.5 mm long). The open 

panicle somewhat resembles that of several species of similar habitat which often co-occur with M. expansa – Sporobolus 

teretifolius, S. pinetorum, S. curtissii, S. floridanus, and Calamovilfa brevipilis, but the panicle of M. expansa is capillary,

POACEAE 896 

flexuous, and fragile, tending to break up over the winter (vs. fine-textured but not capillary, the branches rigid and ascending, 

more likely to persist over the winter in relatively intact condition). The vegetative characters listed above and under 

Calamovilfa brevipilis are also useful. See M. capillaris for discussion of Morden & Hatch (1989) advocating varietal status for 

the three taxa in the M. capillaris-expansa-sericea complex. [= RAB, F, FNA, GW, HC, S; = M. capillaris var. trichopodes 

(Elliott) Vasey – C, K] 

Muhlenbergia frondosa (Poiret) Fernald, Smooth Wirestem Muhly. Mt, Pd (GA, NC, VA), Cp (NC): moist forests and 

disturbed areas; uncommon (rare in Piedmont and Coastal Plain). September-October. This species is widespread in e. North 

America, south to ne. GA and west into the Plains. [= RAB, C, F, FNA, G, GW, HC, K, W; = M. mexicana – S, misapplied] 

Muhlenbergia glabriflora Scribner, Clay-pan Muhly. Pd (NC, VA): in clayey soils (such as those derived from diabase); 

rare (NC Watch List, VA Rare). October-November. VA and NC west to IA, MO, AL, and TX, local and apparently rare in all 

of that range. In NC, only known from one collection, that from Durham County in 1936, with vague habitat data. F describes 

the habitat as "dry exsiccated or baked soils, prairies, gravels or rocky slopes," Pohl (1969) as "mostly on low ground, in shade on 

heavy clay soils." [= C, F, G, HC, K; = M. glabrifloris – FNA, orthographic variant] 

Muhlenbergia glomerata (Willdenow) Trinius, Spiked Muhly. Mt (NC, VA): fens and seeps over mafic (amphibolite) or 

ultramafic (olivine) rocks; rare (NC Rare, VA Rare). August-October. This species is widespread in n. North America, ranging 

south in a scattered and disjunct pattern to NC. [= RAB, C, F, FNA, GW, HC, K, W; < M. racemosa (Michaux) Britton, Sterns, 

& Poggenburg – G, S] 

Muhlenbergia mexicana (Linnaeus) Trinius, Hairy Wirestem Muhly. Mt (NC, VA), Pd (VA): forest edges; rare (NC 

Watch List). September-October. The epithet is a misnomer; the species is largely northern, occurring nearly throughout the 

United States and s. Canada. [= RAB, C, F, G, HC, K, W; > M. mexicana var. filiformis (Torrey) Scribner – FNA; > M. 

mexicana var. mexicana – FNA; = M. foliosa (Roemer & J.A. Schultes) Trinius – S] 

Muhlenbergia schreberi J.F. Gmelin, Nimblewill, Dropseed. Mt, Pd, Cp (GA, NC, SC, VA): bottomland and other moist 

forests, disturbed areas; common. August-October. This species is widespread in e. United States. [= RAB, C, F, FNA, GW, 

HC, K, S, W; > M. schreberi var. schreberi – G; > M. schreberi var. palustris (Scribner) Scribner – G; > M. palustris Scribner] 

Muhlenbergia sericea (Michaux) P.M. Peterson, Dune Hairgrass, Sweet Grass. Cp (GA, NC, SC): maritime dry 

grasslands, maritime wet grasslands, interdune swales, low dunes, sometimes edges of freshwater or brackish marshes, apparently 

limited to the barrier islands (sometimes in close proximity with M. capillaris); uncommon, though sometimes locally abundant 

(SC Rare). October-November. This species is a very conspicuous part of the Outer Banks flora in the autumn, especially showy 

and abundant between Rodanthe (Chicamacomico) and Avon (Kinnakeet), Dare County, NC, and also abundant on Ocracoke 

Island, Hyde County, NC. The capillary pedicels and awns of its purple inflorescences are so light as to be moved by the 

slightest breeze. By December or January they fade to tan, but remain showy. This grass is a major component of baskets made 

in the Low Country of SC by the Gullah, who call it "sweet grass." I agree with Curtis (1843), Blomquist (1948), Pinson & 

Batson (1971), Gould (1975), and others who consider M. sericea (as M. filipes) a species distinct from M. capillaris. Its range is 

from NC (slightly north of Oregon Inlet, Dare County, south of Nags Head) south to FL and west to TX, primarily on barrier 

islands. In addition to a discussion of its relationship to M. capillaris, Pinson and Batson (1971) and Morden & Hatch (1989) 

provide descriptions, not elsewhere available. See M. capillaris for a discussion of a recent paper (Morden & Hatch 1989) 

advocating varietal status for the three taxa in the M. capillaris-expansa-sericea complex. [= FNA; < M. capillaris – RAB, GW; 

= M. capillaris var. filipes (M.A. Curtis) Chapman ex Beal – HC, K, S; = M. filipes M.A. Curtis] 

Muhlenbergia sobolifera (Muhlenberg ex Willdenow) Trinius, Rock Muhly. Mt (GA, NC, VA), Pd (VA): dry wooded 

limestone slopes, rock outcrops and rocky forests; uncommon (GA Special Concern, NC Watch List). July-September. This 

species is widespread in e. United States, south to AL. [= RAB, C, F, FNA, G, HC, K, S, W] 

Muhlenbergia sylvatica Torrey ex A. Gray, Woodland Muhly. Mt (GA, NC, SC, VA), Pd (NC, VA), Cp (VA): bottomland 

and other moist forests, calcareous strembanks; rare (GA SPecial Concern, NC Watch List). September. This species is 

widespread in e. United States, south to ne. GA. [= RAB, C, FNA, K, W; > M. sylvatica var. sylvatica – F, G, GW, HC; = M. 

umbrosa Scribner – S] 

Muhlenbergia tenuiflora (Willdenow) Britton, Sterns, & Poggenburg, Slender Muhly. Mt, Pd (GA, NC, VA), Cp (VA), 

{SC}: moist forests and disturbed areas, up to at least 1400m; uncommon (rare in Piedmont). August-October. This species is 

widespread in e. United States. Two varieties are sometimes recognized: var. tenuiflora, with lemma awn 4-11 mm long and the 

sheaths and stems retrorsely hirsute, especially around the nodes, and var. variabilis (endemic to the Southern Appalachians), 

with lemma awn 1-4 mm long or absent, and the sheaths and stems glabrous or nearly so. The validity of the varieties needs 

further assessment. [= RAB, F, FNA, G, HC, K, S, W; > M. tenuiflora var. tenuiflora – C; > M. tenuiflora var. variabilis 

(Scribner) Pohl – C] 

Muhlenbergia torreyana (J.A. Schultes) A.S. Hitchcock, Pinebarren Smokegrass. Cp (GA, NC): moist soils of depression 

meadows and clay-based Carolina bays, often under or near Taxodium ascendens; rare (GA Special Concern, NC Endangered). 

August-November. NJ to GA in the Coastal Plain, and disjunct in KY and TN; currently known to be extant only in NJ, NC, and 

TN. It was first discovered in NC in 1987. Although it rarely flowers except following fire, it can be recognized in sterile 

condition by its forming clonal patches with evenly spaced, upright, blue-green tufts, each tuft a flattened stem with 5-10 

ascending-erect, rather stiff, usually conduplicate leaves, the summit of each sheath with a pronounced cartilaginous thickening, 

easily felt by running the flattened stem from base to apex between thumb and forefinger. [= C, F, FNA, G, HC, K; = 

Sporobolus torreyanus (J.A. Schultes) Nash – S] 

Muhlenbergia asperifolia (Nees & Meyen ex Trinius) Parodi, Alkali Muhly, Scratchgrass. Alkaline soils, wetlands, lawns. 

Reported east and south to MD, PA, and OH (Kartesz 1999). [= C, F, FNA, G, HC, K] {not keyed at this time}

POACEAE 897 

* Muhlenbergia emersleyi Vasey, Bull Muhly, is reported as introduced in NC (Kartesz 1999), based on a specimen at the 

UNC Herbarium. However, the specimen makes clear that it was cultivated at a Soil Conservation Service test nursery; there is 

no evidence that the species is established in our area. [= FNA, HC, K] {not keyed} 

Muhlenbergia uniflora (Muhlenberg) Fernald. Bogs, wet meadows. South to NJ and se. PA (Rhoads & Klein 1993). [= C, 

FNA, G, HC, K; > M. uniflora var. uniflora – F] {not keyed at this time} 

Nassella (Trinius) Desvaux 

* Nassella leucotricha (Trinius & Ruprecht) Pohl, Texas Needlegrass. Cp (SC): waste areas near wool-combing mill; rare, 

introduced from sc. United States and Mexico. [= K; = Stipa leucotricha Trinius & Ruprecht – HC] 

Neeragrostis Bush 

A monotypic genus of warm temperate North America and tropical Central America and n. South America. References: 

Peterson & Harvey (in prep.)=Z. 

Neeragrostis reptans (Michaux) Nicora, is reported for scattered locations as far east as c. TN by Chester et al. (1993), as 

well as in WV, KY, and possibly GA (Kartesz 1999). [= K, Z; = Eragrostis reptans (Michaux) Nees – C, F, G, GW, HC] 

Oplismenus Palisot de Beauvois (Woods-grass, Basket-grass) 

A genus of about 5 species, widespread in the New World and Old World tropics, subtropics, and warm temperate areas. 

References: Wipff in FNA (2003a); Crins (1991)=Z; Scholz (1981)=Y; Peterson et al. (1999). 

1 Sheath and culm axis glabrous or with a few scattered hairs, the hairs < 1 mm long; lemma (7-) 9-11-veined.......................... 

............................................................................................................................................................. O. hirtellus ssp. setarius 

1 Sheath and culm axis noticeably pilose, the hairs 1-3 mm long; lemma 7-veined ...................................................................... 

................................................................................................................................................ [O. hirtellus ssp. undulatifolius] 

Oplismenus hirtellus (Linnaeus) Palisot de Beauvois ssp. setarius (Lamarck) Mez ex Ekman, Woods-grass. Cp (GA, NC, 

SC), Pd (GA, SC): maritime forests, shell middens, moist forests; uncommon (NC Watch List). August-October. O. hirtellus is 

widespread in tropical and subtropicals areas of the New and Old World; ssp. setarius ranges from e. NC south to FL, west to AR 

and TX, and south through the Caribbean and Central America to central South America. Scholz (1981) recognizes many other 

sspp. This species is undoubtedly native in our area, occurring in undisturbed habitats in natural communities entirely devoid of 

alien species; the basis of Gould's (1975) assertion that Oplismenus is "introduced or adventive in the United States" is unknown. 

Superficially, Oplismenus resembles Arthraxon, but has the leaves only slightly cordate at the base (vs. strongly cordateclasping). 

Crins (1991) favors treating O. setarius as a taxonomically unrecognized component within a polymorphic O. 

hirtellus. [= FNA, K, Y; = O. setarius (Lamarck) Roemer & J.A. Schultes – RAB, HC, S; < O. hirtellus (Linnaeus) Palisot de 

Beauvois – Z] 

* Oplismenus hirtellus (Linnaeus) Palisot de Beauvois ssp. undulatifolius (Ard.) U. Scholz, native to the Eastern Hemisphere, 

has been reported as an introduction in Baltimore Co., MD (Peterson et al. 1999). It can be expected to spread, and may likely be 

found in our area. [= FNA, K, Y; < O. hirtellus (Linnaeus) Palisot de Beauvois – Z] 

Oryza Linnaeus (Rice) 

A genus of about 20 species, native of tropical and warm temperate portions of the Old World. References: Tucker (1988)=Z; 

Judziewicz et al. (2000)=Y; Nanda & Sharma (2003=X. 

* Oryza sativa Linnaeus, Rice. Cp (GA, NC, SC): escaped in marshes (probably not truly naturalized); rare, native of Asia. 

October. Perhaps the single most important food crop in the world, developed as a crop in Asia and cultivated at least since 

10,000 years BP (Hancock 2004). Rice was an important crop before the Civil War in SC and extreme se. NC. [= RAB, C, G, 

GW, HC, K, S, X, Y, Z] 

Oryzopsis Michaux (Ricegrass) 

(also see Piptatherum) 

1 Leaves involute when dry, 1-2 mm wide; glumes 3.5-4.8 mm long ............................................ [see Piptatherum canadense] 

1 Leaves flat, 5-15 mm wide; glume 6-9 mm long.

POACEAE 898 

2 Leaves primarily basal or low-cauline, 2-4 (-5) dm long, 4-10 mm wide; inflorescence a raceme or a racemiform 

panicle; culms prostrate, the upper leaves very reduced, often merely bladeless sheaths ............................. O. asperifolia 

2 Leaves primarily cauline, 1-2.5 dm long, 8-15 mm wide; inflorescence a panicle; culms erect, the upper leaves welldeveloped........................................................................................................................... 

[see Piptatherum racemosum] 

Oryzopsis asperifolia Michaux, Rough-leaved Ricegrass, Whiteseed Mountain-ricegrass. Mt (VA): high elevation pineoak/heath 

barrens and woodlands; rare (VA Rare). Newfoundland west to British Columbia, south to w. VA, WV, n. IN, SD, 

NM, and UT. This grass forms large cespitose clumps, the leaves evergreen and somewhat bicolored (green on the upper surface, 

bluish on the lower). [= C, F, G, HC, K] 

Panicum Linnaeus 1753 (Panic Grass) 

(also see Dichanthelium, Phanopyrum, Setaria, Steinchisma, and Urochloa) 

(contributed by Richard J. LeBlond) 

{INTRODUCTION: Describe differences between Panicum, Dichanthelium, Urochloa (=Brachiaria), and Paspalidium (now in 

Setaria), all of which are treated as Panicum in RAB. Describe collection methods and character analysis.} 

There has been considerable controversy over the generic limits of Panicum. In its broader recent conceptions, it has been 

considered to include (in our area) taxa sometimes and variously segregated as Brachiaria, Dichanthelium, Eriochloa, 

Paspalidium, Phanopyrum, Steinchisma, and Urochloa. All were originally recognized based on morphological characteristics, 

to which have recently been added anatomical, chemical, and other evidence. Crins (1991) recognizes Eriochloa, Urochloa 

(including Brachiaria), Paspalidium, and Panicum as genera, with Panicum subdivided into subgenera Panicum, Agrostoides, 

Dichanthelium, Phanopyrum, and Steinchisma. We prefer to recognize most of the segregates as genera, pending further 

analyses, since there is little evidence that these groups are more closely related to one another than they are to other genera 

recognized in the Paniceae. Phanopyrum and Dichanthelium are the only segregate groups with C 3 photosynthesis. Eriochloa 

and Urochloa (including Brachiaria) have C 4 photosynthesis, with PEP-ck decarboxylation. Panicum and Setaria (Paspalidium) 

have C 4 photosynthesis, with NAD-me or NADP-me decarboxylation. Steinchisma, in addition to its unusual expansion of the 

palea, apparently has a peculiar photosynthetic pathway, described by Crins (1991) as "intermediate between" C 3 and C 4 

photosynthesis; "the leaves have Kranz anatomy, but there are fewer organelles than usual in the outer sheath." 

We agree with Hansen & Wunderlin (1988) that "Dichanthelium is as 'good' a grass genus as many others (e.g. Brachiaria, 

Sacciolepis, and many more in other tribes)." Despite arguments to the contrary, there is little doubt that Dichanthelium is a 

natural group. Zuloaga, Ellis, and Morrone (1993) argue against the recognition of Dichanthelium as a genus, preferring to treat 

it as a subgenus under Panicum. They state, however, "within Panicum, Dichanthelium can be distinguished at the subgeneric 

level by the following set of characters: lax inflorescences; ellipsoid to obovoid spikelets; upper glume and lower lemma usually 

7-11 nerved; upper anthecium apiculate or shortly crested, and simple papillae on the lemma and palea. Anatomically, all species 

are non-Kranz or C 3 , with the outer parenchymatous sheath lacking specialized chloroplasts", etc. The argument that 

Phanopyrum also has C 3 photosynthesis does not materially affect the issue of the taxonomic rank at which to recognize the 

groups. 

We also agree with Hansen & Wunderlin (1988) that "the acceptance of Dichanthelium provides a more consistent generic 

classification." It offers conveniences, as well, in our area, where Dichanthelium and Panicum are readily distinguishable from 

each other, and the combined genus would be very large, indeed. References: Lelong (1986)=Z; Zuloaga & Morrone (1996)=Y; 

Freckmann & Lelong in FNA (2003a). 

1 Spikelets tuberculate ............................................................................................................................................P. verrucosum 

1 Spikelets smooth, not tuberculate. 

2 First glume 5-7.5 mm long, nearly as long as sterile lemma; fertile lemma 1/3 length of sterile lemma ............................ 

............................................................................................................................................. [Phanopyrum gymnocarpon] 

2 First glume shorter, or if this long, then at most 3/4 length of sterile lemma; fertile lemma > ½ the length of the sterile 

lemma. 

3 Sterile palea indurate and expanding the spikelet at maturity, as long as sterile lemma; outer surface of the distal 

palea with compound papillae.....................................................................................................[Steinchisma hians] 

3 Sterile palea membranous, not expanding the spikelet at maturity, usually shorter than sterile lemma or absent; 

outer surface of the distal palea lacking compound papillae. 

4 Panicle < 2 cm wide at maturity. 

5 Spikelets >4.5 mm long; first glume > 2.4 mm long; ligule 4-6 mm long; [of coastal dunes]; [subgenus 

Panicum, section Repentia]................................................................................. P. amarum var. amarum 

5 Spikelets < 4 mm long; first glume < 2.1 mm long; ligule < 2 mm long; [not of coastal dunes]. 

6 Blades involute, 1.5-4 mm wide; culms wiry; [subgenus Agrostoides, section Tenera]...................... 

........................................................................................................................................... P. tenerum 

6 Blades flat, the larger 6-20 mm wide; culms stout.

POACEAE 899 

7 Panicles constricted, 0.3-1.6 cm wide; spikelets subsessile to short-pediceled; summit of fertile 

palea not enclosed by fertile lemma ......................................................................P. hemitomon 

7 Panicles > 1 cm wide; spikelets short to long-pediceled; summit of fertile palea enclosed by 

fertile lemma; [subgenus Agrostoides, section Agrostoidea]. 

8 Plants tufted, without rhizomes; culms strongly compressed below; fertile lemma 1.3-1.5 

mm long................................................................................P. rigidulum var. condensum 

8 Plants rhizomatous; culms slightly compressed below; fertile lemma 1.8-2.2 mm long. 

9 Rhizomes short, usually < 3 cm long; leaves 20-50 cm long, 4-18 mm wide; 

spikelets 2.5-3.9 mm long, acuminate; first glume with 3-5 green nerves................... 

.................................................................................................. P. anceps var. anceps 

9 Rhizomes elongate, often > 4 cm long; leaves 10-30 (-40) cm long, 2-10 mm wide; 

spikelets 2.2-2.8 mm long, acute to short-acuminate; first glume with 1-3 green 

nerves................................................................................P. anceps var. rhizomatum 

4 Panicle > 2 cm wide at maturity. 

10 Plants from a cluster of fibrous roots, without rhizomes or hard knotty crowns, annual. 

11 First glume 1/5 to 1/4 length of spikelet, broadly rounded to truncate; sheaths usually glabrous; 

nodes glabrous; [subgenus Panicum, section Dichotomiflora]............................................................ 

..........................................................................................P. dichotomiflorum var. dichotomiflorum 

11 First glume 1/3 to 1/2 length of spikelet, acute to subacute; sheaths villous or hispid; nodes often 

bearded; [subgenus Panicum, section Panicum]. 

12 Spikelets 4.5-6 mm long; panicle branches often nodding or drooping at maturity..................... 

................................................................................................................................P. miliaceum 

12 Spikelets 1.8-3.6 mm long; panicle branches ascending-spreading at maturity. 

13 Spikelets long-acuminate, (2.6-)3.0-3.6 mm long; mature panicle slender, usually 2-3 

times as long as wide.............................................................................................P. flexile 

13 Spikelets short-pointed to acuminate, 1.8-2.5 (-2.8) mm long; mature panicle usually > ½ 

as wide to wider than long. 

14 Panicle usually equal to or longer than culm; largest blades usually 10-20 mm wide; 

spikelets acuminate, lanceolate to lance-ovoid ..........................................P. capillare 

14 Panicle usually not as long as culm; largest blades usually 10 mm or less wide; 

spikelets short-pointed, ellipsoid, ovoid, or obovoid. 

15 Herbage purple-tinged; blades 2-6 mm wide, ascending; pulvini glabrous to 

sparsely pilose; spikelets 1.8-2.2 mm long, > 2× as long as wide; mature fertile 

lemma blackish .............................................................................P. lithophilum 

15 Herbage yellow-green to green; blades 2-12 mm wide, spreading; pulvini 

glabrous or pilose; spikelets 1.4-2.4 mm long, < 2× as long as wide; mature 

fertile lemma straminous. 

16 Culm blades 5-12 mm wide; blade of flag (inflorescence bract) usually > 

½ as long as panicle; panicle ellipsoid to obovoid; pulvini glabrous; 

secondary panicle branches and pedicels divergent; spikelets 1.9-2.4 mm 

long .......................................................................................... P. gattingeri 

16 Culm blades 2-6 mm wide; blade of flag usually < ½ as long as panicle; 

panicle broadly ovoid to deltoid; pulvini pilose; secondary panicle 

branches and pedicels appressed; spikelets 1.4-2.1 mm long....................... 

........................................................................................P. philadelphicum 

10 Plants with rhizomes or hard knotty crowns, perennial. 

17 Plants with rhizomes; fertile lemma 1.6-4 mm long. 

18 First glume truncate apically ........................................................................................ P. repens 

18 First glume acute to obtuse. 

19 Culms slightly compressed below; ligules 0.5 mm long or less; spikelets subsessile and 

subsecund, usually some obliquely bent above the first glume; fertile lemma 1.8-2.2 mm 

long; [subgenus Agrostoides, section Agrostoidea]. 

20 Rhizomes short, usually < 3 cm long; leaves 20-50 cm long, 4-18 mm wide; 

spikelets 2.5-3.9 mm long, acuminate; first glume with 3-5 green nerves................... 

.................................................................................................. P. anceps var. anceps 

20 Rhizomes elongate, often > 4 cm long; leaves 10-30 (-40) cm long, 2-10 mm wide; 

spikelets 2.2-2.8 mm long, acute to short-acuminate; first glume with 1-3 green 

nerves................................................................................P. anceps var. rhizomatum 

19 Culms terete; ligules 1-6 mm long; spikelets pediceled and not at all secund, essentially 

straight; fertile lemma 2-4 mm long; (subgenus Panicum, section Repentia]. 

21 Panicle narrow, the branches erect; sheaths longer than internodes; spikelets 4.3-7.7 

mm long; fertile lemma 3-4 mm long. 

22 Rhizomes usually elongate; culms solitary to loosely tufted, 0.2-1.5 m tall; 

leaves 0.7-3.6 dm long; panicles 2-6 cm wide, the primary branches usually 1-2

POACEAE 900 

per node, loosely flowered; spikelets 4.7-7.7 mm long; first glumes 2.5-5.5 mm 

long, 2/3-3/4 as long as the spikelet, 7-9 nerved, the nerves thickened and 

raised; fertile lemma 1.3-1.8 mm wide ........................ P. amarum var. amarum 

22 Rhizomes usually short; culms usually tufted, 1-2 (-3) m tall; leaves 2-5 dm 

long; panicles 3-10 cm wide, the primary branches usually 2 or more per node, 

densely flowered; spikelets 4.0-5.9 mm llong; first glumes 2-3.5 mm long, ½- 

2/3 as long as the spikelet, 3-5 (-7) nerved, the nerves thin and wiry; fertile 

lemma 1.0-1.5 mm wide ...................................................................................... 

..................................................................................P. amarum var. amarulum 

21 Panicle with divergent to spreading-ascending branches; upper sheaths shorter than 

internodes; spikelets 2.8-5 mm long; fertile lemma 2-2.6 mm long. 

23 Spikelets 2.8-3.5 mm long; first glume 1/2 length of spikelet, blunt to acute ...... 

.....................................................................................P. virgatum var. cubense 

23 Spikelets 3.5-5 mm long; first glume 2/3 length of spikelet, acuminate .............. 

................................................................................... P. virgatum var. virgatum 

17 Plants with hard crowns, lacking rhizomes; fertile lemma 1.2-1.6 mm long; [subgenus Agrostoides, 

section Agrostoidea]. 

24 Ligule of white hairs 0.5-3 mm long; culms to 1 m long; cauline blades 2-8 mm wide, usually 

pilose adaxially near the base; spikelets 2.0-4.0 mm long. 

25 Ligules 1-3 mm long; spikelets 2.0-2.7 mm long, 2.5-4× as long as wide, often obliquely 

set on the pedicels..............................................................P. longifolium var. longifolium 

25 Ligules 0.5-1.5 mm long; spikelets 2.4-4.0 mm long, 3.5-5× as long as wide, erect on the 

pedicels.................................................................................... P. longifolium var. combsii 

24 Ligule a tawny membrane 0.5-1.0 mm long, often erose or lacerate, or with a minute ciliate 

fringe; culms to 1.8 m long; cauline blades 4-12 mm wide, usually glabrous; spikelets 1.6-2.8 

mm long. 

26 Spikelets 2.4-2.8 mm long, long-acuminate, usually < 0.7 mm wide; fertile lemma often 

conspicuously stipitate........................................................... P. rigidulum var. elongatum 

26 Spikelets 1.6-2.5 mm long, short-acuminate, usually > 0.7 mm wide; fertile lemma 

estipitate to short stipitate. 

27 Culms to 1 m long; mature panicle ½ to nearly as wide as long, the branches 

ascending to spreading; spikelets 1.6-2.2 mm long......... P. rigidulum var. rigidulum 

27 Culms to 1.8 m long; mature panicle < 1/3 as wide as long, the branches erect; 

spikelets 2.0-2.5 mm long.............................................P. rigidulum var. condensum 

Panicum amarum Elliott var. amarulum (A.S. Hitchcock & Chase) P.G. Palmer, Southern Seabeach Grass. Cp (GA, NC, 

SC, VA): coastal dunes and shores, sandflats, and sandhills; rare. July-November. NJ s. to FL and West Indies, w. to TX and 

Mexico; restricted to the Coastal Plain except for WV. Although well-marked individuals of var. amarulum and var. amarum are 

quite distinctive, only the number and structure of first glume nerves appears to be a constant over the range of the two taxa 

(Palmer 1975). Primarily a coastal plant, var. amarulum has been found in the Sandhills of NC (Richmond Co.). Blomquist 

1948 says this taxon "does not seem to grow naturally in North Carolina." [= K, Z; = P. amarulum A.S. Hitchcock & Chase – 

RAB, C, F, G, HC, S; = P. amarum ssp. amarulum (A.S. Hitchcock & Chase) Freckmann & Lelong – FNA; not Panicum] 

Panicum amarum Elliott var. amarum, Bitter Seabeach Grass. Cp (GA, NC, SC, VA): coastal dunes and shores; common. 

August-November. CT s. to FL, w. to TX; restricted to the coast. See note under var. amarulum. [= K, Z; = P. amarum – RAB, 

C, F, G, HC, S; = P. amarum ssp. amarum – FNA; not Panicum] 

Panicum anceps Michaux var. anceps, Beaked Panic Grass. Mt, Pd, Cp (NC, SC, VA), {GA}: moist sandy woods, 

swamps, sloughs, roadsides, fields, waste places; common. June-October. NJ w. to IL, s. to FL and TX. The sheaths of var. 

anceps are glabrous to pilose, while those of var. rhizomatum are often villous; the leaves of var. rhizomatum also tend to be 

hairier. [= RAB, F, G, Z; < P. anceps – C, GW, K, W; = P. anceps ssp. anceps – FNA; = P. anceps – HC, S; not Panicum] 

Panicum anceps Michaux var. rhizomatum (A.S. Hitchcock & Chase) Fernald, Small Beaked Panic Grass. Cp (GA, NC, 

SC, VA): moist to dry sandy or loamy pinelands, ditches; common (VA Watch List). July-October. Se. VA and KY s. to FL 

and TX. See note under var. anceps. [= RAB, F, G, Z; < P. anceps – C, GW, K; = P. anceps ssp. rhizomatum (A.S. Hitchcock 

& Chase) Freckmann & Lelong – FNA; = P. rhizomatum A.S. Hitchcock & Chase – HC, S; not Panicum] 

Panicum capillare Linnaeus, Old-witch Grass, Tumbleweed, Tickle Grass. Mt, Pd (GA, NC, SC, VA), Cp (VA): open 

sandy or stony soil, fields, roadsides, waste places, often weedy in cultivated soil; common (rare in SC). August-November. E. 

to c. Canada, s. to FL and TX; Bermuda. Plants formerly known as P. capillare var. occidentale Rydberg, ranging from Canada 

south to NJ, WV, KY, TX, and CA, are distinguished by long-acuminate spikelets 2.5-4 mm long that are mostly subsessile or 

short-pedicelled. In our region, P. capillare has short-acuminate spikelets 1.8-2.8 mm long, mostly on long pedicels. [= RAB, 

K, S, Z; < P. capillare – C, Y (also see P. gattingeri); > P. capillare var. capillare – F, HC, W; = P. capillare ssp. capillare – 

FNA; = P. capillare var. agreste Gattinger – G; Panicum s.s.] 

Panicum dichotomiflorum Michaux var. dichotomiflorum, Spreading Panic Grass, Fall Panic Grass. Mt, Pd, Cp (GA, NC, 

SC, VA): marshy shores, exposed wet soils, alluvial deposits in floodplain forests, spoil banks, ditches; common. July-October. 

E. Canada w. to SD, s. to FL and TX; also in the Bahamas (Sorrie & LeBlond 1997). P. dichotomiflorum var. puritanorum 

ranges along the coast from s. NH to DE, and occurs inland in n. IN. It is distinguished by ovoid to ellipsoid, abruptly short-

POACEAE 901 

tipped spikelets only 1.8-2.2 mm long, culms to 6 dm long, and leaves 1-8 mm wide. In var. dichotomiflorum, the oblonglanceolate, 

acuminate spikelets are (2.0-) 2.6-3.6 mm long, culms to 2 m long, and leaves 4-20 mm wide. Plants with geniculate 

bases, enlarged lower nodes and sheaths, and panicles with included peduncles and divergent branches have been recognized as 

var. geniculatum (A. Wood) Fernald. Plants with spikelets similar to those of var. puritanorum, but with culm and leaf features 

of var. dichotomiflorum, have been recognized as var. imperiorum Fernald, and are known only from se. VA. Recognition of any 

infraspecific taxa in this morphologically complex species is risky business. [= HC, K; < P. dichotomiflorum – RAB, C, GW, S, 

Z; > P. dichotomiflorum var. dichotomiflorum – F, G, W; > P. dichotomiflorum var. geniculatum – F, G, W; > P. 

dichotomiflorum var. imperiorum – F; Panicum s.s.] 

Panicum flexile (Gattinger) Scribner, Wiry Panic Grass. Pd (GA, NC, SC, VA), Mt (GA, NC,VA): glades and openings 

over mafic rocks, damp sandy meadows, open woods; rare (NC Rare). July-October. NY, sw. Québec, S. Ontario, and ND south 

to FL and TX. First reported for SC by Nelson & Kelly (1997). [= RAB, C, F, FNA, G, HC, K, S, W, Y, Z; Panicum s.s.] 

Panicum gattingeri Nash, Gattinger's Panic Grass. Mt, Pd (NC, VA), {GA}: damp or dry, usually calcareous sandy soils 

of fields, roadsides, shores, and cultivated ground; frequent in Mountains, uncommon in Piedmont (VA Watch List). August- 

October. NY, sw. Québec, and MN south to NC, TN, GA, AL, and AR. [= RAB, F, HC, K, S; < P. capillare – C, Y; = P. 

philadelphicum Bernhardi ex Trinius ssp. gattingeri (Nash) Freckmann & Lelong – FNA; = P. capillare Linnaeus var. campestre 

Gattinger – G, W; Panicum s.s.] 

Panicum hemitomon J.A. Schultes, Maidencane. Cp (GA, NC, SC, VA), Mt (VA): lake, pond, and river shores, swamp 

borders, marshes, ditches, often in shallow water; common (VA Rare). June-July. Coastal Plain from s. NJ south to FL, west to 

TX; also TN; South America. Often forming dense colonies in the low margin and shallow waters of limesink ponds. [= RAB, 

C, F, FNA, G, GW, HC, K, S, W, Z; not Panicum] 

Panicum lithophilum Swallen, Flatrock Panic Grass. Pd (GA, NC, SC), Mt (NC): soil islands on granitic flatrocks and 

domes; rare (NC Rare). August-October. Restricted to granite outcrops in NC, SC, and ec. GA. There is some question about 

the distinctness of this taxon from P. philadelphicum; Zuloaga & Morrone (1996) did not consider it separable from P. 

philadelphicum. [= RAB, HC, K; = P. philadelphicum Bernhardi ex Trinius ssp. lithophilum (Swallen) Freckmann & Lelong – 

FNA; < P. capillare Linnaeus var. sylvaticum Torrey – W; < P. philadelphicum – Y; Panicum s.s.] 

Panicum longifolium Torrey var. combsii (Scribner & Ball) Fernald, Combs Panic Grass. Cp (GA, NC, SC, VA): pond 

shores, depression meadows, cypress savannas, marshes, low woods; uncommon (VA Watch List). July-October. Scattered on 

the outer Coastal Plain from se. MA, NJ, se. VA, se. NC, e. SC, e. GA, and FL, west to se. LA. First glumes of var. combsii 

typically are longer than 1.5 mm long, while those of var. longifolium are shorter than 1.5 mm long. [= RAB, F, G; = P. 

rigidulum Bosc ex Nees ssp. combsii (Scribner & Ball) Freckmann & Lelong – FNA; = P. rigidulum Bosc ex Nees var. combsii 

(Scribner & Ball) Lelong – K, Z; < P. longifolium – C; = P. combsii Scribner & Ball – HC, S; not Panicum] 

Panicum longifolium Torrey var. longifolium, Long-leaved Panic Grass. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): 

wet sandy or peaty soils of bogs, savannas, pond shores, depression meadows; common (uncommon in Piedmont, rare in 

Mountains). July-October. Nova Scotia, NH, MA, PA, and IN south to FL, west to TX. See note under var. combsii. [= RAB, 

G; = P. rigidulum Bosc ex Nees ssp. pubescens (Vasey) Freckmann & Lelong – FNA; = P. rigidulum Bosc ex Nees var. 

pubescens (Vasey) Lelong – K, W, Z; < P. longifolium – C, GW; = P. longifolium – HC, S; > P. longifolium var. longifolium – 

F; > P. longifolium var. pubescens (Vasey) Fernald – F; not Panicum] 

* Panicum miliaceum Linnaeus ssp. miliaceum, Broomcorn Millet, Proso Millet, Hog Millet. Cp (NC), Mt (VA): planted in 

wildlife food plots, sometimes persistent or self-sowing; rare, introduced, native of Eurasia. July-October. [= C, FNA, K; < P. 

miliaceum – F, G, HC, S, Y; Panicum s.s.] 

Panicum philadelphicum Bernhardi ex Trinius, Woodland Panic Grass. Pd, Mt (GA, NC, SC, VA), Cp (VA): glades, 

barrens, desiccated pondshores, riversides, or other rocky or dry sandy soil of open woods and roadsides; frequent (rare in SC). 

Nova Scotia west to WI, south to GA and e. TX. Plants formerly known as P. tuckermanii Fernald, ranging from se. Canada 

south to n. VA and OH, are distinguished by included or short-exerted peduncles less than one-third as long as the panicles. [= 

RAB, C, G, K, S; > P. philadelphicum – F, HC; > P. tuckermanii Fernald – F, HC; = P. philadelphicum Bernhardi ex Trinius 

ssp. philadelphicum – FNA; < P. capillare Linnaeus var. sylvaticum Torrey – W; < P. philadelphicum – Y (also see P. 

lithophilum); Panicum s.s.] 

*? Panicum repens Linnaeus, Torpedo Grass. Cp (GA, NC, SC): disturbed coastal sands, in area where ship's ballast was 

deposited; rare, apparently introduced. First reported for NC by Leonard (1971b). [= FNA, GW, HC, K, S; Panicum s.s.] 

Panicum rigidulum Bosc ex Nees var. condensum (Nash) Mohlenbrock, Dense Panic Grass. Cp (GA, NC, SC, VA): 

marshes, meadows, low woods, ditches, stream and pond shores, freshwater tidal shores; occasional. September-October. 

Coastal Plain south from se. MA to FL, west to se. TX and AR; West Indies. Usually readily identified by its tall stature and 

compact inflorescence, somewhat resembling a large P. hemitomon, with which it occasionally occurs. [= P. agrostoides 

Sprengel var. condensum (Nash) Fernald – RAB, F; < P. rigidulum – C, GW; < P. rigidulum Bosc ex Nees ssp. rigidulum – 

FNA; < P. agrostoides – G; = P. condensum Nash - HC, S; < P. rigidulum var. rigidulum – K, Z; not Panicum] 

Panicum rigidulum Bosc ex Nees var. elongatum (Pursh) Lelong, Tall Flat Panic Grass. Pd, Cp, Mt (GA, NC, SC, VA): 

marshes, low woods, ditches, swamps, shores, meadows; occasional (common in Piedmont). August-October. CT and NY west 

to IN, south to GA, LA, and ne. TX. [= K, W, Z; = P. stipitatum Nash – RAB, F, HC, S; = P. rigidulum Bosc ex Nees ssp. 

elongatum (Pursh) Freckmann & Lelong – FNA; < P. rigidulum – C, GW; = P. agrostoides Sprengel var. elongatum (Pursh) 

Scribner – G; not Panicum] 

Panicum rigidulum Bosc ex Nees var. rigidulum, Redtop Panic Grass. Cp, Pd (GA, NC, SC, VA): wet sandy or peaty 

soils low woods, meadows, marshes, shores, swamps, ditches; frequent. July-October. ME and MI south to FL and TX; also in 

CA and British Columbia; Central America. [= W; = P. agrostoides Sprengel var. agrostoides – RAB, G; < P. rigidulum Bosc

POACEAE 902 

ex Nees ssp. rigidulum – FNA; < P. rigidulum var. rigidulum – K, Z; < P. rigidulum – C, GW; > P. agrostoides var. agrostoides 

– F, HC; > P. agrostoides var. ramosius (C. Mohr) Fernald – F, HC; = P. agrostoides – S; not Panicum] 

Panicum tenerum Beyrich ex Trinius, Southeastern Panic Grass. Cp (GA, NC, SC): limesink ponds, depression meadows, 

cypress savannas, wet pinelands, bogs; rare (GA Special Concern, NC Rare). June-September. Coastal Plain from se. NC to FL, 

west to e. TX; West Indies. The rhizomes produce lines of closely spaced culms. Though 0.5-1 m tall, the culms are narrow and 

inconspicuous. [= RAB, FNA, GW, HC, K, S, Z; not Panicum] 

Panicum verrucosum Muhlenberg, Warty Panic Grass. Cp, Pd, Mt (GA, NC, SC, VA): wet pinelands, marshes, shores, 

ditches; common (occasional in Piedmont, rare in Mountains). August-October. MA and PA west to MI and IN, south to FL and 

se. TX. Spikelets deep green, the warty surface unique among Panicum in our region. [= RAB, C, F, FNA, G, GW, HC, K, S, 

W, Z; not Panicum] 

Panicum virgatum Linnaeus var. cubense Grisebach, Blunt Panic Grass. Cp (GA, NC, SC, VA?): wet to dry sandy 

pinelands; occasional (frequent in the Sandhills). June-October. Coastal Plain from MA to FL, west to MS; also in MI; West 

Indies. [= F, HC, S; < P. virgatum – RAB, C, FNA, G, GW, W, Z; < P. virgatum var. virgatum – K] 

Panicum virgatum Linnaeus var. virgatum, Switchgrass. Cp, Pd, Mt (GA, NC, SC, VA): dry or wet sandy soils of 

pinelands, fresh and brackish marshes, shores; common (occasional in Mountains). June-October. Sw. Québec and ND south to 

FL and TX, west to NV; Bermuda; Central and South America. [= F, HC, S; < P. virgatum – RAB, C, FNA, G, GW, W, Z; < P. 

virgatum var. virgatum – K; not Panicum] 

* Panicum antidotale Retzius. Cp (SC): Native of India. Reported for NC and SC (FNA, Kartesz 1999). {investigate} [= 

FNA, HC, K; not Panicum] {not keyed at this time} 

* Panicum bergii Arechav. Cp (GA): disturbed areas; rare, native of South America. Reported for sc. GA (HC) and AL 

(Kartesz 1999). {investigate} [= FNA, HC, K; Panicum s.s.] {not keyed at this time} 

* Panicum bisulcatum Thunberg. Cp (GA, SC): disturbed areas; rare, native of Asia. Reported introduction in SC, GA, and 

PA (Kartesz 1999), and as a ballast plant for se. PA (Philadelphia) (Rhoads & Klein 1993, as P. acroanthum Steudel). [= FNA, 

K; ? P. acroanthum Thunberg] {not keyed at this time} 

Panicum brachyanthum Steudel. Cp (GA): {habitat unknown}; rare. Sw. GA west to c. TX. [= FNA, HC, K] 

Panicum dichotomiflorum Michaux var. puritanorum Svenson, Puritan Panic Grass. Alleged to be in VA northward 

(FNA). [= F, G, HC, K; < P. dichotomiflorum – C; = P. dichotomiflorum Michaux ssp. puritanorum (Svenson) Freckmann & 

Lelong – FNA] 

* Panicum miliaceum Linnaeus ssp. ruderale (Kitag.) Tzvelev, Panic Millet. [= FNA, K; = P. miliaceum ssp. spontaneum 

(Kit.) Tzvelev – C; < P. miliaceum – F, G, HC] {not keyed at this time} 

Panicum virgatum Linnaeus var. spissum Linder ranges south to PA, MD, and DE (Kartesz 1999). [= F, HC, K; < P. 

virgatum – C, FNA, G; not Panicum] {not keyed at this time} 


Parapholis C.E. Hubbard (Sickle Grass) 

* Parapholis incurva (Linnaeus) C.E. Hubbard, Sickle Grass, Hard Grass, Thin-tail. Cp (NC, VA): sandy and muddy flats, 

brackish or salt marshes; rare, introduced from Europe. [= RAB, C, HC, K, Z; = Pholiurus incurvus (Linnaeus) Schinzius & 

Thellung – F, G; ? Lepturus filiformis (Roth) Trinius] 

Pascopyrum A. Löve (Wheatgrass) 

* Pascopyrum smithii (Rydberg) A. Löve, Western Wheatgrass. Mt (GA): disturbed areas; rare. Reported for ne. GA 

(Rabun County) by Jones & Coile (1988), as Agropyron smithii Rydberg. It is also reported for TN and KY (Kartesz 1999). [= 

K; = Elytrigia smithii (Rydberg) Nevski – C; = Agropyrum smithii Rydberg – F, G, W] 

Paspalidium 

(see Setaria) 

Paspalum Linnaeus 1759 (Paspalum, Crown Grass, Beadgrass) 

(by Alan S. Weakley & Richard J. LeBlond) 

A genus of 300-400 species, of tropical and warm temperate regions. References: Allen & Hall in FNA (2003a); Banks 

(1966)=Z; Silveus (1942)=Y. Key based closely on FNA and on Banks (1966). 

1 Spikelets solitary, not associated with a rudimentary spikelets or naked pedicels. 

2 Panicles with 1-70 branches, if > 1, the branches arranged racemosely.

POACEAE 903 

3 Panicle branches 7-70, the axes extending beyond the outermost spikelets; panicle branches disarticulating at 

maturity...................................................................................................................................................... P. fluitans 

3 Panicle branches 1-6, terminating in a spikelet; panicle branches persistent. 

4 Upper florets olive to dark brown ............................................................................................P. scrobiculatum 

4 Upper florets pale to tan. 

5 Axes of panicle branches not broadly winged, 0.6-1.3 mm wide. 

6 Spikelets orbicular, 2.8-3.2 mm wide...............................................................P. laeve var. circulare 

6 Spikelets slightly longer than broad, 2.0-2.5 mm wide .......................................... P. laeve var. laeve 

5 Axes of panicle branches broadly winged, 1.8-3.3 mm wide. 

7 Spikelets 3.2-4.0 mm long; upper lemmas with a few short hairs at their tips............ P. acuminatum 

7 Spikelets 1.7-2.1 mm long; upper lemmas glabrous........................................................P. dissectum 

2 Panicles usually composed of a terminal pair of brancxhes, sometimes with 1 (-5) additional branches below the 

terminal pair. 

8 Upper glumes pubescent on the back or margins. 

9 Spikelets 1.3-1.9 mm long; upper glumes pilose along the margins .........................................[P. conjugatum] 

9 Spikelets 2.4-3.2 mm long; upper glumes sparsely pubescent on the back .....................................P. distichum 

8 Upper glumes glabrous. 

10 Spikelets elliptic, acute or acuminate at the tip ..............................................................................P. vaginatum 

10 Spikelets ovate to broadly elliptic, obtuse to broadly acute at the tip. 

11 Spikelets 1.9-2.3 mm long; leaf blades flat ................................................................................ [P. minus] 

11 Spikelets 2.5-4.0 mm long; leaf blades flast or longitudinally folded ....................................... P. notatum 

1 Spikelets paired, or at least the second nonfunctional spikelet represented by a naked pedicel. 

12 Spikelets 1.0-1.3 mm long .......................................................................................................................[P. paniculatum] 

12 Spikelets 1.3-4.1 mm long 

13 Margins of upper glumes and lower lemmas pilose. 

14 Panicle branches 2-7; spikelets 2.3-4.0 mm long ............................................................................ P. dilatatum 

14 Panicle branches (4-) 10-30; spikelets 1.8-2.8 mm long ..................................................................... P. urvillei 

13 Margins of upper glumes and lower lemmas neither ciliate-lacerate, winged, nor pilose (if pubescent, the hairs not 

pilose). 

15 Upper florets olive to dark brown. 

16 Panicle branches 10-28 (or more). 

17 Plants annual; axes of panicle branches broadly winged, the wings about as wide as the central 

portion; [common native].............................................................................................. P. boscianum 

17 Plants perennial; axes of panicle branches narrowly winged, the wings narrower than the central 

portion; [rare exotics]. 

18 Axes of panicle branches 0.5-1.2 mm wide; spikelets 1.1-1.8 mm wide ............. P. conspersum 

18 Axes of the panicle branches 1.0-1.7 mm wide; spikelets 1.8-2.4 mm wide ............ P. virgatum 

16 Panicle branches 1-10 (or to 28 in P. boscianum, keyed under both leads). 

19 Plants annual. 

20 Spikelets 1.3-1.8 mm wide, broadly elliptic to orbicular, glabrous; panicles with 1-10 (-28) 

branches, the axes 0.7-2.3 mm wide...................................................................... P. boscianum 

20 Spikelets 1.7-2.4 mm wide, broadly obovate, shortly pubescent; panicles with 1-5 branches, 

the axes 0.8-1.3 mm wide.....................................................................................[P. convexum] 

19 Plants perennial. 

21 Plants cespitose, rhizomes poorly developed; culms 10-20 dm tall; panicle branches 

ascending, divaricate, or reflexed. 

22 Leaves 7-18 mm wide.................................................................................. P. conspersum 

22 Leaves 2.5-4 mm wide................................................................................... P. plicatulum 

21 Plants not cespitose, rhizomatous; culms 1-15 dm tall; panicle branches ascending. 

23 Rhizomes long, evident .......................................................................................P. nicorae 

23 Rhizomes short, indistinct ............................................................................. P. plicatulum 

15 Upper florets white, stramineous, or golden brown. 

24 Lower lemmas with well-developed cross-ribs over the veins; upper glumes absent.....P. malacophyllum 

24 Lower lemmas not ribbed over the veins; upper glumes present. 

25 Panicles with 15-100 branches. 

26 Plants annual; upper glumes and lower lemmas rugose..................................... [P. racemosum] 

26 Plant perennial; upper glumes and lower lemmas smooth. 

27 Plant rhizomatous; panicle branch axes 0.9-1.2 mm wide; panicle branches often arcing... 

.................................................................................................................... P. intermedium 

27 Plant cespitose; panicle branch axes 0.3-0.6 mm wide; panicle branches straight. 

28 Panicle branches spreading to reflexed (rarely ascending); leaf blades 10-23 mm 

wide; axes of panicle branches 0.3-0.4 mm wide ................................P. coryphaeum 

28 Panicle branches erect to ascending; leaf blades 4.9-6.1 mm wide; axes of panicle 

branches 0.5-0.6 mm wide.............................................................. [P. quadrifarium]

POACEAE 904 

25 Panicles with 1-15 branches. 


30 Spikelet pairs barely if at all imbricate; lower glumes usually present ............... P. bifidum 

30 Spikelet pairs imbricate; lower glumes absent or present. 

31 Upper glumes pubescent; lower lemmas usually pubescent. 

32 Lower glumes present......................................................................... [P. langei] 

32 Lower glumes absent .....................................................................P. pubiflorum 

31 Upper glumes glabrous; lower lemmas usually glabrous. 

33 Upper florets golden brown ...........................................................P. floridanum 

33 Upper florets pale to tan. 

34 Terminal panicle branches erect....................................[P. monostachyum] 

34 Terminal panicle branches spreading to ascending. 

35 Plants decumbent, rooting at the lower nodes; spikelets obovate to 

elliptic ............................................................................P. pubiflorum 

35 Plants rhizomatous; spikelets orbicular to elliptic. 

36 Spikelets 2.9-4.1 mm long; 1.9-3.1 mm wide, suborbicular to 

elliptic; upper glumes 5-veined; leaf blades flat.....P. floridanum 

36 Spikelets 2.1-3.1 mm long, 2.0-2.8 mm wide, orbicular or nearly 

so; upper glumes 3-veined; leaf blades laterally folded. 

37 Lower sheaths villous or hirsute........................................... 

................................................ P. praecox var. curtisianum 

37 Lower sheaths glabrous or sparsely papillose pubescent...... 

.......................................................P. praecox var. praecox 


38 Upper glumes (and usually also the lower lemmas) shortly pubescent. 

39 Lower glumes present................................................................................. [P. langei] 

39 Lower glumes absent. 

40 Panicles both terminal and axillary, the axillary panicles partially or completely 

enclosed by the subtending leaf sheath ................................................................ 

........................................................... [see Key to Paspalum setaceum complex] 

40 Panicles all terminal.................................................................. [P. caespitosum] 

38 Upper glumes and lower lemmas glabrous. 

41 Panicles both terminal and axillary, the axillary panicles partially or completely 

enclosed by the subtending leaf sheath ...... [see Key to Paspalum setaceum complex] 

41 Panicles all terminal. 

42 Upper panicle branches erect ................................................[P. monostachyum] 

42 Upper panicle branches spreading to ascending. 

43 Upper glumes and lower lemmas 5-veined .......................... P. caespitosum 

43 Upper glumes and lower lemmas 3-veined. 

44 Lower sheaths villous or hirsute.............. P. praecox var. curtisianum 

44 Lower sheaths glabrous or sparsely papillose pubescent...................... 

.......................................................................P. praecox var. praecox 

Key to Paspalum setaceum complex 

(by Richard J. LeBlond) 

1 Leaves variously pubescent. 

2 Leaves villous to villous-hirsute, 2-10 mm wide; spikelets 1.3-1.9 mm long. 

3 Leaves villous, 2-7 mm wide, not especially crowded toward the base, erect to spreading; [widespread] ................. 

......................................................................................................................................... P. setaceum var. setaceum 

3 Leaves villous-hirsute, 3-10 mm wide, crowded toward the base, recurved; [of n. FL south to Cuba]....................... 

................................................................................................................................[P. setaceum var. villosissimum] 

2 Leaves puberulent, pilose, or hirsute, 3-15 mm wide; spikelets 1.6-2.5 mm long. 

4 Leaves puberulent at least distally on the adaxial surface (and often also pilose in var. stramineum); spikelets 1.6- 

2.2 mm long. 

5 Plants erect to spreading; leaves puberulent and often pilose to nearly glabrous except for the puberulent 

distal adaxial surface; spikelets glabrous to pubescent......................................... P. setaceum var. stramineum 

5 Plants spreading to prostrate; leaves densely puberulent; spikelets pubescent.................................................... 

....................................................................................................................... P. setaceum var. psammophilum 

4 Leaves pilose or hirsute but not puberulent; spikelets 1.8-2.5 mm long. 

6 Plants mostly erect; leaves pilose; spikelets usually glabrous; sterile lemma midnerve usually present............ 

...........................................................................................................................P. setaceum var. muhlenbergii

POACEAE 905 

6 Plants mostly widely spreading; leaves hirsute; spikelets glabrous or pubescent; sterile lemma midnerve 

present or absent........................................................................................................ P. setaceum var. supinum 

1 Leaves glabrous to glabrate (if glabrate, also see var. stramineum in couplet 5). 

7 Blades crowded toward the base, often recurved, 3-8 mm wide; spikelets 1.4-1.9 mm long, usually glabrous.................. 

................................................................................................................................ P. setaceum var. longepedunculatum 

7 Blades not especially crowded toward the base, erect, ascending or spreading, 2-20 mm wide; spikelets 1.6-2.6 mm 

long, pubescent or glabrous. 

8 Blades 3-8 mm wide; spikelets 1.6-1.9 mm long, pubescent, subacute; [of GA and FL southward] .......................... 

......................................................................................................................................................... [P. propinquum] 

8 Blades 2-20 mm wide; spikelets 1.7-2.6 mm long (if < 2.0 then larger leaves usually > 7 mm wide), glabrous or 

pubescent, rounded to blunt; [plants of FL northward and westward]. 

9 Plants stiffly erect; blades 2-6 mm wide; spikelets 2.0-2.6 mm long; [of GA and FL] ....................................... 

............................................................................................................................. P. setaceum var. rigidifolium 

9 Plants erect to spreading; blades 3-20 mm wide; spikelets 1.7-2.6 mm long; [of NJ to TX]............................... 

.............................................................................................................................P. setaceum var. ciliatifolium 

Paspalum acuminatum Raddi, Brook Paspalum, Canoe Grass. Pd (GA): wet areas, often disturbed; rare, possibly only 

adventive in our area. C. GA and ne. TX south to s. FL and s. TX, south through the New World tropics to s. South America. [= 

FNA, HC, K] 

Paspalum bifidum (Bertoloni) Nash, Pitchfork Paspalum, Pitchfork Crown Grass. Cp (GA, NC, SC, VA), Pd (GA, SC): 

mesic to wet longleaf pine savannas and mesic swales in sandhills; uncommon (NC Watch List, VA Rare). August-October. Se. 

VA south to s. FL, west to se. MO, se. OK, and e. TX. [= RAB, C, GW, HC, K, S, Y; > P. bifidum var. bifidum – F, G; > P. 

bifidum var. projectum Fernald – F, G] 

Paspalum boscianum Flügge, Bull Paspalum. Cp, Pd (GA, NC, SC, VA), Mt (GA, SC, VA): low fields, ditches; common 

(VA Watch List). July-October. MD, KY, and TX south through tropical America. [= RAB, C, F, FNA, G, GW, HC, K, S, W, 

Y] 

* Paspalum conspersum Schrad., Scattered Paspalum. Cp (GA): roadsides, other disturbed areas; rare, introduced from a 

native range of Mexico to South America. [= FNA] {synonymy incomplete} 

* Paspalum coryphaeum Trinius, Emperor Crown-grass. Pd (NC): disturbed areas; rare, native of South America. [= FNA, 

K] {synonymy incomplete} 

* Paspalum dilatatum Poiret, Dallis Grass. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, fields, disturbed areas; common, 

introduced from tropical America. May-October. [= RAB, C, F, FNA, G, GW, HC, K, S, W, Y] 

Paspalum dissectum (Linnaeus) Linnaeus, Mudbank Crown Grass, Walter Paspalum. Cp (GA, NC, SC, VA), Pd (NC, SC): 

mud flats, drawdown zones; rare (NC Rare, VA Rare). September. NJ, IL, and KS south to s. FL and e. TX; Cuba. [= RAB, C, 

F, FNA, G, GW, HC, K, S, Y] 

Paspalum distichum Linnaeus, Joint Paspalum, Knotgrass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (NC): brackish 

and freshwater marshes; uncommon (VA Rare). June-August. NJ, KS, and WA south to s. FL, s. TX, s. CA and through the 

New World and Old World tropics. [= RAB, C, F, FNA, G, HC, K, S, W, Y; < P. distichum – GW (also see P. vaginatum); = P. 

paspaloides (Michaux) Scribner] 

Paspalum floridanum Michaux, Florida Paspalum. Cp, Pd, Mt (GA, NC, SC, VA): wet forests, pine savannas; common 

(rare in Mountains). August-October. NJ, IL, and KS south to s. FL and e. TX. [= RAB, C, FNA, GW, K, W; > P. floridanum – 

G; > P. difforme Le Conte – G, HC, S, Y; > P. floridanum var. floridanum – F, HC, S, Y; > P. floridanum var. glabratum 

Engelmann ex Vasey – F, HC, S, Y; > P. giganteum Baldwin ex Vasey – HC, S, Y] 

Paspalum fluitans (Elliott) Kunth, Water Paspalum, Horsetail Crown Grass. Cp (GA, NC, SC, VA), Pd (NC, VA): mucky 

soils in swamp forests; uncommon (NC Watch List). October. MD, IL, and KS south to s. FL and s. TX, and south through 

tropical America to c. South America. [= RAB, C, F, G, HC, K; = P. repens P.J. Bergius – FNA, GW, S, Y] 

* Paspalum intermedium Munro ex Morong. Cp (GA): drainage canals; rare, introduced from South America. Escaped in 

sc. GA (Tift County, where growing along drainage canals in Tifton) (Jones & Coile 1988). [= FNA, HC, K] 

Paspalum laeve Michaux var. circulare (Nash) Stone. {GA, NC, VA}: {need additional herbarium work to fully 

determine range and abundance of varieties} June-August. [= F; < P. laeve – RAB, C, FNA, G, GW, K, W; = P. circulare Nash 

– HC, S, Y] 

Paspalum laeve Michaux var. laeve. Cp, Pd, Mt (GA, NC, SC, VA): forest edges and disturbed areas; common. {need 

additional herbarium work to fully determine range and abundance of varieties} June-August. Overall distribution of P. laeve 

s.l.: MA, NY, MI, and KS south to s. FL and e. TX. [< P. laeve – RAB, C, FNA, G, GW, K, W; > P. laeve var. laeve – F; > P. 

laeve var. pilosum Scribner – F; > P. laeve – HC, S, Y; > P. longipilum Nash – HC, S, Y] 

* Paspalum malacophyllum Trinius, Ribbed Paspalum. Cp (GA): disturbed areas; rare, introduced from a native range of 

Mexico to South America. [= FNA, HC] 

* Paspalum nicorae Parodi, Brunswickgrass. Cp (GA): disturbed areas; rare, introduced from Brazil. [= FNA, HC, K] 

* Paspalum notatum Flügge, Bahia Grass. Cp (GA, NC, SC, VA), Pd (GA, SC, VA), Mt (GA): roadsides and disturbed 

areas; uncommon, introduced from tropical America. June-October. [= FNA, G, GW, Y; > P. notatum var. notatum – HC, K; > 

P. notatum Flügge var. saurae Parodi – RAB, HC, K] 

Paspalum plicatulum Michaux, Brownseed Paspalum. Cp (GA, SC): pine savannas, fields; uncommon. May-July. Se. SC 

south to s. FL, west to s. TX, and south through tropical America to s. South America. [= RAB, FNA, GW, HC, K, S, Y]

POACEAE 906 

Paspalum praecox Walter var. curtisianum (Steudel) Vasey, Curtis’s Crown Grass. Cp (GA, NC, SC, VA): pine savannas; 

rare (NC Watch List, VA Rare). June-October. The variety was named for the Rev. Moses Ashley Curtis (of Hillsborough, NC), 

not Allen Hiram Curtiss (of Jacksonville, FL); the correct spelling of the epithet is therefore "curtisianum." [= RAB, F, G; = P. 

praecox var. curtissianum – C, orthographic error; < P. praecox – FNA, GW, K; = P. lentiferum Lamarck – HC, S, Y] 

Paspalum praecox Walter var. praecox, Early Crown Grass. Cp (GA, NC, SC): pine savannas; rare (NC Watch List). 

May-July. [= RAB, C, F, G; < P. praecox – FNA, GW, K; = P. praecox – HC, S, Y] 

Paspalum propinquum Nash. Cp: {habitat}; uncommon. GA and FL. [= HC, S; < P. setaceum – K] 

Paspalum pubiflorum Ruprecht var. glabrum Vasey, Hairyseed Crown Grass. Mt, Pd (GA, NC, VA), Cp (SC, VA): 

disturbed areas; uncommon (NC Watch List). September-October. PA west to KS and CO, south to FL and s. TX and Mexico; 

Cuba. [= C, F, G, HC, S, Y; < P. pubiflorum – RAB, FNA, GW, K, W] 

* Paspalum scrobiculatum Linnaeus, Indian Paspalum. Cp (GA): disturbed areas; rare, native of India. [= FNA, HC, K] 

Paspalum setaceum Michaux var. ciliatifolium (Michaux) Vasey. Cp, Pd, Mt (GA, NC, SC, VA): dry open areas and 

woodlands, disturbed areas; common. June-September. S. NJ south to s. FL, west to e. TX, interior to s. WV, se. KY, e. TN, n. 

AL, n. MS, c. AR, and e. OK. [= FNA, Z; < P. setaceum – RAB, GW, K, W; < P. setaceum var. ciliatifolium – C (also see var. 

longepedunculatum); = P. ciliatifolium Michaux var. ciliatifolium – F, G; = P. ciliatifolium Michaux – HC, S, Y] 

Paspalum setaceum Michaux var. longepedunculatum (LeConte) A. Wood. Cp (GA, NC, SC): pine flatwoods and pine 

savannas; rare. June-September. Se. NC south to s. FL, west to s. MS. [= F, FNA, Z; < P. setaceum – RAB, GW, K, W; < P. 

setaceum var. ciliatifolium – C; = P. longepedunculatum LeConte – G, HC, S, Y] 

Paspalum setaceum Michaux var. muhlenbergii (Nash) Fernald. Mt, Pd, Cp (GA, NC, SC, VA): dry or moist soils; 

common. June-September. NH west to MI, c. IL, s. IA, and c. KS, south to n. FL, s.AL, s. MS, s. LA, and c. TX. [= C, FNA, Z; 

< P. setaceum – RAB, GW, K, W; > P. setaceum var. calvescens Fernald – F; > P. ciliatifolium Michaux var. muhlenbergii 

(Nash) Fernald – F; = P. ciliatifolium Michaux var. muhlenbergii (Nash) Fernald – G; = P. pubescens Muhlenberg ex Willdenow 

– HC, S, Y] 

Paspalum setaceum Michaux var. psammophilum (Nash) D. Banks. Cp? (VA?): maritime grasslands, sandy disturbed 

areas; rare. June-September. MA south to DC (VA?) in the Coastal Plain. [= C, FNA, Z; < P. setaceum – K; = P. 

psammophilum Nash – F, G, HC, Y] 

Paspalum setaceum Michaux var. rigidifolium (Nash) D. Banks. Cp (GA, NC?, SC?): sandhills; rare. June-September. 

Ne. GA, immediately adjacent to SC (and reported for NC by HC) south to s. FL; Cuba. [= FNA, Z; < P. setaceum – RAB, GW, 

K, W; = P. rigidifolium Nash – HC, S, Y] 

Paspalum setaceum Michaux var. setaceum, Thin Paspalum. Cp, Pd, Mt (GA, NC, SC, VA): sandhills, savannas, dry 

soils; common (uncommon in Piedmont and Mountains). June-September. MA and CT south to s. FL, west to e. TX, inland to 

w. VA, s. WV, s. MO and AR; Cuba. [= C, FNA, Z; < P. setaceum – RAB, GW, K, W; > P. setaceum – G, HC, S, Y; > P. 

debile Michaux – F, HC, S, Y; > P. setaceum var. setaceum – F] 

Paspalum setaceum Michaux var. stramineum (Nash) D. Banks, Yellow Sand Paspalum. Cp (GA, NC): dry sandy soils; 

rare. June-September. MI west to MT, south to LA, and NM; scattered eastwards, especially near the coast, perhaps at least in 

part as introductions [= C, FNA, Z; < P. setaceum – RAB, GW, K, W; = P. ciliatifolium Michaux var. stramineum (Nash) 

Fernald – F, G; = P. stramineum Nash – HC, Y] 

Paspalum setaceum Michaux var. supinum (Bosc ex Poiret) Trinius. Cp (GA, NC, SC, VA?): sandy soils, old fields; 

uncommon. June-September. E. NC (e. VA?) south to s. FL, west to s. MS. Also reported for the Coastal Plain of Virginia by 

Tatnall (1946); needing confirmation of the specimen identification. [= F, FNA, Z; < P. setaceum – RAB, GW, K, W; = P. 

supinum Bosc ex Poiret – HC, S] 

* Paspalum urvillei Steudel, Vasey Grass. Cp, Pd (GA, NC, SC, VA): roadsides, fields, and disturbed areas; common, 

introduced from South America. May-July. [= RAB, C, F, FNA, G, GW, HC, K, S, Y] 

Paspalum vaginatum Swartz, Sand Knotgrass, Seashore Crown Grass. Cp (GA, NC, SC), Pd (NC): brackish marshes, 

rarely inland in disturbed places; rare (NC Watch List). July. NC south to s. FL, west to s. TX, southward through the New 

World tropics; Old World tropics. [= RAB, FNA, HC, K, S, Y; < P. distichum – GW] 

* Paspalum virgatum Linnaeus, Talquezal. Cp (GA): disturbed areas; rare, introduced from Mexico, Central America, and 

South America. [= FNA, K] {synonymy incomplete} 

Paspalum caespitosum Flügge. Pinelands, hammocks. S. AL and n. FL south to s. FL; West Indies, Mexico and Central 

America. [= FNA, GW, HC, K, S] 

Paspalum conjugatum Bergius, Sour Paspalum. Ne. FL, FL Panhandle, and s. AL west to e. TX, south in the New World 

tropics; Old World tropics. [= FNA, HC, K, S] {synonymy incomplete} 

* Paspalum convexum Flügge, Mexican Paspalum. Disturbed areas. MS, LA, and e. TX, introduced from tropical America. 

[= FNA, K] {synonymy incomplete} 

Paspalum langei (E. Fournier) Nash, Rustyseed Paspalum. N. peninsular FL and Panhandle FL west to se. TX, and south 

through the New World tropics to South America. [= FNA, K] {synonymy incomplete} 

Paspalum minus E. Fournier, Matted Paspalum. Disturbed areas. FL Panhandle and s. AL west to e. TX. [= FNA, K] 


Paspalum monostachyum Vasey, Gulfdune Paspalum. Coastal dunes, wet prairies. AL and FL west to TX. [= FNA, HC, 

K, S] {synonymy incomplete} 

* Paspalum paniculatum Linnaeus, Arrocillo. Disturbed areas, native of tropical America. Ec. MS and sw. FL. [= FNA, K] 

{synonymy incomplete}

POACEAE 907 

* Paspalum quadrifarium Lamarck, Tussock Paspalum. Disturbed areas. S. MS. Native of South America. [= FNA] 


* Paspalum racemosum Lamarck, Peruvian Paspalum. Disturbed areas. MS and other widely scattered localities in North 

America, native of n. South America. [= FNA, K] {synonymy incomplete} 

Paspalum setaceum var. villosissimum (Nash) D. Banks. Sandy pine flatwoods and fields. N. FL (very near GA) south to s. 

FL; Cuba. [= FNA, Z; < P. setaceum – GW, K; < P. debile Michaux – HC; = P. villosissimum Nash – S] 

Pennisetum L.C. Richard ex Persoon 

A genus of 80-130 species, perennials and annuals, mainly of the tropics and subtropics. References: Wipff in FNA (2003a). 

Key adapted from FNA. 

1 Primary bristles (immediately subtending each spikelet) scabrous; plants 2-8 m tall ........................................[P. purpureum] 

1 Primary bristles conspicuously long-ciliate; plants 0.1-3 m tall. 

2 Spikelets 9-12 mm long .................................................................................................................................... P. villosum 

2 Spikelets 2.5-7 mm long. 

3 Fascicles not disarticulating from the rachises; fascicles 33-160 per cm of inflorescence; panicles 4-200 cm long; 

leaves 7-70 mm wide ................................................................................................................................P. glaucum 

3 Fascicles disarticulating from the rachises at maturity; fascicles 8-37 per cm of inflorescence; panicles 2-32 cm 

long; leaves 2-13 mm wide. 

4 Spikelets 4.5-7 mm long; leaves 2-3.5 mm wide, folded or conduplicate and superficially appearing even 

narrower; rachis pubescent............................................................................................................ [P. setaceum] 

4 Spikelets 2.5-5.6 mm long; leaves 2-13 mm wide, flat; rachis scabrous. 

5 Inner bristles fused for < ¼ of their length; many outer bristles exceeding the spikelets; terminal bristles 

10.5-23 mm long, noticeably longer than the other bristles in the fascicle................................. [P. ciliare] 

5 Inner bristles fused for ⅓-½ of their length; outer bristles not exceeding the spikelets; terminal bristles 

2.9-6.5 mm long, usually not noticeably exceeding the other bristles in the fascicle ........... [P. setigerum] 

* Pennisetum glaucum (Linnaeus) R. Brown, Pearl Millet. (GA, NC, SC, VA). [= RAB, FNA, HC, K; ? Chaetochloa 

lutescens (Weigel) Stuntz – S; = Setaria glauca (Linnaeus) Palisot de Beauvois] 

* Pennisetum villosum R. Brown ex Fresenius, Feathertop. Reported as an introduction in GA (Kartesz 1999). [= C, FNA, 

HC, K; ? Cenchrus longisetus M.C. Johnston] 

* Pennisetum ciliare (Linnaeus) Link, Buffelgrass. Disturbed areas, native of Africa. Known in our area from ne. FL, s. AL, 

e. TN, and ec. MS. [= FNA, HC; = P. ciliare var. ciliare – K; = Cenchrus ciliaris Linnaeus] 

* Pennisetum purpureum Schumacher, Elephant Grass, Napier Grass. Disturbed areas, native of Africa. Naturalized in FL 

north to the FL-GA border. [= FNA, HC, K] 

* Pennisetum setaceum (Forskål) Chiovenda, Tender Fountaingrass Reported as an introduction in FL, TN, and KY (Wipff in 

FNA). [= FNA, HC, K] {synonymy incomplete} 

* Pennisetum setigerum (Vahl) Wipff. Disturbed areas, native of Africa. Known in our area from ne. FL and ec. MS. [= 

FNA; = P. ciliare (Linnaeus) Link var. setigerum (Vahl) Leeke – K; = Cenchrus setigerus Vahl] 

Phalaris Linnaeus (Canary-grass) 

A genus of about 16 species, north temperate and South American. References: Tucker (1996)=Z. 

1 Perennial, with scaly rhizomes; inflorescence obviously paniculate, 7-25 cm long, with ascending to appressed branches, the 

main branches of the inflorescence apparent, the inflorescence outline thus appearing lobed. 

2 Glumes broadly winged; fertile lemmas ovate-lanceolate, densely pubescent................................................Ph. aquatica 

2 Glumes not winged; fertile lemmas narrowly lanceolate, glabrous to sparsely pubescent....................... Ph. arundinacea 

1 Annual, without rhizomes; inflorescence densely spikelike or almost capitate, 1-6 cm long, the branches not apparent, the 

inflorescence outline a single ovoid, ellipsoid, or lanceolate form. 

3 Keels of the glumes broadly winged (the wing ca. 1 mm wide); sterile lemmas 2.5-4.5 mm long ............ Ph. canariensis 

3 Keels of the glume narrowly winged (the wing < 0.5 mm wide); sterile lemmas 1.5-2.5 mm long. 

4 Nerves of the glumes scabrous; panicle cylindric in outline, 6-18 cm long; glumes 3.5-4.0 mm long ....Ph. angusta 

4 Nerves of the glumes not scabrous; panicle narrowly ovate in outline, usually 2-6 cm long; glumes 5-6 mm long ... 

...........................................................................................................................................................Ph. caroliniana 

* Phalaris angusta Nees ex Trinius. Cp (GA, SC): waterfowl impoundments, marshes; uncommon, introduced from tropical 

America. [= GW, HC, K, Z] 

* Phalaris aquatica Linnaeus, Bulbous Canary-grass. Cp (NC, SC, VA): disturbed areas; rare, introduced from Europe. [= 

K, Z; ? Ph. tuberosa Linnaeus var. stenoptera (Hackel) Hitchcock – HC]

POACEAE 908 

*? Phalaris arundinacea Linnaeus, Reed Canary-grass, Ribbon Grass. Mt, Pd (NC, VA), Cp (VA): moist forests, moist 

disturbed areas, bogs; common (rare in Coastal Plain). June. Newfoundland west to AK, south to NC, TN, AR, NM, CA; 

Mexico; Eurasia. A variegated form, Ph. arundinacea forma variegata (Parn.) Druce, is cultivated for ornament, as Ribbon 

Grass. [= RAB, C, F, GW, K, S, W, Z; > Ph. arundinacea var. arundinacea – G, HC; > Ph. arundinacea var. picta Linnaeus – 

G, HC] 

* Phalaris canariensis Linnaeus, Birdseed Grass, Canary-grass. Cp, Pd (GA, NC, SC, VA), Mt (VA): disturbed areas; rare, 

introduced from Mediterranean Europe. [= RAB, C, F, G, GW, HC, K, S, Z] 

Phalaris caroliniana Walter, Maygrass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): ditches, roadsides, disturbed areas; 

uncommon. May-June. [= RAB, C, F, G, GW, HC, K, S, Z] 

* Phalaris minor Retzius, Lesser Canary Grass. Cp (SC): waste areas near wool-combing mills; rare, introduced from 

Mediterranean Europe. Also reported for other scattered states in e. North America (Kartesz 1999). [= HC, K] {not keyed at 

this time; synonymy incomplete} 

* Phalaris paradoxa Linnaeus, Mediterranean Canary Grass, is reported for MD, NJ, and PA (Kartesz 1999). [= K; > Ph. 

paradoxa var. paradoxa – HC; > Ph. paradoxa var. praemorsa (Lamarck) Coss. & Durieu – HC] {not keyed at this time; 


Phanopyrum (Rafinesque) Nash (Phanopyrum) 

Circumscription of this genus is currently in flux. Phanopyrum is variously treated as a distinct genus or as a subgenus of 

Panicum. Panicum verrucosum perhaps belongs here as well. References: Crins (1991)=Z; Webster (1988)=Y; Freckmann & 

Lelong in FNA (2003a). 

Phanopyrum gymnocarpon (Elliott) Nash, Swamp Phanopyrum, Savanna Phanopyrum. Cp (GA, NC, SC, VA): swamps, 

seasonally flooded soils of cypress-gum sloughs, tidal (freshwater) cypress-gum swamps, disturbed wet soils, low woods, ditches, 

muddy banks of streams and lakes, sinks, floodplains, and marshes; uncommon (NC Watch List, VA Rare). August-October. 

Se. VA south to FL, west to TX and AR. [= K, Y; = Panicum gymnocarpon Elliott – RAB, FNA, GW, HC, S, Z] 

Phleum Linnaeus (Timothy) 

References: Tucker (1996)=Z; Stace (1997)=Y. Key based on Stace (1997). 

1 Spikelets 2.0-3.5 mm long, including the 0.2-1.0 (-1.2) mm long awns; panicle 3-6 mm wide; leaves 2-6 mm wide; ligule 

usually acute ...................................................................................................................................Phl. pratense ssp. nodosum 

1 Spikelets (3.5-) 4-5.5 mm long, including the (0.8-) 1.0-2.0 mm long awns; panicle 6-10 mm wide; leaves 3-9 mm wide; 

ligule usually obtuse ........................................................................................................................ Phl. pratense ssp. pratense 

* Phleum pratense Linnaeus ssp. nodosum (Linnaeus) Arcangeli, Small Timothy. (NC) {included based on Fernald's report 

– corroboration and additional information needed} [< Ph. pratense – RAB, C, G, HC, K, S, W, Z; = Ph. pratense var. nodosum 

(Linnaeus) Hudson – F; = Ph. bertolonii Augustin de Candolle – Y] 

* Phleum pratense Linnaeus ssp. pratense, Timothy. Mt, Pd, Cp (GA, NC, SC, VA): meadows, pastures, roadsides, 

disturbed areas; common, introduced from Europe. June-October. The American common name comes from the name of the 

man who is believed to have introduced it into the United States in 1720, Timothy Hanson; in England, Phleum is called "cat'stail." 

[< Ph. pratense – RAB, C, G, HC, K, S, W, Z; = Ph. pratense var. pratense – F; = Ph. pratense – Y] 

* Phleum subulatum (Savi) Ascherson & Graebner, Italian Timothy, is reported as introduced in MD and PA (Kartesz 1999). 

Not keyed. [= K, Y] 

Phragmites Adanson (Common Reed) 

A genus with one species and 2 or more varieties, nearly worldwide in distribution. References: Allred in FNA (2003a); 

Saltonstall, Peterson, & Soreng (2004)=Z; Saltonstall (2002). Key based on Z. 

Key 1 

1 Ligules 1.0-1.7 mm long; lower glumes 3.0-6.5 mm long; upper glumes 5.5-11.0 mm long; lemmas 8.0-13.5 mm long; leaf 

sheaths caducous with age; culms exposed in the winter smooth and shiny; [native south to WV and NC] .............................. 

................................................................................................................................................... Phr. australis ssp. americanus 

1 Ligules 0.4-0.9 mm long; lower glumes 2.5-5.0 mm long; upper glumes 4.5-7.5 mm long; lemmas 7.5-12.5 mm long; leaf 

sheaths not caducous with age; culms not exposed in the winter, either smooth and shiny or ridged and not shiny.

POACEAE 909 

2 Culms smooth and shiny; [native on the Gulf Coast]....................................................... [Phr. australis var. berlandieri] 

2 Culms ridged and not shiny; [introduced and weedy]............................................................. Phr. australis ssp. australis 

Key 2 

1 First glume 2.3-4.2 mm long; stems at base of plant in summer and late fall mostly tan-brown or yellow; [alien and weedy].. 

........................................................................................................................................................Phr. australis var. australis 

1 First glume 3.5-6.4 mm long; stems at base of plant in summer and late fall reddish-brown or reddish-purple; [native]........... 

....................................................................................................................................................Phr. australis var. berlandieri 

* Phragmites australis (Cavanilles) Trinius ex Steudel var. australis, Common Reed. Cp (GA, NC, SC, VA), Pd, Mt (NC, 

VA): marshes, dredge-spoil deposit islands, ditches; common in outer Coastal Plain (rare elsewhere). September-October. 

Nearly worldwide in distribution. Fox, Godfrey, & Blomquist (1950) report its first collection in NC (in 1948). In most of our 

area, reed is of relatively recent introduction, reported from only nine counties in RAB, but now becoming a serious weed in 

coastal areas, where it aggressively colonizes freshwater and brackish marshes, excluding the native species. [< Ph. australis – 

C, FNA, GW, K; < Ph. communis Trinius – RAB, G, HC; = Ph. communis var. communis – F; < Ph. phragmites (Linnaeus) 

Karsten – S] 

Phragmites australis (Cavanilles) Trinius ex Steudel var. berlandieri (Fornier) C.F. Reed, North American Reed. Cp (VA): 

freshwater marshes; rare. September-October. [< Ph. australis – C, FNA, GW, K; < Ph. communis Trinius – RAB, G, HC; = 

Ph. communis var. berlandieri (Fournier) Fernald – F; < Ph. phragmites (Linnaeus) Karsten – S] 

Phyllostachys Siebold & Zuccarini (Bamboo) 

References: Duncan & Duncan [in prep.]=Z; Judziewicz et al. (2000)=Y. Key adapted from Z. 

1 Internodes at the base of principal culms dissimilar in length, the lowermost internode 1-12 cm long, the next 3 internodes 

distinctly longer, with nodal junctions mostly straight across..................................................................................... Ph. aurea 

1 Internodes at the base of principal culms all similar in length, mostly 4-8 cm, with nodal junctions oblique. 

2 Groove on internode (above the branch) yellowish-green, the rest of the culm dull greenish..................Ph. aureosulcata 

2 Groove on internode (above the branch) the same color as the rest of the culm. 

3 Internodes of principal culms densely velvety; outer surface of culm sheaths with abundant erect brown hairs; 

lowest internode of principal culms ca. 5 cm long; culms pale green atfirst, becoming gray with accumulated waxy 

powder in age..................................................................................................................................... Ph. heterocycla 

3 Internodes of principal culms glabrous or slightly hairy; outer surface of culm sheaths lacking erect brown hairs; 

lowest internode of principal culms ca. 8.5-12 cm long; culms various (see below, but not as decribed in first 

lead). 

4 Largest culms to 15 cm in diameter and 25 m tall; upper culm sheaths with auricles; outer surface of culm 

sheaths usually with a green streak down the middle, flanked by streaks of purple and buff; culms medium to 

dark glossy green at first (some cultivars golden yellow or yellow streaked), remaining so in age .................... 

.................................................................................................................................................Ph. bambusoides 

4 Largest culms to 3.2 (-4) cm in diameter and 10 m tall (rarely taller); upper culm sheaths with or without 

auricles; outer surface of culm sheaths variously streaked, spotted, or mottled with brown or red (but not as 

above); culms pale green to green at first, usually becoming purple spotted, gray, or yellow in age. 

5 Lowest internode of principal culms ca. 8.5 cm long; culm sheaths with auricles, usually sparsely 

pubescent with erect, pale hairs, usually pinkish-brown at maturity, marked with numerous brown spots 

near the tip; culms green at first, usually becoming speckled and then more-or-less completely darkened 

with purplish spots (remaining green in some cultivars) ..............................................................Ph. nigra 

5 Lowest internode of principal culms ca. 12 cm long; culm sheaths lacking auricles, glabrous, usually 

green to buff at maturity, striped and marginally bordered with red; culms pale green at first, becoming 

gray to yellowish in age...............................................................................................Ph. rubromarginata 

* Phyllostachys aurea Carrière ex A. & C. Rivière, Golden Bamboo, Fishpole Bamboo. Cp, Pd (NC, SC, VA): suburban 

woodlands; uncommon, native of China and Japan. Not known to flower in our area. This is the usual large bamboo cultivated 

and naturalizing in our area, forming dense stands, up to 15 m tall. [= RAB, K, Y, Z] 

* Phyllostachys aureosulcata McClure, Yellowgroove Bamboo. Cp (SC), Pd (VA), {GA}: cultivated as an ornamental, 

persistent or spreading from plantings; rare, native of China. [= K, Y, Z] 

* Phyllostachys bambusoides Siebold & Zuccarini, Giant Timber Bamboo. Cp, Pd, Mt (NC, SC): cultivated as an 

ornamental, persistent or spreading from plantings; rare, native of China. [= K, Y, Z] 

* Phyllostachys heterocycla (Carrière) S. Matsum, Moso Bamboo. Cp (SC): cultivated as an ornamental, persistent or 

spreading from plantings; rare, native of China. [= Y; ? Ph. edulis (Carrière) Houzeau de Lehaie – K; ? Ph. pubescens Mazel ex 

Houzeau de Lehaie – Z]

POACEAE 910 

* Phyllostachys nigra (Loddiges) Munro, Black Bamboo. Pd (SC), Cp (VA): cultivated as an ornamental, persistent or 

spreading from plantings; rare, native of China and Japan. [= K, Y, Z] 

* Phyllostachys rubromarginata McClure. Pd (SC): cultivated as an ornamental, persistent or spreading from plantings; 

rare, native of China. [= K, Y, Z] 

* Phyllostachys meyeri McClure is reported as introduced in NC and SC (Kartesz 1999). {investigate} [= K] {not keyed at 

this time} 

A number of other species are sometimes cultivated in our area, and may be encountered. Bamboos are seriously underrepresented 

in herbaria, since they rarely flower and are impractical to press. All of the species above should be anticipated in 

other physiographic provinces and states than those listed. 

Piptatherum Palisot de Beauvois 

1 Leaves involute when dry, 1-2 mm wide; glumes 3.5-4.8 mm long ................................................................... [P. canadense] 

1 Leaves flat, 5-15 mm wide; glume 6-9 mm long. 

2 Leaves primarily basal or low-cauline, 2-4 (-5) dm long, 4-10 mm wide; inflorescence a raceme or a racemiform 

panicle; culms prostrate, the upper leaves very reduced, often merely bladeless sheaths .........[see Oryzopsis asperifolia] 

2 Leaves primarily cauline, 1-2.5 dm long, 8-15 mm wide; inflorescence a panicle; culms erect, the upper leaves welldeveloped..................................................................................................................................................... 

P. racemosum 

Piptatherum racemosum Ricker ex A.S. Hitchcock, Blackseed Ricegrass. Mt (VA): calcareous woodlands and forests; 

common. Québec and Ontario west to ND, south to w. VA, KY, MO, and NE. [= K; = Oryzopsis racemosa (Smith) Ricker ex 

A.S. Hitchcock – C, F, G, HC, W] 

Piptatherum canadense (Poiret) Barkworth ined., Mountain Ricegrass, ranges south to Panther Knob, Pendleton County, 

WV; it may occur in our primary area as well. [= K; = Oryzopsis canadensis (Poiret) Torrey – C, F, G, HC] 

* Piptatherum miliaceum (Linnaeus) Cosson, Smilo Grass, is reported as an introduction in MD, NJ, and PA (Kartesz 1999). 

[= K; = Oryzopsis miliacea (Linnaeus) Bentham & Hooker – HC; = Agrostis miliacea Linnaeus] {not keyed at this time} 

Piptatherum pungens (Torrey ex Sprengel) Barkworth ined. ranges south to e. PA (Rhoads & Klein 1993), NJ, and WV 

(Kartesz 1999). [= K; = Oryzopsis pungens (Torrey ex Sprengel) A.S. Hitchcock – C, F, G, HC] {not keyed at this time} 

Piptochaetium J. Presl (Needlegrass) 

A genus of about 36 species, of temperate North and South America, and montane tropical South America (Cialdella & Giussani 

2002). P. avenacioides (Nash) Valencia & Costa, endemic to FL, is the only other species in the genus in eastern North America. 

References: Cialdella & Giussani (2002). 

Piptochaetium avenaceum (Linnaeus) Parodi, Eastern Needlegrass, Black Oatgrass. Cp, Pd, Mt (GA, NC, SC, VA): 

upland woodlands and forests, sometimes abundant or even dominant in xeric woodlands over granitic or mafic rocks in the 

Piedmont; common (uncommon in the Mountains). April-June. [= C, K; = Stipa avenacea Linnaeus – RAB, F, G, HC, S, W] 

Pleioblastus Nakai 1925 

* Pleioblastus simonii (Carrière) Nakai. Reported for GA (Kartesz 1999). {investigate} [= Arundinaria simonii (Carrière) 

A.& C. Rivière – K] 

Poa Linnaeus 1753 (Bluegrass) 

A genus of about 500 species, cosmopolitan. References: Tucker (1996)=Z; Haines (2004)=Y; Soreng (1998). 

1 Plants with well-developed rhizomes; perennial. 

2 Upper stems strongly flattened; [section Tichopoa].......................................................................................P. compressa 

2 Upper stems terete or nearly so. 

3 Lower nodes of the panicle with 1-3 branches........................................................................................ P. cuspidata 

3 Lower nodes of the panicle with 4 or more branches; [section Poa]........................................................ P. pratensis 

1 Plants lacking rhizomes; perennial or annual. 

4 Plants dioecious, the florests imperfect; lemmas and glumes scarious and silvery; [rare introduction in our area]; 

[section Dioicopoa]......................................................................................................................................P. arachnifera

POACEAE 911 

4 Plants not dioecious, the florets perfect; lemmas and glumes not notably scarious and silvery; [collectively common 

and widespread in our area]; [subgenus Poa]. 

5 Lemmas not webbed at the base. 

6 Annual; culms decumbent to ascending and 1-3 dm long; inflorescence 2-8 cm long, the ascending branches 

bearing crowded spikelets above the middle; lemmas 2.4-3.4 mm long; [section Ochlopoa].............. P. annua 

6 Perennial; culms erect, 3-6 dm long; inflorescence 6-15 cm long, the widely spreading branches bearing a 

few spikelets near the end; lemmas 3.2-4.4 mm long; [section Sylvestres] ...................................P. autumnalis 

5 Lemmas webbed at the base. 

7 Spikelets (most or all) modified into purplish bulblets; culm bulbous-thickened at ground level; [section 

Bolbophorum] .................................................................................................................................... P. bulbosa 

7 Spikelets normal; culm not bulbous-thickened. 

8 Annual; [section Homalopoa] ...........................................................................................P. chapmaniana 

8 Perennial. 

9 Marginal veins of the lemma glabrous. 

10 Nodes of the panicle mostly with 4-8 branches; lemmas pubescent or scabrous on the keel. 

11 Sheaths glabrous; ligule 0.7-2.2 (-3.0) mm long; [section Sylvestres]................. P. alsodes 

11 Sheaths scabrous; ligule (2.5-) 3-7 mm long; [section Pandemos] ......................P. trivialis 

10 Nodes of the panicles mostly with 2 branches; lemmas glabrous on the keel; [section 

Sylvestres]. 

12 Anthers 0.6-0.9 (-1.0) mm long; lemmas broad-acute, obtuse or truncate at the apex, the 

keel and lateral margins of the lemma forming an apical angle of 42-82 degrees, firm at 

the tip, the scarious tip absent or up to 0.25 mm long....................................... P. languida 

12 Anthers 0.9-1.5 mm long; lemmas acute to acuminate at the apex, the keel and lateral 

margins of the lemma forming an apical angle of 10-47 degrees, pliable at the tip, the 

scarious tip prominent and 0.25-0.5 mm long..................................................P. saltuensis 

9 Marginal veins of the lemma pubescent, at least basally. 

13 Lower nodes of the panicles mostly with (1-) 2-3 branches. 

14 Ligule truncate, 0-1 mm long; first glume 1.7-2.2 mm long, second glume 2.0-2.8 mm 

long; anthers 0.5-0.7 mm long.......................................................................P. paludigena 

14 Ligule rounded-ovate, 1-2 mm long; first glume 2.5-3.5 mm long, second glume 3.0-3.8 

mm long; anthers 1.1-1.4 mm long.........................................................................P. wolfii 

13 Lower nodes of the panicles mostly with (4-) 5 or more branches. 

15 Lemmas 5-veined (intermediate veins well-developed); ligule ca. 1 mm long; [section 

Sylvestres]......................................................................................................... P. sylvestris 

15 Lemmas 3-veined (intermediate veins obscure); ligule either (2-) 3-5 mm long or 0.2-1 (- 

1.5) mm long. 

16 Ligule 0.2-1 (-1.5) mm long, truncate; culms 4-8 dm tall; anthers 1.2-1.6 mm long; 

[section Stenopoa] ...................................................................................P. nemoralis 

16 Ligule (2-) 3-5 mm long, ovate-triangular; culms 5-15 dm tall; anthers 0.8-1.2 mm 

long; [section Pandemos]...........................................................................P. palustris 

Poa alsodes A. Gray, Woodland Bluegrass. Mt (NC, VA), Pd (NC): rich forests; uncommon. May-June. Nova Scotia west 

to SD, south to NC and IL; also in w. United States. [= RAB, C, F, G, HC, K, W, Z] 

* Poa annua Linnaeus, Speargrass, Six-weeks Grass, Annual Bluegrass. Cp, Pd, Mt (GA, NC, SC, VA): fields, roadsides, 

disturbed areas; common, introduced from Eurasia. April-May. [= RAB, C, F, G, GW, HC, K, W, Z] 

* Poa arachnifera Torrey, Texas Bluegrass. Pd (GA, NC, SC): disturbed areas; rare, introduced from South America and w. 

United States. April. [= RAB, HC, K] 

Poa autumnalis Muhlenberg ex Elliott. Cp, Pd, Mt (GA, NC, SC, VA): moist or dry nutrient-rich forests; common. April- 

May. NJ west to MI, south to FL and TX. [= RAB, C, F, G, GW, HC, K, W, Z] 

* Poa bulbosa Linnaeus, Bulbous Bluegrass. Cp (NC, VA), Pd (GA, NC, VA): lawns; rare, introduced from Europe. April- 

May. [= RAB, C, F, G, HC, K, Z] 

Poa chapmaniana Scribner. Pd, Cp (GA, NC, SC, VA): low fields, roadsides, disturbed areas; common (VA Watch List). 

April-May. DE west to IA, south to FL and LA. [= RAB, C, F, G, HC, K, W, Z] 

* Poa compressa Linnaeus, Canada Bluegrass. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): fields, roadsides, disturbed 

areas; common, introduced from Europe. May-August. [= RAB, C, F, G, HC, K, W, Z] 

Poa cuspidata Nuttall. Mt, Pd (GA, NC, SC, VA), Cp (GA, NC, VA): moist forests; common. March-April. NJ west to s. 

IN, south to sw. GA. [= RAB, C, F, G, HC, K, W, Z] 

Poa languida Hitchcock, Drooping Bluegrass. Mt (VA): ultramafic outcrop woodlands, barrens, and glades; rare (VA Rare 

as P. saltuensis). April-May. VT and MA west to MN, south to PA, w. VA, KY, and IA. See comments under P. saltuensis. [= 

C, F, G, HC, W; < P. saltuensis – K; = P. saltuensis Fernald & Wiegand ssp. languida (Hitchcock) A. Haines – Y] 

* Poa nemoralis Linnaeus, Wood Bluegrass. Mt (NC, VA), Pd (VA): sandy creek bottoms; rare, introduced from Europe 

(NC Watch List). [= C, F, G, HC; ? P. nemoralis ssp. nemoralis – K] 

Poa paludigena Fernald & Wiegand, Bog Bluegrass. Mt (NC, VA): mountain bogs, especially in deep shade under shrubs; 

rare (US Species of Concern, NC Endangered, VA Rare). April-May. NY west to MN, south to PA, w. NC, and IL. This

POACEAE 912 

species withers and disintegrates shortly after flowering; its ephemeral habit may be responsible for its being overlooked in our 

area for many years. [= C, F, G, HC, K] 

Poa palustris Linnaeus, Fowl Bluegrass, Fowl Meadow-grass. Mt (NC, SC, VA), Pd (VA): meadows, moist areas; rare 

(NC Rare, VA Rare). June-July. Circumboreal, south in North America to VA, w. NC, MO, and NM. [= RAB, C, F, G, HC, K, 

W, Z] 

* Poa pratensis Linnaeus, Kentucky Bluegrass, Junegrass, Speargrass. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): lawns, 

roadsides, disturbed areas; common, introduced from Europe. April-August. [= RAB, C, F, G, HC, W, Z; P. pratensis ssp. 

pratensis – K] 

Poa saltuensis Fernald & Wiegand, Old-pasture Bluegrass. Mt (NC, VA): northern hardwood forests, ultramafic outcrop 

woodlands, barrens, and glades; rare (NC Rare, VA Rare). April-May. Newfoundland west to MN, south to PA, w. VA, and w. 

NC. The NC occurrences (on serpentinized olivine barrens) reported as P. languida are actually P. saltuensis. The taxonomic 

distinctions between P. saltuensis and P. languida have been controversial; Haines (2004) provides a detailed and valuable 

discussion. [= C, F, G, HC, W; < P. languida – RAB, Z, misidentification; = P. saltuensis var. saltuensis – F; < P. saltuensis – 

K (also see P. languida); = P. saltuensis ssp. saltuensis – Y] 

Poa sylvestris A. Gray, Forest Bluegrass. Mt, Pd, Cp (GA, NC, SC, VA): moist forests; common. April-May. NY west to 

WI and IA, south to FL and TX. [= RAB, C, F, G, GW, HC, K, W, Z] 

* Poa trivialis Linnaeus, Rough Bluegrass. Mt, Pd (NC, VA), Cp (VA), {GA}: moist forests, disturbed areas, bottomlands; 

common, introduced from Europe. April-June. [= RAB, C, F, G, GW, HC, K, W, Z] 

Poa wolfii Scribner. Mt (NC, VA): moist rich forests; rare (VA Rare). {} OH west to MN, south to n. VA, MO, and e. 

NE. The NC occurrence is based on material from Great Smoky Mountains National Forest (Haywood County) (K. Langdon, 

pers. comm.. 2006). [= C, F, G, HC, K, W, Z] 

* Poa infirma Kunth. Introduced in SC (Kartesz 1999), but not attributed to any e. North American area in FNA (in prep.). 

{investigate} [= K] {not keyed at this time; synonymy incomplete} 

References: Tucker (1996)=Z; Barkworth in FNA (in prep.). 

Polypogon Desfontaines 

1 Inflorescence verticillate, the rachis visible between the verticils; glumes 1.6-2.3 mm long, without awns; spikelets 

disarticulating near base of pedicel; stoloniferous perennial.........................................................................................P. viridis 

1 Inflorescence dense, cylindrical, and spikelike; glumes 2-3 mm long, with prominent awns 3.5-7 mm long; spikelets 

disarticulating near apex of pedicel; annual. 

2 Glumes deeply lobed, the awn borne between the lobes; glume ciliate-fringed; lemma 0.4-0.7 mm long, awnless ........... 

.............................................................................................................................................. P. maritimus var. maritimus 

2 Glumes slightly notched at the tip, the awn borne from near the tip; glume not ciliate-margined; lemma 0.7-1.1 mm 

long, awned ............................................................................................................................................. P. monspeliensis 

* Polypogon maritimus Willdenow var. maritimus, Meditteranean Beardgrass. Cp (GA, SC): brackish marshes; rare, 

introduced from Meditteranean Europe. P. maritimus Willdenow is reported as introduced to GA (Small 1933). [< P. maritimus 

– HC, K, S, Z] 

* Polypogon monspeliensis (Linnaeus) Desfontaines, Rabbitfoot Grass, Beardgrass, Annual Beardgrass. Cp (GA, NC, SC, 

VA), Pd (GA): brackish marshes, disturbed areas; uncommon, introduced from s. Europe and w. Asia. May-July. [= RAB, C, F, 

G, GW, HC, K, S, Z] 

* Polypogon viridis (Gouan) Breistr., Water Bent-grass. Cp (SC): introduced on ballast around old ports, probably not 

persistent; rare, introduced from the Old World. Distinguished from Agrostis in having the spikelet falling as a whole, 

disarticulating below the glumes. [= K, Z; = Agrostis viridis Gouan – C; ? Agrostis verticillata Villars – F; ? Agrostis 

semiverticillata (Forskål) C. Christensen – G, HC] 

Pseudosasa Makino ex Nakai (Arrow Bamboo) 

References: Duncan & Duncan [in prep.]=Z; Judziewicz et al. (2000)=Y. Key adapted from Z. 

* Pseudosasa japonica (Siebold & Zuccarini ex Steudel) Makino ex Nakai, Arrow Bamboo. Cp (VA): cultivated as an 

ornamental, persistent or spreading from plantings; rare, native of Japan. [= K, Y, Z; = Sasa japonica (Siebold & Zuccarini ex 

Steudel) Makino] 

A genus of about 80 species, north temperate. 

Puccinellia Parlatore (Alkali Grass, Goosegrass)

POACEAE 913 

1 Lemmas 3.0-4.5 mm long; spikelets 5-11-flowered.............................................................................................. [P. maritima] 

1 Lemmas 1.5-2.5 mm long; spikelets 2-6-flowered. 

2 Inflorescence diffuse, the lower branches with spikelets restricted to the distal portions; lower inflorescence branches 

spreading horizontal to deflexed at maturity; lemma 1.5-2.1 mm long, the midnerve not reaching the apex ......P. distans 

2 Inflorescence compact, the lower branches bearing spikelets nearly to the base; lower inflorescence branches ascending 

at maturity; lemma 2.0-2.5 mm long, the midnerve reaching the apex, and often excurrent as a mucro....... P. fasciculata 

* Puccinellia distans (Jacquin) Parlatore, European Alkali Grass, Goosegrass. Cp (VA): coastal sands; rare, introduced from 

Europe. [= P. distans – C, G, HC; > P. distans var. distans – F; > P. distans ssp. distans – K] 

Puccinellia fasciculata (Torrey) Bicknell, Eastern Alkali Grass, Saltmarsh Goosegrass. Cp (VA): salt or brackish marshes; 

rare (VA Rare). Nova Scotia south to VA; Europe; and in sw. United States. [= C, F, G, HC, K] 

* Puccinellia maritima (Hudson) Parlatore, Seaside Alkali Grass, Seaside Speargrass, salt marshes and ballast near ports, is 

introduced south to se. PA (Philadelphia), NJ (Camden), and DE, especially on ballast. [= C, F, G, HC; > P. americana 

Sorenson – K] {synonymy incomplete} 

A genus of about 4 species, of the New World tropics. 

Reimarochloa A.S. Hitchcock 

Reimarochloa oligostachya (Munro ex Bentham) A.S. Hitchcock. AL, FL; Cuba. [= HC, K] 

Rhynchelytrum Nees 

(see Melinis) 

References: Hodkinson et al. (2002). 

Ripidium Trinius (Ravenna-grass) 

* Ripidium ravennae (Linnaeus) Trinius, Ravenna-grass, Plume-grass. Cp (GA): cultivated as an ornamental and rarely 

escaping or persisting; rare, introduced from s. Europe. In sw. GA, TN, and MD (Kartesz 1999) and DC (Steury 2004a). [= 

Saccharum ravennae (Linnaeus) Linnaeus – FNA, K; = Erianthus ravennae (Linnaeus) Palisot de Beauvois – F; > Erianthus 

ravennae var. ravennae – HC; > Erianthus ravennae var. purpurascens (Anderss.) Hackel – HC] 

Rostraria Trinius 

* Rostraria cristata (Linnaeus) Tzvelev. Cp (SC): waste areas near wool-combing mills; rare, introduced. It also occurs at 

scattered other sites in eastern United States, such as on ballast in se. PA (Rhoads & Klein 1993), and reported for MD, AL, and 

FL (Kartesz 1999). Not keyed. [= K; = Lophochloa cristata (Linnaeus) Hylander; ? Koeleria phleoides (Vill.) Persoon – HC] 

Rottboellia Linnaeus f. (Itch-grass) 

A genus of about 5 species, native to tropical Asia and Africa. References: Wipff in FNA (2003a); Wipff & Rector (1993)=Z. 

* Rottboellia cochinchinensis (Loureiro) Clayton, Itch-grass. Cp (GA, NC): disturbed ground; rare, native of tropical Asia. 

August-October. This grass, considered a noxious weed, was found in at least 13 GA counties by 1985 (Duncan 1985) and on a 

farm in Robeson County, NC in 1984. [= FNA, K, Z; = Rottboellia exaltata Linnaeus f. – HC; = Manisuris exaltata (Linnaeus 

f.) Kuntze – S] 

Saccharum Linnaeus (Plume Grass) 

(also see Ripidium) 

A genus of uncertain circumscription at this time. Clayton & Renvoize (1986) have pointed out that the "traditional division [of 

Saccharum] into awned (Erianthus) and awnless species seems wholly artificial;" Hodkinson et al. (2002) develop molecular 

evidence which suggests that our species are not congeneric with Saccharum, however. Further study is needed, but likely our 

native southeastern species will be merged into Miscanthidium Stapf, while the introduced S. ravennae will be placed in the 

genus Ripidium Trinius (Hodkinson et al. 2002). Sugarcane (Saccharum officinarum Linnaeus, S. sinense Roxburgh, S. barberi 

Jeswiet, S. spontaneum Linnaeus, and cultivars and hybrids derived from those four species) is cultivated further south, notably in

POACEAE 914 

FL and LA. References: Webster in FNA (2003a); Webster & Shaw (1995)=Z; Gandhi & Dutton (1993); Hodkinson et al. 

(2002). 

1 Lowermost inflorescence node densely hairy; callus hairs (ring of hairs beneath the spikelet) (7-) 9-25 mm long, equal to or 

longer than the spikelet; stem appressed-pubescent below the inflorescence, on the internodes as well as the nodes. 

2 Lemma awn flattened and spirally twisted at base; callus hairs 9-14 mm long, silvery or tinged with purple; leaves 

usually glabrous on the upper surface at maturity; [of moist to dry sites, rarely in wetlands]...................S. alopecuroides 

2 Lemma awn nearly terete, straight or slightly flexuous; callus hairs (7-) 15-20 (-25) mm long, tawny or brown; leaves 

usually pilose on the upper surface at maturity; [of moist to wet sites, rarely in uplands].............................S. giganteum 

1 Lowermost inflorescence node glabrous; callus hairs (ring of hairs beneath the spikelet) 0-6.5 mm long, shorter than or equal 

to the spikelet (or absent in S. brevibarbe); stem glabrous below the inflorescence, except sometimes on the nodes. 

3 Callus hairs (ring of hairs beneath the spikelet) absent, or of few hairs 0-2 mm long (much shorter than the spikelet); 

panicle branches closely appressed, the panicle usually 1-3 cm broad; panicle branches glabrous ................. S. baldwinii 

3 Callus hairs (ring of hairs beneath the spikelet) present, dense, 3-6.5 mm long (from about half as long to nearly as long 

as the spikelet); panicle branches ascending, the panicle usually 4-10 cm broad; panicle branches pubescent. 

4 Awn of the lemma of the upper floret terete at the base, and not spiraled; spikelets dark brown; spikelet pair 

dissimilar in size, the lemma of the upper floret 0.7-0.8× as long as the lemma of the lower floret; lemma of the 

lower floret typically 3-nerved .............................................................................................................S. coarctatum 

4 Awn of the lemma of the upper floret flattened at the base, either spiraled or not; spikelets straw-colored or 

purplish; spikelet pair homomorphic, the upper lemma 0.9-1.0× as long as the lower lemma; lemma of the lower 

floret not distinctly nerved. 

5 Awn of the lemma of the upper floret not basally spiraled, 10-18 mm long; lemma of the upper floret entire... 

............................................................................................................................. S. brevibarbe var. brevibarbe 

5 Awn of the lemma of the upper floret basally spiraled, 15-22 mm long; lemma of the upper floret bifid, the 

tooth on either side of the lemma 2.0-2.5 mm long...............................................S. brevibarbe var. contortum 

Saccharum alopecuroides (Linnaeus) Nuttall, Silver Plume Grass. Pd, Mt, Cp (GA, NC, SC, VA): fields, roadsides, 

woodland borders; common (rare in Mountains). October. NJ west to IN, IL, MO, and OK, south to FL and TX. [= FNA; = 

Saccharum alopecuroideum (Linnaeus) Nuttall – Z, orthographic variant; = Erianthus alopecuroides (Linnaeus) Elliott – RAB, 

C, F, G, GW, HC, W; = Saccharum alopecuroidum – K, orthographic variant; = Erianthus divaricatus (Linnaeus) A.S. 

Hitchcock – S; = Miscanthidium species 1] 

Saccharum baldwinii Sprengel, Narrow Plume Grass. Cp (GA, NC, SC, VA): marshes, clay-based Carolina bays, ditches; 

common. July-October. E. VA south to FL, west to TX, AR, scattered northward inland to TN and MO. [= FNA, K, Z; = 

Erianthus strictus Elliott – RAB, C, F, G, GW, HC, S; = Miscanthidium species 2] 

Saccharum brevibarbe (Michaux) Persoon var. brevibarbe, Short-bearded Plume Grass. Cp (NC): marshes, ditches; rare. 

September-October. MS, AL, and TN west to TX, AR, and OK; disjunct in e. NC. [= FNA, K, Z; < Erianthus brevibarbis 

Michaux – RAB, C, G, GW, S (also see S. coarctatum); = E. brevibarbis – F; >< Erianthus coarctatus Fernald var. coarctatus – 

HC; >< Erianthus coarctatus var. elliottianus Fernald – HC; = Miscanthidium species 3] 

Saccharum brevibarbe (Michaux) Persoon var. contortum (Elliott) R. Webster, Bent-awn Plume Grass. Cp, Pd, Mt (GA, 

NC, SC, VA): open woodlands and forests, woodland borders; common, rare in Mountains. Late July-October. DE and MD 

south to panhandle FL, west to TX and AR, with scattered occurrences north to TN. [= FNA, K, Z; = Erianthus contortus Elliott 

– RAB, C, F, G, GW, HC, S, W; = Saccharum contortum (Elliott) Nuttall; = Erianthus brevibarbis Michaux var. contortus 

(Elliott) D.B. Ward; = Miscanthidium species 4] 

Saccharum coarctatum (Fernald) R.D. Webster, Brown Plume Grass. Cp (GA, NC, SC, VA): marshes, ditches, clay-based 

Carolina bays, swamps; common (rare in VA). September-October. DE and MD south to FL, west to TX (Brown & Marcus 

1998). [= FNA, K, Z; < Erianthus brevibarbis Michaux – RAB, C, G, GW, S; >< Erianthus coarctatus Fernald – F, HC; >< 

Erianthus coarctatus var. coarctatus – HC; >< Erianthus coarctatus var. elliottianus Fernald – HC; = Miscanthidium species 5] 

Saccharum giganteum (Walter) Persoon, Sugarcane Plume Grass, Giant Plume Grass. Cp, Pd, Mt (GA, NC, SC, VA): 

marshes, ditches; common, rare in Mountains. September-October. NY south to FL, west to se. TX and AR; inland in TN and 

KY. [= FNA, K, Z; = Erianthus giganteus (Walter) Palisot de Beauvois – RAB, C, G, GW, HC, W; > Erianthus giganteus var. 

giganteus – F; > Erianthus giganteus var. compactus (Nash) Fernald – F; = Erianthus saccharoides Michaux – S; = 

Miscanthidium species 6] 

Sacciolepis Nash 

A genus of about 30 species, primarily in the tropics and subtropics. References: Wipff in FNA (2003a). 

1 Annual, cespitose; spikelets 2.5-3.5 mm long; [rare alien] ........................................................................................... S. indica 

1 Perennial, from creeping stolons; spikelets (3-) 4-5 mm long; [common native] .........................................................S. striata 

* Sacciolepis indica (Linnaeus) Chase. Cp (GA, NC, SC): low fields, ditches; rare, introduced from India. October. [= 

RAB, FNA, GW, HC, K]

POACEAE 915 

Sacciolepis striata (Linnaeus) Nash, American Cupscale. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): marshes, 

interdune swales, ditches, swamps; common (rare in Piedmont and Mountains). July-October. S. NJ south to FL, west to e. TX 

and OK, nearly limited to the Coastal Plain, but occasionally inland as in w. NC and TN; also native in the West Indies and n. 

South America. [= RAB, C, F, FNA, G, GW, HC, K, W] 

Schedonorus Palisot de Beauvois 

The correct generic placement of the introduced species Schedonorus arundinaceus (= Festuca elatior; = Festuca arundinacea; 

= Lolium arundinaceum) and Sch. pratense has been disputed. The traditional placement in Festuca has been defended by Aiken 

et al. (1997); Darbyshire (1993) transferred them to Lolium; and Soreng & Terrell (1998) place them in the genus Schedonorus. 

NOTE: apparently, the Soreng & Terrell (1998) paper was preceded by another paper which renders Soreng & Terrell's 

combinations superfluous. References: Darbyshire (1993)=X; Aiken & Darbyshire (1990)=Y; Tucker (1996)=Z; Soreng & 

Terrell (1998)=V; Darbyshire in FNA (in prep.). Key based in part on C and Y. 

1 Auricles ciliate (sometimes only very sparsely so – check several at 10-20× magnification); spikelets with 3-6 (-9) florets; 

old sheaths pale straw-colored, often remaining intact; internodes of the rachilla antrorsely scabrous ........Sch. arundinaceus 

1 Auricles glabrous; spikelets with (2-) 4-10 (-12) florets; old sheaths brown, decaying to fibers; internodes of the rachilla 

glabrous (smooth) or nearly so..............................................................................................................................Sch. pratensis 

* Schedonorus arundinaceus (Schreber) Dumortier, Tall Fescue, Alta Fescue. Cp, Pd, Mt (GA, NC, SC, VA): fields, 

roadsides, pastures, disturbed areas; common, introduced from Eurasia. May-July. [= FNA, V; < Festuca elatior Linnaeus – 

RAB, F, S, W, misapplied; = Festuca arundinacea Schreber – HC, Y; = Festuca elatior Linnaeus – C; = Festuca elatior var. 

arundinacea (Schreber) Wimmer – G; < Festuca pratensis Hudson – GW; = Lolium arundinaceum (Schreber) Darbyshire – K, 

X, Z; ? Schedonorus phoenix (Scopoli) Holub] 

* Schedonorus pratensis (Hudson) Palisot de Beauvois, Meadow Fescue. Mt (VA): fields, roadsides, pastures, disturbed 

areas; rare, introduced from Eurasia. May-July. [= FNA, V; < Festuca elatior Linnaeus – F, S, W, misapplied; = Festuca 

pratensis Hudson – C, Y; = Festuca elatior var. pratensis (Hudson) A. Gray – G; < Festuca pratensis Hudson – GW; = Festuca 

elatior – HC, misapplied; = Lolium pratense (Hudson) Darbyshire – K, X, Z ] 

A monotypic genus, circumboreal in Asia and North America. 

Schizachne Hackel (False Melic) 

Schizachne purpurascens (Torrey) Swallen, Purple Oatgrass, False Melic. Mt (VA): moist, rocky northern hardwood and 

spruce forests; rare (VA Rare). Newfoundland west to AK, south to MD, w. VA, WV, KY, IL, NM, and Mexico; also in ne. 

Asia. May-July. [= F, G, HC, K; > S. purpurascens var. purpurascens – C] 

Schizachyrium Nees (Little Bluestem) 

A genus of about 60 species, widespread in tropical, subtropical, and warm temperate regions of the World. References: Wipff 

(1996a)=Z; Gandhi (1989)=Y; Wipff in FNA (2003a). Key based in part on Wipff in FNA (2003a). 

1 Leaf blades 0.5-1.5 mm wide, with a lighter-colored zone in the center of the upper surface; sessile spikelet ca. 4 mm long ... 

...............................................................................................................................................................................Sch. tenerum 

1 Leaf blades >1.5 mm wide, lacking a distinct lighter zone on the upper surface; sessile spikelet 5-11 mm long. 

2 First glume of sessile spikelet pubescent ......................................................................Sch. sanguineum var. hirtiflorum 

2 First glume of sessile spikelet glabrous. 

3 Plants rhizomatous, with internodes 6 mm long or longer; sessile spikelet 5-7 mm long ........................................... 

.............................................................................................................................. Sch. scoparium var. stoloniferum 

3 Plants tufted, rhizome internodes absent or < 3 mm long, the stem sometimes decumbent at the base and rooting at 

the lower nodes (appearing nearly rhizomatous); sessile spikelet 6-10 mm long. 

4 Leaf sheaths broad and strongly keeled, hairs of the raceme internodes ca. 5 mm long; stems decumbent at 

base, rooting at the lower nodes ..................................................................................................... Sch. littorale 

4 Leaf sheaths rounded or weakly keeled; hairs of the raceme internodes 1-3 (-4) mm long; stems erect, not 

rooting at the lower nodes. 

5 Pedicellate spikelets of the proximal spikelet units on each rame staminate, 5-10 mm long, with a 

lemma, the pedicellate spikelets of the distal units usually smaller (1-4 mm long) and sterile; sheaths and 

blades densely tomentose to glabrate .......................................................[Sch. scoparium var. divergens] 

5 Most pedicellate spikelets sterile, 1-6 mm long, without a lemma; sheaths and blades usually glabrous, 

occasionally pubescent..............................................................................Sch. scoparium var. scoparium

POACEAE 916 

Schizachyrium littorale (Nash) Bicknell, Seaside Little Bluestem. Cp (GA, NC, VA): coastal dunes and maritime dry 

grasslands, often with Uniola paniculata, Panicum amarum, and other dune plants; common. August-October. E. MA south to 

NC (or SC?), and inland on the shores of the Great Lakes. In NC, Sch. littorale is present and abundant on dunes of barrier 

islands from Shackleford Banks, Carteret County south to Brunswick County, near the SC border, and entirely absent from the 

Outer Banks (from Cape Lookout, Carteret County, north through Hyde County to Dare County). [= FNA, GW, K; < 

Andropogon scoparium Michaux – RAB; = Sch. scoparium var. littorale (Nash) Gould – C, Z; = Andropogon scoparius 

Michaux var. littoralis (Nash) A.S. Hitchcock – F, G; = Andropogon littoralis Nash – HC, S; < Sch. scoparium (Michaux) Nash 

ssp. littorale (Nash) Gandhi & Smeins – Y] 

Schizachyrium sanguineum (Retzius) Alston var. hirtiflorum (Nees) Hatch, Hairy Crimson Bluestem. Cp (GA): 

{habitat}; rare. Sw. GA and FL west to AZ and south through Central America to South America. [= FNA, K; = Andropogon 

hirtiflorus (Nees) Kunth – HC, S; ? Sch. sanguineum var. brevipedicellatum (Beal) Hatch] 

Schizachyrium scoparium (Michaux) Nash var. scoparium, Common Little Bluestem. Cp, Pd, Mt (GA, NC, SC, VA): in a 

wide range of moist to dry habitats; common. (June-) August-October. New Brunswick west to Alberta, south to FL and 

Mexico. One of the most ubiquitous plants in the modern landscape of our area, occurring throughout in the majority of habitats. 

This species is extremely variable, some of the variability correlated with habitat and geography; the recognition of infraspecific 

taxa is warranted. [= C, FNA, Z; < Andropogon scoparius Michaux – RAB (also see Sch. littorale); = Sch. scoparium – GW; > 

Andropogon scoparius var. scoparius – F, G, HC; > Andropogon praematurus Fernald – F, G; > Andropogon scoparius var. 

polycladus Scribner & Ball – F; > Andropogon scoparius var. frequens F.T. Hubbard – F; = Sch. scoparium ssp. scoparium – K, 

Y; < Andropogon scoparius – S, W] 

Schizachyrium scoparium (Michaux) Nash var. stoloniferum (Nash) J. Wipff, Creeping Little Bluestem. Cp (GA, SC): 

fall-line sandhills in the inner Coastal Plain, perhaps in other dry habitats, the habitat and range in our area requiring further 

study; uncommon? August-October. SC and GA south to FL and west to MS. See Wipff (1996a) for additional discussion. [= 

FNA, K, Z; = Sch. stoloniferum Nash – GW; = Andropogon stolonifer (Nash) A.S. Hitchcock – HC, S; < Sch. scoparium ssp. 

littorale (Nash) Gandhi & Smeins – Y] 

Schizachyrium tenerum Nees, Slender Bluestem. Cp (GA): longleaf pine savannas; uncommon. GA west to e. TX. [= 

FNA, K; = Andropogon tener (Nees) Kunth – HC, S] 

Schizachyrium maritimum (Chapman) Nash. AL, FL west to LA. [= K; = Andropogon maritimus Chapman] {not keyed at 

this time; add to synonymy} 

Schizachyrium niveum (Swallen) Gould, Pinescrub Bluestem, is reported for Lowndes Co. in sc. GA (Kral 1973), but the 

report has been discounted by later authors (Wipff in FNA 2003a). Not keyed. [= FNA, K; = Andropogon niveus Swallen – HC, 

S] 

Schizachyrium scoparium (Michaux) Nash var. divergens (Hackel) Gould, Pinehill Bluestem. East to c. TN, AL, KY. [= 

FNA, K; = Andropogon scoparius Michaux var. divergens Hackel; = Andropogon divergens – HC; < Andropogon scoparius – 

S] 

Sclerochloa Palisot de Beauvois 

References: Tucker (1996)=Z; Brandenburg, Estes, & Thieret (1991). 

* Sclerochloa dura (Linnaeus) Palisot de Beauvois. Mt (VA): disturbed areas; rare, introduced from Mediterranean Europe. 

A monotypic genus, native to s. Europe. [= C, HC, K, Z] 


Secale Linnaeus 1753 (Rye) 

* Secale cereale Linnaeus, Rye. Cp, Pd, Mt (GA, NC, SC, VA): fields; commonly cultivated, rarely persistent or 

volunteering following cultivation, introduced from Eurasia. May-June. An important crop. The lemmas are awned from 2-6 

cm long. [= RAB, C, F, G, HC, K, Z] 

Setaria Palisot be Beauvois 1807 (Foxtail Grass) 

(also see Pennisetum) 

A genus of about 110-140 species, of tropicala nd warm temperate regions. References: Webster (1993)=Z; Webster (1995)=Y; 

Crins (1991)=X; Webster (1988); Rominger in FNA (2003a); Allen in FNA (2003a). Key adapted from FNA. 

Setaria corrugata (Elliott) J.A. Schultes. Cp (GA, NC, SC): pinelands, disturbed areas; common. From ne. NC south to s. 

FL, west to e. TX; Cuba; Dominican Republic. [= RAB, FNA, HC, K, Z; = Chaetochloa corrugata (Elliott) Lamson-Scribner – 

S]

POACEAE 917 

* Setaria faberi R.A.W. Herrmann, Nodding Foxtail Grass, Giant Foxtail-grass. Mt, Pd, Cp (GA, NC, SC, VA): disturbed 

areas; uncommon, native of China. [= RAB, C, FNA, G, K, W; = S. faberii – F, HC, Z, orthographic variant] 

Setaria geminata (Forsskål) Veldkamp var. paludivaga (A.S. Hitchcock & Chase) R.D. Webster, Alligator Grass, 

Paspalidium. Cp (GA, SC): in shallow water; rare. December. S. SC south to FL, west to TX; also in Central and South 

America. This taxon is sometimes considered an introduction from the Old World, but its occurrence in undisturbed wetlands 

remote from extensive human activity suggests that it is native. Webster (1995) has merged Paspalidium into Setaria. [= Y; = 

Panicum paludivagum A.S. Hitchcock & Chase – RAB, HC, S; < Paspalidium geminatum – FNA, GW, X; = Paspalidium 

geminatum (Forsskål) Stapf var. paludivagum (A.S. Hitchcock & Chase) Gould – K; = Paspalidium paludivagum (A.S. 

Hitchcock & Chase) Parodi] 

* Setaria italica (Linnaeus) Palisot de Beauvois, Foxtail-millet, Italian-millet. Pd (GA, NC, SC, VA), Cp, Mt (VA): 

disturbed areas, rare, native of Eurasia. Probably derived via cultivation from S. viridis, and cultivated as a food crop in China 

since at least 6000 BP and later in Europe (Hancock 2004). [= RAB, C, F, FNA, G, HC, K, W, Z; = Chaetochloa italica 

(Linnaeus) Lamson-Scribner – S] 

Setaria macrosperma (Lamson-Scribner & Merrill) K. Schumann, Coral Bristlegrass. Cp (GA, SC): hammocks and 

maritime forests, also disturbed areas; rare. SC south to FL; Bahamas, Mexico. [= RAB, FNA, HC, K, Z; = Chaetochloa 

macrosperma Lamson-Scribner & Merrill – S] 

Setaria magna Grisebach, Saltmarsh Foxtail-grass, Giant Foxtail-grass. Cp (GA, NC, SC, VA), Pd* (GA*): interdune 

swales, near-coastal marshes; uncommon. NJ south to s. FL, west to e. TX; disjunct inland in GA, AR, LA, TX, and NM; West 

Indies, Bermuda, Costa Rica. [= RAB, C, F, FNA, G, HC, K, Z; = Chaetochloa magna (Grisebach) Lamson-Scribner – S] 

Setaria parviflora (Poiret) Kerguélen, Knotroot Bristlegrass, Perennial Foxtail-grass. Cp, Pd, Mt (GA, NC, SC, VA): 

marshes, ditches, moist disturbed areas; common. MA to IA south to s. FL and s. TX, south through Mexico to Central America; 

CA and NV; West Indies. Gandhi & Barkworth (2003) provide a detailed discussion of the reasons for the nomenclatural 

change. [= FNA, K, Z; = S. geniculata Palisot de Beauvois – RAB, C, F, G, HC, W; = Chaetochloa geniculata (Palisot de 

Beauvois) Millspaugh & Chase – S] 

* Setaria pumila (Poiret) Roemer & Schultes ssp. pumila, Yellow Foxtail. Mt, Pd, Cp (GA, NC, SC, VA): disturbed areas, 

lawns, fields; common, native of Europe. [= FNA; = Setaria glauca (Linnaeus) Palisot de Beauvois – RAB, C, F, G, W, 

misapplied; >< Setaria lutescens (Weigel) Hubb. – HC, misapplied; >< S. pumila ssp. pallidifusca – K, treatment apparently 

garbled; = Chaetochloa lutescens (Weigel) Stuntz – S] 

* Setaria verticillata (Linnaeus) Palisot de Beauvois, Hooked Bristlegrass. Mt (VA): disturbed areas; uncommon, introduced 

from Europe. [= FNA, G, K; = S. verticillata var. verticillata – C, F, HC; = Chaetochloa verticillata (Linnaeus) Lamson- 

Scribner – S; < S. verticillata – Z] 

* Setaria viridis (Linnaeus) Palisot de Beauvois var. viridis. Cp, Pd, Mt (GA, NC, SC, VA): fields, disturbed areas; common, 

introduced from Eurasia. [= C, FNA, K, Z; < S. viridis – RAB, HC, W; > S. viridis var. viridis – F, G; > S. viridis var. 

weinmannii (Roemer & J.A. Schultes) Bolbás – F; > S. viridis var. breviseta (Doell) A.S. Hitchcock – G; = Chaetochloa viridis 

(Linnaeus) Lamson-Scribner – S] 

Setaria adhaerans (Forsskål) Chiovenda. Distributed widely throughout the tropics and subtropics, in North America from 

s. AL west to CA (perhaps only adventive in portions of that distribution). [= FNA, K, Z] {synonymy incomplete} 

Setaria geminata (Forsskål) Veldkamp var. geminata. AL, FL. [= Y; < Paspalidium geminatum – FNA, GW, X; = 

Panicum geminatum Forsskål – HC, S; = Paspalidium geminatum (Forsskål) Stapf var. geminatum – K] 

* Setaria verticilliformis Dunart. is reported for NJ, PA, MD, and AL (FNA 2003a, Kartesz 1999). [= FNA, K; = S. 

verticillata (Linnaeus) Palisot de Beauvois var. ambigua (Guss.) Parlatore – C, F, HC; = S. viridis (Linnaeus) Palisot de 

Beauvois var. ambigua (Guss.) Coss. & Durieu – G; = Chaetochloa ambigua Guss. – S; < S. verticillata – Z] 

* Setaria viridis (Linnaeus) Palisot de Beauvois var. major (Gaudin) Pospichal, Giant Green Foxtail, is reported as introduced 

in TN, MD, and PA (Kartesz 1999). [= C, FNA, G, K, Z; < S. viridis – RAB, HC] 

Sorghastrum Nash (Indiangrass) 

A genus of about 18-20 species, of tropical and subtropical America and Africa, rarely extending into temperate areas. 

References: Hall (1982)=Z; Dávila Aranda & Hatch in FNA (2003a). Key adapted from Z. 

1 Awns 10-22 (-30) mm long, once-geniculate; plants rhizomatous; surfaces of the glumes tan to slightly brown basally; ligule 

3-10 mm long, prominently auricled............................................................................................................................ S. nutans 

1 Awns 16-46 mm long, twice-geniculate; plants cespitose; surfaces of the glumes brown; ligule 1-5 mm long, truncate. 

2 Axis of the panicle straight, erect, the branchlets appressed to ascending, the spikelets drooping-secund; spikelets 0.8- 

1.2 mm wide ...................................................................................................................................................S. secundum 

2 Axis of the panicle arching, usually strongly so, the branchlets ascending to spreading, the spikelets not droopingsecund; 

spikelets 1.1-1.8 mm wide. 

3 Axis of the panicle straight, with the branches distributed no more than 180 degrees around the axis (as viewed 

from above)................................................................................................................................ [S. apalachicolense] 

3 Axis of the panicle arching, with the branchlets distributed through 360 degrees around the axis (as viewed from 

above)...........................................................................................................................................................S. elliottii

POACEAE 918 

Sorghastrum elliottii (C. Mohr) Nash, Slender Indiangrass. Cp, Pd (GA, NC, SC, VA), Mt (GA, NC, SC): woodlands and 

forests, river-scour areas, including oak-hickory forests and woodlands over mafic rocks; uncommon. September-October. MD 

south to FL and west to TX, inland to TN, AR, and OK, mainly on the Coastal Plain, but extending inland to other physiographic 

provinces. [= RAB, C, F, G, HC, K, S, W, Z; < S. elliottii – FNA (also see S. apalachicolense)] 

Sorghastrum nutans (Linnaeus) Nash, Yellow Indiangrass. Mt, Pd, Cp (GA, NC, SC, VA): xeric and mesic woodlands 

and forests of a wide variety, powerline rights-of-way, roadbanks; common. September-October. ME and Québec west to s. 

Manitoba, south to FL, TX, UT, AZ, and Mexico. Along with Andropogon gerardii, Schizachyrium scoparium, and Panicum 

virgatum, Sorghastrum nutans is one of the dominant grasses of the tall-grass prairie. It is also common in a variety of open 

habitats (natural and altered) in the forested landscape of eastern North America. [= RAB, C, F, FNA, G, HC, K, S, W, Z; = S. 

avenaceum (Michaux) Nash] 

Sorghastrum secundum (Elliott) Nash, Lopsided Indiangrass. Cp (GA, SC): sandhills; uncommon. September-October. 

S. SC south to FL and west to s. AL (Sorrie & Leonard 1999). [= RAB, FNA, HC, K, S, Z] 

Sorghastrum apalachicolense D.W. Hall, Apalachicola Indiangrass, Open Indiangrass. Flatwoods and sandhills. Panhandle 

FL west to s. MS (Sorrie & Leonard 1999). It may well occur as well in GA. July-August. [= K, Z; < S. elliottii – FNA] 

Sorghum Moench (Sorghum, Milo, Johnson Grass) 

A genus of about 25 species, of tropical and subtropical Old World (1 species in Mexico). References: Barkworth in FNA 

(2003a); de Wet (1978)=Z. 

1 Rhizomatous perennial; leaves 1-2 cm wide...........................................................................................................S. halepense 

1 Fibrous-rooted annual; leaves (2-) 3-5 cm wide. 

2 Inflorescence dense, compact; plants 0.5-1.3 m tall......................................................................... S. bicolor var. bicolor 

2 Inflorescence open, with spreading branches; plants 1.0-3.0 m tall ........................................ S. bicolor var. drummondii 

* Sorghum bicolor (Linnaeus) Moench var. bicolor, Sorghum, Milo, Broomcorn, Sorgo. Cp, Pd (GA, NC, SC, VA): 

cultivated, rarely persistent; common in cultivation, rare as an escape. October. [= C; < Sorghum vulgare Persoon – RAB; < 

Sorgum vulgare – F, orthographic variant; = S. vulgare var. vulgare – HC; = S. bicolor ssp. bicolor – FNA, K; < Holcus 

sorghum Linnaeus – S] 

* Sorghum bicolor (Linnaeus) Moench var. drummondii (Nees ex Steudel) Mohlenbrock, Shattercane. Cp, Pd (GA, NC, SC, 

VA): cultivated, rarely persistent; common in cultivation, rare as an escape. October. This is the taller variety with open 

inflorescences, usually sporadically present in sorghum fields. [= C; < Sorghum vulgare Persoon – RAB; < Sorgum vulgare – F, 

orthographic variant; = Sorghum bicolor ssp. ×drummondii (Nees ex Steudel) de Wet – FNA; = Sorghum vulgare Persoon var. 

drummondii (Nees ex Steudel) Hackel ex Chiov. – HC; = Sorghum bicolor ssp. drummondii (Nees ex Steudel) de Wet & Harlan 

– K; < Holcus sorghum Linnaeus – S] 

* Sorghum halepense (Linnaeus) Persoon, Johnson Grass. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, fields, waste places; 

common, introduced from Eurasia. A serious weed, difficult to eradicate. [= RAB, C, FNA, GW, HC, K, W; = Sorgum 

halepense – F, G, orthographic variant; = Holcus halepensis Linnaeus – S] 

Spartina Schreber (Cordgrass) 

A genus of ca. 15 species, of temperate America, Europe, and Africa. References: Barkworth in FNA (2003a). 

1 Leaf blades mostly 5-40 mm wide, flat toward the base, generally somewhat involute toward the tip, or involute after drying; 

plants 0.3-4.0 m tall. 

2 Glumes with smooth keels; spikelets spaced 3-8 mm apart on each face of the rachis (the rachis triangular in crosssection, 

with spikelets attached on two faces); [of salt to brackish coastal marshes] ....................................S. alterniflora 

2 Glumes with scabrous keels; spikelets spaced 1-3 mm apart on each face of the rachis; [of fresh to brackish coastal 

marshes, or inland]. 

3 Second glume acute, not awned; first glume averaging ca. 1/2 as long as the lemma; spikes (6-) 20-50 (-more) per 

inflorescence; [of fresh to brackish coastal marshes] .........................................................................S. cynosuroides 

3 Second glume with an awn 3-10 mm long; first glume averaging ca. 7/8 as long as the lemma; spikes (5-) 7-27 per 

inflorescence; [of fresh marshes, either inland or coastal] ....................................................................... S. pectinata 

1 Leaf blades mostly 1-4 (-6) mm wide, involute; plants 0.3-1.0 m tall (to 2.0 m tall in the distinctly clumped S. bakeri). 

4 Plants forming large clumps with numerous culms and leaves; spikelets 6-9 mm long; spikes usually 9-14 per 

inflorescence; [of se. SC southward] ...................................................................................................................S. bakeri 

4 Plants with creeping rhizomes, culms arising singly or few together from a point; spikelets 7-13 mm long; spikes 

usually 1-9 per inflorescence; [widely distributed in coastal parts of our area]. 

5 Spikelets 7-10 mm long; second glume acute to obtuse (rarely acuminate); spikes (2-) 4-9 per inflorescence; 

second highest leaf blade on the stem (1-) avg. 2 (-5) dm long; plants to 15 dm tall; culms to 6 mm in diameter at 

base ..................................................................................................................................... S. patens var. monogyna

POACEAE 919 

5 Spikelets 9-13 mm long; second glume acuminate; spikes 1-4 per inflorescence; second highest leaf blade on the 

stem (0.5-) avg. 1 (-2) dm long; plants to 8 dm tall; culms to 3 mm in diameter at base ...........S. patens var. patens 

Spartina alterniflora Loiseleur, Saltmarsh Cordgrass, Smooth Cordgrass. Cp (GA, NC, SC, VA): salt marshes; common. 

August-October. Newfoundland south to FL, west to TX; e. South America; introduced in n. Europe. S. alterniflora is the 

dominant plant (often essentially a monoculture) of intratidal salt marshes in our area. [= RAB, C, FNA, GW, K; > S. 

alterniflora var. alterniflora – F, G, HC, S; > S. alterniflora var. glabra (Muhlenberg ex Bigelow) Fernald – F, G, HC, S; > S. 

alterniflora var. pilosa (Merrill) Fernald – F, G, HC] 

Spartina bakeri Merrill, Sand Cordgrass. Cp (GA, SC): brackish marshes, marsh edges, wet coastal hammocks, under 

Sabal palmetto, Quercus virginiana, and Juniperus virginiana var. silicicola; rare. June. Se. SC south to s. FL, west to 

panhandle FL. Distinctive among our species in its densely clumped growth form. [= FNA, GW, HC, K, S] 

Spartina cynosuroides (Linnaeus) Roth, Giant Cordgrass. Cp (GA, NC, SC, VA): brackish and freshwater tidal marshes, 

especially along margins of tidal creeks; common. June-September. MA south to FL, west to e. TX. [= RAB, C, FNA, G, GW, 

HC, K, S; > S. cynosuroides var. cynosuroides – F] 

Spartina patens (Aiton) Muhlenberg var. monogyna (M.A. Curtis) Fernald, Large Saltmeadow Cordgrass. Cp (GA, NC, 

SC, VA): sandy shores, overwash flats; common. June-September. MA south to FL, west to TX. Whether var. monogyna is 

worthy of recognition is a matter of debate; there appear to morphological differences correlated with geography and, according 

to some authors, habitat, but positive identification to variety is sometimes difficult. [= F, G, HC; < S. patens – RAB, C, FNA, 

GW, K, S] 

Spartina patens (Aiton) Muhlenberg var. patens, Small Saltmeadow Cordgrass, Salt Hay, Marsh-hay Cordgrass. Cp (NC, 

VA): dunes, sand flats, upper edges of marshes, maritime wet grasslands; common. June-September. Newfoundland south to 

NC, and perhaps further. [= F, G, HC; < S. patens – RAB, C, FNA, GW, K, S] 

Spartina pectinata Link, Prairie Cordgrass, Slough Grass. Mt, Cp (NC, VA): spray cliffs below waterfalls, rocky or sandy 

flood-scoured riverside grasslands, tidal freshwater (oligohaline) marshes, calcareous oak flatwoods and prairies; rare (GA Rare, 

NC Rare, VA Rare). July-September. Newfoundland west to WA, south to ne. NC, sw. NC, AR, TX, and NM. [= RAB, C, F, 

FNA, G, GW, HC, K, W; > S. pectinata var. pectinata – F; > S. pectinata var. suttiei (Farwell) Fernald – F; = S. michauxiana 

A.S. Hitchcock – S] 

Spartina spartinae (Trinius) Merr. ex A.S. Hitchcock, Gulf Cordgrass. Brackish marshes and inland saline situations. AL 

and FL west to TX. [= FNA, GW, HC, K, S] {not keyed at this time} 

Sphenopholis Scribner (Wedgegrass) 

A genus of 5 species, North American. References: Tucker (1996)=Z. Key based in part on C. 

1 Spikelets 5-9.5 mm long; second lemma with an awn 3.5-7 mm long............................................................... S. pensylvanica 

1 Spikelets 1.5-5 mm long; second lemma awnless, or with an awn up to 3.5 mm long. 

2 Lower leaf blades mostly (10-) 15-45 cm long, involute, < 2 mm wide .......................................................... S. filiformis 

2 Lower leaf blades mostly less han 10 cm long, flat, 2-8 mm wide. 

3 First glume 1/3 to 2/3 as wide as the second glume; second glume strongly scabrous...................................S. nitida 

3 First glume less tha 1/3 as wide as the second glume; second glume smooth to slightly scabrous. 

4 First lemma with an awn up to 3.5 mm long ..................................................................................... S. ×pallens 

4 First lemma unawned. 

5 Panicle open; second glume 3-6× as long as wide, acute at the tip; lowermost rachilla internode 0.8-1.0 

mm long .................................................................................................................................S. intermedia 

5 Panicle densely cylindrical; second glume 2-3× as long as wide, rounded or truncate at the tip; 

lowermost rachilla internode 0.5-0.7 mm long.......................................................................... S. obtusata 

Sphenopholis filiformis (Chapman) Scribner. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): pine savannas, sandy woodlands; 

uncommon (VA Rare). April-May. Se. VA south to c. FL, west to e. TX. [= RAB, C, F, K, S, Z] 

Sphenopholis intermedia (Rydberg) Rydberg, Slender Wedgegrass. Mt, Pd, Cp (GA, NC, SC, VA): moist nutrient-rich 

forests; rare. May-June. Newfoundland west to c. AK, south to panhandle FL, c. TX, and AZ. [= RAB, F, K, S; = S. obtusata 

(Michaux) Scribner var. major (Torrey) K.S. Erdman – C, Z; < S. intermedia – G (also see S. ×pallens); < S. obtusata – GW, W] 

Sphenopholis nitida (Biehler) Scribner. Cp, Pd, Mt (GA, NC, SC, VA): moist forests, bottomlands; common. April-May. 

MA west to IL, south to FL and TX. [= RAB, C, F, K, W, S, Z; > S. nitida var. glabra (Nash) Scribner – G; > S. nitida var. 

nitida – G] 

Sphenopholis obtusata (Michaux) Scribner, Prairie Wedgegrass. Cp, Pd, Mt (GA, NC, SC, VA): forests, roadsides, 

disturbed areas; common (uncommon in Mountains). April-May. ME west to MN and British Columbia, south to s. FL, TX, c. 

Mexico, and s. CA. [= RAB, G, K, S; = S. obtusata var. obtusata – C, Z; > S. obtusata var. obtusata – F; > S. obtusata var. 

pubescens (Lamson-Scribner & Merrill) Lamson-Scribner – F; < S. obtusata – GW, W (also see S. intermedia)] 

Sphenopholis ×pallens (Biehler) Scribner (pro sp.) [S. obtusata × pensylvanica]. Cp (NC, SC, VA): ditches, wet forests; 

rare. Not always with its parents. May. [= C, K; = S. pallens – RAB, F, S; < S. intermedia (Rydberg) Rydberg – G]

POACEAE 920 

Sphenopholis pensylvanica (Linnaeus) A.S. Hitchcock, Swamp-oats. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): bogs, 

ditches, wet forests; uncommon. April-June. MA west to OH and se. MO, south to n. FL and LA. [= C, K, Z; = Trisetum 

pensylvanicum (Linnaeus) Palisot de Beauvois ex Roemer & J.A. Schultes – RAB, F, G, S; = S. pennsylvanica – GW, 

orthographic variant] 

Sporobolus R. Brown 1810 (Dropseed) 

A genus of about 160 species, perennials and annuals, of tropical, subtropical, and warm-temperate parts of the New World and 

Old World. References: Riggins (1977)=Z; Weakley & Peterson (1998)=Y; Peterson, Hatch, & Weakley in FNA (2003a). 

1 Inflorescence an open panicle, > 2 cm broad, the branches ascending to spreading. 

2 Branches of the panicle verticillate, whorled; spikelets 2.5-4 mm long............................................................. S. junceus 

2 Branches of the panicle alternate (some occasionally rather randomly subopposite or opposite, but never regularly 

whorled); spikelets either 4-6.5 mm long, or 1.5-2.5 (2.7) mm long. 

3 Spikelets 1.5-2.5 (-2.7) mm long. 

4 Panicle branches bare of spikelets in the lower 1/4 to 1/8 of their length.....................................S. cryptandrus 

4 Panicle branches bearing spikelets to the base ............................................................................S. domingensis 

3 Spikelets 4-6.5 mm long. 

5 First glume scaberulous, acuminate or awn-like; spikelets dark gray; base of plant relatively fibrous; grain 

spherical; [of rocky barrens of the Mountains of NC and VA] ..................................................... S. heterolepis 

5 First glume glabrous, acute to acuminate; spikelets purplish (fading tan); base of plant smooth and hard, made 

up of the indurated leaf bases; grain oblong (when present, usually abortive); [of pine savannas and seeps of 

the Coastal Plain of NC, SC, and southward]. 

6 Leaves terete or subterete (wiry), oval in cross-section, sometimes irregularly channelled for portions of 

their lengths (never with any portion above the sheath flat), < 1 mm wide, tending to senesce and turning 

tan in autumn, the margins generally smooth; culms (including the inflorescence) (2-) 4-7 (-10) dm tall; 

culms (from base to first inflorescence branch) 1.5-5 dm tall; first glume averaging about 0.7× as long as 

the second glume (though variable, ranging from 0.5-0.75×).................................................S. teretifolius 

6 Leaves flat (folded when dry), plane or V-shaped in cross-section, with free margins their entire length, 

1.2-2 (-2.7) mm wide, tending to remain green into the winter (at least until December), the margins 

scabrous (except in S. curtissii); culms (including the inflorescence) 3-22 dm tall; culms (from base to 

first inflorescence branch) (4-) 6-10 dm tall; first glume averaging 0.75-1× as long as the second glume 

(though variable, collectively ranging from about 0.6-1.2×). 

7 First glume averaging 0.95-1.1× as long as the second glume (though variable, ranging from 0.8- 

1.3×); pedicels mostly 1-3 mm long (a few sometimes as long as 10 mm long), appressed; culms 

(including the inflorescence) 3-7 dm tall; inflorescence branches stiffly ascending; leaves 0.5-1.5 

mm wide (or to 2.0 mm wide when unburned), mostly 1.5-4 dm long (rarely to 5 dm long), smooth 

on the margins; [of e. SC southward] ................................................................................. S. curtissii 

7 First glume averaging 0.6-0.9× as long as the second glume (though variable, ranging from 0.6- 

0.95×); pedicels mostly 4-15 mm long, spreading; culms (including the inflorescence) (3-) 7-16 (- 

22) dm tall; inflorescence branches initially ascending, later loosely ascending to spreading; leaves 

1.2-10.0 mm wide, mostly (3-) 4-8 dm long, upwardly scabrous on the margins; [of e. NC 

southward]. 

8 Leaves (2.0-) 3-10 mm wide, pale bluish-green (often with some yellowish leaves as well); 

first glume averaging 0.75-0.9× as long as the second glume (though variable, ranging from 

0.6-0.95×); culms (including the inflorescence) usually 12-22 dm tall; inflorescence usually 

3.5-5 dm long; [of se. SC southward]..................................................................... S. floridanus 

8 Leaves 1.2-2.0 (-3.0) mm wide, dark green; first glume averaging 0.6-0.8× as long as the 

second glume (though variable, ranging from 0.6-0.8×); culms (including the inflorescence) 

usually 6-12 (-18) dm tall; inflorescence usually 2-3.5 dm long; [of e. NC south to e. GA] ....... 

................................................................................................................................ S. pinetorum 

1 Inflorescence a contracted, spike-like panicle, < 2 cm broad, the branches appressed. 

9 Plant a geniculate annual; most inflorescences enclosed by sheaths (or most or all exserted); inflorescence 2-5 cm long. 

10 Spikelets (1.3-) 1.6-2.8 mm long; grain falling free of the lemma and palea; lemma glabrous................S. neglectus 

10 Spikelets 2.3-5 mm long; grain falling enclosed in the lemma and palea; lemma strigose (use 10× or more) or 

glabrous. 

11 Lemma and palea shorter than the glumes; palea usually shorter than the lemma; lemma glabrous or strigose 

with hairs < 0.2 mm long; spikelets 2.3 -3.3 (-3.8) mm long; floret (lemma, palea and enclosed grain) 1.6-3.3 

(-3.8)× as long as wide ................................................................................................................... S. ozarkanus 

11 Lemma and palea longer than the glumes; palea usually longer than the lemma; lemma strigose with hairs > 

0.2 mm long; spikelets 2.8-5 mm long; floret (lemma, palea and enclosed grain) 2.2-5.7 (-7.5)× as long as 

wide............................................................................................................................................. S. vaginiflorus 

9 Plant a rhizomatous or tufted perennial; most inflorescences exserted to partly enclosed; inflorescence 5-15 cm long.

POACEAE 921 

12 Plant creeping extensively by slender rhizomes; leaf blades cauline, distichous, to 12 cm long.............S. virginicus 

12 Plant loosely tufted, from short rhizomes; leaf blades basal or cauline, not distichous, 10-100 cm long. 

13 Spikelets 1.5-2.2 mm long; first glume 0.5-0.8 mm long; leaves primarily basal ............................... S. indicus 

13 Spikelets 4-8 mm long; first glume 2-5 mm long; leaves cauline and basal. 

14 Lemma pubescent, usually conspicuously shorter than the palea; pericarp loose when moist .................... 

............................................................................................................................................S. clandestinus 

14 Lemma glabrous, about as long as the palea; pericarp gelatinous when moist. 

15 Culms (1.4-) 2.0-5.0 mm thick; terminal sheath (1.3-) 1.5-6.0 mm wide; panicles with 12-35 

primary branches, crowded, dense.......................................................S. compositus var. compositus 

15 Culms 1.0-2.0 (-2.5) mm thick; terminal sheath 0.8-2.0 (-2.5) mm wide; panicles with 8-18 primary 

branches, lax, loosely flowered ......................................................[S. compositus var. drummondii] 

Sporobolus clandestinus (Biehler) A.S. Hitchcock, Rough Dropseed. Pd, Cp (GA, NC, SC, VA), Mt (VA): glades, 

barrens, and thin soil of woodlands, also in dry sands; uncommon. September-October. This species is widespread in e. United 

States. Wipff & Jones (1995) recommend reducing this taxon to a variety under S. compositus, because of its morphologic 

similarity. While S. clandestinus and S. compositus are undoubtedly closely related, I prefer to retain the two as species. [= 

RAB, C, FNA, F, G, HC, K, S, W, Z; = S. compositus (Poiret) Merrill var. clandestinus (Biehler) J. Wipff & S.D. Jones] 

Sporobolus compositus (Poiret) Merrill var. compositus, Tall Dropseed. Pd (NC?, VA), Mt, Cp (VA): diabase glades and 

barrens, limestone glades and barrens, disturbed areas over diabase or calcareous rocks; rare (NC Watch List, VA Rare). 

September-November. This species and variety are reported for NC in a recent revision of the S. asper group (Riggins 1977); 

little is known about the occurrence of this species in NC. The general range is centered in the Plains, but extending east into ne. 

United States. The name S. compositus has nomenclatural priority over the more familiar S. asper (Kartesz & Gandhi 1995). [= 

FNA, K; = S. asper (Michaux) Kunth var. asper – C, G, HC, Z; = S. asper – F, S] 

* Sporobolus cryptandrus (Torrey) A. Gray, Sand Dropseed. Cp? (NC?): disturbed areas; rare, probably adventive from c. 

and w. North America. This species is reported for NC by HC, F, and S. [= C, FNA, G, K, HC, S, X; > S. cryptandrus var. 

cryptandrus – F] 

Sporobolus curtissii (Vasey ex Beal) Small ex Scribner, Curtiss's Dropseed. Cp (GA, SC): moist, gummy-clay flatwoods; 

uncommon (rare north of GA). September-November. E. SC south to c. FL. First positively documented for our area in 1993. 

Earlier attributions of S. curtissii to NC and SC were apparently based on misapplication or confusion with S. teretifolius and/or 

Sporobolus pinetorum. S. curtissii differs from other "bunchgrass" Sporobolus of our area in having the spikelets shortpedicelled 

and appressed against the panicle branches (as opposed to long-pedicelled and spreading in S. teretifolius and 

Sporobolus pinetorum). [= FNA, HC, K, S, Y] 

*? Sporobolus domingensis (Trinius) Kunth, Coral Dropseed. Cp (GA): coastal sands?; rare, uncertain whether native or 

introduced. Se. GA south to s. FL; West Indies, Mexico. The e. GA record (Glynn County) is at Univ. of Georgia (Sorrie, pers. 

comm.). [= FNA, HC, K, S] 

Sporobolus floridanus Chapman, Florida Dropseed. Cp (GA, SC): wet savannas; uncommon (rare north of GA). June- 

September. Se. SC south to ne. FL, west to Panhandle FL. First positively documented for our area in 1995. Earlier attributions 

of S. floridanus to NC and SC were apparently based on misapplication or confusion with Sporobolus pinetorum. [= FNA, K, Y; 

< S. floridanus – GW, HC, S (also see S. pinetorum); the inclusion of S. floridanus in RAB was based on a misidentification of S. 

pinetorum] 

Sporobolus heterolepis (A. Gray) A. Gray, Prairie Dropseed. Mt (GA, NC, VA): barrens and glades over mafic, 

ultramafic, and calcareous rocks (olivine, serpentine, limestone); rare (NC Endangered, VA Rare). August-September. The 

primary distribution of S. heterolepis is in the Plains, with outliers east to nw. GA (Jones & Coile 1988), c. TN (Estes & Beck 

2005), w. NC, w. VA, se. PA, ne. United States, and adjacent Canada in calcareous, mafic, or ultramafic glades, barrens, and 

prairies. [= RAB, C, F, FNA, G, HC, K, W, Y] 

* Sporobolus indicus (Linnaeus) R. Brown, Smut Grass, Blackseed. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, lawns, 

disturbed situations; common, introduced from the tropics of Asia. July-October. [= C, FNA, GW, W; > S. poiretii (Roemer & 

J.A. Schultes) A.S. Hitchcock – RAB, F, G, HC; > S. indicus – HC; = S. berteroanus (Trinius) A.S. Hitchcock & Chase – S; = S. 

indicus var. indicus – K] 

Sporobolus junceus (Palisot de Beauvois) Kunth, Sandhills Dropseed. Cp (GA, NC, SC), Pd (GA, NC, SC, VA): sandhills, 

other dry, open areas; common, uncommon in Piedmont (VA Rare). September-October. Se. VA south to FL and west to se. 

TX. [= RAB, C, F, FNA, G, HC, K, Y; = S. gracilis (Trinius) Merrill – S] 

Sporobolus neglectus Nash, Barrens Dropseed. Mt (VA): dry rocky barrens and outcrops, over calcareous rocks (such as 

limestone or dolomite); rare (VA Rare). August-September. ME west to ND, south to NJ, w. VA, TN, LA, and TX; apparently 

disjunct in WA and AZ. S. ozarkanus, S. neglectus, and S. vaginiflorus form a still very poorly understood complex. [= C, F, 

FNA, G, HC, K, S, W] 

Sporobolus ozarkanus Fernald, Ozark Dropseed. Pd (NC): diabase glades; rare (NC Rare). September-October. KY west 

to KS, south to e. TN, AR, and TX; disjunct in c. NC. In Granville County, NC, it is associated (on glades of diabase, a mafic 

rock) with other taxa with affinities to midwestern glades and prairies: Oligoneuron rigidum, Oligoneuron album, Baptisia 

australis var. aberrans, Symphyotrichum depauperatum, Silphium terebinthinaceum, Parthenium auriculatum, Ruellia humilis, 

and others. S. ozarkanus, S. neglectus, and S. vaginiflorus form a still very poorly understood complex. [= C, F, G, HC, K; = S. 

vaginiflorus (Torrey ex A. Gray) Wood var. ozarkanus (Fernald) Shinners – FNA, K] 

Sporobolus pinetorum Weakley & P.M. Peterson, Carolina Dropseed, Savanna Dropseed. Cp (GA, NC, SC): wet 

savannas, savanna-pocosin ecotones, sandhill-pocosin ecotones, and extending upslope into mesic flatwoods or loamy or clayey

POACEAE 922 

shelves in the fall-line sandhills; rare (GA Rare, NC Watch List, SC Rare). June-September (and into December in response to 

growing-season fire). The identity of this taxon has been obscure; it is now clear that it is a previously unrecognized species, 

endemic to NC, SC, and adjacent e. GA. RAB included it in their concept of S. teretifolius, though it does not key well (keying 

imperfectly to either S. floridanus or S. heterolepis); in S and HC, it will key to S. floridanus, but the leaves are much narrower. 

Additionally, S. floridanus is a taller and coarser plant, the culms often averaging about 1.5 meters in height and 2-3 mm in 

diameter basally (vs. 1 meter high and 1 mm in diameter for Sporobolus pinetorum). In wet savannas of Columbus County, NC, 

S. species 1 occurs with true S. teretifolius (the two codominant over many hectares!), and the two taxa are manifestly distinct. 

The leaves of S. pinetorum are not terete; after lengthy drought in the field (or dry on an herbarium sheet), the leaves become 

tightly folded to involute and can appear wiry. Like many Southeastern pineland grasses, S. pinetorum flowers only following 

fire. In vegetative condition it may be distinguished from Aristida stricta and A. beyrichiana, with which it often grows, by the 

leaf pubescence (S. pinetorum with scaberulous margins, best felt by running a finger along the margin near the base, from apex 

toward base, A. stricta and A. beyrichiana not scaberulous, and with a sparse line of pilose hairs running more or less the length 

of the leaf in A. stricta and sometimes in A. beyrichiana) and base (much more indurated and polished in Sporobolus than in 

Aristida). [= FNA, K, Y; >< S. teretifolius – RAB, misapplied; > S. floridanus – RAB, misapplied; < S. floridanus Chapman – 

HC, S] 

Sporobolus teretifolius Harper, Wireleaf Dropseed. Cp (GA, NC, SC): wet savannas, pitcherplant bogs; rare (US Species 

of Concern, GA Rare, NC Threatened). July-September (and later in response to growing-season fire). Very similar vegetatively 

to Aristida stricta, S. teretifolius can be distinguished by its tuft of hairs at the base of the otherwise glabrous blade (as opposed to 

line of pilose hairs the length of the blade in A. stricta). This very rare species is known only from se. NC, ne. SC, s. GA, and se. 

AL (Houston County). Many of the counties reported for this species in RAB actually are based on misidentified specimens of 

Sporobolus pinetorum. In a few very wet savannas of Columbus and Brunswick counties, NC, S. teretifolius is dominant or 

codominant over many hectares. Like many savanna grasses, S. teretifolius generally flowers only following fire. [= FNA, HC, 

K, S, Y; < S. teretifolius – RAB (also see S. pinetorum)] 

Sporobolus vaginiflorus (Torrey ex A. Gray) Wood, Poverty Dropseed. Pd, Mt (GA, NC, SC, VA), Cp (VA): glades, 

barrens, open disturbed sites; uncommon. September-October. The species occurs nearly throughout e. United States. S. 

ozarkanus, S. neglectus, and S. vaginiflorus form a still very poorly understood complex. [= RAB, C, G, HC, W; = S. 

vaginiflorus var. vaginiflorus – F, FNA, K; = S. vaginaeflorus – S, orthographic variant] 

Sporobolus virginicus (Linnaeus) Kunth, Seashore Dropseed, Coastal Dropseed. Cp (GA, NC, SC): salt marshes, tidal 

mud flats, and low dunes in the outer Coastal Plain; rare (NC Watch List). September-October. This species occurs from se. NC 

along the coast to TX, in the West Indies and into n. South America (its alleged occurrence in se. VA is apparently incorrect). 

Sporobolus virginicus is similar in aspect and growth form to Distichlis spicata, with which it occurs in tidal flats. Sporobolus 

virginicus is more delicate, and typically has long hairs on either side of the collar of the sheath; Distichlis spicata is generally a 

coarser plant, and lacks long hairs around the collar of the sheath. [= RAB, C, F, FNA, G, GW, HC, K, S] 

* Sporobolus airoides (Torrey) Torrey, Alkali Sacaton. Cp (SC): waste areas near wool-combing mills; rare, intoduced from 

, not known to be established or persistent. [= FNA, HC, K] {not keyed} 

Sporobolus compositus (Poiret) Merrill var. drummondii (Trinius) Kartesz & Gandhi, ranges east to the Ridge and Valley 

province of e. TN (Chester et al. 1993), occurring over limestone, and allegedly to KY and GA (Kartesz 1999). It could very 

likely occur in sw. VA, as it is in Hawkins County, TN, immediately adjacent to VA (Chester et al. 1993). [= FNA, K; = S. 

asper (Michaux) Kunth var. drummondii (Trinius) Vasey – C, Z; = S. drummondii (Trinius) Vasey – F, S; = S. asper var. hookeri 

(Trinius) Vasey – G, HC, misapplied] 

* Sporobolus fimbriatus (Trinius) Nees. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a waif, intoduced 

from Africa. [= FNA, HC, K] {not keyed} 

* Sporobolus flexuosus (Thurb. ex Vasey) Rydberg. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a 

waif, intoduced from sw. United States and n. Mexico. [= FNA, HC, K] {not keyed} 

* Sporobolus tenuissimus (Martius ex Schrank) Kuntze. Cp (SC): waste areas near wool-combing mills; rare, perhaps only a 

waif, introduced from the tropical Old World and New World. [= FNA, K] {not keyed} 

* Sporobolus wrightii Munro ex Scribner, Giant Sacaton. Cp (SC): waste areas near wool-combing mills; rare, perhaps only 

a waif, intoduced from sw. United States. [= FNA, HC, K] {not keyed} 

Steinchisma Rafinesque 1830 (Gaping Panic Grass) 

A genus of about 6 species, perennial herbs, of s. North America, Central America, and South America. See discussion following 

Panicum regarding generic concepts. References: Zuloaga et al. (1998)=Z; Freckmann & Lelong in FNA (2003a). 

Steinchisma hians (Elliott) Nash, Gaping Panic Grass. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (GA): stream, pond, 

and lake shores, low woods, cypress-gum ponds, floodplains, marshes, ditches, seepage slopes; common in Piedmont, occasional 

in Coastal Plain. May-October. Se. VA south to FL, west to TX and OK, and south through Mexico and Central America to 

Colombia; also in s. South America. The large, thickened, pale sterile palea of this species is unique among panicoids of our 

region; it is one of several characters that has led to the segregation of Steinchisma as a genus, or as a subgenus of Panicum. The 

enlargement of the sterile palea causes the spikelet to spread open, or "gape." [= FNA, K, Z; = Panicum hians Elliott – RAB, C, 

F, G, GW, HC, S, W]

POACEAE 923 

Stenotaphrum Trinius (St. Augustine Grass) 

A genus of about 7 species, tropical and subtropical. References: Allred in FNA (2003a); Sauer (1972)=Z. 

Stenotaphrum secundatum (Walter) Kuntze, St. Augustine Grass, Carpet Grass. Cp (GA, NC, SC): brackish marshes, 

roadsides, lawns; common. July-October. A pioneer species of beaches and shores, S. secundatum was known from the 

Carolinas prior to 1800. It has been interpreted as native or introduced in our area; its original range is probably now impossible 

to determine. Sauer (1972) maps it as widespread along the coasts of s. North America, Central America, South America, the 

West Indies, Africa, Australia, and sw. Pacific Islands. In our area it is certainly now more frequently encountered as a lawn or 

roadside grass than in anything that could be construed as a natural habitat. The other 6 species in the genus are Asian, or on 

islands of the sw. Pacific or Indian Oceans. [= RAB, FNA, HC, K, S, Z] 

Stipa 

(see Nassella and Piptochaetium) 

References: Tucker (1996)=Z; Barkworth (1997)=Y. 

Thinopyrum (Prat) Á. Löve 

* Thinopyrum intermedium (Host) Barkworth & D.R. Dewey. Pd (GA): waif in railroad yards; rare, introduced from {}. 

Tucker (1996) states that the record is as a waif in railroad yards. [= K, Z; = Elytrigia intermedia (Host) Nevski; = Agropyron 

intermedium (Host) Palisot de Beauvois] {not keyed at this time; add to synonymy} 

* Thinopyrum ponticum (Podp.) Z.W. Liu & R. R.-C. Wang, Tall Wheatgrass. Cp (SC): waste areas near wool-combing 

mills; rare, intoduced from {}, not known to be established or persistent. [= K; ? Agropyron elongatum (Host) Palisot de 

Beauvois] {not keyed at this time; add to synonymy} 

Torreyochloa Church (Pale Mannagrass) 

A genus of 4 species, with a classic Tertiary moist temperate disjunct pattern; Torreyochloa is distributed in e. North America 

(our taxa), one species in the Pacific Northwest, and two in e. Asia (Tucker 1996). References: Davis (1991)=Y; Tucker 

(1996)=Z. 

1 Leaf blades 1-3 mm wide; anthers 0.2-0.5 mm long...........................................................................[T. pallida var. fernaldii] 

1 Leaf blades 4-8 mm wide; anthers ca. 1 mm long.................................................................................... T. pallida var. pallida 

Torreyochloa pallida (Torrey) Church var. pallida, Pale Mannagrass. Mt (GA, NC, VA), Cp (NC, VA), Pd (SC): bogs, 

mucky wetlands such as old beaver-ponds, pools in cypress swamps, drawdown shores of natural ponds; rare (GA Special 

Concern, NC Rare). June-July. The species as a whole is widespread in e. North America. Var. pallida ranges from Nova 

Scotia west to MN, south to e. VA, se. NC (Columbus County), nw. NC (Avery County), and nw. GA (Jones & Coile 1988). 

Var. pauciflora (J. Presl) J.J. Davis is distributed in w. North America. Intermediates occur between the varieties. [= K, Y, Z; < 

Glyceria pallida (Torrey) Trinius – RAB, GW, HC, W; < Puccinellia pallida (Torrey) Clausen – C; = G. pallida – F; = G. 

pallida var. pallida – G; = Panicularia pallida (Torrey) Kuntze – S] 

Torreyochloa pallida (Torrey) Church var. fernaldii (A.S. Hitchcock) Dore ex Koyama & Koyama ranges from 

Newfoundland west to MN, south to WV and TN. [= K, Y, Z; < Glyceria pallida (Torrey) Trinius – RAB, GW, HC, W; < 

Puccinellia pallida (Torrey) Clausen – C; = G. fernaldii (A.S. Hitchcock) St. John – F; = G. pallida var. fernaldii A.S. Hitchcock 

– G] 

Tragus Haller 

* Tragus racemosus (Linnaeus) Allioni, Texas Bur. Cp (NC): on ballast near old seaports; rare, introduced from the Old 

World, probably no longer present. [= HC, C, F, G, K] 

* Tragus australianus S.T. Blake. Cp (SC): waste areas around wool-combing mills; rare, perhaps only a waif, introduced. 


* Tragus berteronianus J.A. Schultes. Cp (SC): waste areas around wool-combing mills; rare, perhaps only a waif, 

introduced. [= K] {not keyed} 

* Tragus heptaneuron W.D. Clayton. Cp (SC): waste areas around wool-combing mills; rare, perhaps only a waif, 

introduced. [= K] {not keyed}

POACEAE 924 

Tridens Roemer & J.A. Schultes (Triodia, Redtop, Tridens, Fluffgrass) 

A genus of about 14 species, native to the Western Hemisphere. References: Valdés-Reyna in FNA (2003a). 

1 Panicle dense and spike-like, > 4× as long as wide, the branches ascending to appressed. 

2 Plants from elongate rhizomes; lemma 4-5 mm long; spikelet 7-9 mm long ............................................. T. carolinianus 

2 Plants cespitose; lemma 2.5-3 mm long; spikelet 4-6 mm long .......................................................................... T. strictus 

1 Panicle open and spreading, < 4× as long as wide, the branches well-developed and spreading-ascending to reflexed. 

3 Spikelets 4-5 mm long, 2.5-3.5 mm wide .......................................................................................................T. ambiguus 

3 Spikelets 6-8 mm long, 1.5-2.2 mm wide. 

4 Primary pulvini densely pubescent, the hairs encircling the base of the panicle branch; secondary pulvini 

pubescent; spikelets mostly on pedicels 3-20 mm long; main branches of the inflorescence stiffly spreading........... 

...............................................................................................................................................................T. chapmanii 

4 Primary pulvini glabrous to sparsely pubescent, tufted only in the axil (the upper surface of the panicle branch); 

secondary pulvini glabrous; spikelets on pedicels mostly < 3 mm long; main branches of the inflorescence 

spreading, ascending or drooping.................................................................................................................. T. flavus 

Tridens ambiguus (Elliott) J.A. Schultes, Pineland Triodia, Flatwoods Fluffgrass. Cp (GA, NC, SC): wet savannas, claybased 

Carolina bays; uncommon (rare north of GA) (NC Rare). August-October. S. NC south to FL, west to e. TX. [= RAB, 

FNA, GW, HC, K; = Triodia elliottii Bush – S] 

Tridens carolinianus (Steudel) Henrard, Carolina Triodia, Carolina Fluffgrass. Cp (GA, NC, SC): mesic swales in 

sandhills; rare (GA Rare, NC Rare). August-October. S. NC south to FL, west to LA. [= RAB, FNA, HC, K; = Triodia 

drummondii Scribner & Kearney – S] 

Tridens chapmanii (Small) Chase, Chapman's Triodia. Cp (GA, NC, SC, VA): loamy sands of disturbed longleaf pine 

woodlands, roadsides; rare (NC Watch List). August-October. NJ south to FL, west to TX and OK. [= HC; = Tridens flavus 

(Linnaeus) A.S. Hitchcock var. chapmanii (Small) Shinners – RAB, C, FNA, K; = Triodia chapmanii (Small) Bush – F, G; < 

Triodia flava (Linnaeus) Smyth – S] 

Tridens flavus (Linnaeus) A.S. Hitchcock, Redtop, Tall Redtop, Purpletop Tridens, Greasy Grass. Cp, Pd, Mt (GA, NC, 

SC, VA): roadsides, disturbed areas, glades; common. July-October. NH west to NE, south to FL and TX. [= HC; = Tridens 

flavus var. flavus – RAB, C, FNA, K; = Triodia flava (Linnaeus) Smyth – F, G; < Triodia flava (Linnaeus) Smyth – S (also see 

Tridens chapmanii); < Tridens flavus – W] 

Tridens strictus (Nuttall) Nash, Spike Triodia, Longspike Fluffgrass, Longspike Tridens. Cp (GA, NC, SC, VA), Pd (GA, 

SC, VA): sandhills, moist pine savannas, roadsides; rare (NC Rare, VA Rare). August-October. S. VA south to AL, west to TX, 

north in the interior to IL and KS. It is possible that this grass is introduced only north and east of GA (thus in our entire area). 

Rhoads & Klein (1993) report an old specimen from w. PA. [= RAB, FNA, GW, HC, K; = Triodia stricta (Nuttall) Bentham ex 

Vasey – F, G, S] 

Triplasis Palisot de Beauvois (Sandgrass) 

A genus of 2 species, of eastern and central North America south through Mexico to Costa Rica. References: Hatch in FNA 

(2003a). 

Identification notes: The foliage of both of our species has a sour taste. 

1 Lemma awn 4.5-8 mm long; culm internodes appressed pilose or puberulent; perennial......................................T. americana 

1 Lemma awn 0.5-1.5 mm long; culm internodes glabrous to minutely scaberulous; annual (or rarely perennial) ....................... 

......................................................................................................................................................... T. purpurea var. purpurea 

Triplasis americana Palisot de Beauvois, Southern Sandgrass. Cp (GA, NC, SC): open sandy areas; common. August- 

October. A Southeastern Coastal Plain endemic: NC south to FL, west to e. LA. [= RAB, FNA, HC, K, S] 

Triplasis purpurea (Walter) Chapman var. purpurea, Purple Sandgrass. Cp (GA, NC, SC, VA): dunes, maritime dry 

grasslands, open sandy areas; common. September-October. NH south to FL, and west to TX, along the coast; also around the 

Great Lakes, and in central United States. Var. caribensis R.W. Pohl is in the New World tropics. [= FNA; < T. purpurea – 

RAB, C, F, G, HC, K; > T. intermedia Nash – S; > T. purpurea – S] 

Tripsacum Linnaeus (Gamma Grass) 

A genus of about 12 species, tropical and subtropical American. References: Barkworth in FNA (2003a); DeWet, Harlan, & 

Brink (1982)=Z.

POACEAE 925 

Tripsacum dactyloides (Linnaeus) Linnaeus var. dactyloides, Gamma Grass. Pd, Mt, Cp (GA, NC, SC, VA): roadsides, 

moist areas, disturbed areas, moist riverbanks; common. Late May-November. T. dactyloides is widespread in e. North America 

north to MA, MI, IA, and NE, ranging south into tropical Central and South America; var. dactyloides is North American. This 

important species of moist and wetland areas in the Great Plains is generally seen in disturbed habitats in our area; its original 

habitats in our area (if indeed it was native in the flora area) are poorly understood. [= FNA, Z; < T. dactyloides – RAB, C, G, 

HC, K, S, W; > T. dactyloides var. dactyloides – F; > T. dactyloides var. occidentale Cutler & Anderson – F] 

Trisetum Persoon (Oat-grass) 

(also see Sphenopholis) 

A genus of about 85 species, north and south temperate. References: Randall & Hilu (1986)=Z; Tucker (1996)=Y. 

Trisetum spicatum (Linnaeus) K. Richter, Alpine Oat-grass, Spike Trisetum. Mt (NC, VA): mountain cliffs at high 

elevations; rare (NC Endangered, VA Rare). June-August. A circumboreal species, widespread and common in arctic and alpine 

areas, south in e. North America to New England, NY, and, rarely, PA, and disjunct to Hawksbill Mountain, Page County, VA 

(where extant) and Roan Mountain, Mitchell County, NC (where not seen since the nineteenth century). The species is also 

known from the West Indies, Mexico, and s. South America. T. spicatum, as broadly treated here, following Randall & Hilu 

(1986), is polymorphic and consists of several ploidies. [= C, HC, K, S, Y, Z; > T. spicatum var. molle (Michaux) Beal – RAB, 

F, G; > T. triflorum (Bigelow) Löve & Löve ssp. molle (Michaux) Löve & Löve – W; > T. spicatum var. maidenii (Gandoger) 

Fernald – F] 

References: Tucker (1996)=Z; Zohary & Hopf (1994). 

Triticum Linnaeus 1753 (Wheat) 

* Triticum aestivum Linnaeus, Bread Wheat. Cp, Pd, Mt (GA, NC, SC, VA): fields; frequently cultivated, rarely persistent 

or volunteering following cultivation, introduced from Eurasia. May-June. One of the most important crops in the world. The 

lemmas can either be awnless or with long awns (to 8 cm long). [= RAB, C, F, G, HC, K, Z] 

Uniola Linnaeus (Sea Oats) 

(also see Chasmanthium) 

A genus of 2 species. The only other species of the genus ranges from Baja California south along the Pacific Ocean to Ecuador; 

other species previously treated in Uniola have been shown to be only distantly related and are now treated as Chasmanthium. 

References: Yates in FNA (2003a); Yates (1966a, 1966b)=Z. 

Uniola paniculata Linnaeus, Sea Oats. Cp (GA, NC, SC, VA): abundant on unforested primary and secondary dunes on 

barrier islands, and on dry to mesic sand flats and interdune swales; common (VA Watch List). June-November. Se. VA south 

to FL and west to TX and Mexico; West Indies. This is the most important sand-binding grass on ocean dunes from NC south, 

playing a critical role in primary succession on dunes. It is against the law in NC to pick or destroy Uniola paniculata. [= RAB, 

C, F, FNA, G, HC, K, S, Z] 

Urochloa Palisot de Beauvois (Para-grass, Signal-grass) 

A genus of about 100 species, pantropical and subtropical. References: Crins (1991)=Z; Webster (1988)=Y; Wipff & Thompson 

in FNA (2003a). Key adapted in part from GW. 

1 Spikelets suffused with purple, borne in pairs (or threes) in each row........................................................................ U. mutica 

1 Spikelets green, borne singly in each row. 

2 Upper half of second glume and first lemma with evident transverse veins connecting the longitudinal veins; spikelets 

3.5-4.7 mm long............................................................................................................................................U. platyphylla 

2 Upper half of second glume and first lemma without evident transverse veins, or with very obscure cross-veins; 

spikelets either 2-4 mm or 5-6 mm long. 

3 Spikelets 2-4 mm long ............................................................................................................................... U. ramosa 

3 Spikelets 5-6 mm long .................................................................................................................................U. texana 

* Urochloa mutica (Forskål) Nguyen, Para-grass. Cp (SC): margin of pond; rare, introduced from Africa. August. [= FNA, 

K, Z; ? Panicum purpurascens Raddi – RAB, HC; ? B. purpurascens (Raddi) Henrard – GW; = Brachiaria mutica (Forskål) 

Stapf]

POACEAE 926 

* Urochloa platyphylla (Munro ex Wright) R. Webster, Broadleaf Signal-grass. Cp (GA, NC, SC, VA), Pd (GA, SC, NC): 

disturbed wet or seasonally moist areas; rare, presumably introduced from South America. E. NC south to FL, west to TX, north 

in the interior to AR, OK, and se. MO; also in MD (Terrell & Reveal 1996). [= FNA, K, Y, Z; = Brachiaria platyphylla (Munro 

ex Wright) Nash – RAB, GW, HC; ? B. extensa Chase – S] 

* Urochloa ramosa (Linnaeus) Nguyen, Browntop Millet, Dixie Signalgrass. Pd, Cp (GA, NC, SC, VA): disturbed areas; 

rare, introduced from tropical Africa and Asia. This species has apparently been widely planted for wildlife food and erosion 

control in southeastern states. [= FNA, K, Z; = Panicum ramosum Linnaeus – HC; = Brachiaria ramosa (Linnaeus) Stapf] 

* Urochloa texana (Buckley) R. Webster, Texas Millet, Texas Signalgrass. Cp (GA, NC, SC, VA), Pd (GA, SC): disturbed 

areas, fields, gardens; uncommon, introduced from TX. First reported for South Carolina by Hill & Horn (1997). [= K, Y, Z; = 

Panicum texanum Buckley – RAB, C, HC, S; = Brachiaria texana (Buckley) S.T. Blake] 

Urochloa fusca (Swartz) B.F. Hansen & Wunderlin var. reticulata (Torrey) B.F. Hansen & Wunderlin, east to GA (Kartesz 

1999). [< Urochloa fusca – FNA; ? Urochloa fasciculata (Sw.) R. Webster – K; ? Panicum fasciculatum Swartz – HC] {not 

keyed at this time; synonymy incomplete} 

* Urochloa maxima (Jacquin) R. Webster var. maxima, Guinea Grass. Introduced in the Gulf states, east to GA (FNA). 

Native of Africa. [= FNA; < U. maxima – K; ? Panicum maximum Jacquin] {not keyed at this time; synonymy incomplete} 

* Urochloa plantaginea (Link) R. Webster. Cp (GA): Reported for s. GA by Jones & Coile (1988), as Brachiaria 

plantaginea. [= FNA, K, Y, Z; = Brachiaria plantaginea (Link) A.S. Hitchcock] {not keyed at this time; synonymy 

incomplete} 

Urochloa reptans (Linnaeus) Stapf. Cp (GA): [= FNA] {not keyed at this time; synonymy incomplete} 

Vulpia C. Gmelin (Annual Fescue) 

A genus of about 23 species, north and south temperate. References: Tucker (1996)=Z. Key based in part on C. 

1 First glume < half as long as the second glume ..........................................................................................................V. myuros 

1 First glume > ½ as long as the second. 

2 Lemma pubescent; lowest lemma 2.5-3.5 mm long; grains 1.5-2 mm long.......................................................V. elliottea 

2 Lemma glabrous or scabrous; lowest lemma 2.7-7 mm long; grains 1.7-3.3 mm long. 

3 First glume 1.7-4.5 mm long; lemma awns 3-12 mm long; spikelets with 4-7 loosely imbricate florets; rachilla 

internodes mostly 0.9-1.1 mm long........................................................................................................V. bromoides 

3 First glume 3.5-5 mm long; lemma awns 0.3-6 (-9) mm long; spikelets with 5-11 (-more) closely imbricate florets; 

rachilla internodes mostly 0.5-0.7 mm long. 

4 Spikelets 4-5.5 mm long; awn of the lowest lemma 2.5-6 (-9) mm long.......................V. octoflora var. glauca 

4 Spikelets 5.5-10 mm long; awn of the lowest lemma 0.3-3 mm long ....................... V. octoflora var. octoflora 

* Vulpia bromoides (Linnaeus) S.F. Gray, European Squirreltail Fescue, Brome Fescue. Cp (VA): sandy disturbed areas; 

rare, introduced from Eurasia. [= C, K, Z; Festuca dertonensis (Allioni) Ascherson & Graebner – G, HC] 

Vulpia elliotea (Rafinesque) Fernald, Squirreltail Fescue. Cp (GA, NC, SC, VA), Pd (GA): sandy roadsides, fields, 

disturbed areas; common. April-May. S. NJ south to FL, west to TX, and north in the interior to MO. [= C, F, K; = Vulpia 

sciurea (Nuttall) Henrard – Z; = Festuca sciurea Nuttall – RAB, G, HC, S] 

* Vulpia myuros (Linnaeus) K.C. Gmelin, Rat-tail Fescue. Cp (NC, SC, VA), Pd (GA, NC, SC, VA), Mt (GA, NC, SC): 

roadsides, fields, disturbed areas; common, introduced from Eurasia. May-June. [= C, F, K, Z; = Festuca myuros Linnaeus – 

RAB, G, HC, S, W] 

Vulpia octoflora (Walter) Rydberg var. glauca (Nuttall) Fernald, Northern Six-weeks Fescue. {Cp, Pd, Mt (GA, NC, SC, 

VA): fields, roadsides, disturbed areas; common.} April-June. S. ME west to British Columbia, south to GA, AR, TX, and CA. 

[= C, K; < Festuca octoflora Walter – RAB, GW, S, W; = Festuca octoflora var. tenella (Willdenow) Fernald – F, G, HC; < 

Vulpia octoflora – Z] 

Vulpia octoflora (Walter) Rydberg var. octoflora, Southern Six-weeks Fescue. {Cp, Pd, Mt (GA, NC, SC, VA): fields, 

roadsides, disturbed areas; common.} April-June. S. NJ south to FL, west to TX, north in the interior to MO and OK. [= C, F, 

K; < Festuca octoflora Walter – RAB, GW, S, W; > Festuca octoflora var. aristulata Torrey ex L.H. Dewey – G; = Festuca 

octoflora var. octoflora – HC; < Vulpia octoflora – Z] 

Zea Linnaeus (Corn, Maize) 

A genus of about 5 species, native of Mexico and Central America. References: Iltis in FNA (2003a). 

1 Pistillate spikelets (kernels) borne on a spongy rachis (cob) in rows.............................................................Z. mays spp. mays 

1 Pistillate spikelets embedded in a hardened rachis. 

2 Annual.......................................................................................................................................... [Z. mays ssp. mexicana] 

2 Perennial from creeping rhizomes.....................................................................................................................Z. perennis

POACEAE 927 

* Zea mays Linnaeus ssp. mays, Corn, Maize. Cp, Pd, Mt (GA, NC, SC, VA): very commonly cultivated, rarely volunteering 

in old fields or around trashpiles; common in cultivation, rare as a short-lived escape. June-October. Zea is one of the most 

important cultivated plants in the world, originating in Mexico, probably from Zea mays ssp. parviglumis Iltis & Doebley. It was 

initially cultivated in sw. Mexico (before 8000 BP), spreading to the sw. United States before 5000 BP, and to the e. United 

States by 2000 years BP. At the time of European contact, Zea mays ssp. mays was an important staple crop from s. Canada 

south to s, South America (Hancock 2004). [= FNA, K; < Z. mays – RAB, F, HC, S] 

* Zea perennis (A.S. Hitchcock) Reeves & Manglesdorf, Mexican Teosinte. Cp (SC): disturbed areas; rare, apparently 

established at least formerly. Z. perennis was considered by HC to be "established on James Island, S.C." [= K; = Euchlaena 

perennis A.S. Hitchcock – HC] 

* Zea mays (Schrader) Kuntze ssp. mexicana (Schrader) H.H. Iltis, Chalco Teosinte, Nobogame Teosinte. Reported for AL 

and FL (Kartesz 1999). HC state that this taxon is "occasionally cultivated in the Southern States for green forage" and is similar 

to Z. perennis, except in being, like Z. mays ssp. mays, a coarse annual. It is considered to be an ancestor of Zea mays. [= FNA; 

= Z. mexicana (Schrader) Kuntze – K; = Euchlaena mexicana Schrader – HC, S] 

Zizania Linnaeus (Wild-rice) 

A genus of 4 species (and 6 taxa) of northern and eastern North America. References: Terrell et al. (1997)=Y; Tucker (1988)=Z; 

Judziewicz et al. (2000)=X. 

Zizania aquatica Linnaeus var. aquatica, Northern Wild-rice. Cp (GA, NC, SC, VA): freshwater marshes, usually tidal; 

common (uncommon in NC). May-October. Var. aquatica ranges from ME west to WI, south to FL and LA; var. brevis Fassett 

is restricted to the St. Lawrence River in Québec. Zizania was formerly an important food for Amerindians; it is now gathered as 

a specialty grain, commanding high prices. [= C, F, G, HC, K, X, Y, Z; < Z. aquatica – RAB, GW, S] 

Zizaniopsis Döll & Ascherson (Southern Wild-rice) 

A genus of about 5 species, of tropical and subtropical America. References: Tucker (1988)=Z; Judziewicz et al. (2000)=Y. 

Identification notes: Superficially similar to Zizania in its habitat and large size, Zizaniopsis may be distinguished by its very 

different inflorescence and by its stout horizontal rhizomes (Zizania is annual or perennial, but not rhizomatous). 

Zizaniopsis miliacea (Michaux) Döll & Ascherson, Southern Wild-rice, Water-millet, Giant Cutgrass. Cp (GA, NC, SC, 

VA): brackish and freshwater marshes; common. May-July. MD south to FL, west to TX, north in the interior to MO, and 

disjunct in w. Mexico. The other species of the genus are South American. [= RAB, C, F, G, GW, HC, K, S, Y, Z; = Zizania 

miliacea Michaux] 

Zoysia Willdenow (Zoysia) 

* Zoysia japonica Steudel, Zoysia, is used as a lawn grass. Reported for VA (Kartesz 1999). It is not known to naturalize in 

our area. [= C, HC, K] 

* Zoysia matrella (Linnaeus) Merr. var. matrella, Zoysia, is used as a lawn grass. Reported for GA (Kartesz 1999). It is not 

known to naturalize in our area. [= K; = Z. matrella – HC] 

PONTEDERIACEAE Kunth 1816 (Pickerelweed Family) 

A family of about 9 genera and 33 species, primarily of the tropics, but with some temperate representatives. References: 

Rosatti (1987a); Cook in Kubitzki (1998b); Horn in FNA (2002a). 

1 Plant floating (or stranded by dropping water levels), the petioles expanded into air-filled floats; perianth lobes 3-4 cm long. 

..................................................................................................................................................................................Eichhornia 

1 Plant rooted, the petioles not adapted as floats; perianth lobes 0.4-1.0 cm long. 

2 Leaves lanceolate to ovate, 1.5-10× as long as wide, the base cordate, truncate, or cuneate; flowers 2-lipped; corolla 

blue, marked with yellow; stamens 6 (3 each of 2 different lengths) .................................................................Pontederia 

2 Leaves either reniform, 0.5-1.5× as long as wide, the base cordate, or narrowly linear, 20-50× as long as wide, the base 

attenuate; flowers radially symmetrical; corolla white, pale blue, or yellow; stamens 3............................... Heteranthera 

Eichhornia Kunth 1842 (Water Hyacinth)

PONTEDERIACEAE 928 

A genus of 7-8 species, native of tropical America and Africa, but now introduced widely in warm regions. References: Cook in 

Kubitzki (1998b); Horn in FNA (2002a). 

* Eichhornia crassipes (Martius) Solms-Laub, Water Hyacinth. Cp (GA, NC, SC, VA), Pd (GA, NC): ponds, ditches, 

sluggish water; uncommon, introduced from South America. June-September. E. crassipes is "generally considered the world's 

most serious aquatic weed" (Rosatti 1987). Originally native to tropical South America, E. crassipes is now a widespread 

naturalized weed throughout the tropics and subtropics. In the northern part our area, water hyacinth is rare, probably not long 

persisting. Further south, it can be an aggressive aquatic weed. [= RAB, C, F, FNA, G, GW, K; = Piaropus crassipes (Martius) 

Rafinesque – S] 

Heteranthera Ruiz & Pavón 1794 (Mud-plantain) 

(also see Zosterella) 

A genus of 10-12 species, of tropical and temperate America and tropical Africa. References: Cook in Kubitzki (1998b); Horn 

(1998)=Z; Horn in FNA (2002a). Key based in part on FNA. 

1 Leaves narrowly linear, 20-50× as long as wide, the base attenuate; flowers solitary, the corolla yellow; stamens and anthers 

all alike ......................................................................................................................................................................... H. dubia 

1 Leaves reniform, 0.5-1.5× as long as wide, the base cordate; flowers 1-several, the corolla white or pale blue; stamens and 

anthers dimorphic. 

2 Spathe with 2-several flowers; perianth tube 3-12 mm long. 

3 Anthers and filaments with dark purple hairs; internode below the spathe < 1 cm long; spike with (3-) 7-16 

flowers, typically elongating well out of the spathe...............................................................................H. multiflora 

3 Anthers and filaments with white hairs; internode below the spathe > 1 cm long; spike with 2-8 flowers, typically 

mostly included within the spathe......................................................................................................... H. reniformis 

2 Spathe with solitary flower; perianth tube 11-45 mm long. 

4 Vegetative stems elongating only in water deeper than 5 cm; blades of petiolate leaves oblong to ovate, the base 

truncate to cuneate; perianth tube 15-45 mm long ....................................................................................[H. limosa] 

4 Vegetative stems commonly elongating; blades of petiolate leaves round to oblong, the base cordate to truncate; 

perianth tube 11-29 mm long ........................................................................................................... [H. rotundifolia] 

Heteranthera dubia (Jacquin) MacMillan, Water Stargrass. Mt (NC, VA), Pd (VA), Cp (VA): streams, rivers; uncommon 

(rare in NC). August-September. Québec west to WA, south to Cuba and Central America, but rare or absent in much of the se. 

United States. The attribution of this species to SC is in error (as by Kartesz 1999), based on a misidentified specimen (C. Horn, 

pers. comm.). [= RAB, F, FNA, GW, K, W; = Zosterella dubia (Jacquin) Small – C, G, S] 

Heteranthera multiflora (Grisebach) Horn. Cp (NC, VA): in shallow, stagnant water in floodplains, or emersed on mud; 

rare (NC Watch List). June-October. IL west to NE, south to MS; also on the Atlantic Coastal Plain from NJ south through PA 

to ne. NC; also in South America (Brazil, Paraguay, Argentina, and Venezuela). [= C, FNA, K, Z] 

Heteranthera reniformis Ruíz & Pavón. Cp (NC, VA), Pd (GA, NC, SC, VA): in shallow, stagnant water in floodplains, 

or emersed on mud; uncommon (SC Rare). June-October. CT west to NE, south to FL and TX and into South America. First 

reported for South Carolina by Hill & Horn (1997). [= RAB, C, F, FNA, G, GW, K, S, W, Z] 

Heteranthera limosa (Swartz) Willdenow occurs east to TN, KY, and FL (Kartesz 1999); it is attributed to VA in Small 

(1933), but the documentation is not known. [= FNA, C, F, G, K, S, Z] 

Heteranthera rotundifolia (Kunth) Grisebach. Ponds. Midwestern, as a rare disjunct east to c. KY (Larue County) (Medley 

1993). [= FNA, C, K, Z] 

Pontederia Linnaeus 1753 (Pickerelweed) 

A genus of 3-6 species, from North America to South America. References: Lowden (1973)=Z; Cook in Kubitzki (1998b); Horn 

in FNA (2002a). 

1 Floral tube villous when young, essentially glabrous to sparsely glandular in maturity; leaves primarily ovate to triangularlanceolate, 

2.2-21 cm wide, the base generally cordate or truncate (rarely cuneate).............................. P. cordata var. cordata 

1 Floral tube persistently pubescent with short glandular hairs; leaves lanceolate, 0.4-8.3 cm wide, the base generally cuneate 

to truncate ..........................................................................................................................................P. cordata var. lancifolia 

Pontederia cordata Linnaeus var. cordata, Heartleaf Pickerelweed. Cp, Pd (GA, NC, SC, VA), Mt (NC): marshes, pondshores, 

lake-shores; common, uncommon in Piedmont, rare in Mountains. May-October. Nova Scotia west to MN, south to FL 

and TX; Belize; s. Brazil, Argentina, Paraguay, and Uruguay. The recognition of infraspecific taxa in Pontederia cordata is 

controversial and requires additional study. P. cordata exhibits tristyly, an interesting breeding system. Each plant has one of 3

PONTEDERIACEAE 929 

types of flowers: (a) a short style, 3 medium and 3 long stamens, (b) a medium style, 3 short and 3 long stamens, or (c) a long 

style, 3 short and 3 medium stamens. [= GW, Z; < P. cordata – RAB, C, FNA, K, W; = P. cordata – F, G, S] 

Pontederia cordata Linnaeus var. lancifolia (Muhlenberg ex Elliott) Torrey, Lanceleaf Pickerelweed. Cp (GA, NC, SC): 

marshes, pond-shores, lake-shores; rare. May-October. S. MA (alleged to occur as far north as ME, but these reports may be 

entirely based on misidentifications of var. cordata) to s. FL, west to e. TX, mostly on the Coastal Plain, with a few records 

around the Great Lakes; Cuba; s. Brazil, Argentina, Paraguay, and Uruguay. A third variety of P. cordata, var. ovalis (Martens 

in Roemer & Schultes) Solms in Augustin de Candolle, is restricted to South America. [= GW, Z; < P. cordata – RAB, C, FNA, 

K, W; = P. lanceolata Nuttall – F, G, S] 

Zosterella Small 1913 

(see Heteranthera) 

POTAMOGETONACEAE Dumortier 1829 (Pondweed) 

A family of 2-3 genera and about 90 species, aquatic herbs, nearly cosmopolitan. References: Haynes & Hellquist in FNA 

(2000); Haynes (1978); Les & Haynes (1996); Haynes, Les, & Holm-Nielsen in Kubitzki (1998b); Wiegleb & Kaplan (1998)=Z. 

1 Stipules not adnate, or adnate to the blade < ½ the length of the stipule; peduncle stiff, the flowering spike elevated above 

the water's surface; submersed leaves translucent, flat, flexible; floating leaves present or absent .......................Potamogeton 

1 Stipules adnate to the blade for at least 2/3 the length of the stipule; peduncle flexible, the flowering spike submersed; 

submersed leaves opaque, channeled, stiff; floating leaves absent ............................................................................. Stuckenia 

Potamogeton Linnaeus 1753 (Pondweed) 

(also see Stuckenia) 

A genus of about 80 species, aquatic herbs, nearly cosmopolitan. References: Haynes & Hellquist in FNA (2000); Haynes & 

Hellquist (1996); Wiegleb & Kaplan (1998). Treatment adapted from Haynes & Hellquist in FNA (2000). 

1 Stipular sheaths of submersed leaves adnate with leaf blade base, the tip usually projecting as a ligule..........................Key A 

1 Stipular sheaths of submersed leaves free from the leaf blade base, or with only a few adnate, the ligule not obvious. 

2 Submersed leaves broadly linear-oblong to lanceolate to elliptic or nearly orbicular, 10-58 mm wide (occasional 

stranded forms lack submersed leaves).....................................................................................................................Key B 

2 Submersed leaves linear, thread-like or ribbon-like, 0.1-10 mm wide......................................................................Key C 

Key A 

1 Leaves stiffish, conspicuously 2-ranked, auriculate-lobed to rounded at the junction with the stipule, with 20-60 fine veins ... 

................................................................................................................................................................................. P. robbinsii 

1 Leaves lax, not conspicuously 2-ranked, lacking basal lobes, with fewer than 20 veins. 

2 Tips of submersed leaves obtuse to acute; floating leaves rounded at apex. 

3 Tips of submersed leaves acute; fruit 1-2 mm wide, the lateral keel with acute tips, beak minute .....P. diversifolius 

3 Tips of submersed leaves obtuse; fruit 1.3-2.4 mm wide, the lateral keel with blunt tips, beak lacking ..................... 

...................................................................................................................................................................P. spirillus 

2 Tips of submersed leaves acute to long-tapering; floating leaves acute at apex. 

4 Submersed leaves 0.1-0.6 mm wide, without obvious lacunae; floating leaves 3-7 veined .................P. bicupulatus 

4 Submersed leaves 0.2-1 (-2) mm wide, with abundant lacunae; floating leaves 9-23 veined............ P. tennesseensis 

Key B 

1 Leaf margins conspicuously serrate; stem flattened; fruit beak 2-3 mm long; turions commonly formed, hard......... P. crispus 

1 Leaf margins entire; stem terete; fruit beak < 1 mm long; turions rarely formed. 

2 Submersed leaves clasping the stem; floating leaves absent. 

3 Rhizomes spotted rusty-red; leaf tips cucullate, splitting when pressed; stipules usually persistent and conspicuous 

........................................................................................................................................................... [P. praelongus] 

3 Rhizomes unspotted; leaf tips flat, not splitting when pressed; stipules deciduous or deteriorating into fibers. 

4 Leaves orbicular to ovate, often lanceolate in soft water, 1-6 cm long, with 3-25 delicate veins; stipules 

deteriorating and deciduous, absent on lower portions of stem..................................................... P. perfoliatus 

4 Leaves ovate-lanceolate to narrowly lanceolate, 1.6-13 cm long, with 3-35 coarse veins; stipules 

disintegrating to persistent fibers, even on lower portions of stem........................................... [P. richardsonii]

POTAMOGETONACEAE 930 

2 Submersed leaves petioled or sessile, not clasping the stem; floating leaves absent or present. 

5 Submersed leaves 19-49 veined, distinctly arcuate ...............................................................................P. amplifolius 

5 Submersed leaves with fewer than 29 veins, not arcuate. 

6 Stems conspicuously black-spotted; submersed leaves crisped along the margin; floating leaves 15-21 veined 

........................................................................................................................................................... P. pulcher 

6 Stems inconspicuously spotted or lacking spots; submersed leaves flat along the margin; floating leaves 7-29 

veined. 

7 Submersed leaves with petioles 1-13 cm long. 

8 Larger submersed leaves acute at the apex with a sharp awl-like tip; fruit gray-green to olive-green, 

with well-developed lateral ridges.................................................................................. P. illinoensis 

8 Larger submersed leaves acute at the apex but lacking a sharp awl-like tip; fruit red to reddishbrown, 

with muricate lateral ridges ....................................................................................P. nodosus 

7 Submersed leaves sessile. 

9 Stipules blunt; submersed leaves 7 veined; fruit plump, stalked, tawny-olive.................. [P. alpinus] 

9 Stipules acute; submersed leaves 3-19 veined; fruit laterally compressed, not stalked, reddishbrown, 

gray-green, or olive-green. 

10 Fruit reddish-brown, with obsolete or rounded keel; submersed leaves with (3-) 5-9 veins........ 

................................................................................................................................P. gramineus 

10 Fruit gray-green or olive-green, with well-developed keel; submersed leaves with 7-19 veins .. 

................................................................................................................................ P. illinoensis 

Key C 

1 Fruit with a prominent keel 0.2-1.2 mm broad; floating leaves often present; lacunae prominent in submersed leaves. 

2 Submersed leaves 3-13 veined; stipules of submersed leaves not adnate to the leaf base; floating leaves rounded at apex 

........................................................................................................................................................................P. epihydrus 

2 Submersed leaves 1-3 (-7) veined; stipules of at least some submersed leaves adnate to the leaf base; floating leaves 

acute at apex................................................................................................................................................P. tenneseensis 

1 Fruit with a keel < 0.2 mm broad; floating leaves absent or present; lacunae present in some species, but generally not 

prominent. 

3 Floating leaves present, at least in some plants of the population. 

4 Floating leaves 0.3-1.6 cm long; peduncle 0.6-1.5 cm long; fruit < 2.5 mm long...................................... [P. vaseyi] 

4 Floating leaves 1.5-12 cm long; peduncle 2.5-3.5 mm long; fruit 2.5-5 mm long. 

5 Petiole junction with leaf distinctly pale in color; floating leaves ovate, oblong-ovate, cordate at base, rarely 

tapering ................................................................................................................................................P. natans 

5 Petiole junction with leaf lacking pale color; floating leaves elliptical, ovate-elliptical, or oblong-ellliptical. 

6 Floating leaves 7-12 mm wide, tapering at both ends; fruit apparently not produced ......... [P. floridanus] 

6 Floating leaves 10-20 (-30) mm wide, obtuse, round or tapering at the base; fruit often produced............. 

..............................................................................................................................................P. oakesianus 

3 Floating leaves absent from all plants in the population. 

7 Rhizomes obvious; peduncle 5-25 cm long; leaves thread-like, 0.1-0.5 mm wide............................. P. confervoides 

7 Rhizomes absent or not apparent; peduncle 0.3-7 cm long, often curved; leaves usually not thread-like, 0.1-5 mm 

wide. 

8 Nodal glands absent. 

9 Leaves 15-35 veined, > 2 mm wide; stem conspicuously flattened; peduncles terminal, usually straight... 

...........................................................................................................................................P. zosteriformis 

9 Leaves 3-5 veined, usually < 2 mm wide; stem terete; peduncles usually axillary, recurved. 

10 Leaves acute, 3 (-5) veined, 0.3-1.5 (-2.3) mm wide; fruits 1-keeled, 1.4-2.3 (-2.7) mm long............ 

....................................................................................................................... P. foliosus var. foliosus 

10 Leaves usually bristle-tipped, occasionally apiculate to blunt, 3 veined, 1-2.2 (-4) mm wide; fruits 

3-keeled, 2.3-4 mm long.......................................................................................................... P. hillii 

8 Nodal glands present. 

11 Stipules fibrous, often whitish. 

12 Leaf apex acute or apiculate; leaves 5-7 (-9) veined; turions with inner leaves at a right angle to 

outer leaves..........................................................................................................................[P. friesii] 

12 Leaf apex usually bristle-tipped, acute or rarely obtuse to apiculate; leaves 3-5 (-7) veined; turions 

flattened with inner and outer leaves in same plane ......................................................P. strictifolius 

11 Stipules not fibrous, usually delicate, green, brown, or white. 

13 Leaf apex bristle-tipped (rarely apiculate); peduncles recurved, axillary or axillary and terminal, 

0.5-6.6 cm long........................................................................................................................ P. hillii 

13 Leaf apex blunt, acute, or apiculate, but not bristle-tipped; peduncles straight, terminal, 0.5-6.6 cm 

long.


14 Mature fruit obovate, sides concave, beak mostly forward; peduncle filiform to cylindrical, 

usually 1-3 per plant; inflorescence usually interrupted; leaves with up to 2 rows of lacunae 

along midrib, apex acute, rarely apiculate; stipules mostly connate ...... P. pusillus var. pusillus 

14 Mature fruit mostly widest at middle, or ovate, sides rounded, beak mostly central; peduncle 

cylindrical, usually > 3 per plant; inflorescence continuous; leaves with 1-5 rows of lacunae 

along midrib, apex acute to obtuse; stipules mostly convolute........P. pusillus var. tenuissimus 

Potamogeton amplifolius Tuckerman, Bigleaf Pondweed. Cp (NC, VA), Pd (VA), Mt (GA, VA): ponds, lakes, sluggish 

streams; rare (GA Special Concern, VA Rare). June-September. Newfoundland west to British Columbia, south to e. NC, nw. 

GA (Jones & Coile 1988), n. AL, OK, and CA. [= RAB, C, F, FNA, G, K, S, W, Z] 

Potamogeton bicupulatus Fernald. Mt (VA): quiet waters; rare. July-September. ME west to WI, south to VA and se. TN. 

[= FNA, F, K, Z; = P. diversifolius Rafinesque var. trichophyllus Morong – C, GW] 

Potamogeton confervoides Reichenbach, Alga Pondweed, Conferva Pondweed, Tuckerman's Pondweed. Cp (NC, SC): 

acidic blackwater pools and streams; rare (NC Rare). April-September. Newfoundland west to Ontario, south to NJ and PA; 

disjunct in sc. NC and nc. SC (fall-line sandhills). [= RAB, C, F, FNA, G, K, Z] 

* Potamogeton crispus Linnaeus, Curled Pondweed, Curly Pondweed. Cp, Pd (GA, NC, VA), Mt (NC, VA): uncommon. 

May-September. ME, MN, s. Saskatchewan and s. British Columbia, south to NC, Panhandle FL, TX, AZ, and CA. [= RAB, C, 

F, FNA, G, GW, K, W, Z] 

Potamogeton diversifolius Rafinesque, Common Snailseed Pondweed. Cp, Pd, Mt (GA, NC, SC, VA): pools, ponds, and 

lakes; common. June-September. MA and NY west to MN, MT, and OR, south to FL, TX, and CA. [= RAB, FNA, G, K, S, W, 

Z; = P. diversifolius var. diversifolius – C, GW; > P. diversifolius – F; > P. capillaceus Poiret var. capillaceus – F; > P. 

capillaceus Poiret var. atripes Fernald – F] 

Potamogeton epihydrus Rafinesque, Ribbonleaf Pondweed. Mt, Pd (NC, VA), Cp (NC, SC, VA): uncommon. June- 

September. Newfoundland west to AK, south to GA, s. MS (Sorrie & Leonard 1999), LA, CO, and CA. [= RAB, C, FNA, K, S, 

W; > P. epihydrus var. epihydrus – F, G; > P. epihydrus var. nuttallii (Chamisso & Schlechtendahl) Fernald – F, G; < P. 

epihydrus – Z (also see P. tennesseensis)] 

Potamogeton foliosus Rafinesque var. foliosus, Leafy Pondweed. Mt (GA, NC, VA), Pd (SC, VA), Cp (NC, SC, VA): 

uncommon. May-October. Newfoundland west to AK, south to SC, w. FL, TX, and Mexico. [= C; < P. foliosus – RAB, G, 

GW, S, W, Z; > P. foliosus var. foliosus – F; > P. foliosus var. macellus Fernald – F; = P. foliosus ssp. foliosus – FNA, K; > P. 

curtissii Morong – S; > P. foliosus – S] 

Potamogeton gramineus Linnaeus, Variable Pondweed. Cp (VA): estuarine waters; rare. Greenland and AK, south to sc. 

PA (Rhoads & Klein 1993), NJ, WV (Kartesz 1999), n. VA, MI, WI, CO, UT, and CA. Reported for VA (Fairfax County); 

specimen identification needing confirmation. [= C, FNA, G, K, Z; > P. gramineus var. maximus Morong – F] 

Potamogeton hillii Morong, Hill's Pondweed. Mt (VA): spring-fed oxbow pond, rare (VA Rare). VT, MA, Ontario, and 

WI south to PA, VA, and OH. [= C, FNA, G, K, Z; > P. hillii – F; > P. porteri Fernald – F] 

Potamogeton illinoensis Morong, Illinois Pondweed. Cp (GA, NC, SC, VA), Mt (GA, VA), Pd (VA): calcareous waters of 

streams, lakes, and ponds; rare. May-September. Québec west to Nortwest Territories and s. British Columbia, south to FL, TX, 

Mexico, and CA. [= RAB, C, F, FNA, G, GW, K, W, Z; > P. angustifolius Berchtold & K. Presl – S; > P. heterophyllus 

Schreber – S; > P. lucens Linnaeus – S, misapplied] 

Potamogeton natans Linnaeus, Floating Pondweed. Mt (NC): lakes and slow streams; rare. June-September. 

Newfoundland west to AK, south to e. WV, w. NC, KS, NM, AZ, and CA. [= RAB, C, F, FNA, G, K, S, W; < P. natans – Z 

(also see P. floridanus)] 

Potamogeton nodosus Poiret, Longleaf Pondweed, American Pondweed. Cp (GA, NC, VA), Mt (NC, VA), Pd (VA), 

{SC}: ponds, streams; uncommon. May-September. ME and Québec west to British Columbia, south to FL, TX, Mexico, and 

CA. [= RAB, C, F, FNA, G, GW, K, W, Z; ? P. fluitans Roth – S; P. oblongifolium Forster, proposed for nomenclatural 

rejection (Reveal et al. 2003)] 

Potamogeton oakesianus J.W. Robbins, Oakes Pondweed. Cp, Mt (VA): lakes and streams; rare (VA Rare). 

Newfoundland west to MN, south to VA and n. IL; apparently disjunct in MT, and British Columbia, and possibly in s. AL 

(Sorrie, pers. comm.). [= C, F, FNA, G, K, W, Z] 

Potamogeton perfoliatus Linnaeus, Perfoliate Pondweed, Redhead Grass. Cp (NC, VA): rare (VA Watch List). June- 

October. Newfoundland, Labrador west to MI, south to ne. NC, and n. OH; apparently disjunct in w. FL, s. AL, and se. LA, and 

in SD. [= FNA, G, K, S, Z; > P. perfoliatus var. bupleuroides (Fernald) Farwell – RAB, F, GW; P. bupleuroides Fernald] 

Potamogeton pulcher Tuckerman, Spotted Pondweed. Cp, Pd, Mt (GA, NC, SC, VA): ponds, pools, ditches, streams; 

common. June-September. Nova Scotia west to WI, south to FL and e. TX. [= RAB, C, F, FNA, G, GW, K, S, W, Z; = P. 

rotundifolium Forster, proposed for nomenclatural rejection (Reveal et al. 2003)] 

Potamogeton pusillus Linnaeus var. pusillus. Cp (GA, NC, SC, VA), Pd, Mt (NC, SC, VA): acid and alkaline waters; 

uncommon? May-September. Nova Scotia west to AK, south to Mexico. [< P. berchtoldii Fieber – RAB; > P. pusillus var. 

pusillus – F; > P. pusillus var. minor (Bivona-Bernardi) Fernald & Schubert – F; = P. pusillus ssp. pusillus – FNA, K; < P. 

pusillus var. pusillus – C; < P. pusillus – G, GW, S, Z] 

Potamogeton pusillus Linnaeus var. tenuissimus F.K. Mertens & W.D.J. Koch, Slender Pondweed. Cp, Pd, Mt (NC, SC, 

VA), {GA}: millponds, other quiet waters; rare? (GA Special Concern). May-September. Newfoundland west to AK, south to 

w. FL, TX, NM, and CA. Reported from SC by Gaddy & Rayner (1980). [= W; < P. berchtoldii Fieber – RAB; < P. pusillus 

var. pusillus – C; > P. berchtoldii var. acuminatus Fieber – F; > P. berchtoldii var. berchtoldii – F; > P. berchtoldii var. 

lacunatus (Hagström) Fernald – F; > P. berchtoldii var. acuminatus Fieber – F; > P. berchtoldii var. polyphyllus (Morong)


Fernald – F; > P. berchtoldii var. tenuissimus (Mertens & Koch) Fernald – F; < P. pusillus – G, GW, S, Z; = P. pusillus ssp. 

tenuissimus (Mertens & Koch) R.R. Haynes & C.B. Hellquist – FNA, K] 

Potamogeton robbinsii Oakes, Fern Pondweed. Pd (VA): muddy waters; rare (VA Rare). August-September. Nova Scotia 

and Prince Edward Island west to Keewatin and AK, south to n. VA, n. IL, s. MN, CO, UT, and CA; disjunct in s. AL. [= C, F, 

FNA, G, K, Z] 

Potamogeton spirillus Tuckerman, Northern Snailseed Pondweed. Cp (VA): quiet waters; rare (VA Rare). July- 

November. Newfoundland west to Manitoba, south to e. VA, n. OH, n. IA, and se. NE. [= C, F, FNA, G, K, Z] 

Potamogeton strictifolius A. Bennett, Straightleaf Pondweed. Mt (VA): calcareous waters; rare (VA Rare). July- 

September. Newfoundland west to Yukon, south to w. VA, n. IL, WY, and n. UT. [= C, FNA, G, K, W, Z; > P. strictifolius var. 

strictifolius – F; > P. strictifolius var. rutiloides Fernald – F; > P. longiligulatus Fernald – F] 

Potamogeton tennesseensis Fernald, Tennessee Pondweed. Mt (VA), {NC?}: quiet or flowing water; rare (VA Rare). 

Late May-September. PA and OH south to w. VA, and se. TN. [= F, FNA, K, W; < P. epihydrus – Z, in part] 

Potamogeton zosteriformis Fernald, Flatstem Pondweed. Cp, Mt (VA): quiet waters; rare (VA Rare). July-September. 

Newfoundland west to AK, south to n. VA, n. IL, KS, UT, and CA. [= C, F, FNA, G, K, Z] 

Potamogeton alpinus Balbis, Red Pondweed, south to e. PA (Rhoads & Klein 1993). [= FNA, G, K, Z; > P. alpinus var. 

tenuifolius (Rafinesque) Ogden – C, F; > P. tenuifolius Rafinesque] 

Potamogeton floridanus Small, Florida Pondweed, is apparently endemic to blackwater rivers of the Panhandle of FL. 

Considering the under-collection of Potamogeton, it should be sought elsewhere. [= FNA, S; < P. natans – Z] 

Potamogeton friesii Ruprecht, Fries's Pondweed, south to c. PA (Rhoads & Klein 1993). (VA Watch List) [= C, F, FNA, 

G, K, Z] 

Potamogeton obtusifolius Mertens & Koch, south to MD, NJ, and PA. [= C, F, FNA, G, K] {not keyed at this time; 


Potamogeton praelongus Wulfen, Whitestem Pondweed, south to MD and nw. PA (Rhoads & Klein 1993). [= C, F, FNA, 

G, K, Z] 

Potamogeton richardsonii (Bennett) Rydberg, Richardson Pondweed, south to DE, MD, and PA. [= C, F, FNA, G, K, Z] 

Potamogeton vaseyi J.W. Robbins, Vasey Pondweed, south to se. and sc. PA (Rhoads & Klein 1993). [= C, F, FNA, G, K, 

Z] 

Stuckenia C. Börner 1912 (Sago-pondweed) 

A genus of about 10 species, nearly cosmopolitan. This genus should be called Stuckenia, which has priority over Coleogeton. 

References: Haynes & Hellquist in FNA (2000); Les & Haynes (1996)=Z; Haynes, Les, & Král (1998)=Y; Wiegleb & Kaplan 

(1998)=X. 

1 Leaves minutely acute (use 10× magnification); achenes usually 3-4.5 mm long, with the style persistent as a very short 

beak ......................................................................................................................................................................... S. pectinata 

1 Leaves minutely rounded or retuse (use 10× magnification); achenes usually 2-3 mm long, with the sessile stigma persistent 

as a bump............................................................................................................................................ [S. filiformis ssp. alpine] 

Stuckenia pectinata (Linnaeus) C. Börner, Sago-pondweed. Cp (NC, SC, VA), Mt, Pd (VA), {GA}: calcareous or brackish 

waters of ponds, lakes, estuaries, sounds; uncommon. June-September. The species is irregularly cosmopolitan. [= FNA, K, Y; 

= Potamogeton pectinatus Linnaeus – RAB, C, F, G, GW, S, W, X; = Coleogeton pectinatus (Linnaeus) D.H. Les & R.R. 

Haynes – Z] 

Stuckenia filiformis (Persoon) C. Börner ssp. alpina (Blytt) R.R. Haynes, D.H. Les, & M. Král, Threadleaf Pondweed, 

approaches our area in se. and sc. PA. [= FNA, K, Y; = Potamogeton filiformis Persoon var. alpinus (Blytt) Ascherson & 

Graebner; > Potamogeton filiformis Persoon var. borealis (Rafinesque) H. St. John – C, F, G; < Potamogeton filiformis – X; = 

Coleogeton filiformis (Persoon) D.H. Les & R.R. Haynes ssp. alpinus (Blytt) D.H. Les & R.R. Haynes – Z] 

RUPPIACEAE Horaninow ex Hutchinson 1934 (Wigeon-grass Family) 

A family of a single genus and 1-10 species. References: Haynes (1978)=Z; Haynes in FNA (2000); Haynes, Holm-Nielsen, & 

Les in Kubitzki (1998b). 

Ruppia Linnaeus (Wigeon-grass) 

A genus of 1-10 species, nearly cosmopolitan. References: Haynes (1978)=Z; Haynes in FNA (2000); Haynes, Holm-Nielsen, & 

Les in Kubitzki (1998b).


Identification notes: Separable from superficially similar species of Potamogeton by the stipules adnate their entire length (vs. 

separate at least at the tip in Potamogeton). 

Ruppia maritima Linnaeus, Wigeon-grass, Ditch-grass. Cp (GA, NC, SC, VA): brackish estuaries, rivers, marsh pools; 

common. July-October. Nearly cosmopolitan. [= RAB, C, FNA, GW, K, S, Z; > R. maritima var. maritima – F, G; > R. 

maritima var. longipes Hagström – F; > R. maritima var. rostrata Agardh – F, G] 

RUSCACEAE M. Roemer 1840 (Ruscus Family) 

As here circumscribed, a family of about 28 genera and 500 species, of North America, Central America, Europe, and Asia. The 

Convallariaceae has been supported by molecular studies (Judd 2003, Bogler & Simpson 1995). Molecular studies show that 

Nolina is much more closely related to Convallaria, Polygonatum, etc. than to the Agavaceae (Yucca and Manfreda in our flora), 

with which it has often been associated. References: Bogler & Simpson (1995); Bogler in Kubitzki (1998a); Yeo in Kubitzki 

(1998a); Conran & Tamura in Kubitzki (1998a); Yamashita & Tamura (2000). 

1 Plant with an upright stem with alternate leaves. 

2 Shrub; "leaves" (actually phylloclades) coriaceous, evergreen, glossy; [exotic, rarely naturalized]; [tribe Rusceae]......... 

..................................................................................................................................................................................Danae 

2 Herb; leaves herbaceous, deciduous, dull or slightly glossy; [native]; [tribe Polygonatae]. 

3 Inflorescence terminal, a raceme or panicle; tepals separate............................................................... Maianthemum 

3 Inflorescence of 1-several axillary flowers; tepals fused........................................................................Polygonatum 

1 Plant tufted, the leaves essentially basal (although the sheathing bases form a 'false' stem in Convallaria). 

4 Leaves 2-3, narrowly elliptic; tepals fused, white or greenish; [tribe Convallarieae]...................................... Convallaria 

4 Leaves many, linear; tepals separate (or fused basally), white or violet. 

5 Fruit dehiscent, quickly exposing berry-like seeds with a fleshy seed coat; inflorescence spikelike, to 3 dm tall; 

[aliens, scarcely naturalized from horticultural plantings]; [tribe Ophiopogoneae] ........................................ Liriope 

5 Fruit indehiscent, dry and capsular; inflorescence a panicle or raceme, to 15 dm tall; [natives, of longleaf pine 

woodlands of SC, GA, and FL]; [tribe Nolineae]............................................................................................. Nolina 

Convallaria Linnaeus 1753 (Lily-of-the-Valley) 

A genus of 3 species, of north temperate n. Eurasia and e. North America. References: Utech in FNA (2002a); Judd (2003)=Z; 

Conran & Tamura in Kubitzki (1998a). 

1 Leaf blades averaging 10-15 cm long, 3-5 cm wide; rhizomes short-creeping, the "individual" plants spaced closely 

(typically 5-10 cm apart); flowering scape > ½ as long as the leaves; longer bracts of the inflorescence 4-10 mm long; 

[introduced, persistent around old home sites and other plantings] ............................................................................C. majalis 

1 Leaf blades averaging 15-35 cm long, 5-13 cm wide; rhizomes absent or long-creeping, the individual plants spaced widely 

(usually at least 15 cm apart); flowering scape < ½ as long as the leaves; longer bracts of the inflorescence 8-20 mm long; 

[native, of forests of the Mountains and upper Piedmont] .....................................................................................C. majuscula 

* Convallaria majalis Linnaeus, European Lily-of-the-Valley. Pd, Cp, Mt (NC, VA): persistent after cultivation; rare, 

introduced from Eurasia. April-May; July-August. [= F, K, W; = C. majalis var. majalis – RAB, FNA; < C. majalis – C, G, S, Z 

(also see C. majuscula)] 

Convallaria majuscula Greene, American Lily-of-the-Valley. Mt (GA, NC, SC, VA), Pd (NC, VA): mountain forests, 

particularly in rocky woodlands or forests on or near ridgetops under northern red oak at about 1000 to 1500 m elevation, 

sometimes at lower elevations (down to at least 700 m) and under Quercus montana; uncommon, rare in Piedmont (SC Rare). 

April-June; August. Endemic to the Southern Appalachians: WV and VA through NC and TN to ne. GA (Jones & Coile 1988) 

and nw. SC. Cronquist's (1991) statement that Southern Appalachian plants "may reflect an early escape of a different phase of 

the species from cultivation" can be discounted; there is no doubt that C. majuscula is both native and taxonomically distinct, at a 

varietal level at least. Utech in FNA (2002a) states that our plants are more closely related to the Asian taxon, variously treated 

as C. keiskei Miquel or C. majalis var. keiskei (Miquel) Makino, than to the European C. majalis s.s. As best as can be 

determined, Rafinesque's name Convallaria montana does not apply to this species. [= K; = C. majalis Linnaeus var. montana 

(Rafinesque) Ahles – RAB, FNA, apparently misapplied; < C. majalis – C, G, S, Z; = C. montana Rafinesque – F, W, apparently 

misapplied] 

A monotypic genus, a shrub, of sw. Asia. 

Danae Medikus (Alexandrian Laurel, Danaë)

RUSCACEAE 934 

* Danae racemosa (Linnaeus) Moench, Alexandrian Laurel. Pd (NC): suburban forests; rare, uncoomon in cultivation, arrely 

escaping to suburban forests, native of sw. Asia. The "leaves" are actually odd structures called phylloclades, and represent 

modified stems. 

Liriope Loureiro 1790 (Liriope, Lilyturf) 

A genus of 8 species, herbs, of e. and se. Asia. References: Conran & Tamura in Kubitzki (1998a); Judd (2003)=Z. 

* Liriope muscari (Dcne.) Bailey, Liriope, Big Blue Lilyturf. Pd (NC), {SC}: planted and persistent; commonly cultivated, 

rarely persistent. [= K, Z] {not keyed at this time} 

* Liriope spicatum Loureiro, Creeping Lilyturf. Reported for AL, FL, MD (Kartesz 1999). [= K; L. spicata Loureiro – Z, 

orthographic variant] {not keyed at this time} 

Maianthemum G.H. Weber ex Wiggers 1780 (Mayflower, Solomon's-plume) 

A genus of about 28 species, herbs, of n. Europe, e. Asia, North America, and Central America. The inclusion of the traditional 

Smilacina in Maianthemum will cause considerable consternation; LaFrankie's (1986) reasoning, however, seems very strong, 

and has been additionally supported by more recent evidence (Conran & Tamura in Kubitzki 1998a). The only consistent 

difference between the two previously accepted genera is whether the flowers are dimerous (Maianthemum) or trimerous 

(Smilacina). LaFrankie cites research that shows that the dimerous flowers of Maianthemum (sensu stricto) are the result of 

reduction from trimerous flowers, as indicated by vestigial vascular traces. Consideration of the many close similarities, 

particularly as compared to similar genera such as Prosartes, Polygonatum, Streptopus, and Clintonia, may convince the 

skeptical (see LaFrankie 1986 and Therman 1956). As an example, the fruits of M. canadense and M. racemosum are closely 

similar in form, coloration, and size; much more similar than the fruits of our 2 species of Prosartes. References: LaFrankie 

(1986)=Z; Judd (2003)=Y; LaFrankie in FNA (2002a); Conran & Tamura in Kubitzki (1998a). 

1 Flowers in a terminal panicle................................................................................................... M. racemosum ssp. racemosum 

1 Flowers in a simple raceme. 

2 Perianth segments 4 (flowers 2-merous); leaves (1-) 2 (-3) ..........................................................................M. canadense 

2 Perianth segments 6 (flowers 3-merous); leaves 6 or more (or 1-4 in M. trifolium). 

3 Leaves 6 or more, pubescent beneath......................................................................................................M. stellatum 

3 Leaves 1-4, glabrous beneath................................................................................................................[M. trifolium] 

Maianthemum canadense Desfontaines, Canada Mayflower, False Lily-of-the-valley. Mt (GA, NC, VA), Pd (VA): moist 

forests, especially at high elevations; common. Mid May-early July; August-September. Labrador and Newfoundland west to 

Mackenzie, south to MD, NC, n. GA (Jones & Coile 1988), KY and SD. Two varieties have been described, but their 

recognition is not strongly supported. Var. canadense, with leaves glabrous beneath, the margins entire or minutely crenulate, 

cross-veins of the leaf well-developed, is widepread in the distribution of the species. Var. interius Fernald has the leaves 

pubescent beneath, the leaf margins distinctly ciliate, and cross-veins obscure; it is not known from our area, but extends east and 

south as far as MA, NY, and OH. Further study of these varieties is needed. Weller (1970) reports equivocal results on the 

recognition of varieties, based on a study in n. MI. [= RAB, FNA, K, W, Y, Z; > Maianthemum canadense Desfontaines var. 

canadense – C, F, G; = Unifolium canadense (Desfontaines) Greene – S] 

Maianthemum racemosum (Linnaeus) Link ssp. racemosum, Eastern Solomon's-plume, False Solomon's-seal. Mt, Pd, Cp 

(GA, NC, SC, VA): forests; common. Mid April-June; August-October. The species ranges from Nova Scotia west to British 

Columbia, south to GA and AZ. A variety of chromosome races are known (2n = 36, 72, 144). The eastern ssp. racemosum is 

tetraploid; ssp. amplexicaule (Nuttall) LaFrankie is diploid and more western; these are perhaps more appropriately treated as 

species. Under the generic name Smilacina, two varieties had been described for our area, Smilacina racemosa var. racemosa 

and S. racemosa var. cylindrata Fernald, the former larger in nearly all respects and more northern than the latter, smaller, and 

more southern form (see F for details). If these varieties are determined to have merit (further research is needed), the 

appropriate transfer to Maianthemum will need to be made. [= FNA, K, Y, Z; < Smilacina racemosa (Linnaeus) Desfontaines – 

RAB, C, G, W; > S. racemosa var. racemosa – F; > S. racemosa var. cylindrata Fernald – F; > Vagnera racemosa (Linnaeus) 

Morong – S; > Vagnera australis Rydberg – S] 

Maianthemum stellatum (Linnaeus) Link, Starry Solomon's-plume. Mt, Pd (VA): alluvial forests; rare (VA Rare). April- 

June; August-October. Newfoundland west to British Columbia, south to NJ, w. VA, e. TN, IN, MO, and CA. [= FNA, K, Y, Z; 

= Smilacina stellata (Linnaeus) Desfontaines – C, F, G, W] 

Maianthemum trifolium (Linnaeus) Sloboda occurs in bogs and moist sphagnous forests, south to sc. PA. [= FNA, K, Z; = 

Smilacina trifolia (Linnaeus) Desfontaines – C, F, G] 

Nolina Michaux 1803 (Beargrass)

RUSCACEAE 935 

A genus of about 30 species, rosette shrubs and trees, of s. United States and Mexico. References: Hess in FNA (2002a); Judd 

(2003)=Z; Bogler in Kubitzki (1998a). 

1 Leaves 3-4 (-5) mm wide; fruit 4-4.5 mm long, strongly asymmetrical; [of moist flatwoods of the FL Coastal Plain].............. 

...........................................................................................................................................................................[N. atopocarpa] 

1 Leaves 4-10 mm wide, glaucous; fruit 6.5-8 mm long, symmetrical; [of dry to dry-mesic sandhills of the GA and SC Coastal 

Plain]...................................................................................................................................................................... N. georgiana 

Nolina georgiana Michaux, Georgia Beargrass, Sandhills Lily. Cp (GA, SC): sandhills, sometimes locally common on 

slightly less xeric lower sandhill slopes; rare (SC Rare). Late May-June; late June-August. Nc. SC south to sc. GA. This species 

has been attributed to FL (Small 1933), but is not included in either Clewell (1985) or Wunderlin (1982, 1998). [= RAB, FNA, 

K, S, Z] 

Nolina atopocarpa Bartlett, Florida Beargrass. Pine flatwoods and savannas. Endemic to panhandle FL (Liberty and 

Franklin counties) and e. peninsular FL (St. Johns and Brevard counties). [= FNA, K, S, Z] 

Polygonatum P. Miller 1754 (Solomon's-seal) 

A genus of about 57 species, of temperate Eurasia and North America (most diverse in e. Asia). The P. biflorum complex is in 

need of further study. References: Ownbey (1944)=Z; Judd (2003)=Y; Utech in FNA (2002a); Eigsti (1942); Therman (1950, 

1953); Kawano & Iltis (1963); Conran & Tamura in Kubitzki (1998a). 

1 Leaves pubescent on the veins beneath; flowers 7-13 mm long............................................................................. P. pubescens 

1 Leaves glabrous; flowers 12-21 mm long. 

2 Stem slender, 1.5-5 mm in diameter; plants to 9 dm tall; lower axillary peduncles terete or nearly so, with (1-) 2-3 (-5) 

flowers; lowest peduncle in the axil of the (1st-) 3rd (-5th) axil; larger leaves 5.5-15 cm long, 1.2-6 cm wide; lower 

leaves clasping to 90 (-180) degrees .......................................................................................... P. biflorum var. biflorum 

2 Stem robust, 5-13 mm thick below the leaves; plants to 20 dm tall; lower axillary peduncles strongly flattened, with (2- 

) 3-6 (-15) flowers; lowest peduncle in the axil of the (3rd-) 4th-5th (-8th) leaf; larger leaves 9-25 cm long, 3.5-13 cm 

wide; lower leaves clasping to 300 degrees ........................................................................P. biflorum var. commutatum 

Polygonatum biflorum (Walter) Elliott var. biflorum, Small Solomon's-seal. Mt, Pd, Cp (GA, NC, SC, VA): moist to dry 

forests; common. April-June; August-October. CT, NY, and s. Ontario west to MI, NE, and IN, south to n. FL and s. AL. In 

addition to the two varieties recognized for our area, P. biflorum includes several additional varieties: var. hebetifolium R.R. 

Gates of panhandle FL (Apalachicola River area), var. melleum (Farwell) R. Ownbey of MI and Ontario, and var. necopinum R. 

Ownbey from the Black Hills of SD. The complex needs additional study. See var. commutatum for discussion of its distinction 

from var. biflorum. [= Z; < P. biflorum – RAB, C, FNA, W, Y; = P. biflorum – F, G, in the narrow sense; < P. biflorum var. 

commutatum (J.A. & J.H. Schultes) Morong – K; < P. commutatum (J.A. & J.H. Schultes) A. Dietrich – S] 

Polygonatum biflorum (Walter) Elliott var. commutatum (J.A. & J.H. Schultes) Morong, Large Solomon's-seal, King 

Solomon's-seal. Mt (GA, NC, SC?, VA), Pd (NC, VA): moist forests, roadbanks; common. May-June; September-October. 

NH west to s. Manitoba, south to SC, GA, LA, and TX. There has been a wide divergence of opinion regarding the merits (and 

practicality) of distinguishing this taxon from typical P. biflorum, and the characters considered most reliable; the two taxa may 

differ in chromosome number and geographical distribution; they are not, however, always readily distinguished 

morphologically. I prefer to recognize this taxon as a variety. See references for additional discussion. [< P. biflorum – RAB, 

C, FNA, W, Y; = P. canaliculatum (Muhlenberg ex Willdenow) Pursh – F, G, misapplied; < P. biflorum var. commutatum (J.A. 

& J.H. Schultes) Morong – K; < P. commutatum (J.A. & J.H. Schultes) A. Dietrich – S; = P. commutatum – Z] 

Polygonatum pubescens (Willdenow) Pursh. Mt (GA, NC, SC?, VA): moist forests, especially cove forests; common. 

Late April-June; August-October. S. Québec west to s. Manitoba, south to nw. SC, WV, KY, IN, WI, and IA. [= RAB, C, F, 

FNA, G, K, W, Y, Z; = P. biflorum – S, misapplied] 

SCHEUCHZERIACEAE F. Rudolphi 1830 (Scheuchzeria Family) 

A monotypic family, circumboreal in arctic and cold temperate regions. References: Nienaber in FNA (2000); Haynes, Les, & 

Holm-Nielsen in Kubitzki (1998b). 

Scheuchzeria Linnaeus (Scheuchzeria, Pod-grass) 

A monotypic genus, circumboreal in arctic and cold temperate regions. References: Nienaber in FNA (2000); Haynes, Les, & 

Holm-Nielsen in Kubitzki (1998b).

RUSCACEAE 936 

Scheuchzeria palustris Linnaeus var. americana Fernald, Pod-grass. Sphagnum bogs. South to sc. PA (Rhoads & Klein 

1993), NJ, and Pocahontas County, WV. [= F; < Sch. palustris – C, FNA, G; = Sch. palustris ssp. americana (Fernald) Hultén – 

K; = Sch. americana (Fernald) G.N. Jones] 

SMILACACEAE Ventenat 1799 (Greenbrier Family) 

A family of 3-12 genera and about 375 species, widespread in tropical, subtropical, and temperate regions. References: Holmes 

in FNA (2002a); Judd (1998); Conran in Kubitzki (1998a). 

Smilax Linnaeus 1753 (Greenbrier, Carrionflower, Smilax) 

A genus of about 300 species, woody vines and herbs, subcosmopolitan in temperate and tropical regions. Our deciduous species 

are a monophyletic group within Smilax, with a classic eastern North American - east Asian disjunction, and are treated as section 

Nemexia or subgenus Luiste (Wilbur 2004, Fu et al. 2005). Smilax berries and shoots provide important food sources for many 

wildlife species, including black bears (Ursus americanus). References: Mangaly (1968)=Z; Judd (1998)=Y; Holmes in FNA 

(2002a); Wilbur (2004); Fu et al. (2005); Duncan (1967); Godfrey (1988). Key for the woody species based in part on Godfrey 

(1988). 

1 Stem herbaceous, lacking prickles; ovules 2 per carpel; peduncles usually > 4 cm long; [section Nemexia]. 

2 Plants erect, 0.2-1.0 m tall, even when well-developed with < 20 leaves [note that immature or depauperate individuals 

(nonflowering) of S. pseudochina, S. herbacea, S. lasioneura, and S. pulverulenta often have this aspect]; tendrils 

absent or rudimentary; peduncles usually few (usually 1-4), the lowest often from bract axils. 

3 Leaves glabrous and glaucous beneath, thick in texture, base cordate, tip acute or acuminate; lowest peduncle from 

a leaf axil (very rarely from bract axils), upper peduncles from leaf axils; leaves 5-7, clustered together near the 

summit of the stem ............................................................................................................................. S. biltmoreana 

3 Leaves pubescent and green (or glaucous) beneath, usually thin in texture, base cordate, truncate, or rounded, tip 

acuminate, acute, or obtuse; lowest peduncles from axils of bracts below the lowest leaves, upper peduncles also 

often from bracts (the uppermost often from leaf axils); leaves either clustered together near the summit of the 

stem or well distributed. 

4 Leaves relatively many, (7-) 10-13 (-20), often well distributed in the upper half of the stem, notably reduced 

in size from lower to upper, mostly with the base cordate and the tip acuminate; berry 3-5 seeded ................... 

........................................................................................................................................................... S. ecirrata 

4 Leaves few, usually 4-8, usually clustered together near the summit of the stem (rarely well distributed), 

about the same size, mostly with the base ovate (to subcordate), the tip acute to obtuse; berry 2-3 seeded........ 

............................................................................................................................................................. S. hugeri 

2 Plants vine-like, climbing or sprawling, to 3 m tall, when well-developed with > 30 leaves; tendrils present and 

numerous; peduncles usually many, from leaf axils. 

5 Leaf bases hastate, the leaf margins straight or concave in outline; longest fruiting pedicels < 2× as long as the 

fruit; anthers equaling or longer than the filaments; perianth 1.5-2.5 mm long; leaves glabrous and glaucous 

beneath................................................................................................................................................S. pseudochina 

5 Leaf bases cordate, the leaf margins convex in outline; longest fruiting pedicels 2× or more as long as the fruit; 

anthers shorter than the filaments; perianth 3.5-6 mm long; leaves either puberulent beneath (at least along the 

veins), or glabrous and glaucous beneath. 

6 Leaves glabrous and glaucous on the lower surface; fruit dark blue and glaucous; peduncles 5-8× as long as 

the subtending petioles..................................................................................................................... S. herbacea 

6 Leaves puberulent on the lower surface, at least on the veins; fruit dark blue and glaucous or black and not 

glaucous; peduncles 1-10× as long as the subtending petioles. 

7 Leaves bright green and shiny beneath; fruit black, not glaucous; peduncles 1-2 (-3)× as long as the 

subtending petioles..............................................................................................................S. pulverulenta 

7 Leaves pale green and dull below; fruit dark blue, glaucous; peduncles (3-) 5-10× as long as the 

subtending petioles.................................................................................................................S. lasioneura 

1 Stem woody, usually with prickles; ovules 1 per carpel; peduncles usually < 3 cm long; [section China]. 

8 Stems and petioles tomentose, lacking prickles; leaves densely tomentose beneath; berries red; plant trailing or 

ascending, rarely > 0.5 m tall (with determinate growth).....................................................................................S. pumila 

8 Stems and petioles stellate-scurfy or glabrous, generally with prickles; leaves glabrous or papillate beneath; berries red, 

black, or dark blue; plant climbing, ascending, or trailing, mature plants generally well over 0.5 m tall (with 

indeterminate growth). 

9 Lower surfaces of leaves strongly glaucous................................................................................................. S. glauca 

9 Lower surfaces of leaves green (rarely very slightly glaucous). 

10 Prickles of the stem abundant, thin and needle-like, shiny brown or black......................................... S. hispida 

10 Prickles of the stem fewer, broad-based and awl-like or catclaw-like, green, brown, or black.

SMILACACEAE 937 

11 Midvein (as seen on the lower surface) much more pronounced than the principal lateral veins, which 

are scarcely raised; leaves evergreen, thick, coriaceous ...........................................................S. laurifolia 

11 Midvein (as seen ion the lower surface) little if any more pronounced than the principal lateral veins; 

leaves evergreen or deciduous, thin, subcoriaceous. 

12 Leaves mostly lanceolate, the base cuneate, the tip acute to acuminate; berries dull red ..... S. smallii 

12 Leaves mostly ovate, oblong, pandurate, or hastate, the base cordate, truncate, rounded, or cuneate, 

the tip rounded to acute; berries various in color. 

13 Margin of the leaf blade prominently thickened with a marginal vein (this appearing as a 

thickening, a visible vein, or an apparent revolute margin); berries with 1-3 seeds. 

14 Inflorescence peduncle (stalk of the umbel) as long as or shorter than the subtending leaf 

petiole; stems and prickles glabrous; leaves evergreen; berries usually with 2-3 seeds; 

[generally of xeric or less commonly mesic sands] ........................................ S. auriculata 

14 Inflorescence peduncle (stalk of the umbel) > 1.5×as long as the subtending leaf petiole; 

stems (especially the lower) and prickles brownish stellate-scurfy; leaves semi-evergreen 

to evergreen; berries usually with 1 seed; [of a wide variety of habitats] .........S. bona-nox 

13 Margin of the leaf blade thin, sometimes revolute; berries with (1-) 2-4 seeds. 

15 Berries blue-black; perianth green; leaves semi-evergreen to evergreen, margins of 

mature leaves generally not revolute, the margins of the leaves and the petioles often with 

minute, flattish, toothlike projections; berries with (1-) 2-3 seeds; [of a wide variety of 

upland and wetland habitats] ........................................................................S. rotundifolia 

15 Berries bright red; perianth brownish-yellow; leaves deciduous, margins of mature leaves 

usually revolute, the margins of the leaves and the petioles lacking minute, flattish, 

toothlike projections; berries with 2-4 seeds; [of swamp forests, bogs, often where 

submersed for at least part of the year].................................................................S. walteri 

Smilax auriculata Walter, Dune Greenbrier. Cp (GA, NC, SC): dunes on barrier islands, dry sandy openings in maritime 

forests or sandhills near the coast; common. May-July; October-November (and persisting). E. NC (Dare County) south to s. 

FL and west to LA; Bahama Islands. [= RAB, FNA, GW, K, S, Y] 

Smilax biltmoreana (Small) J.B.S. Norton ex Pennell, Biltmore Carrionflower. Mt, Pd (GA, NC, SC), Cp (GA): dry forests 

(such as dry pine ridges and chestnut oak forests) and moist forests; rare (NC Rare). April-May; August-October. The species is 

apparently limited to to NC, SC, and GA, primarily in the Blue Ridge Escarpment region, with disjunct occurrences in panhandle 

FL, s. AL, and sc. KY. [= FNA, K, W, Y, Z; = Smilax ecirrata (Engelmann ex Kunth) S. Watson var. biltmoreana (Small) Ahles 

– RAB; < S. ecirrhata – G, in part (concept interpreted from stated geographic range); = Nemexia biltmoreana Small – S] 

Smilax bona-nox Linnaeus, Catbrier, Tramp's-trouble. Cp, Pd, Mt (GA, NC, SC, VA): in a wide variety of upland and 

wetland habitats; common. Late April-May; September-November. MD and MO south to c. FL and TX, and also in Mexico. [= 

RAB, C, FNA, G, GW, K, S, W, Y; > S. bona-nox var. hastata (Willdenow) Alphonse de Candolle – F; > S. bona-nox var. 

exauriculata Fernald – F; > S. bona-nox var. hederifolia (Beyrich) Fernald – F; > S. bona-nox var. bona-nox – F] 

Smilax ecirrata (Engelmann ex Kunth) S. Watson. Mt (VA): forests; rare (VA Rare). Mid May-early June; August- 

September. N. OH MI, WI, and s. MN south to w. VA, TN, s. IL, MO, and e. OK. [= K, Y, Z; = S. ecirrhata – C, F, FNA, 

orthographic variant; < S. ecirrhata – G, broader sense (apparently also including in statement of range S. hugeri and/or S. 

biltmoreana); = Nemexia ecirrhata (Engelmann ex Kunth) Small – S] 

Smilax glauca Walter, Whiteleaf Greenbrier, Wild Sarsaparilla. Mt, Pd, Cp (GA, NC, SC, VA): in a wide variety of upland 

and wetland habitats; common. Late April-early June; September-November (and persisting). NJ, c. PA, OH, IN, MO, and KA 

south to c. FL and TX, and also in Mexico. [= RAB, C, FNA, GW, S, W, Y; > S. glauca var. glauca – F, G, K; > S. glauca var. 

leurophylla Blake – F, G, K] 

Smilax herbacea Linnaeus, Common Carrionflower. Mt (GA, NC, SC, VA), Pd (GA, NC, SC, VA), Cp (GA, VA): moist 

deciduous forests; common. May-June; August-October. Centered in the Appalachian Mountains, from Québec and ME west to 

OH, south to AL, GA, and TN. Young, non-flowering plants closely resemble S. biltmoreana. [= F, FNA, K, W, Y, Z; = S. 

herbacea var. herbacea – RAB, C, G; = Nemexia herbacea (Linnaeus) Small – S] 

Smilax hispida Rafinesque, Bristly Greenbrier, Hellfetter. Mt, Pd, Cp (GA, NC, SC, VA): moist to wet forests; common. 

CT, NY, MN, and NE south to c. FL and TX. Wilbur (2003) discusses the complicated nomenclatural problems involving this 

plant and concludes that S. hispida Raf. is the correct name. [= Smilax tamnoides Linnaeus – FNA, GW, K, W, Y, misapplied; = 

S. hispida Muhlenberg – RAB, C, G, S; > S. tamnoides var. hispida (Muhlenberg) Fernald – F; > S. tamnoides var. tamnoides – 

F; > S. hispida var. australis Small – S; > S. hispida var. hispida – S] 

Smilax hugeri (Small) J.B.S. Norton ex Pennell, Huger's Carrionflower. Mt (GA, NC, SC), Pd (GA, NC, SC), Cp (GA, 

SC): moist deciduous forests; rare (NC Watch List). March-April; August-October. S. NC and e. TN south through SC, GA, 

and AL to panhandle FL. [= FNA, K, W, Y, Z; = S. ecirrata (Engelmann ex Kunth) S. Watson var. hugeri (Small) Ahles – RAB; 

= Nemexia hugeri Small – S] 

Smilax lasioneura Hooker, Midwestern Carrionflower. Mt (NC, SC, VA?), Pd (NC), {GA}: moist deciduous forests, bluff 

forests, pine-oak hickory submesic forests, perhaps only or primarily over mafic rocks; rare (GA Rare, NC Rare). April-May; 

August-September. Ontario and MT south to w. VA (?), w. NC, n. FL, OK, and CO. Material from VA is ambiguous. [= F, 

FNA, K; = S. herbacea var. lasioneura (Hooker) Alphonse de Candolle – C, G; = Nemexia lasioneuron (Hooker) Rydberg – S; = 

S. lasioneuron – Y, orthographic variant]

SMILACACEAE 938 

Smilax laurifolia Linnaeus, Blaspheme-vine, Bamboo-vine. Cp, Pd, Mt (GA, NC, SC, VA): pocosins, swamp forests, 

mountain bogs in sw. NC; common (rare in Piedmont and Mountains). July-August; September-October of the second year (and 

persisting). Primarily a Southeastern Coastal Plain species: NJ south to s. FL, west to w. TN, AR, and e. TX, and also in the 

Bahama Islands and Cuba. [= RAB, C, F, FNA, G, GW, K, S, W, Y] 

Smilax pseudochina Linnaeus, Coastal Carrionflower. Cp (GA, NC, SC, VA): pocosins, swamp forests, edges of pine 

savannas; uncommon. May; August-October. An Atlantic Coastal Plain endemic: NJ, se. PA, and DE south to e. GA. [= C, 

FNA, K, Y; = S. tamnifolia Michaux – RAB, G; = S. pseudo-china – F, W, Z, orthographic variant; > Nemexia tamnifolia 

(Michaux) Small – S; > Nemexia leptanthera (Pennell) Small – S] 

Smilax pulverulenta Michaux. Mt (GA, NC, SC, VA), Pd, Cp (NC, SC, VA): moist deciduous forests: common. May- 

June; August-October. Se. NY, se. and sc. PA, IN, MO, and e. KS south to NC, TN, and AR. [= F, FNA, K, W, Y, Z; = S. 

herbacea var. pulverulenta (Michaux) A. Gray – RAB, C, G; = Nemexia pulverulenta (Michaux) Small – S] 

Smilax pumila Walter, Sarsaparilla-vine, Dwarf Smilax. Cp (GA, SC): mesic to dryish hammocks and bluffs, northward 

primarily in maritime-influenced mainland forest, with Magnolia grandiflora and Tilia americana var. caroliniana; rare (NC 

Watch List). October-November; January-April (and persisting). Ne. SC (within a few hundred meters of Brunswick County, 

NC) to FL and west to TX. It occurs on Colkins Neck, along the NC-SC border, in maritime-influenced forests with southern 

affinities, now largely destroyed by golf-course development. This unusual Smilax is sometimes cultivated as an ornamental 

ground-cover. [= RAB, FNA, K, S, Y] 

Smilax rotundifolia Linnaeus, Common Greenbrier, Bullbrier, Horsebrier. Mt, Pd, Cp (GA, NC, SC, VA): in a wide 

variety of upland and wetland habitats; common. April-May; September-November (and persisting). Nova Scotia and s. Ontario 

south to n. FL and e. TX. [= RAB, C, F, FNA, G, GW, K, S, W, Y; > S. rotundifolia var. quadrangularis (Muhlenberg ex 

Willdenow) Wood] 

Smilax smallii Morong, Jackson-brier. Cp (GA, NC, SC, VA), Pd, Mt (GA): bottomland forests; uncommon, rare in VA 

(VA Rare). June-July; April-June of the next year. Se. VA to c. FL, west to s. AR and e. TX, primarily on the Coastal Plain. [= 

RAB, FNA, G, GW, K, W, Y; = S. lanceolata Linnaeus – S, misapplied] 

Smilax walteri Pursh, Coral Greenbrier, Red-berried Swamp Smilax. Cp, Pd (GA, NC, SC, VA): swamp forests, bogs, 

often where submersed for at least part of the year; common (rare in Piedmont). Late April-May; September-November (and 

persisting). NJ south to c. FL and west to TN, AR, and TX. [= RAB, C, F, FNA, G, GW, K, S, W, Y] 

Smilax leptanthera Pennell. See Pennell (1916) for additional information. Treated as valid and rare by GAHP. [= 

Nemexia leptanthera (Pennell) Small – S; < S. pseudochina] {investigate} 

SPARGANIACEAE (Bur-reed Family) 

(see TYPHACEAE) 

STEMONACEAE Engler 1887 (Stemona Family) 

A family of 3-4 genera and 30-35 species, herbs and shrubs, of Asia, Australia, and se. North America. References: Whetstone 

in FNA (2002a); Rogers (1982)=Z; Kubitzki in Kubitzki (1998a). 

Croomia Torrey 1840 (Croomia) 

A genus of 3 species, 2 in China and Japan and 1 in se. North America. Sometimes segregated into the Croomiaceae. 

References: Whetstone in FNA (2002a); Rogers (1982)=Z; Kubitzki in Kubitzki (1998a). 

Croomia pauciflora (Nuttall) Torrey, Croomia. Cp (GA): moist forests, often with beech and basswood; rare (GA 

Threatened). April-May. AL (or perhaps LA) to sw. GA, Panhandle FL, and allegedly se. GA (Whetstone in FNA 2002, Jones 

& Coile 1988). [= FNA, K, S, Z] 

THEMIDACEAE Salisbury 1866 

A family of 12 genera and about 60 species, herbs, of w. North America south to Guatemala. References: Rahm in Kubitzki 

(1998a). 

Dichelostemma Kunth 1843 

A genus of 5 species, of w. United States and Mexico. References: Pires in FNA (2002a); Rahm in Kubitzki (1998a). 

* Dichelostemma congestum (Smith) Kunth, native of the Pacific northwestern North America, is cultivated and apparently 

escaped in the Piedmont of nc. GA (Jones & Coile 1988). {further investigate} [= FNA, K; = Brodiaea congesta Smith]

STEMONACEAE 939 

TOFIELDIACEAE Takhtajan 1994 (False-asphodel Family) 

A family of 5 genera and about 30 species, of disjunct distribution in north temperate and subarctic areas, and in the Guayana 

Shield and northern Andes areas of n. South America. Reveal & Zomlefer (1998) place the Tofieldiaceae in the monotypic order 

Tofieldiales, only distantly related to the Liliaceae. Tamura in Kubitzki (1998a) treats this group as subfamily Tofieldioideae of 

the Nartheciaceae. References: Zomlefer (1997c, 1999); Tamura in Kubitzki (1998a). 

1 Inflorescence 1-flowered; tepals yellow; seeds yellowish; [endemic to Panhandle FL] ........................... [Harperocallis flava] 

1 Inflorescence a raceme or thyrse; tepals white to pale cream (fading to yellowish on dried specimens); seeds brown; 

[collectively widespread]. 

2 Bracts of the inflorescence large, spathelike, acuminate-aristate at the tip; tepals 9-17 mm long; stamens (6-) 9 (-12) ..... 

................................................................................................................................................................................... Pleea 

2 Bracts of the inflorescence minute; tepals 2.5-5 mm long; stamens 6. 

3 Inflorescence a raceme (the flower pedicels attached to the scape singly); scape smooth; flowering (late August-) 

late September-October.................................................................................................................................Tofieldia 

3 Inflorecence a thyrse (flower pedicels attached to the scape in groups of 3-7); scape scurfy-scabrous; flowering 

June-August ..................................................................................................................................................Triantha 

Harperocallis McDaniel (Harper's Beauty) 

A monotypic genus, perennial, of southeastern United States. References: McDaniel (1968)=Y; Zomlefer (1997c)=Z; Utech & 

Anderson in FNA (2002a). 

Harperocallis flava McDaniel, Harper's Beauty. Pineland bogs. Endemic to FL Panhandle (Franklin and Liberty counties). 

[= FNA, K, Y, Z] 

Pleea Michaux 1803 (Rush-featherling) 

A monotypic genus, of se. North America, sometimes included in Tofieldia. References: Zomlefer (1997c)=Z; Tamura in 

Kubitzki (1998a); Packer in FNA (2002a). 

Pleea tenuifolia Michaux, Rush-featherling. Cp (GA?, NC, SC): locally abundant in wet savannas, pocosin margins, 

usually in peaty soil, locally abundant in a few counties in se. NC, rare inland (very rarely as far as Cumberland County, NC); 

uncommon (SC Rare). September-October; October-November. A Southeastern Coastal Plain endemic: se. NC and ne. SC 

south to sw. GA, n. FL and s. AL, but apparently absent from s. SC and ne. GA. When in flower in wet savannas and powerline 

rights-of-way in Brunswick County, Pleea visually dominates areas up to hundreds of hectares. In sterile condition, it is 

recognizable by its leathery equitant leaves, bright red at their bases. [= RAB, FNA, GW, K, S; = Tofieldia tenuifolia (Michaux) 

Utech – Z] 

Tofieldia Hudson 1778 (Bog Asphodel) 

A genus of about 7-8 species, of temperate to subarctic North America and Eurasia. There is controversy about the 

circumscription of Tofieldia.relative to the related genera Pleea and Triantha (here recognized, but sometimes subsumed into 

Tofieldia. Some believe that Tofieldia, Triantha, and Pleea should be treated together in a broadly circumscribed Tofieldia 

(Utech 1978, Zomlefer 1997c); others that all three should be treated separately (Ambrose 1980; Packer 1993; Cruden 1991). 

Packer in FNA (2002a) has recently recognized Triantha, Pleea, and Tofieldia as separate genera, a conclusion followed here in 

part because of the ancient, relictual nature of these units. References: Zomlefer (1997c)=Z; Packer (1993); Ambrose (1980); 

Utech (1978); Hitchcock (1944)=Y; Tamura in Kubitzki (1998a); Packer in FNA (2002a); Cruden (1991). 

Identification notes: In sterile condition, Tofieldia glabra can be distinguished from Iris verna by its minutely upwardlyscabrous 

margins (Iris has smooth margins). 

Tofieldia glabra Nuttall, Carolina Bog Asphodel, White Asphodel. Cp (NC, SC): savanna-pocosin ecotones, wet savannas, 

seepage bogs; rare (US Species of Concern, NC Rare, SC Rare). (Late August-) late September-October; October-November. 

Endemic to the coastal plain and sandhills of NC and northern SC (the reports from GA are dubious). [= RAB, FNA, GW, K, S, 

Z] 

Triantha (Nuttall) Baker

TOFIELDIACEAE 940 

A genus of ca. 4 species, herbs, of North America and Japan. References: Cruden (1991). 

1 Perianth equal to or longer than the capsule; seeds with tails 1/2 or less as long as the body................................Tr. racemosa 

1 Perianth shorter than the capsule; seeds with at least 1 tail equal to or longer than the body ................................ Tr. glutinosa 

Triantha glutinosa (Michaux) Baker, Sticky Bog Asphodel. Mt (GA, NC, VA), Pd (NC): bogs and seeps, especially over 

mafic or calcareous rocks; rare (GA Rare, NC Rare, VA Rare). July-August; September-October. Newfoundland west to British 

Columbia, south to w. NC, ne. GA (Jones & Coile 1988), WV, OH, n. IN, WI, MT, and OR. [= FNA, K, S; = Tofieldia 

racemosa var. glutinosa (Michaux) Ahles – RAB; = Tofieldia glutinosa (Michaux) Persoon – F, G, W; > Tofieldia glutinosa ssp. 

glutinosa – GW, Y, Z; > Tofieldia glutinosa var. glutinosa – C] 

Triantha racemosa (Walter) Small, Coastal Plain Bog Asphodel. Cp (GA, NC, SC, VA), Mt (VA), Pd (GA): savannas, 

savanna-pocosin ecotones, seepage bogs, sinkhole ponds (dolines) in the mountains of VA; uncommon (VA Rare). June-early 

August; late September-October. NJ south to nw. FL, west to e. TX; disjunct in c. TN. [= FNA, K, S; = Tofieldia racemosa var. 

racemosa – RAB; = Tofieldia racemosa (Walter) Britton, Sterns, & Poggenburg – C, F, G, GW, W, Z] 

References: Farmer & Schilling (2002). 

TRILLIACEAE Lindley 1846 (Trillium Family) 

Trillium Linnaeus 1753 (Trillium, Toadshade, Wake-robin) 

A genus of about 50 species, of e. North America, w. North America, and e. Asia (especially se. North America). The genus 

Trillium in our area is difficult and complex. Trillium is now usually separated from the Liliaceae (along with Eurasian genera 

such as Paris) into the Trilliaceae (Zomlefer 1996, Kato et al. 1995, Kawano & Kato 1995, and others) or less drastically as part 

of the Melanthiaceae (Chase et al. 2000; Tamura et al. 2004). The traditonal division of the genus into two well-marked 

subgenera, subgenus Trillium, the pedicellate trilliums, and subgenus Phyllantherum, the sessile-flowered trilliums, has been 

partly supported by molecular and morphological phylogenetic studies (Kawano & Kato 1995, Kato et al. 1995). These studies 

support the monophyly of subgenus Phyllantherum, but suggest that subgenus Trillium consists of several groups which are only 

rather distantly related (Kawano & Kato 1995, Kazempour Osaloo et al. 1999; Farmer & Schilling 2002). Most species are slowgrowing 

perennials; seedlings, juveniles, and depauperate or "tired" plants are one-leaved ("monilliums"), recognizable by the 

similar color, texture and venation of the single leaf to the three leaves of mature plants. In some species, such as T. pusillum, 

individual plants remain in the single-leaf stage for long periods of time, and populations may consist largely of juvenile plants. 

References: Patrick (1986)=Z; Freeman (1975)=Y; Case & Case 1997=X; Patrick in Wofford (1989); Case in FNA (2002a); 

Mitchell (1990); Kato et al. (1995); Kawano & Kato (1995); Tamura in Kubitzki (1998a); Zomlefer (1996); Farmer & Schilling 

(2002). Key adapted from Z, unpublished keys of J.D. Freeman, and other sources. 

Identification notes: Teratological forms are frequent in some species, as, for instance, leaves, sepals, and stamens in 2's or 4's, 

petals sepaloid, or sepals petaloid, and so forth. What are called “leaves” in Trillium are actually bracts. 

1 Leaves mottled with 2-3 different shades of green (very rarely the mottling not apparent); flower sessile; [subgenus 

Phyllantherum] .................................................................................................................................................................Key A 

1 Leaves solid green; flower on a pedicel (the pedicel sometimes very short or essentially absent in some varieties of T. 

pusillum); [subgenus Trillium]. 

2 Petals relatively thick in texture, straight-margined, maroon or white, rarely yellow or green (if white, turning brown 

with age); stigmas thicker at base, tapering gradually toward tip, distinct; ovary purple-black, maroon, pink, or white, 

6-angled; [Erectum group] ........................................................................................................................................Key B 

2 Petals relatively delicate in texture, wavy-margined, white to deep pink (if white, generally fading to pink with age); 

stigmas thin, uniform in thickness from base to apex, somewhat fused at the base into a short style; ovary greenishwhite 

to white, 3- or 6-angled or-lobed.....................................................................................................................Key C 

Key A – trilliums with sessile flowers and mottled leaves (subgenus Phyllantherum) 

1 Scape more-or-less decumbent in a gentle S-shape, the leaves lying on the ground, or nearly so; flower fragrance putrid; [T. 

sessile group]. 

2 Anther dehiscence extrorse (toward the outside of the flower); stamens about 0.25× as long as the petals; upper stem 

puberulent; petals 4-10 cm long; filaments 2-5 mm long..............................................................................T. decumbens 

2 Anther dehiscence introrse (toward the inside of the flower); stamens about 0.5× as long as the petals; upper stem 

glabrous; petals 2.5-5.5 cm long; filaments 1-2 mm long ................................................................................ T. reliquum 

1 Scape erect, straight, the leaves bone well above the ground (the leaf tips sometimes nearly touching the ground); flower 

fragrance various.

TRILLIACEAE 941 

3 Sepals abruptly deflexed between and below the leaves, distinctly descending below the approximately horizontal plane 

of the leaves; filaments about as long as incurved anthers; [T. recurvatum group]. 

4 Leaves sessile or subsessile, borne in a descending or drooping manner (similar to the sepals); petals usually > 4× 

as long as wide..................................................................................................................................... T. lancifolium 

4 Leaves distinctly petiolate, borne in an ascending manner (strongly contrasting in position with the strongly 

deflexed sepals); petals usually ca. 2× as long as wide ........................................................................ T. recurvatum 

3 Sepals erect, ascending, or spreading, usually borne at or above the approximately horizontal plane of the leaves; 

filaments much shorter than the upright anthers. 

5 Petals spreading to horizontal, with 1-2 spiral twists (looking something like an airplane propellor); anther 

dehiscence extrorse (toward the outside of the flower); [T. sessile group]......................................... [T. stamineum] 

5 Petals erect to slightly spreading, not spirally twisted; anther dehiscence introrse (toward the inside of the flower), 

or latrorse (toward the side). 

6 Petals broadly spatulate, clawed, broadly rounded (though sometimes with an apiculus) at the tip; petals pale 

lemon-yellow (the claws greenish or maroon); flower fragrance clove-like; [of the Savannah River drainage, 

from sw. NC southeastward along the GA-SC border]; [T. sessile group]......................................... T. discolor 

6 Petals lanceolate, elliptic, obovate, or oblanceolate, but not broadly spatulate and distinctly clawed, generally 

acute at the tip; petals maroon-red, purplish-brown, yellow, or green; flower fragrance various; [collectively 

widespread in our area]. 

7 Stigmas > 1.5× as long as the ovary; stamens about 0.5× as long as the petals; anther connectives 

prominently prolonged into a beak 1.0-5.0 mm long (beyond the anther sacs); [T. sessile group].............. 

......................................................................................................................................................T. sessile 

7 Stigmas as long as the ovary or shorter; stamens < 0.5× as long as the petals; anthers blunt, the 

connectives extended < 1.0 mm beyond the anther sacs. 

8 Ovary ellipsoid; leaves acute, the margins of the outer 1/3 more or less straight; leaf blade mottled 

with 3 or more shades of green, the palest shade forming a very conspicuous pale green streak 

along the midvein; [of the Coastal Plain and fall-line area of GA, AL, and FL panhandle]; [T. 

sessile group]. 

9 Stem 2.5-3× as long as the leaves; petals oblanceolate-obovate, usually 1.5-3× as long as wide 

..................................................................................................................................T. decipiens 

9 Stem 1-2× as long as the leaves; petals narrowly elliptic to oblanceolate-obovate, usually 3-5× 

as long as wide ....................................................................................................T. underwoodii 

8 Ovary ovoid; leaves acute to acuminate, the margins of the outer 1/3 convex; leaf blade mottled 

with 2-3 shades of green, paler shades sometimes prominent along the midvein, but not as above; 

[collectively widespread in our area]; [T. maculatum group]. 

10 Petals < 4× as long as wide, elliptic-oblanceolate to oblanceolate; [of inland provinces, rarely 

in the Coastal Plain]. 

11 Flower fragrance fruity-spicy, like green apples or Calycanthus (rarely musky); petals 

maroon, bronze, green, yellow; portions of ovary and stamens purplish during anthesis.... 

.........................................................................................................................T. cuneatum 

11 Flower fragrance lemon-like; petals greenish-yellow darkening to yellow; ovary and 

stamens greenish-white during anthesis................................................................T. luteum 

10 Petals > 4.5× as long as wide, narrowly oblanceolate-spatulate to linera oblnaceolate; [of the 

Coastal Plain, rarely further inland]. 

12 Ovary 3-angled at base of stigmas (rarely hexagonal); petals 7-17 mm wide, narrowly 

spatulate (appearing clawed); outer whorl of stamens broader, anther dehiscence introrse; 

flower fragrance faintly spicy-fragrant, banana-like; [of AL, n. FL, GA, and e. SC] .......... 

.......................................................................................................................T. maculatum 

12 Ovary 6-angled; petals 3-8 mm wide, linear-oblanceolate, narrowly elliptic, to linearlanceolate 

(weakly or not clawed); flower fragrance putrid, like rotting meat; [of MS and 

LA]. 

13 Petals 3-5 mm wide; anther dehiscence introrse; anther connective extending 1-1.5 

mm beyond the anther sacs.............................................................[T. foetidissimum] 

13 Petals 4-8 mm wide; anther dehiscence latrorse; anther connective scarcely 

extending beyond the anther sacs ...................................................[T. ludovicianum] 

Key B – trilliums with unmottled leaves and pedicellate flowers, 

of the Erectum Group 

1 Pedicel abruptly declined below the leaves; leaves petiolate to subsessile (or even sessile); petals recurved between the 

sepals. 

2 Stamens far exceeding the pistil, filaments as long as the ovary or longer, at least partly maroon, the anther sacs yellow 

to maroon; ovary small, globose, 3-12 mm long; flower fragrance pungent, rose-like; pedicel long, 3-13 cm long; petals 

strongly overlapping, usually maroon (rarely white or whitish) ........................................................................... T. vaseyi


2 Stamens at most 1.5× longer than the pistil, filaments shorter than the ovary, white, the anther sacs lavender to vivid 

purple (or albino); ovary large, ovoid, 10-17 mm long; flower fragrance weak, like green apples; pedicel short, 1.5-3 

cm long; petals not strongly overlapping, usually white (rarely maroon). 

3 Anthers 7.5 mm long or less, about as long as the filaments or shorter; petals narrowly elliptic to obovate, often 

scarcely larger than the sepals, delicate, occasionally margined in pink or green; [of damp forests south to n. VA] . 

................................................................................................................................................................. T. cernuum 

3 Anthers 7.0 mm long or more, longer than the filaments; petals ovate to elliptic, much broader than the sepals; [of 

mesic forests north to n. NC] ....................................................................................................................... T. rugelii 

1 Pedicel inclined, erect or declined (the flower generally borne above the leaves); leaves sessile to subsessile; petals variously 

disposed, generally spreading in the same plane as the sepals or forming a cup concealing the ovary in side view (then 

recurved toward the apex). 

4 Anthers creamy-white, 2-5× as long as the filaments; ovary white to pink (rarely darker), ovoid (widest near the base); 

stigmas prominent, nearly as long as the ovary; flower fragrance weakly sweet to musty ................................ T. flexipes 

4 Anthers creamy-white, yellowish, or purplish, at most 2.2× as long as the filaments; ovary purple-black to maroon (or 

albino), subglobose; stigmas smaller, < 0.3× as long as the ovary; flower fragrance variable (see below). 

5 Petals lanceolate to narrowly ovate or elliptic, spreading from base in the same plane as the sepals, rarely > 2× as 

broad as the sepals; sepals 0.5-0.8× as long as the pedicel. weakly sulcate-tipped (keeled and upturned near apex); 

flower fragrance unpleasant, musty............................................................................................................T. erectum 

5 Petals ovate, overlapping in some instances and forming a cup-shaped base, variably recurved apically, > 2× as 

broad as the sepals; sepals < 0.5 as long as the pedicel, sulcate-tipped; fragrance pleasant, sweet to fungal. 

6 Sepals 0.4-0.7× as long as the pedicel; leaves broadly elliptic; stamens 1.2-1.8× pistil height; flowers 

generally large, petals much longer than the sepals; sepals green; petals usually white (rarely maroon); flower 

fragrance sweet, like green apples; [of sw. NC and nw. SC in our area]............................................... T. simile 

6 Sepals 0.2-0.4× as long as the pedicel; leaves broadly obovate; stamens 0.9-1.6× pistil height; flowers 

relatively small, petals only slightly longer than the sepals; sepals suffused with purple; petals usually maroon 

(rarely white); flower fragrance fungal, like fresh mushrooms; [of sw. VA and nw. NC in our area] ................ 

......................................................................................................................................................... T. sulcatum 

Key C – trilliums with unmottled leaves and pedicellate flowers, of various affinities 

1 Petals white with triangular red blaze (rarely entirely white or pinkish); anther sacs lavender to white, dehiscence extrorse; 

fruit a red berry; leaves long-acuminate; [of acidic sites in the Mountains, generally strongly associated with either Pinus, 

Tsuga, Picea, Rhododendron, or other heaths] .....................................................................................................T. undulatum 

1 Petals white to deep pink, lacking a red blaze; anther sacs yellow, dehiscence introrse; fruit a white to greenish-white, 

fleshy, irregularly dehiscent capsule; leaves obtuse to acute (or somewhat acuminate in T. grandiflorum); [of less distinctly 

acidic sites, collectively widespread in our area]. 

2 Pedicel declined below the leaves (rarely erect); sepals arcuate-recurved; anthers irregularly twisted outward; pollen 

egg-yolk yellow ............................................................................................................................................... T. catesbaei 

2 Pedicel inclined above leaves to strictly erect; sepals not arcuate-recurved; anthers erect, regular; pollen light yellow. 

3 Sepals about as broad as the petals or broader, obtuse; leaves obtuse; anthers purplish-green between anther sacs; 

pedicel erect through fruiting. 

4 Pedicel 1-4 mm long ............................................................................................ T. pusillum var. virginianum 

4 Pedicel 5-30 mm long ................................................................................................ T. pusillum var. pusillum 

3 Sepals narrower than the petals, acute; anthers white to greenish-white between the anther sacs; leaves obtuse, 

acute, or acuminate; pedicel somewhat angled from the vertical. 

5 Ovary obscurely 3-lobed; leaves < 5 cm long, blue-green, obtuse ........................................................ T. nivale 

5 Ovary sharply 6-angled (-winged); leaves > 5 cm long, green, acute to acuminate. 

6 Petals obovate, tightly rolled at base, abruptly flared near the apex; leaves broadly elliptic, acuminate; 

style minute, < 1.0 mm long.............................................................................................. T. grandiflorum 

6 Petals elliptic, loose, gradually separating; leaves ovate, acute; style conspicuous, > 1.5 mm long............ 

................................................................................................................................................T. persistens 

Trillium catesbaei Elliott, Catesby's Trillium, Bashful Trillium, Rosy Wake-robin. Pd (GA, NC, SC), Mt (GA, NC, SC), Cp 

(GA, NC): bottomland forests, mesic slopes, cove forests; common (uncommon in Mountains). Late March-early June; July- 

August. Nc. NC south to sw. GA and se. AL, north in the interior to n. AL and se. TN, centered in the Piedmont from NC to GA, 

but extending into the Mountains and Coastal Plain. Petals white to pink. [= RAB, FNA, K, S, W, X, Z] 

Trillium cernuum Linnaeus, Northern Nodding Trillium. Mt (VA): damp forest with Fraxinus nigra and Ulmus 

americana; rare (VA Rare). Late April-May. Newfoundland, Hudson Bay area, and se. Saskatchewan south to n. VA, ne. WV, 

n. IN, n. IL, n. IA, and SD. Petals white, pink, maroon, or green. [= FNA, K, W, X, Z; < T. cernuum – RAB, F, S (apparently 

also including T. rugelii); > T. cernuum var. cernuum – C, G; >< T. cernuum var. macranthum A.J. Eames & Wiegand – C, G] 

Trillium cuneatum Rafinesque, Sweet Betsy, Purple Toadshade, Large Toadshade, Wedge-petal Trillium, Bloody Butcher. 

Pd, Mt (GA, NC, SC), Cp (GA): in rich soils of cove forests, moist slopes, and bottomlands, usually over mafic or calcareous 

rocks; uncommon, but locally abundant. Mid March-April; late May-June. Centered in the Southern Appalachians (but is more


abundant in adjacent physiographic provinces), extending north to the Highland Rim of KY, west to the Interior Low Plateau of 

TN, south to the Coastal Plain of MS and AL, and east to the Piedmont of GA, SC, and NC. Petals maroon, yellow, green, or 

various intermediate shades. [= C, FNA, K, W, X, Y, Z; = T. cuneatum var. cuneatum – RAB; > T. cuneatum – F; >< T. viride 

Beck – F, misapplied with respect to NC material; < T. viride var. luteum (Muhlenberg) Gleason – G, misapplied (also see T. 

luteum); > T. hugeri – S; >< T. underwoodii – S, misapplied] 

Trillium decipiens J.D. Freeman, Chattahoochee Trillium, Deceptive Trillium. Cp (GA): moist forests; uncommon. Late 

January-early April. W. FL and sc. AL east to ec. GA, and might be expected in SC, near the Fall Line. It is similar to T. 

underwoodii. [= FNA, K, X, Y, Z] 

Trillium decumbens Harbison, Decumbent Trillium. Mt, Pd, Cp (GA): moist forests; uncommon (rare in Piedmont and 

Coastal Plain). Mid-March-April. Se. TN (Chester et al. 1993) south and west to nw. GA and nc. AL, and disjunct in Houston 

County, in central GA Coastal Plain; it should be sought in extreme sw. NC, an extremely "under-botanized" area. [= FNA, K, S, 

X, Y, Z] 

Trillium discolor Wray ex Hooker, Pale Yellow Trillium, Pale Trillium, Small Yellow Toadshade. Mt (GA, NC, SC), Pd 

(GA, SC): rich cove forests, restricted to the Savannah River drainage; rare (GA Special Concern, NC Threatened, SC Rare). 

Late March-early May; June-July. Endemic to the Savannah River drainage of nw. SC, ne. GA, and sw. NC, occurring in the 

Blue Ridge and Piedmont. In NC it is restricted to a few sites along the Whitewater and Thompson Rivers. Petals pale yellow, 

with maroon or greenish claws. [= RAB, FNA, K, S, W, X, Y, Z] 

Trillium erectum Linnaeus, Red Trillium, Purple Trillium, Stinking Willie, Stinking Benjamin, Wake-robin. Mt (GA, NC, 

SC, VA): wooded slopes, usually at middle to high elevations; common. April-early June; July-August. New Brunswick, 

Québec, and MI south to w. NC, nw. SC, n. GA, e. TN, IN, and se. WI. Petals maroon, white, yellow, green, or various 

intermediate shades. [= C, K, W, X, Z; < T. erectum var. erectum – RAB (also see T. sulcatum); < T. erectum – F, G, S (also see 

T. sulcatum); > T. erectum var. erectum – FNA; > T. erectum var. album (Michaux) Pursh – FNA] 

Trillium flexipes Rafinesque, Bent White Trillium. Mt (GA, NC, VA): moist coves over mafic or calcareous rocks; rare 

(GA Special Concern, NC Rare, VA Rare). April. E. PA, s. Ontario and s. MN south to w. NC, nw. GA, n. AL, n. MS, mostly 

west of the Blue Ridge, but scattered in the Blue Ridge of NC, and disjunct east of the Blue Ridge in DE, PA, and MD. Petals 

white or maroon. [= C, F, FNA, K, W, X, Z; < T. erectum var. vaseyi – RAB; = T. gleasoni Fernald – G; = T. declinatum (A. 

Gray) Gleason – S, misapplied] 

Trillium grandiflorum (Michaux) Salisbury, Large-flowered Trillium, White Trillium, Great White Trillium. Mt (GA, NC, 

SC, VA), Pd (NC, SC, VA): rich coves and mesic slopes, also less typically on ridges over "rich" rock types; common (SC 

Rare). April-May; July-August. S. Québec, s. Ontario, MI, and MN, south to NJ, c. NC, nw. SC, n. GA, n. AL, s. IL, and IA. 

Petals white to pink. [= RAB, C, F, FNA, G, K, S, W, X, Z] 

Trillium lancifolium Rafinesque, Lanceleaf Trillium, Narrowleaf Trillium. Pd (SC), Mt (GA), Cp (GA): rich forests over 

marble, limestone, and other calcareous substrates, floodplain forests; rare (GA Special Concern, SC Rare). Late March-April. 

Nc. SC and se. TN south through w. GA and AL to panhandle FL and se. AL. Petals purple, green, or greenish-purple. [= FNA, 

K, Y, X, Z; = T. lanceolatum (S. Watson) Boykin ex Small – RAB, S] 

Trillium luteum (Muhlenberg) Harbison, Yellow Trillium, Yellow Toadshade, Wax Trillium, Lemon-scented Trillium. Mt 

(GA, NC, VA*): moist coves over mafic or calcareous rocks, restricted to the vicinity of the Great Smokies; uncommon (but 

locally abundant). Mid March-April; late May-June. Nearly endemic to the Southern Appalachians: w. NC, e. TN, nw. GA, and 

se. KY, allegedly disjunct in c. AL (planted and naturalized in Frederick County, VA). Petals yellow. [= C, F, FNA, K, W, X, Y, 

Z; = T. cuneatum var. luteum (Muhlenberg) Ahles – RAB; < T. viride Beck var. luteum (Muhlenberg) Gleason – G (also see T. 

cuneatum); < T. sessile – S, misapplied] 

Trillium maculatum Rafinesque, Mottled Trillium, Spotted Trillium. Cp, Pd (GA, SC): rich forests and floodplains, over 

calcareous materials such as coquina limestone ("marl") or on shell middens; uncommon. Early February-mid April. S. SC 

south to n. FL, west to sc. AL. Petals maroon or yellow. [= FNA, K, X, Y, Z; < T. viride – RAB, misapplied; < T. sessile – S, 

misapplied] 

Trillium nivale Riddell, Snow Trillium, Dwarf White Trillium. Mt (VA): rocky, calcareous forests; rare (VA Rare). Early 

March-early April. MA, sw. PA, MI, WI, s. MN, and e. SD south to n. VA, KY, s. IN, s. IL, s. MO, and se. NE. Petals white to 

pink. [= C, F, FNA, G, K, X, Z] 

Trillium persistens Duncan, Persistent Trillium. Mt (GA, SC): acidic forests with hemlocks and heaths; rare (US 

Endangered, GA Endangered, SC Rare). April. Endemic to a short stretch of the Tallulah-Tugaloo river system in nw. SC and 

ne. GA. Petals white to pink. [= FNA, K, W, X, Z] 

Trillium pusillum Michaux var. monticulum Bodkin & Reveal. Mt (VA): dry to dry-mesic forests and woodlands, moist 

forests along small mountain streams; rare. Endemic to nw. VA, e. WV, w. MD and disjunctly in sw. VA (at The Glades, 

Grayson County). NC. Var. monticulum Bodkin & Reveal has been controversial; see Cabe (1995) and Cabe & Werth (1995) 

for additional discussion of variation within T. pusillum in Virginia and elsewhere. Petals white to pink. [< T. pusillum – Z; < T. 

pusillum var. virginianum – C, K; < T. pusillum var. pusillum – FNA; = T. pusillum var. monticola Bodkin & Reveal – X, 

orthographic error] {not keyed at this time} 

Trillium pusillum Michaux var. ozarkanum (Palmer & Steyermark) Steyermark. Mt (NC): dry to dry-mesic slopes, in NC 

under Quercus coccinea and Kalmia latifolia; rare (NC Endangered). KY, TN, w. NC, AR, MO, and s. MS. Petals white to 

pink. [= K, X; < T. pusillum – RAB, G, S, Z; < T. pusillum var. pusillum – FNA; = T. ozarkanum Palmer & Steyermark] {not 

keyed at this time} 

Trillium pusillum var. 2. Mt (GA): {habitat}; rare. Endemic to n. GA. Apparently most closely related to Trillium 

texanum. Petals white to pink. {not keyed at this time}


Trillium pusillum Michaux var. pusillum, Carolina Least Trillium, Carolina Dwarf Trillium. Cp (NC, SC), Mt (GA, NC): 

bottomland forests along small streams in the upper Coastal Plain, ecotones of calcareous savannas and swamp forests in the 

lower Coastal Plain; rare (US Species of Concern, GA Special Concern, NC Endangered, SC Rare). Late March-May; June-July. 

Endemic to the outer Coastal Plain of e. NC and e. SC. T. pusillum is somewhat reminiscent of a tiny T. grandiflorum. The 

species as a whole has a highly disjunct and fragmented range, involving most of the Southeastern states. In addition to the taxa 

treated here, the complex includes T. texanum Buckley (of e. TX). The Trillium pusillum complex is currently undergoing study 

by Susan Farmer (Univ. of Tennessee); preliminary analysis shows that the published varieties are "good" and that the 

recognition of additional taxa may be warranted. Petals white to pink. [= X; < T. pusillum var. pusillum – C, F, FNA, K; < T. 

pusillum – RAB, G, S, Z; = T. pusillum (sensu stricto)] 

Trillium pusillum Michaux var. virginianum Fernald, Virginia Least Trillium, Virginia Dwarf Trillium. Cp (NC, VA), Mt 

(VA): bottomland forests along small streams in the upper Coastal Plain, swamps and bottomland forests, also mesic beech 

islands in swamp forests, moist mafic areas in Grayson Co. VA; rare (US Species of Concern, NC Endangered, VA Rare). Late 

March-May; June-July. Var. virginianum occurs in the Coastal Plain of se. VA and ne. NC; disjunct in Grayson County in the 

Mountains of sw. VA. The Grayson County site is within a kilometer of the Alleghany County, NC border, and the plant may be 

found to occur in nw. NC. Petals white to pink. [= F, FNA, X ; < T. pusillum – RAB, G, S, Z; < T. pusillum var. virginianum – 

C, K (also see var. monticulum); = T. virginianum (Fernald) C.F. Reed] 

Trillium recurvatum Beck, Prairie Trillium, Prairie Wake-robin. Mt, Pd (NC): rich soils of cove over calcareous rock; rare 

(NC Rare). W. OH west to s. MI, s. WI, and e. IA, south to c. TN, c. AL, c. MS, n. LA, and e. TX; disjunct in the Cumberland 

Plateau of e. TN, e. KY, and the Blue Ridge and w. Piedmont of NC. The two known NC occurrences (Catawba and Madison 

counties) appear to be native. Petals maroon or yellow. [= C, F, FNA, G, K, S, X, Y, Z] 

Trillium reliquum J.D. Freeman, Relict Trillium. Pd (GA, SC), Cp (GA): rich forests on bluffs and ravine slopes; rare (US 

Endangered, GA Endangered, SC Rare). Mid March-late April. Known from two disjunct areas, along the Savannah River in 

the vicinity of Augusta, on the border of SC (Aiken County) and GA (Richmond counties), and along the Chattahootchee River 

in sw. GA (Clay and Early counties). [= FNA, K, X, Y, Z] 

Trillium rugelii Rendle, Southern Nodding Trillium. Mt, Pd (GA, NC, SC): rich woodlands and forests over mafic or 

calcareous rocks; rare (NC Watch List, SC Rare). April-early May. W. NC and e. TN south to c. GA, and c. AL. Petals white or 

maroon. [= FNA, K, W, X, Z; < T. cernuum – RAB, F, S; < T. cernuum var. macranthum A.J. Eames & Wiegand – C, G] 

Trillium sessile Linnaeus, Sessile Trillium, Sessile Toadshade, Toad Trillium. Mt (VA), Pd (VA), Cp (NC, VA): rich 

forests, in NC limited to very rich soils of natural levees and lower slopes along the Roanoke River; uncommon in VA, rare in 

NC (NC Rare). March-April. Primarily a species of the northern Midwest, T. sessile ranges from MD, w. PA, w. NY, s. MI, n. 

IL and n. MO, south to e. VA, ne. NC, c. TN, n. AL, and n. AR. The easternmost occurrences are disjunct populations east of the 

Blue Ridge, in MD, VA, and along the Roanoke River in ne. NC. Petals maroon or green. [= C, F, FNA, G, K, W, X, Y, Z] 

Trillium simile Gleason, Sweet White Trillium. Mt (GA, NC, SC): very rich soils of slopes and coves over mafic or 

calcareous rocks, often also in or near seepage; rare (GA Special Concern, NC Rare, SC Rare). Late March-early May; June- 

July. A Southern Appalachian endemic: Blue Ridge of w. NC, nw. SC, e. TN, and n. GA. Petals white or very rarely maroon. 

[= FNA, K, S, W, X, Z; < T. erectum var. vaseyi – RAB] 

Trillium sulcatum T. Patrick, Southern Red Trillium, Barksdale Trillium. Mt (GA, NC, VA): coves and moist slopes; 

uncommon (GA Special Concern). April-May. Primarily a species of the sedimentary rock Appalachians, T. sulcatum ranges 

from s. WV, sw. VA, and e. KY south to nw. NC (where it enters the Blue Ridge), w. TN, nw. GA, and ne. AL. This species 

seems quite distinctive for its small, generally maroon flowers (with strongly sulcate sepals purplish as well), borne on very long 

pedicels. Petals maroon or white. [= C, FNA, K, W, X, Z; < T. erectum var. erectum – RAB; < T. erectum – F, G, S] 

Trillium underwoodii Small, Underwood's Trillium. Cp (GA): moist forests; uncommon. Late February-mid April. N. FL 

north to wc. GA and c. and s. AL; it is the only erect trillium with the stems < 2× as long as the leaves. [= FNA, K, S, X, Y, Z] 

Trillium undulatum Willdenow, Painted Trillium, Striped Wake-robin. Mt (GA, NC, SC, VA): acidic soils of ridges, 

slopes, and bog margins, mostly at high elevations and often associated with Rhododendron, Tsuga, Pinus, or Picea; common 

(SC Rare). Late April-May; late July-August. New Brunswick, e. Québec, s. Ontario, and MI, south to w. NC, nw. SC, n. GA, e. 

TN, and ne. OH. Of all our species, this is the species best adapted to acidic soils. Petals white with a red blaze. [= RAB, C, F, 

FNA, G, K, S, W, X, Z] 

Trillium vaseyi Harbison, Sweet Trillium, Vasey Trillium, Sweet Beth. Mt (GA, NC, SC), Pd (GA): cove forests; 

uncommon. Late April-early June. This species is a Southern Appalachian endemic: w. NC and e. TN south to nw. SC, n. GA, 

and ne. AL. Perhaps the largest trillium species, with the stems to 7 dm tall. Petals maroon or white. [= FNA, K, S, W, X, Z; < 

T. erectum var. vaseyi (Harbison) Ahles – RAB (also see T. simile and T. flexipes)] 

Trillium foetidissimum J.D. Freeman, Stinking Wake-robin. Bluffs, ravines, bottomlands. Late February-early April. MS 

west to LA. [= FNA, K, X, Y, Z] 

Trillium ludovicianum Harbison, Louisiana Wake-robin. Floodplains, streambanks, ravine forests. Early March-April. MS 

west to LA. Reports of this species for AL are based on specimens of Trillium species 2. [= FNA, K, X, Y, Z] 

Trillium pusillum var. 1. AL, TN, KY. Petals white to pink. [< T. pusillum var. pusillum – C, F, FNA, K; < T. pusillum – 

G, S, Z; = T. pusillum var. alabamicum – X (nomen nudum)] {not keyed at this time} 

Trillium species 1. Pd (SC): rich forests; rare. Under study by L.L. Gaddy. Somewhat similar to T. lancifolium and T. 

recurvatum. With clawed, bicolored petals (the claw maroon and the blade green) and green sepals reflexed somewhat (in the 

same plane as the drooping leaves). [previously misidentified as T. lancifolium] {not keyed at this time} 

Trillium species 2. Mt (GA): rich forests; rare. Under study by Susan Farmer. {not keyed at this time}


Trillium species 3. Lookout Mountain area, reesembles T. ludovicianum. Under study by Susan Farmer. {not keyed at this 

time} 

Trillium stamineum Harbison, Twisted Trillium. Floodplains, slopes, especially over limestone. Late March-mid May. C. 

TN (Chester et al. 1993) south to c. AL, and e. MS. [= FNA, K, S, X, Y, Z] 

TYPHACEAE A.L. de Jussieu 1789 (Cattail Family) 

A family of 2 genera with 16-30 species, wetland herbs, cosmopolitan. References: Kaul in FNA (2000); Smith in FNA (2000); 

Thieret & Luken (1996); Kubitzki in Kubitzki (1998b). 

1 Inflorescences headlike, globular............................................................................................................................ Sparganium 

1 Inflorescences spikelike, cylindrical ................................................................................................................................. Typha 

Sparganium Linnaeus 1753 (Bur-reed) 

A genus of about 14 species, wetland and aquatic herbs, primarily circumboreal in arctic and temperate regions, but also in the 

tropics of Asia, and temperate Australia. References: Kaul in FNA (2000); Thieret (1982)=Z; Beal (1960)=Y; Crow & Hellquist 

(2000b)=X; Kubitzki in Kubitzki (1998b). 

1 Stigmas 2; fruits truncate at apex, obpyramidal, very abruptly beaked, 4-8 mm broad ...................................... S. eurycarpum 

1 Stigmas 1; fruits rounded or acuminate to a beak at the apex, elliptic, fusiform, or obovate, 1-3 (-4) mm broad. 

2 Pistillate heads (primarily those upward) supra-axillary (borne distinctly above the axils of the subtending leaf-like 

bracts); tepals lacking subapical dark spot ....................................................................................................... S. emersum 

2 Pistillate heads (all) axillary (borne in the axils of the subtending leaf-like bracts) or several on axillary branches which 

lack leaf-like bracts; tepals with prominent subapical dark spot. 

3 Mature fruits dull, finely pitted, the body 3-5 mm long; fruiting heads 1.5-2.5 cm in diameter; branches of the 

inflorescence with (0-) 1-3 pistillate heads (in addition to staminate heads); stigma 0.8-1.9 (-2.8 in the Coastal 

Plain) mm long....................................................................................................................................S. americanum 

3 Mature fruits shiny, smooth, the body 5.5-7 mm long; fruiting heads 2.5-3.5 cm in diameter; branches of the 

inflorescence with 0 (-1) pistillate heads (in addition to staminate heads); stigma 1.5-3 mm long ............................. 

.......................................................................................................................................................... S. androcladum 

Sparganium americanum Nuttall, American Bur-reed. Cp, Mt, Pd (GA, NC, SC, VA): streams, marshes, ponds, pools, 

often submerged; common (rare in NC and SC Piedmont). May-September. Newfoundland west to MN, south to c. peninsular 

FL and c. TX. Beal (1960) discusses the interesting variation in S. americanum, perhaps worthy of taxonomic recognition. The 

"Appalachian Race" has stigmas 0.6-0.9 mm long, inflorescence branches 0-3, and relatively narrow leaves; in our area it is 

montane in distribution, and in general is Appalachian, Ozarkian, and northern. The "Coastal Race" has stigmas 1.5-2.8 mm 

long, 2-5 inflorescence branches, and relatively wide leaves; in our area it is primarily of the Coastal Plain, disjunct to the 

mountains of NC and SC south of the Asheville Basin (like many Coastal Plain taxa), and in general is nearly limited to the 

Coastal Plain, ranging from MA south to FL, west to e. TX, and north in the interior to sc. TN, s. IN, and s. MO. The 

"Ubiquitous Race" is intermediate, with stigmas 1.0-1.4 m long; it occurs throughout the range of the species. The pattern is 

suggestive of imperfect evolutionary separation of two taxa. [= RAB, C, F, FNA, G, GW, K, W, X, Y, Z; > S. americanum – S; 

> S. eurycarpum – S, misapplied] 

Sparganium androcladum (Engelmann) Morong. Cp, Pd (VA): marshes, shores; rare (VA Rare). May-September. ME 

and Québec west to MN, south to se. VA, e. TN, s. MO, and ne. OK. [= C, F, FNA, G, K, W, X, Y, Z] 

Sparganium emersum Rehmann, Greenfruit Bur-reed. Mt (NC, VA): bogs, stream margins; rare (NC Rare, VA Rare). 

May-September. Newfoundland and c. Québec west to s. Alberta and WA, south to w. NC, IN, IA, CO, and CA. [= FNA, X; > 

S. chlorocarpum Rydberg – RAB, C, F, G, W, Y, Z; > S. chlorocarpum var. acaule (Beeby) Fernald – F; ? S. angustifolium 

Michaux – K, misapplied; > S. acaule (Beeby) Rydberg] 

Sparganium eurycarpum Engelmann ex A. Gray, Giant Bur-reed. Cp, Pd, Mt (VA): marshes, shores; uncommon (VA 

Watch List). Nova Scotia west to British Columbia, south to w. VA, IN, OK, and CA. [= C, F, FNA, G, K, W, X; = S. erectum 

Linnaeus ssp. stoloniferum (Graebner) C.D.K. Cook & M.S. Nicholls] 

Sparganium angustifolium Michaux, Narrow-leaved Bur-reed. South to n. NJ and n. PA; attributed to VA and WV by 

Kartesz (1999), apparently erroneously. [= C, FNA, K, X] {not keyed; synonymy incomplete} 

Typha Linnaeus 1753 (Cattail) 

A genus of 8-13 species, wetland herbs, cosmopolitan. References: Smith in FNA (2000); Kubitzki in Kubitzki (1998b).

TYPHACEAE 946 

1 Staminate and pistillate portions of spike normally contiguous; pistillate portion of spike (1.5-) 2.0-3.5 cm in diameter at 

maturity; leaves (8-) 10-24 mm wide, flat on one side; stigmas lance-ovate, fleshy, persistent; pollen grains in 4's; [in acid to 

alkaline waters, widespread in our area] .................................................................................................................... T. latifolia 

1 Staminate and pistillate portions of spike normally separated by a gap; pistillate portion of spike 0.5-2.2 (-2.5) cm in 

diameter at maturity; leaves 4-15 mm wide, biconvex (or flat on one side in T. ×glauca); stigmas linear to lance-linear, not 

fleshy (or slightly so in T. ×glauca), either quickly deciduous or persistent; pollen grains single; [in circumneutral to alkaline 

waters, primarily in tidal situations in the outer Coastal Plain, but scattered inland, especially in VA]. 

2 Pith at base of stem yellowish buff; stigmas lance-linear, slightly fleshy; pistillate portion of spike 1.6-2.5 cm in 

diameter at maturity; pistillate bracteoles usually absent (or present on a few flowers) ....................................T. ×glauca 

2 Pith at base of stem white; stigmas linear, not fleshy; pistillate portion of spike 0.5-2.5 cm in diameter at maturity; 

pistillate bracteoles present on all flowers. 

3 Leaves 4-7 (-11) mm wide, auriculate at the junction of the blade and the sheath; pistillate portion of spike 0.5-2.0 

cm in diameter and dark brown at maturity; plants 1-1.5 m tall; pistillate bracteoles rounded to blunt at the tip ....... 

............................................................................................................................................................ T. angustifolia 

3 Leaves 6-12 (-15) mm wide, most or all not auricled at the junction of the blade and the sheath; pistillate portion 

of spike (0.8-) 1.3-2.5 cm in diameter and light cinnamon brown at maturity; plants (1-) 2-4 m tall; pistillate 

bracteoles acute to acuminate at the tip...............................................................................................T. domingensis 

Typha angustifolia Linnaeus, Narrowleaf Cattail. Cp (GA?, NC, SC, VA), Pd, Mt (VA): brackish to fresh waters of 

marshes and swamps, usually tidal; common (rare in Piedmont and Mountains). May-July; June-November. Nova Scotia west to 

ND, south to SC, FL (?), LA, and TX (?); Eurasia. [= C, F, FNA, G, GW, K, W; < T. angustifolia – S (also see T. domingensis)] 

Typha domingensis Persoon, Southern Cattail. Cp (GA, NC, SC, VA): brackish to nearly fresh waters of marshes and 

swamps, usually tidal; common. June-July; July-November. DE south to s. FL, west to TX; north inland to NE and UT; and 

south into tropical America; Eurasia; Africa; Oceania. [= C, F, FNA, G, GW, K; < T. angustifolia – S] 

Typha ×glauca Godron (pro sp.), Hybrid Cattail. Cp (GA?, NC, SC, VA), Mt (VA): fresh to brackish waters of lakes, 

ponds, and rivers; rare. May-July; June-November. Both C and K apply this name to two different hybrids: T. angustifolia × 

latifolia and T. domingensis × latifolia. The name properly applies to T. angustifolia × latifolia (Smith in FNA 2000). [= C, GW, 

K; = T. glauca Godron – RAB, F] 

Typha latifolia Linnaeus, Common Cattail. Mt, Pd, Cp (GA, NC, SC, VA): fresh waters of ponds, lakes, ditches, marshes, 

including in tidal freshwater marshes; common. May-July; June-November. Newfoundland west to AK, south to FL, TX, CA, 

and Mexico; Central America; South America; Eurasia. [= C, F, FNA, G, GW, K, S, W] 

UVULARIACEAE A. Gray ex Kunth 1843 (Bellwort Family) 

[see CALOCHORTACEAE (Prosartes, Streptopus, Tricyrtis), COLCHICACEAE (Uvularia), LILIACEAE (Clintonia)] 

XYRIDACEAE C. Agardh 1823 (Yellow-eyed Grass Family) 

A family of 5 genera and about 325-350 species, nearly cosmopolitan (most diverse in tropical and subtropical regions, and 

especially South America). References: Kral in FNA (2000); Kral in Kubitzki (1998b). 

Xyris Linnaeus 1753 (Yellow-eyed Grass) 

A genus of about 300 species, nearly cosmopolitan (most diverse in tropical and subtropical regions, and especially South 

America). This "technical" genus is known well by only a few botanists, and additional undescribed taxa are possible. 

References: Kral in FNA (2000); Bridges & Orzell (2003)=X; Kral (1966a)=Z; Kral (1983b, 1999); Kral in Kubitzki (1998b). 

Key adapted from X, GW, and Z. 

Identification notes: In vegetative condition, Xyris is often confused with other monocots with equitant leaves, such as Iris spp. 

(Iridaceae), Lachnanthes caroliniana (Haemodoraceae), and Tofieldia spp. (Tofieldiaceae). 

1 Keel of the lateral sepals shortly ciliate-scabrid (or sometimes entire in X. brevifolia, and then the bract tips purplish-tinged). 

2 Plants small, usually < 30 cm tall; principal leaves usually < 10 cm long; mature spikes < 1 cm long when mature. 

3 Leaves filiform, with expanded brownish lustrous bases, usually exceeding the sheath of the scape; [plants of s. 

AL and the FL Panhandle] ................................................................................................................... [X. isoetifolia] 

3 Leaves linear, the bases not expanded, shorter than, equaling, or slightly exceeding the sheath of the scape; [plants 

collectively more widespread]. 

4 Keel of the lateral sepals straight to slightly curved, remotely ciliate or entire; spikes broadly ovoid to 

subglobose, the bracts loose, bicolored, the outer portions maroon or purplish and often with erose margins ... 

........................................................................................................................................................ X. brevifolia 

4 Keel of the lateral sepals strongly curved, densely ciliate; spikes lance-ovoid to ovoid, the bracts entire, not 

purple-tinged, and lacking erose borders.

XYRIDACEAE 947 

5 Plants perennial; leaves ascending, green with a distinct brown patch at the base; fruiting spikes ovoid, 

blunt, somewhat 2-edged from the strongly keeled outer bracts .........................................X. drummondii 

5 Plants annual; leaves flabellate arranged, spreading to recurved against the substrate, usually maroon; 

fruiting spikes often elongated and acute, not 2-edged......................................................X. flabelliformis 

2 Plants large, usually > 30 cm tall; principal leaves > 10 cm long; mature spikes > 1 cm long when mature. 

6 Leaves ascending, twisted, strongly grooved; spikes ovoid, the bracts and lateral sepals with a small tuft of short, 

reddish-brown hairs; bases of leaves abruptly expanded, pinkish or purplish (dark brown in age), the outermost 

leaves often scale-like, the plant base therefore appearing bulbous; [of the Mountains, Piedmont, and Coastal 

Plain]............................................................................................................................................................... X. torta 

6 Leaves spreading, not twisted or only slightly so; spikes narrowly ovoid, ellipsoidal, or oblong; bracts and sepals 

without a small apical tuft of hairs; bases of leaves whitish, tan, pink, purplish, maroon, or dark brown, the 

outermost leaves not scale-like, the plant base not appearing bulbous; [typically of the Coastal Plain, rarely 

disjunct inland]. 

7 Seeds lustrous, translucent, broadly ovoid; spike pale brown or tan, the scales loosely imbricate; plant bases 

pinkish, purplish, or tan, with dark longitudinal striations on the inner leaf bases; leaves 3-20 mm wide; petal 

blades obovate, 6-7 mm long, opening in early morning, usually closing by mid-day..................... X. ambigua 

7 Seeds farinose, dark brown (X. stricta) or pale (X. louisianica) at maturity, narrowly ellipsoid to ovoid; spike 

dark brown, the scales tightly imbricate; plant bases maroon, purplish, dark-brown, or reddish-brown; leaves 

2-5 mm wide; petal blades triangular-cuneate, 3-5 mm long, opening at mid-day. 

8 Seeds pale when mature; plant bases maroon to maroon-brown, solitary or in small clumps; upper end of 

scape somewhat flattened, but not nearly as broad as the spike; spike narrowly ovoid to ellipsoid, 

slightly pointed......................................................................................................................X. louisianica 

8 Seeds dark brown when mature; plant bases dark maroon to dark brown, desnely cespitose; upper end of 

the scape conspicuously flattened, almost as broad as the spike; spike oblong-cylindrical, obtuse............. 

......................................................................................................................................................X. stricta 

1 Keel of the lateral sepals irregularly lacerate or fimbriate, or if entire then the the bract tips not purplish. 

9 Leaves narrowly linear to filiform, 0.5-2.0 (-2.5) mm wide, not twisted (or scarcely so); leaf bases expanded, lustrous, 

hard, tan to brown, neither bulbous nor deeply set in the substrate; spikes ovoid or ellipsoid, 4-15 mm long. 

10 Leaves filiform, terete or elliptic in cross-section, 0.5-1.0 mm wide, without a paler, hardened margin; scape as 

broad as or broader than the leaf blades; scales smooth-edged to denticulate, not curled away from the head, the 

head thus appearing smooth; staminodia beardless.............................................................................X. baldwiniana 

10 Leaves linear, flattened in cross-section, 1-2 (-2.5) mm wide, with a pale, hardened margin; scape usually 

narrower than the leaf blades; scales ragged-lacerate, the tips curling away from the head, giving it a ragged 

appearance; staminodia bearded.................................................................................................................. X. elliottii 

9 Leaves broader, (1.5-) 2.0-25 mm wide, strongly twisted to straight, the leaf bases either not expanded, lustrous, hard, 

and tan to brown, or, if so, then the base also either bulbous and/or deeply seated in the substrate; spikes narrowly 

lanceolate, ellipsoid, to broadly ovoid, 4-40 mm long. 

11 Keel of the lateral sepals long-fimbriate toward its apex, the fimbriate tip conspicuously exserted from the 

subtending bract (sometimes eroded and less conspicuous on older spikes). 

12 Leaves strongly twisted, 2-5 mm wide; leaf bases hardened, swollen, bulbous, dark lustrous brown; scape 

ridges smooth; petal blades white or yellow; [of moist to dry pinelands] ....................................X. caroliniana 

12 Leaves not twisted, 5-25 mm wide; leaf bases soft, not swollen, not bulbous, pale; scape ridges strongly 

scabrous; petal blades yellow; [of aquatic to very wet peaty, mucky, or sandy ponds, marshes, or other 

wetlands]..........................................................................................................................................X. fimbriata 

11 Keel of the lateral sepals lacerate, or if very shortly fimbriate, then not conspicuously exserted from the 

subtending bract. 

13 Lateral sepals longer than and exserted from the subtending bracts; scapes 5-15 dm tall. 

14 Leaf blades 1-2 (-3) mm wide, 6-30 cm long; spikes 10-16 mm long; seeds 0.4-0.6 mm long; [endemic 

to Panhandle FL and s. AL]............................................................................................... [X. longisepala] 

14 Leaf blades 5-15 mm wide; (20-) 30-50 (-60) cm long; spikes 10-20 (-25) mm long; seeds (0.6-) 0.7 (- 

0.8) mm long; [more widespread in our area] .........................................................................X. smalliana 

13 Lateral sepals shorter than the subtending bracts, and therefore hidden (except when the spikes open to shed 

seeds); scapes 1.5-12 dm tall. 

15 Scapes flexuous, usually spirally twisted; upper portion of leaf blades conspicuously twisted; plant bases 

pinkish, purplish, or dark brown, bulbous or deeply set in the substrate. 

16 Base of plant deeply set in the substrate, without distinct outer scale leaves; leaf bases not 

noticeably expanded, the plant base therefore not bulbous; leaves smooth, 2-4 mm wide; petal 

blades ca. 3 mm long..................................................................................................... X. chapmanii 

16 Base of plant shallowly set on the substrate, often with short, black outer scale leaves; leaf bases 

noticeably expanded, the plant base therefore appearing bulbous; leaves either smooth and 5-10 

mm wide, or scabrous and 2-10 mm wide; petal blades ca. 5 mm long. 

17 Leaf and scape surfaces prominently papillose or tuberculate-scabrid; petal blades 

suborbicular, yellow; seeds narrowly ovoid or narrowly ellipsoidal, ca. 1.0 mm long................ 

............................................................................................................................... X. scabrifolia


17 Leaf and scape surfaces smooth (or scabrous only along margins and ridges); petal blades 

obovate, white or yellow; seeds ovoid or ellipsoid, 0.5-0.6 mm long. 

18 Seeds translucent; leaf margins smooth; [plants of acidic sites of the Coastal Plain] .......... 

......................................................................................................................... X. platylepis 

18 Seeds opaque; leaf margins slightly scabrous; [plants of calcareous seeps and fens of the 

Ridge and Valley]......................................................................................X. tennesseensis 

15 Scapes usually not flexuous, usually not spirally twisted; upper portion of leaf blades not conspicuously 

twisted; plant bases variously colored, flabellate or equitant and set at ground level. 

19 Summit of the scape distinctly flattened and broad relative to the spike; scape ridges 2-3, the 2 most 

prominent comprising the flattened edge of the scape. 

20 The 2 principal scape ridges noticeably and abruptly flattened and winglike below the spike, 

their combined width (on fresh material) broader than the scape proper; fruiting spikes mostly 

8-15 mm long; seeds 0.4-0.6 mm long, translucent, ovoid or ellipsoidal, about 1.5× as long as 

wide, with lines of very fine papillae, not farinose...................................................X. difformis 

20 The 2 principal scape ridges not abruptly flattened and winglike below the spike, their 

combined width < the scape proper, which is itself flattened (narrowly elliptic in crosssection); 

fruiting spikes mostly (10-) 20-25 mm long; seeds 0.8-1.0 mm long, dark when ripe, 

fusiform to narrowly elliptic, 2-3× as long as wide, with lines of very fine papillae, these 

however obscured by a farinose covering................................................................. X. iridifolia 

19 Summit of the scape nearly terete or somewhat flattened, much narrower than the spike; scape 

ridges several (usually > 3), at least on the mid to lower portion of the scape. 

21 Seeds farinose, very dark; surfaces of leaves tuberculate-scabrid, the leaves strongly 

ascending, linear, generally > 10 cm long; leaves generally dull-colored. 

22 Mature spikes ovoid, sharply acute; plants solitary or in small clumps; leaves 10-30 (-50) 

cm long, 1.5-6.0 mm wide, dark maroon or purplish at the base ......................X. floridana 

22 Mature spikes ovoid to ellipsoid, acute to obtuse; plants typically in large dense tufts; 

leaves 20-50 cm long, 3-12 mm wide, the older ones with dark-brown to gray bases, the 

younger with tan bases........................................................................................X. serotina 

21 Seeds translucent, not farinose; surfaces of leaves smooth (or sparsely tuberculate-scabrid in 

X. curtissii, which also has leaves linear-curvate and generally < 10 cm long); leaves generally 

a bright yellowish-green above the base. 

23 Leaves broadly linear-curvate, spreading, typically < 10 cm long, 2-4.5 mm wide; plants 

perennial, in tufts (rarely solitary); mature spikes acute, with < 10 flowers; leaf bases 

pinkish or purplish; old flowers fugacious, not persisting on spikes...................X. curtissii 

23 Leaves linear, ascending, 10-60 cm long, 5-10 mm wide; plants annual, solitary or in 

small tufts; mature spikes obtuse, many-flowered; leaf bases tan (very rarely pinkish); old 

flowers often persistent on the spikes, drying blackish.........................................X. jupicai 

Xyris ambigua Beyrich ex Kunth. Cp (GA, NC, SC, VA), Pd (NC), Mt (VA): wet savannas and flatwoods, pinelands, 

edges of depression ponds; common. June-August. Se. VA south to FL, west to AL and ec. TX, primarily on the Coastal Plain. 

[= RAB, C, F, FNA, G, K, W, X; < X. ambigua – GW, S, Z (also see X. louisianica)] 

Xyris baldwiniana J.A. Schultes, Grassleaf Yellow-eyed Grass. Cp (GA, NC, SC), Pd (NC): wet savannas, seepage bogs, 

sandhill seeps, wet savanna ecotones; rare (NC Watch List). June-July. Se. NC south to n. FL, west to s. AR and ec. TX, 

primarily on the Coastal Plain. [= RAB, FNA, GW, K, S, X, Z] 

Xyris brevifolia Michaux, Shortleaf Yellow-eyed Grass. Cp (GA, NC, SC): wet sands of pinelands, especially seasonally 

wet, open, white sands of spodosol longleaf pine flatwoods (Leon series soils), margins of Carolina bay sandrims; rare (NC Rare, 

SC Rare). June-August. Se. NC south to s. FL, west to s. AL and w. FL; West Indies and South America. [= RAB, FNA, GW, 

K, S, X, Z] 

Xyris caroliniana Walter, Pineland Yellow-eyed Grass. Cp (GA, NC, SC, VA): dry to moist pine flatwoods, moist 

savannas, scrub oak sandhills; common (VA Rare). June-July. Se. VA south to FL, west to se. TX, and disjunct northward in s. 

NJ. White-petaled populations of X. caroliniana occurring in the East Gulf Coastal Plain need additional study. [= RAB, C, 

FNA, GW, K, X, Z; > X. flexuosa Muhlenberg ex Elliott – F, G, S; > X. pallescens (C. Mohr) Small – S] 

Xyris chapmanii Bridges & Orzell, Chapman's Yellow-eyed Grass. Cp (GA?, NC, SC): sandhill seepage bogs in areas of 

copious lateral seepage in deep muck soils; rare (NC Rare). Sc. NC south to panhandle FL, west to e. TX. This taxon is 

abundantly distinct from X. scabrifolia. [= X; < X. scabrifolia – FNA, K, Z] 

Xyris curtissii Malme, Curtiss's Yellow-eyed Grass. Cp (GA, NC, SC, VA), Pd (SC): savannas; rare (NC Watch List, VA 

Rare). July-August. Se. VA south to n. FL and west to s. AR and ec. TX, primarily on the Coastal Plain; disjunct in s. NJ. [= 

RAB, G; = X. difformis Chapman var. curtissii (Malme) Kral – C, FNA, GW, K, X, Z; > X. bayardii Fernald – F; > X. curtissii – 

F; ? X. neglecta Small – S] 

Xyris difformis Chapman. Cp (GA, NC, SC, VA), Pd (GA, NC, SC), Mt (NC): savannas, roadside ditches, pond margins, 

other wet habitats; common (VA Watch List). August-October. New England and s. Canada south to n. FL and ec. TX. [= X; = 

X. difformis Chapman var. difformis – C, FNA, GW, K, Y, Z; < X. difformis – RAB, F, G, S, W (also see X. curtissii)] 

Xyris drummondii Malme, Drummond's Yellow-eyed Grass. Cp (GA): wet pine flatwoods, ditches; uncommon. Se. GA 

south to Panhandle FL, west to s. MS. [= FNA, GW, K, X, Z]


Xyris elliottii Chapman, Elliott's Yellow-eyed Grass. Cp (GA, SC): margins of drawdown zones of clay-based Carolina 

bays, limesinks and flatwoods swales, wet savannas; uncommon. May-June. E. SC south to the Gulf Coastal Plain. [= RAB, 

FNA, GW, K, S, X, Z] 

Xyris fimbriata Elliott, Giant Yellow-eyed Grass. Cp (GA, NC, SC, VA): in mucky or sandy soils of upland depression 

ponds, also along sandhill streams, impoundments and in deep muck of sandhills seepage slopes often just below the zone 

occupied by Xyris chapmanii; common (VA Rare). September-October. Se. VA south to c. FL, west (interruptedly) to se. TX; 

disjunct in s. NJ, DE, and c. TN. [= RAB, C, F, FNA, G, GW, K, S, X, Z] 

Xyris flabelliformis Chapman, Savanna Yellow-eyed Grass. Cp (GA, NC, SC): wet sands of pinelands, especially 

seasonally wet, open, white sands of spodosol longleaf pine flatwoods (Leon series soils), margins of Carolina bay sandrims; rare 

(NC Rare, SC Rare). May-June. Se. NC south to s. FL, west to se. LA, on the Coastal Plain. [= RAB, FNA, GW, K, S, X, Z] 

Xyris floridana (Kral) Bridges & Orzell, Florida Yellow-eyed Grass. Cp (GA, NC, SC): savannas, wet pine flatwoods, 

ditches; rare (NC Rare, SC Rare). August. Se. NC south to s. FL, west to se. LA. [= X; = Xyris difformis Chapman var. 

floridana Kral – FNA, GW, K, Y, Z] 

Xyris iridifolia Chapman, Irisleaf Yellow-eyed Grass. Cp (GA, NC, SC, VA): marshes, upland pond margins, blackwater 

river channels, floodplain pools, other wet habitats; uncommon (NC Watch List, VA Rare). July-September. Se. VA south to n. 

FL, west to e. TX; disjunct in c. TN. [= RAB, C, GW, S, Z; = X. laxifolia Mart. var. iridifolia (Chapman) Kral – FNA, K, X] 

Xyris jupicai L.C. Richard. Cp (GA, NC, SC, VA), Pd (NC), Mt (VA): ditches, various wet habitats; common. July- 

September. NJ south to s. FL, west to TN, AR, OK, and TX, and in Latin America. Sometimes weedy and considered by some 

to be adventive from further south. At least some populations in our area are native and may additionally be worthy of 

taxonomic recognition as distinct from "true" X. jupicai (P. McMillan, pers. comm., 2003). [= RAB, C, FNA, GW, K, W, X, Z; 

= F. caroliniana – F, misapplied; > X. elata Chapman – G, S; > X. communis Kunth – S; > X. caroliniana – G, S, misapplied] 

Xyris louisianica Bridges & Orzell. Cp (GA): pine savannas, bogs, ditches and disturbed areas; rare. FL Panhandle and 

GA west to se. TX. [= K, X; = X. stricta Chapman var. obscura Kral – FNA; < X. ambigua – GW, S, Z] 

Xyris platylepis Chapman. Cp (GA, NC, SC, VA): sandhill seeps, savannas, ditches; common (VA Watch List). July- 

September. Se. VA south to s. FL, west to se. LA; disjunct in sw. LA and se. TX. [= RAB, C, F, FNA, G, GW, K, S, X, Z] 

Xyris scabrifolia Harper, Roughleaf Yellow-eyed Grass. Cp (GA, NC, SC): sandhill seepage bogs and wet pine savannas; 

rare (US Species of Concern, NC Rare). Sc. and se. NC south to panhandle FL, west to s. AL and s. MS; disjunct in sw. LA-se. 

TX. X. chapmanii is a taxon distinct from X. scabrifolia. [= GW, S, X; < X. scabrifolia – FNA, K, Z (also see X. chapmanii)] 

Xyris serotina Chapman. Cp (GA, NC, SC): depression meadows, ultisol savannas (Lynchburg/Rains complex or 

Eulonia/Oketee), ditches; rare (NC Rare, SC Rare). September. Se. NC south to c. FL, west to s. MS, in the Coastal Plain. 

Reported for our area by Kral (1966b) and relocated by B.A. Sorrie. [= RAB, FNA, GW, K, S, X, Z] 

Xyris smalliana Nash, Small's Yellow-eyed Grass. Cp (GA, NC, SC), Mt (SC): pond margins, ditches; uncommon (NC 

Watch List). July-August. ME south to FL, west to s. MS; disjunct to se. TX. [= RAB, C, FNA, GW, K, S, W, X, Z; > X. 

congdonii Small – F; > X. smalliana – F; > X. smalliana var. smalliana – G; > X. smalliana var. olneyi (Wood) Gleason – G] 

Xyris stricta Chapman. Cp (GA, NC, SC): depression ponds, depression meadows, borrow pits, ultisol savannas and 

ditches; rare. July-September. SC south to n. FL, west to s. MS and se. LA. Reported for our area by Kral (1966b). P. 

McMillan (pers. comm.) reports this species from a number of locations in the outer Coastal Plain of NC and SC. [= GW, K, S, 

X, Z; = X. stricta var. stricta – FNA] 

Xyris tennesseensis Kral, Tennessee Yellow-eyed Grass. Mt (GA): seepy, fenlike areas over limestone; rare (US 

Endangered, GA Endangered). TN, AL, and nw. GA (Jones & Coile 1988). See Kral (1978b). [= FNA, K] 

Xyris torta J.M. Smith, Mountain Yellow-eyed Grass. Mt, Pd (GA, NC, SC, VA), Cp (SC, VA): mountain bogs, marshes, 

ditches; uncommon (SC Rare). June-August. NH west to WI, south to e. VA, e. NC, w. SC, c. GA, LA, OK, and TX. This is 

our only species of Xyris not strongly associated with the Coastal Plain. [= RAB, C, FNA, GW, K, S, W, Z; > X. torta var. 

macropoda Fernald – F, G; > X. torta var. torta – F, G] 

Xyris isoetifolia Kral. Bogs, savannas, and depression pond margins. FL Panhandle and s. AL. [= FNA, GW, K, X, Z] 

Xyris longisepala Kral. Depression pond margins. FL Panhandle and s. AL. [= FNA, GW, K, X, Z] 

ZANNICHELLIACEAE Dumortier 1829 (Horned Pondweed Family) 

A family of 4 genera and about 10-12 species, aquatic herbs, nearly cosmopolitan. Probably better included in the 

Potamogetonaceae (Angiosperm Phylogeny Group 2003). References: Haynes & Hellquist in FNA (2000); Haynes & Holm- 

Nielsen (1987)=Z; Haynes, Les, & Holm-Nielsen in Kubitzki (1998b). 

Zannichellia Linnaeus 1753 (Horned Pondweed) 

A genus of about 5 species, aquatic herbs, nearly cosmopolitan. References: Haynes & Hellquist in FNA (2000); Haynes & 

Holm-Nielsen (1987)=Z. 

Identification notes: Zannichellia is sometimes confused with other aquatics, such as Ruppia and narrow-leaved Potamogeton. 

Potamogeton has at least some leaves alternate; Zannichellia and Ruppia are opposite-leaved. Zannichellia lacks the abruptly

ZANNICHELLIACEAE 950 

broadened sheath of Najas. Also, the seeds are flattened in Zannichellia, and toothed down one side; Najas has a cylindric or 

elliptic fruit. Zannichellia has longer leaves (3-10 cm long) than Najas (< 4 cm long). 

Zannichellia palustris Linnaeus, Horned Pondweed. Cp (NC, VA), Mt (VA), {GA}: fresh or brackish water; common. 

February-October. The species occurs throughout much of the world. [= RAB, C, FNA, G, GW, K, S, W, Z; > Z. palustris var. 

major (Hartman) W.D.J. Koch – F; > Z. palustris var. palustris – F] 

ZOSTERACEAE Dumortier 1829 (Eelgrass Family) 

A family of 3 genera and about 18 species, nearly cosmopolitan in distribution. References: Haynes in FNA (2000); Kuo & 

McComb in Kubitzki (1998b). 

Zostera Linnaeus 1753 (Eelgrass) 

A genus of about 12 species, aquatic herbs, of nearly cosmopolitan distribution. References: Haynes in FNA (2000); Crow & 

Hellquist (2000)=Y; Kuo & McComb in Kubitzki (1998b); Green & Short=Z. 

Zostera marina Linnaeus var. stenophylla Ascherson & Graebner, Eelgrass. Cp (NC, VA): estuarine waters; common. 

February-March. The species occurs in Eurasia and North America. Var. stenophylla is North American, and ranges south along 

the Atlantic coast to NC and allegedly to FL (though reports from that state are apparently not substantiated and may be based on 

misidentification of other aquatics). [= F, G, Y; < Z. marina – RAB, C, FNA, K, S, Z] 

BIBLIOGRAPHY 

Ackerfield, J., and J. Wen. 2002. A morphometric analysis of Hedera L. (the ivy genus, Araliaceae) and its taxonomic 

implications. Adansonia 24: 197-212. 

Adams, P. 1961. Observations on the Sagittaria subulata complex. Rhodora 63: 247-265. 

Adams, R.M. II, and W.J. Dress. 1982. Nodding Lilium species of eastern North America (Liliaceae). Baileya 21: 165-188. 

Adams, R.P. 1986. Geographic variation in Juniperus silicicola and J. virginiana of the Southeastern United States: 

multivariant analyses of morphology and terpenoids. Taxon 35: 31-75. 

–––. 1995. Revisionary study of Caribbean species of Juniperus (Cupressaceae). Phytologia 78: 134-150. 

–––, and T. Demeke. 1993. Systematic relationships in Juniperus based on random amplified polymorphic DNAs (RAPDs). 

Taxon 42: 553-571. 

Adams, W.P. 1957. A revision of the genus Ascyrum (Hypericaceae). Rhodora 59: 73-95. 

–––. 1962. Studies in the Guttiferae. I. A synopsis of Hypericum section Myriandra. Contr. Gray Herbarium Harv. 182: 1-51. 

–––. 1973. Clusiaceae of the southeastern United States. J. Elisha Mitchell Sci. Soc. 89: 62-71. 

–––, and N.K.B. Robson. 1961. A re-evaluation of the generic status of Ascyrum and Crookea (Guttiferae). Rhodora 63: 10-16. 

Adler, L. 1999. Polygonum perfoliatum (mile-a-minute weed). Chinquapin 7: 4. 

Aedo, C., J.J. Aldasoro, and C. Navarro. 1998. Taxonomic revision of Geranium sections Batrachioidea and Divaricata 

(Geraniaceae). Ann. Missouri Bot. Gard. 85: 594-630. 

Affolter, J.M. 1985. A monograph of the genus Lilaeopsis (Umbelliferae). Systematic Bot. Monographs 6. 

Ahles, H.E., and A.E. Radford. 1959. Species new to the flora of North Carolina. J. Elisha Mitchell Sci. Soc. 75: 140-147. 

Ahrendt, L.W.A. 1961. Berberis and Mahonia: a taxonomic revision. J. Linn. Soc., Bot. 57: 1-410. 

Aiken, S.G. 1981. A conspectus of Myriophyllum (Haloragaceae) in North America. Brittonia 33: 57-69. 

–––, and S.J. Darbyshire. 1990. Fescue grasses of Canada. Agriculture Canada Publication 1844/E. 

–––, M.J. Dallwitz, C.L. McJannet, and L.L. Consaul. 1997. Biodiversity among Festuca (Poaceae) in North America: 

diagnostic evidence from DELTA and clustering programs, and an INTKEY package for interactive, illustrated 

identification and information retrieval. Can. J. Bot. 75: 1527-1555. 

Akiyama, S. 1988. A revision of the genus Lespedeza section Macrolespedeza (Leguminosae). Univ. of Tokyo Press. 

–––, and H. Ohba. 1985. The branching of the inflorescence and vegetative shoot and taxonomy of the genus Kummerowia 

(Leguminosae). Bot. Mag. Tokyo 98: 137-150. 

Al-Shehbaz, I.A. 1984. The tribes of Cruciferae (Brassicaceae) in the southeastern United States. J. Arnold Arb. 65: 343-373. 

–––. 1985a. The genera of Thelypodieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 66: 95- 

111. 

–––. 1985b. The genera of Brassiceae (Crucifrae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 66: 279-351. 

–––. 1986a. The genera of Lepidieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 67: 265-311. 

–––. 1986b. New wool-alien Cruciferae (Brassicaceae) in eastern North America: Lepidium and Sisymbrium. Rhodora 88: 347- 

355. 

–––. 1987. The genera of Alysseae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 68: 185-240. 

–––. 1988a. The genera of Arabideae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 69: 85-166.

BIBLIOGRAPHY 951 

–––. 1988b. The genera of Sisymbrieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arb. 69: 213- 

237. 

–––. 1988c. Cardamine dissecta, a new combination replacing Dentaria multifida (Cruciferae). J. Arnold Arb. 69: 81-84. 

–––. 1988d. The genera of Anchonieae (Hesperideae) (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold 

Arb. 69: 193-212. 

–––. 1991. The genera of Boraginaceae in the southeastern United States. J. Arnold Arb. Suppl. Series 1: 1-169. 

–––. 2003. Transfer of most North American species of Arabis to Boechera (Brassicaceae). Novon 13: 381-391. 

–––, and V. Bates. 1987. Armoracia lacustris (Brassicaceae), the correct name for the North American lake cress. J. Arnold 

Arb. 68: 357-359. 

–––, and S.L. O'Kane, Jr. 2002. Lesquerella is united with Physaria (Brassicaceae. Novon 12: 319-329. 

–––, and R.A. Price. 1998. Delimitation of the genus Nasturtium (Brassicaceae). Novon 8: 124-126. 

–––, and B.G. Schubert. 1989. The Dioscoreaceae in the southeastern United States. J. Arnol Arb. 70: 57-95. 

–––, K. Mummenhoff, and O. Appel. 2002. Cardaria, Coronopus, and Stroganowia are united with Lepidium (Brassicaceae). 

Novon 12: 5-11. 

Allan, G.J., and J.M. Porter. 2000. Tribal delimitations and phylogenetic relationships of Loteae and Coronilleae (Faboideae: 

Fabaceae) with special reference to Lotus: evidence from nuclear ribosomal ITS sequences. Amer. J. Botany 87: 1871- 

1881. 

Albach, D.C., and M.W. Chase. 2001. Paraphyly of Veronica (Veroniceae; Scrophulariaceae): evidence from the Internal 

Transcribed Spacer (ITS) sequences of nuclear ribosomal DNA. J. Plant Res. 114: 9-18. 

–––, H.M. Meudt, and B. Oxelman. 2005. Piecing together the "new" Plantaginaceae. Amer. J. Bot. 92: 297-315. 

Aldasoro, J.J., C. Aedo, F.M. Garmendia, F. Pando de la Hoz, and C. Navarro. 2004. Revision of Sorbus subgenera Aria and 

Torminaria (Rosaceae-Maloideae). Systematic Botany Monographs 69: 1-148. 

Alexander, E.J. 1934. Parnassia caroliniana, Carolina grass-of-Parnassus, native of southeastern United States. Addisonia 18: 

43-46. 

–––. 1941. Two new species from the southern Appalachians. Castanea 6: 30-32. 

Alice, L.A., and C.S. Campbell. 1999. Phylogeny of Rubus (Rosaceae) based on nuclear ribosomal DNA internal transcribed 

spacer region sequences. Amer. J. Bot. 86: 81-97. 

Allard, H.A. 1940. Phacelia ranunculacea (Nutt.) Constance, its length of day, temperature reactions and seasonal adaptations. 

Castanea 5: 94-97. 

Allison, J.R. 2006. Big-fruited buckthorn, Sideroxylon macrocarpum (Sapotaceae), a long-forgotten Georgia endemic. Sida 22: 

243-264. 

–––, and T.E. Stevens. 2001. The endemic flora of Ketona dolomite outcrops in Bibb County, Alabama. Castanea 66: 154-205. 

–––, M.W. Morris, and A.N. Egan. 2006. A new species of Pediomelum (Fabaceae) from the lower Piedmont Plateau of 

Georgia and South Carolina. Sida 22: 227-241. 

Allred, K.W. 1984. Studies in the genus Aristida (Gramineae) of the southeastern United States. I. Spikelet variation in A. 

purpurescens, A. tenuispica, and A. virgata. Rhodora 86: 73-77. 

–––. 1985. Studies in the Aristida (Gramineae) of the southeastern United States. III. Nomenclature and a taxonomic 

comparison of A. lanosa and A. palustris. Rhodora 87: 147-155. 

–––. 1986. Studies in the Aristida (Gramineae) of the southeastern United States. IV. Key and conspectus. Rhodora 88: 367- 

387. 

–––, and F.W. Gould. 1983. Systematics of the Bothriochloa saccharoides complex (Poaceae: Andropogoneae). Systematic 

Botany 8: 168-184. 

Ambrose, J.D. 1980. A re-evaluation of the Melanthoideae (Liliaceae) using numerical analyses. Pp. 65-81 in C.D. Brickell, 

D.F. Cutler, & M. Gregory (editors), Petaloid monocotyledons. Linn. Soc. Symp. Ser. 8. Academic Press, London. 

–––. 1985. Lophiola, familial affinity with the Liliaceae. Taxon 34: 149-150. 

Amoroso, J.L., compiler. 1997. Natural Heritage Program list of the rare plants of North Carolina. North Carolina Natural 

Heritage Program, Raleigh, NC. 

Anderberg, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104. 

–––, and X. Zhang. 2002. Phylogenetic relationships of Cyrillaceae and Clethraceae (Ericales) with special emphasis on the 

genus Purdiaea Planch. Org. Divers. Evol. 2: 127-137. 

Anderson, E., and R.E. Woodson. 1935. The species of Tradescantia indigenous to the United States. Contr. Arnold Arb. 9: 

132. 

Anderson, E.F. 2001. The cactus family. Timber Press, Portland, OR. 

Anderson, L.C. 1970. Studies in Bigelowia (Astereae, Compositae) 1. Morphology and taxonomy. Sida 3: 451-465. 

–––. 1983. Hydrocotyle bowlesioides in Georgia – new to United States. Castanea 48: 317. 

–––. 1985. Forestiera godfreyi (Oleaceae), a new species from Florida and South Carolina. Sida 11: 1-5. 

–––. 1987. Boltonia apalachicolensis (Asteraceae): a new species from Florida. Systematic Bot. 12: 133-138. 

–––. 1988. Status of endangered Rhynchospora crinipes (Cyperaceae). Systematic Bot. 13: 407-410. 

–––. 1996. New geographical and morphological data for Sideroxylon thornei (Sapotaceae). Sida 17: 343-348. 

–––. 1998. Arnoglossum album (Asteraceae): new species from northern Florida. Sida 18: 377-384. 

–––, and D.W. Hall. 1993. Luziola bahiensis (Poaceae): new to Florida. Sida 15: 619-622.


Anderson, L.E., and T.T. Bannister. 1952. An addition to the fern flora of North Carolina. J. Elisha Mitchell Sci. Soc. 68: 81- 

83. 

––––––, H.A. Crum, and W.R. Buck. 1990. List of the mosses of North America north of Mexico. The Bryologist 93: 448-499. 

Angiosperm Phylogeny Group. 1998. An ordinal classification for the families of flowering plants. Ann. Mo. Bot. Garden 85: 

531-553. 

–––. 2003. An update of the Angiosperm Phylogeny Group clssification for the orders and families of flowering plants: APG II. 

Bot. J. Linn. Soc. 141: 399-436. 

Anonymous. 1999. Harmful aquatic weed discovered in several North Carolina counties. Wildlife in North Carolina 63: 32. 

–––. 2003. Rare plant relocated after 64 years. BotSoc News 77: 2-3. 

Aplet, G.H., R.D. Laven, M.B. Falkner, and R.B. Shaw. 1994. Population and site characteristics of a recently discovered 

disjunct population of Croton alabamensis (Euphorbiaceae). Sida 16: 37-55. 

Appel, O. 1998. The status of Teesdaliopsis and Teesdalia (Brassicaceae). Novon 8: 218-219. 

Arbo, M.M. 1990. Turneraceae: novedades para la Guayana Venezolana. Ann. Missouri Bot. Gard. 77: 340-352. 

–––. 1995. Turneraceae – parte 1: Piriqueta. Flora Neotropica Monograph 67. 

Argus, G.W. 1986. The genus Salix (Salicaceae) in the southeastern United States. Systematic Bot. Monographs 9: 1-170. 

–––. 1997. Infrageneric classification of Salix (Salicaceae) in the New World. Systematic Botany Monographs 52: 121. 

Armstrong, J.E. 1985. The delimitation of Bignoniaceae and Scrophulariaceae based on floral anatomy, and the placement of 

problem genera. Am. J. Bot. 72: 755-766. 

Arriagada, J.E. 1998. The genera of Inuleae (Compositae; Asteraceae) in the southeastern United States. Harvard Papers in 

Botany 3: 1-48. 

Arriagada, J.E., and N.G. Miller. 1997. The genera of Anthemidae (Compositae; Astaeraceae) in the southeastern United States. 

Harvard Papers in Botany 2: 1-46. 

Ashe, W.W. 1922. The eastern shrubby species of Robinia. J. Elisha Mitchell Sci. Soc. 37: 175-177. 

Austin, D.F., and Z. Huáman. 1996. A synopsis of Ipomoea (Convolvulaceae) in the Americas. Taxon 45: 3-38. 

–––, and R.S. Bianchini. 1998. Additions and corrections in American Ipomoea (Convolvulaceae). Taxon 47: 833-838. 

–––, G.M. Diggs, Jr., and B.L. Lipscomb. 1997. Calystegia (Convolvulaceae) in Texas. Sida 17: 837-840. 

Averett, J.E., and D.E. Boufford. 1985. The flavonoids and flavonoid systematics of Circaea (Circaeeae, Onagraceae). 

Systematic Bot. 10: 363-373. 

Azuma, H., J.G. García-Franco, V. Rico-Gray, and L.B. Thien. 2001. Molecular phylogeny of the Magnoliaceae: the 

biogeography of tropical and temperate disjunctions. Am. J. Bot. 88: 2275-2285. 

–––, L.B. Thien, and S. Kawano. 1999. Molecular phylogeny of Magnolia (Magnoliaceae) inferred from cpDNA sequences and 

evolutionary divergence of the flora scents. J. Plant Res. 112: 291-306. 

Baas, P. 1984. Vegetative anatomy and the taxonomic status of Ilex collina and Nemopanthus (Aquifoliaceae). J. Arnold Arb. 

65: 243-250. 

Bacigalupo, N.M., and E.L. Cabral. 1999. Revisión de las especies americanas del género Diodia (Rubiaceae, Spermacoceae). 

Darwiniana 37: 153-165. 

Backlund, A., and N. Pyck. 1998. Diervillaceae and Linnaeaceae, two new families of caprifolioids. Taxon 47: 657-661. 

Backlund, M., B. Oxelman, and B. Bremer. 2000. Phylogenetic relationships within Gentianales based on ndhF and rbcL 

sequences, with particular reference to the Loganiaceae. Am. J. Bot. 87: 1029-1043. 

Bailey, V.A. 1962. Revision of the genus Ptelea (Rutaceae). Brittonia 14: 1-45. 

Ballard, H.E., Jr. 1992. Summary: systematics of Viola section Viola in north America north of Mexico. Unpublished 

manuscript. 

–––, and D.E. Wujek. 1994. Evidence for the recognition of Viola appalachiensis. Systematic Bot. 19: 523-538. 

–––, K.J. Sytsma, and R.R. Kowal. 1999. Shrinking the violets: phylogenetic relationships of infrageneric groups in Viola 

(Violaceae) based on internal transcribed spacer DNA sequences. Systematic Botany 23: 439-458. 

–––, D.A. Casamatta, Jr., M.M. Hall, R.A. McCauley, M.C. Segovia-Salcedo, and R.G. Verb. 2001. Phenetic analysis shows 

conspecifity between Hispaniolan Viola domingensis Urban and North American Viola macloskeyi sensu lato (Violaceae). 

Brittonia 53: 122-136. 

Ballard, R. 1986. Bidens pilosa complex (Asteraceae) in North and Central America. Amer. J. Bot. 73: 1452-1465. 

Banks, D.J. 1966. Taxonomy of Paspalum setaceum (Gramineae). Sida 2: 269-284. 

Barber, S.C. 1982. Taxonomic studies in the Verbena stricta complex (Verbenaceae). Systematic Bot. 7: 433-456. 

Barden, L.S. 1987. Invasion of Microstegium vimineum (Poaceae), an exotic, annual, shade-tolerant, C4 grass, into a North 

Carolina floodplain. Amer. Midland Naturalist 118: 40-45. 

Barkley, F.A. 1937. A monographic study of Rhus and its immediate allies in North and Central America, including the West 

Indies. Ann. Missouri Bot. Gard. 24: 265-499. 

Barkley, T.M. 1962. A revision of Senecio aureus Linn. and allied species. Trans. Kan. Acad. 65: 318-408. 

–––. 1968. Taxonomy of Senecio multilobatus and its allies. Brittonia 20: 267-284. 

–––. 1978. Senecio. N. Amer. Fl. II 10: 50-139. 

–––. 1980. Taxonomic notes on Senecio tomentosus and its allies (Asteraceae). Brittonia 32: 291-308. 

–––. 1999. The segregates of Senecio, s.l., and Cacalia, s.l., in the flora of North America north of Mexico. Sida 18: 661-672. 

Barkworth, M.E. 1997. Taxonomic and nomenclatural comments on the Triticeae in North America. Phytologia 83: 302-311. 

Barneby, R.C. 1964. Atlas of North American Astragalus. Mem. New York Bot. Gard. 13: 1-1188.


–––. 1977. Daleae imagines, an illustrated revision or Errazurizia Philippi, Psorothamnus Rydberg, Marina Liebmann, and 

Dalea Lucanus emend. Barneby, including all species of Leguminosae tribe Amorpheae Borissova ever referred to Dalea. 

Mem. N.Y. Bot. Garden 27: 1-892. 

–––. 1991. Sensitivae Censitae: a description of the genus Mimosa Linnaeus (Mimosaceae) in the New World. Mem. New 

York Bot. Garden 65. 

–––, and E.L. Bridges. 1987. A new species of Astragalus (Fabaceae) from Tennessee's Central Basin. Brittonia 39: 358-363. 

Barnes, P.G. 1990. A summary of the genus Shortia. The Plantsman 12: 23-34. 

Barringer, K. 1993. New combinations in North American Asarum (Aristolochiaceae). Novon 3: 225-227. 

Bartgis, R.L. 1992. The endangered sedge Scirpus ancistrochaetus and the flora of sinkhole ponds in Maryland and West 

Virginia. Castanea 57: 46-51. 

–––. 1993. The limestone glades and barrens of West Virginia. Castanea 58: 69-89. 

–––, G.P. Fleming, and R. Wiegand. 1997. The prairie-redroot (Ceanothus herbaceus Raf.) in the mid-Atlantic United States. 

Castanea 62: 127-128. 

Baskin, J.M., and C.C. Baskin. 1998. Seed dormancy and germination in the rare plant species Amaranthus pumilus. Castanea 

63: 493-494. 

–––, C.C. Baskin, & M.E. Medley. 1983. The historical geographical distribution of Onosmodium molle Michaux subsp. molle 

(Boraginaceae). Bull. Torrey Bot. Club 110: 73-76. 

–––, K.M. Snyder, and C.C. Baskin. 1993. Nomenclatural history and taxonomic status of Echinacea angustifolia, E. pallida, 

and E. tennesseensis (Asteraceae). Sida 15: 597-604. 

–––, R.W. Tyndall, M. Chaffins, and C.C. Baskin. 1998. Effect of salinity on germination and viability of nondormant seeds on 

the federal-threatened species Aeschynomene virginica (Fabaceae). J. Torrey Bot. Soc. 125: 246-248. 

Bassett, I.J. 1966. Taxonomy of North American Plantago L., section Micropsyllium Decne. Can J. Bot. 44: 467-479. 

–––. 1967. Taxonomy of Plantago L. in North America: sections Holopsyllium Pilger, Palaeopsyllium Pilger, and 

Lamprosantha Decne. Can. J. Bot. 45: 565-577. 

–––, and C.W. Crompton. 1982. The genus Chenopodium in Canada. Can. J. Bot. 60: 586-610. 

Bateman, R.M., A.M. Pridgeon, and M.W. Chase. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideaea, Orchidaceae) 

based on nuclear ITS sequences. 2. Infrageneric relationships and reclassification to achieve monophyly of Orchis sensu 

stricto. Lindleyana 12: 113-141. 

Bates, D.M. 1967. A reconsideration of Sidopsis Rydberg and notes on Malvastrum A. Gray (Malvaceae). Rhodora 69: 9-28. 

Bates, V.M., and E.T. Browne. 1981. Azolla filiculoides new to the southeastern United States. Amer. Fern J. 71: 33-34. 

Batson, W.T. 1977. A guide to the genera of native and commonly introduced ferns and seed plants of eastern North America 

from the Atlantic to the Great Plains from Key West-southern Texas into the Arctic. John Wiley & Sons, New York, N.Y. 

Baum, B.R. 1978. The genus Tamarix. Israel Acad. of Sciences and Humanities, Jerusalem. 

Bayer, C., M.F. Fay, A.Y. de Bruijn, V. Savolainen, C.M. Morton, K. Kubitzki, W.S. Alverson, and M.W. Chase. 1999. Support 

for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atpB 

and rbcL DNA sequences. Bot. J. Linnean Society 129: 267-303. 

Bayer, R.J. 1984. Chromosome numbers and taxonomic notes for North American species of Antennaria (Asteraceae: Inuleae). 


–––. 1985. Investigations into the evolutionary history of the polyploid complexes in Antennaria (Asteraceae: Inuleae). II. The 

A. parlinii complex. Rhodora 87: 321-339. 

–––, and G.L. Stebbins. 1982. A revised classification of Antennaria (Asteraceae: Inuleae) of the eastern United States. 


–––, and G.L. Stebbins. 1987. Chromosome numbers, patterns of distribution, and apomixis in Antennaria (Asteraceae: 

Inuleae). Systematic Bot. 12: 305-319. 

–––, and G.L. Stebbins. 1993. A synopsis with keys for the genus Antennaria (Asteraceae: Inuleae: Gnaphaliinae) of North 

America. Can J. Bot. 71: 1589-1604. 

Beadle, C.D. 1913. Crataegus. Pp. 532-569 in J.K. Small, Flora of the southeastern United States, being descriptions of the 

seed-plants, ferns and fern-allies growing naturally in North Carolina, South Carolina, Georgia, Florida, Tennessee, 

Alabama, Mississippi, Arkansas, Louisiana, and in Oklahoma and Texas east of the one hundredth meridian. Published by 

the author, New York, NY. 1394 pp. 

–––, and F.E. Boynton. 1901. Revision of the species of Marshallia. Biltmore Bot. Studies 1: 3-10. 

Beal, E.O. 1956. Taxonomic revision of the genus Nuphar Sm. of North America and Europe. J. Elisha Mitchell Sci. Soc. 72: 

317-346. 

–––. 1960. Sparganium (Sparganiaceae) in the southeastern United States. Brittonia 12: 176-181. 

–––, and R.M. Southall. 1977. The taxonomic significance of experimental selection by vernalization in Nuphar 

(Nymphaeaceae). Systematic Bot. 2: 49-60. 

–––, J.W. Wooten, and R.B. Kaul. 1982. Review of the Sagittaria engelmanniana complex (Alismataceae) with environmental 

correlations. Systematic Bot. 7: 417-432. 

Beard, L.S. 1963. A taxonomic study of Mimosa quadrivalvis L. (Schrankia Willd. nom. cons.). Ph.D. thesis, Univ. of North 

Carolina at Chapel Hill. 

Beardsley, P.M., and R.G. Olmstead. 2002. Redefining Phrymaceae: the placement of Mimulus, tribe Mimuleae, and Phryma. 

Amer. J. Bot. 89: 1093-1102.


Beckmann, R.L., Jr. 1979. Biosystematics of the genus Hydrophyllum L. (Hydrophyllaceae). Can J. Bot. 66: 1053-1061. 

Beitel, J.M. 1979. Clubmosses (Lycopodium) in North America. Fiddlehead Forum 6: 1-8. 

–––, and J.T. Mickel. 1992. The Appalachian firmoss, a new species in the Huperzia selago (Lycopodiaceae) complex in eastern 

North America, with a new combination for the western firmoss. Amer. Fern J. 82: 41-46. 

Belden, A., Jr., and N.E. Van Alstine. 2003. Status of the federally listed Aeschynomene virginica (L.) BSP. on the James River 

in Virginia. Castanea 68: 179-181. 

Belden, A., Jr., G.R. Fleming, J.C. Ludwig, J.F. Townsend, N.E. Van Alstine, and T.F. Wieboldt. 2004. Noteworthy collections: 

Virginia. Castanea 69: 144-153. 

Bell, A.D. 1974. Rhizome organization in relation to vegetative spread in Medeola virginiana. J. Arnold Arb. 55: 458-468. 

Bell, C.D. 2004. Preliminary phylogeny of Valerianaceae (Dipsacales) inferred from nuclear and chloroplast DNA sequence 

data. Molecular Phylogenetics and Evolution 31: 340-350. 

Bell, C.R. 1949. A cytotaxonomic study of the Sarraceniaceae of North America. J. Elisha Mitchell Sci. Soc. 65: 137-166. 

–––. 1952. Natural hybrids in the genus Sarracenia. I. History, distribution, and taxonomy. J. Elisha Mitchell Sci. Soc. 68: 55- 

80. 

–––, and F.W. Case. 1956. Natural hybrids in the genus Sarracenia. II. Current notes on distribution. J. Elisha Mitchell Soc. 

72: 142-152. 

–––. 1963. The genus Eryngium in the southeastern United States. Castanea 28: 73-79. 

Benson, L. 1982. The cacti of the United States and Canada. Stanford Univ. Press, Stanford, California. 

Bicknell, E.P. 1896. The blue-eyed grasses of the eastern United States (genus Sisyrinchium). Bull. Torrey Bot. Club 23: 130- 

133. 

–––. 1899a. Studies in Sisyrinchium – I. Sixteen new species from the southern states. Bull. Torrey Bot. Club 26: 217-231. 

–––. 1899b. Studies in Sisyrinchium – VI. Additional new species from the southern states. Bull. Torrey Bot. Club 26: 605- 

616. 

Bierner, M.W. 1972. Taxonomy of Helenium sect. Tetrodus and a conspectus of North American Helenium (Compositae). 

Brittonia 24: 331-355. 

–––. 1989. Taxonomy of Helenium sect. Amarum (Asteraceae). Sida 13: 453-459. 

Binns, S.E., B.R. Baum, and J.T. Arnason. 2002. A taxonomic revision of Echinacea (Asteraceae: Heliantheae). Syst. Bot. 27: 

610-632. 

Bishop, M., A. Davis, and J. Grimshaw. 2001. Snowdrops: a monograph of cultivated Galanthus. Griffin Press, Maidenhead, 

UK. 364 pp. 

Blackwell, W.H., Jr., M.D. Baechle, and G. Williamson. 1978. Synopsis of Kochia (Chenopodiaceae) in North America. Sida 

7: 248-254. 

–––, and K.P. Blackwell. 1974. The taxonomy of Peltandra (Araceae). J. Elisha Mitchell Sci. Soc. 90: 137-140. 

Blattner, F.R. 2004. Phylogenetic analysis of Hordeum (Poaceae) as inferred by nuclear rDNA ITS sequences. Molecular 

Phylogenetics and Evolution 33: 289-299. 

Blomquist, H.L. 1948. The grasses of North Carolina. Duke University Press, Durham, N.C. 

–––. 1957. A revision of Hexastylis of North America. Brittonia 8: 255-281. 

Bodo Slotta, T.A., and D.M. Porter. 2006. Genetic variation within and between Iliamna corei and I. remota (Malvaceae): 

implications for species delimitation. Bot. J. Linn. Soc. 151: 345-354. 

Boetsch, J.R. 2002. The Aizoaceae and Molluginaceae of the southeastern United States. Castanea 67: 42-53. 

–––-, and E. Nielsen. 2003. Notes on the distribution of the Southern Appalachian endemic, Ilex collina Alexander. Castanea 

68: 232-235. 

Bogin, C. 1955. Revision of the genus Sagittaria (Alismataceae). Memoirs N.Y. Botanical Garden 9: 179-233. 

Bogle, A.L. 1969. The genera of Portulacaceae and Basellaceae in the southeastern United States. J. Arnold Arb. 50: 566-598. 

–––. 1974. The genera of Nyctaginaceae in the southeastern United States. J. Arnold Arb. 55: 1-37. 

Bogler, D.J., and B.B. Simpson. 1995. A chloroplast DNA study of the Agavaceae. Systematic Bot. 20: 191-205. 

–––-, and B.B. Simpson. 1996. Phylogeny of Agavaceae based on ITS rDNA sequence variation. Amer. J. Bot. 83: 1225-1235. 

Bohm, B.A., J.Y. Yang, J.E. Page, and D.S. Soltis. 1999. Flavonoids, DNA and relationships of Itea and Pterostemon. 

Biochemical Systematics and Ecology 27: 79-83. 

Bohs, L., and R.G. Olmstead. 1997. Phylogenetic relationships in Solanum (Solanaceae) based on ndhF sequences. Systematic 

Bot. 22: 5-18. 

Bolle, F. 1933. Eine Übersicht über die Gattung Geum L. und ihr nahestehenden Gattungen. Feddes Repertorium 72: 1-119. 

Bolli, R. 1994. Revision of the genus Sambucus. Dissertationes Botanicae 223. J. Cramer, Berlin. 227 pp. 

Bolmgren, K., and B. Oxelman. Generic limits in Rhamnus L. s.l. (Rhamnaceae) inferred from nuclear and chloroplast DNA 

sequences. Taxon 53: 383-390. 

Boom, B.M. 1982. Synopsis of Isoetes in the southeastern United States. Castanea 47: 38-59. 

Boufford, D.E. 1977. Ammoselinum butleri (Umbelliferae), new to North Carolina. Sida 7: 220. 

–––. 1982. Notes on Peperomia (Piperaceae) in the southeastern United States. J. Arnold Arb. 63: 317-325. 

–––. 1983 ["1982"]. The systematics and evolution of Circaea (Onagraceae). Ann Mo. Bot. Gard. 69: 804-994. 

–––. 2005. Circaea lutetiana sensu lato (Onagraceae) reconsidered. Harvard papers in Botany 9: 255-256. 

–––, and S.A. Spongberg. Calycanthus floridus (Calycanthaceae) – a nomenclatural note. J. Arnold Arb. 62: 265-266. 

–––, and E.W. Wood. 1977. An unusual plant community in South Carolina. Castanea 42: 116-119.


Bounds, R.R. 1987. Rare species of Rhexia L. Castanea 52: 304-308. 

Bown, D. 2000. Aroids: plants of the Arum Family. Timber Press, Portland, OR. 392 pp. 

Boyce, P. 2006. The genus Arum. Kew Magazine Monograph. Royal Botanic Gardens, Kew. 196 pp. 

Bozeman, J.R., and J.F. Logue. 1968. A range extension for Hudsonia ericoides in the southeastern United States. Rhodora 70: 

289-292. 

–––, and G.A. Rogers. 1986. "This very curious tree." Tipularia 1: 9-15. 

Brandenburg, D.M., and J.W. Thieret. 2000. Cinna and Limnodea (Poaceae): not congeneric. Sida 19: 195-2000. 

–––, W.H. Blackwell, and J.W. Thieret. 1991. Revision of the genus Cinna (Poaceae). Sida 14: 581-596. 

–––, J.R. Estes, & S.L. Collins. 1991. A revision of Diarrhena (Poaceae) in the United States. Bull. Torrey Bot. Club 118: 128- 

136. 

–––, J.R. Estes, & J.W. Thieret. 1991. Hard grass (Sclerochloa dura, Poaceae) in the United States. Sida 14: 369-376. 

Braun, E.L. 1942. A new species and a new variety of Solidago from Kentucky. Rhodora 44: 1-4. 

Bremer, B., and L. Struwe. 1992. Phylogeny of the Rubiaceae and the Loganiaceae: congruence or conflict between 

morphological and molecular data? Am. J. Bot. 79: 1171-1184. 

–––, K. Bremer, N. Heidari, P. Erixon, R.G. Olmstead, A.A. Arneberg, M. Källersjö, & E. Barkhordarian. 2002. Phylogenetics 

of asterids based on 3 coding and 3 non-coding chloroplast DNA markers and the utility of non-coding DNA at higher 

taxonomic levels. Molecular Phylogenetics and Evolution 24: 274-301. 

Bremer, K. 1994. Asteraceae: cladistics and classification. Timber Press, Portland, OR. 752 pp. 

Bretting, P.K., and S. Nilsson. 1988. Pollen morphology of the Martyniaceae and its systematic implications. Systematic Bot. 

13: 51-59. 

Bridges, E.L., and S.L. Orzell. 1989. Evolvulus sericeus (Convolvulaceae) in Georgia, with floristic and ecological notes. Sida 

13: 509-512. 

–––––, and S.L. Orzell. 1992. The rediscovery of Rhynchospora solitaria Harper (Cyperaceae) in Georgia. Phytologia 72: 369- 

372. 

–––, and S.L. Orzell. 2002. Euphorbia (Euphorbiaceae) section Tithymalus subsection Inundatae in the southeastern United 

States. Lundellia 5: 59-78. 

–––, and S.L. Orzell. 2003. Two new species and a new combination in southeastern United States Xyris (Xyridaceae) from 

Florida. Novon 13: 16-25. 

–––, S.L. Orzell, and J.R. Burkhalter. 1993. Cladium mariscoides (Cyperaceae) in the western Florida panhandle and its 

phytogeographic significance. Phytologia 74: 35-42. 

–––. [in prep.] Xyridaceae. In: Flora of Florida, Volume 2 – monocots. 

Brizicky, G.K. 1964a. The genera of Celastrales in the southeastern United States. J. Arnold Arb. 45: 206-234. 

–––. 1964b. A further note on Ceanothus herbaceus versus C. ovatus. J. Arnold Arb. 45: 471-473. 

–––. 1964c. The genera of Rhamnaceae in the southeastern United States. J. Arnold Arb. 45: 439-463. 

–––. 1966. The genera of Sterculiaceae in the southeastern United States. J. Arnold Arb. 47: 60-74. 

Brooks, R.E., and A.T. Whittemore. 1999. Juncus anthelatus (Juncaceae, Juncus subg. Poiophylli), a new status for a North 

American taxon. Novon 9: 11-12. 

Brouillet, L., and J.C. Semple. 1981. A propos du status taxonomique de Solidago ptarmicoides. Can. J. Bot. 59: 17-21. 

Brown, C.A. 1959. Vegetation of the Outer Banks of North Carolina. Louisiana State University Studies, Coastal Studies Series 

No. 4. La. State Univ. Press, Baton Rouge, LA. 179 pp. 

Brown, L.E., and S.J. Marcus. 1998. Notes on the flora of Texas with additions and other significant records. Sida 18: 315-324. 

Brown, P.M. 1999. Recent taxonomic and distributional notes from Florida. 1. North American Native Orchid Journal 5: 3-16. 

–––. 2001. Recent taxonomic and distributional notes from Florida 11. Spiranthes sylvatica P.M. Brown, a new species of 

ladies'-tresses from the southeastern United States. North American Native Orchid Journal 7: 193-205. 

–––. 2003. The wild orchids of North America, north of Mexico. University Press of Florida, Gainesville, FL. 

–––. 2004. Understanding Platanthera chapmanii (Orchidaceae), its origins and hybrids. Sida 21: 853-859. 

–––. 2006a. Resurrection of the genus Gymnadeniopsis Rydberg. North American Native Orchid Journal 12: 33-40. 

–––. 2006b. Revalidation of Platanthera conspicua, the southern white fringed orchis. North American Native Orchid Journal 

12: 41-50 

–––, and R.B. Pike. 2006. Triphora trianthophora var. texensis (Orchidaceae) a new variety endemic to Texas. North American 

Native Orchid Journal 12: 5-10. 

Bruederle, L.P. 1999. Genetic differentiation of geographicaly marginal populations in Carex mitchelliana (Cyperaceae): 

implications for conservation. J. Torrey Bot. Soc. 126: 1-8. 

–––, and D.E. Fairbrothers. 1986. Allozyme variation in populations of the Carex crinita complex (Cyperaceae). Systematic 

Bot. 11: 583-594. 

–––, D.E. Fairbrothers, and S.L. Hanks. 1989. A systematic circumscription of Carex mitchelliana (Cyperaceae) with reference 

to taxonomic status. Am. J. Bot. 76: 124-132. 

Brummitt, R.K. 1965. New combinations in North American Calystegia. Ann. Missouri Bot. Gard. 52: 214-216. 

–––. 1980. Further new names in the genus Calystegia (Convolvulaceae). Kew Bull. 35: 327-334. 

–––. 1988. Report of the Committee for Spermatophyta: 34. Taxon 37: 139-140. 

–––. 1999. Proposals to conserve or reject. Report of the Committee on Spermatophyta. Taxon 48: 367. 

–––. 2001. Report of the Committee for Spermatophyta: 52. Taxon 50: 1179-1182.


–––. 2005. Report of the Committee for Spermatophyta: 57. Taxon 54: 1093-1103. 

Brunsfeld, S.J., P.S. Soltis, D.E. Soltis, P.A. Gadek, C.J. Quinn, D.D. Strenge, T.A. Ranker. 1994. Phylogenetic relationships 

among the genera of Taxodiaceae and Cupressaceae: evidence from rbcL sequences. Systematic Bot. 19: 253-262. 

Brunton, D.F., and D.M. Britton. 1996a. Noteworthy collections: Alabama and Georgia. Castanea 61: 398-399. 

–––, and D.M. Britton. 1996b. The status, distribution, and identification of Georgia Quillwort (Isoetes georgiana; Isoetaceae). 

American Fern Journal 86: 105-113. 

–––, and D.M. Britton. 1997. Appalachian quillwort (Isoetes appalachiana, sp. nov.; Isoetaceae), a new pteridophyte from the 

eastern United States. Rhodora 99: 118-133. 

–––, and D.M. Britton. 1998. Isoetes microvela (Isoetaceae), a new quillwort from the coastal plain of the southeastern United 

States. Rhodora 100: 261-275. 

–––, and D.M. Britton. 1999. Rush quillwort (Isoetes junciformis, sp. nov.), a new pteridophyte from southern Georgia. 

American Fern Journal 89: 187-197. 

–––, and D.M. Britton. 2006. Isoetes melanopoda ssp. sylvatica (subsp. nov.), a new quillwort (Isoetaceae) from eastern North 

America. Castanea 71: 15-30. 

–––, D.M. Britton, and W.C. Taylor. 1994. Isoetes hyemalis, sp. nov. (Isoetaceae): a new quillwort from the southeastern 

United States. Castanea 59: 12-21. 

–––, D.M. Britton, and T.F. Wieboldt. 1996. Taxonomy, identity, and status of Isoetes virginica (Isoetaceae). Castanea 61: 145- 

160. 

–––, W.H. Wagner, Jr., and J.M. Beitel. 1992. Pacific firmoss (Huperzia miyoshiana) (Lycopodiaceae) in eastern North 

America at Gros Morne National Park, Newfoundland. Amer. Fern J. 82: 63-67. 

Bryan, F.A., and D.E. Soltis. 1987. Electrophoretic evidence for allopolyploidy in the fern Polypodium virginianum. Systematic 

Bot. 12: 553-561. 

Bryson, C.T. 1980. A revision of the North American Carex section Laxiflorae (Cyperaceae). Ph.D. dissertation, Mississippi 

State Univ. 

–––, R. Kral, and J.R. Manhart. 1987. A new species of Carex (Cyperaceae: section Oligocarpae) from the southeastern United 

States. Rhodora 89: 357-363. 

–––, J.R. MacDonald, R. Carter, and S.D. Jones. 1996. Noteworthy Carex, Cyperus, Eleocharis, Kyllinga, and Oxycaryum 

(Cyperaceae) from Alabama, Arkansas, Georgia, Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17: 

501-518. 

Buck, W.R. 1977. A new species of Selaginella in the S. apoda complex. Can. J. Bot. 55: 366-371. 

Buddell, G.F. II, and J.W. Thieret. 1985. Notes on Erigenia bulbosa (Apiaceae). Bartonia 51: 69-76. 

Bunsawat, J., N.E. Elliott, K.L. Hertweck, E. Sproles, and L.A. Alice. 2004. Phylogenetics of Mentha (Lamiaceae): evidence 

from chloroplast DNA sequences. Systematic Botany 29: 959-964. 

Burckhalter, R.E. 1992. The genus Nyssa (Cornaceae) in North America: a revision. Sida 15: 323-342. 

Burk, C.J. 1961. Distribution records and range extensions from the North Carolina Outer Banks. Castanea 26: 138-139. 

Burks, K.C. 2002. Nymphoides cristata (Roxb.) Kuntze, a recent adventive expanding as a pest plant in Florida. Castanea 67: 

206-211. 

Cabe, P.R. 1995. The Trillium pusillum Michaux (Liliaceae) complex in Virginia. I. Morphological investigations. Castanea 

60: 1-14. 

–––, and C. Werth. 1995. The Trillium pusillum Michaux (Liliaceae) complex in Virginia. II. Isozyme evidence. Castanea 60: 

15-29. 

Calie, P.J. 1981. Systematic studies in Sedum section Ternata (Crassulaceae). Brittonia 33: 498-507. 

–––, E.E. Schilling, and D.H. Webb. 1983. Flavonoid chemistry of the generic segregates Ascyrum and Crookea of Hypericum. 

Biochem. Syst. and Ecology 11: 107-109. 

Callahan, H.S. 1997. Infraspecific differentiation in the Amphicarpaea bracteata (Fabaceae) species complex: varieties and 

ecotypes. Rhodora 99: 64-82. 

Camelbeke, K., A.A. Reznicek, and P. Goetghebeur. 2003. Proposal to conserve the name Scleria reticularis with a conserved 

type (Cyperaceae). Taxon 52: 355-356. 

Cameron, K.M., and M.W. Chase. 1999. Phylogenetic relationships of Pogoniinae (Vanilloideae, Orchidaceae): an herbaceous 

example of the eastern North America - eastern Asia phytogeographic disjunction. J. Plant Res. 112: 317-329. 

–––, K.J. Wurdack, and R.W. Jobson. 2002. Molecular evidence for the common origin of snap-traps among carnivorous plants. 

Amer. J. Bot. 89: 1503-1509. 

–––, M.W. Chase, W.M. Whitten, P.J. Kores, D.C. Jarrell, V.A. Albert, T. Yukawa, H.G. Hills, and D.W. Goldman. 1999. A 

phylogenetic analysis of the Orchidaceae: evidence from rbcL nucleotide sequences. Amer. J. Bot. 86: 208-224. 

Camp, W.H. 1935. Studies in the Ericales. I. The genus Gaylussacia in North America north of Mexico. Bull. Torrey Bot. 

Club 62: 129-132. 

–––. 1938. Studies in the Ericales. III. The genus Leiophyllum. Bull. Torrey Bot. Club 65: 99-104. 

–––. 1945. The North American blueberries with notes on other groups of Vacciniaceae. Brittonia 5: 203-275. 

Campbell, C.S. 1983. Systematics of the Andropogon virginicus complex (Gramineae). J. Arnold Arb. 64: 171-254. 

–––. 1985. The subfamilies and tribes of Gramineae (Poaceae) in the southeastern United States. J. Arnold Arb. 66: 123-199. 

–––. 1986. Phylogenetic reconstructions and two new varieties in the Andropogon virginicus complex (Poaceae: 

Andropogoneae). Syst. Bot. 11: 280-292.


–––, P.E. Garwood, and L.P. Specht. 1986. Bambusoid affinities of the north temperate genus Brachyelytrum (Gramineae). 

Bull. Torrey Bot. Club 113: 135-141. 

Campbell, G.R. 1952. The genus Myosurus L. (Ranunculaceae) in North America. El Aliso 2: 389-403. 

Campbell, J.J.N. 2000. Notes on North American Elymus species (Poaceae) with paired spikelets: I. E. macgregorii sp. nov. 

and E. glaucus ssp. mackenzii comb. nov. J. Ky. Acad. Sci. 61: 88-98. 

Canne, J.M. 1979. A light and scanning electronic microscope study of seed morphology in Agalinis (Scrophulariaceae) and its 

taxonomic significance. Systematic Bot. 4: 281-296. 

Cantino, P.D. 1982. A monograph of the genus Physostegia (Labiatae). Contr. Gray Herb. 211. 

–––. 1985. Facultative autogamy in Synandra hispidula (Labiatae). Castanea 50: 105-111. 

–––, and S.J. Wagstaff. 1998. A reexamination of North American Satureja s.l. (Lamiaceae) in light of molecular evidence. 

Brittonia 50: 63-70. 

Caplen, C.A., and C.R. Werth. 2000a. Isozymes of the Isoetes riparia complex, I. Genetic variation and relatedness of diploid 

species. Syst. Bot. 25: 235-159. 

–––, and C.R. Werth. 2000b. Isozymes of the Isoetes riparia complex, II. Ancestry and relationships of polyploids. Syst. Bot. 

25: 260-280. 

Carter, R. 1991. Cyperus entrerianus (Cyperaceae), an overlooked species in temperate North America. Sida 14: 69-77. 

–––, and C.T. Bryson. 2000. Cyperus sanguinolentus (Cyperaceae) new to the southeastern United States, and its relationship to 

the supposed endemic Cyperus louisianensis. Sida 19: 325-343. 

Carulli, J.P., and D.E. Fairbrothers. 1988. Allozyme variation in three eastern United States species of Aeschynomene 

(Fabaceae), including the rare A. virginica. Systematic Bot. 13: 559-566. 

–––, A.O. Tucker, and N.H. Dill. 1988. Aeschynomene rudis Benth. (Fabaceae) in the United States. Bartonia 54: 18-20. 

Carvell, W.N., and W.H. Eshbaugh. 1982. A systematic study of the genus Buckleya (Santalaceae). Castanea 47: 17-37. 

Case, F.W., Jr. 2005. Correction to the type citation of Sarracenia alabamensis and validation of the name Sarracenia 

alabamensis subsp. wherryi (Sarraceniaceae). Sida 21: 2169-2170. 

–––, and R.B. Case. 1976. The Sarracenia rubra complex. Rhodora 78: 270-325. 

–––, and R.B. Case. 1997. Trilliums. Timber Press, Portland, OR. 285 pp. 

Case, M.A., H.T. Mlodozeniec, L.E. Wallace, and T.W. Weldy. 1998. Conservation genetics and taxonomic status of the rare 

Kentucky lady's slipper: Cypripedium kentuckiense (Orchidaceae). Amer. J. Bot. 85: 1779-1786. 

Catalani, M. 2004. A field study of Sarracenia oreophila. Carnivorous Plant Newsletter 33: 6-12. 

Catling, P.M. 1983a. Spiranthes ochroleuca (Rydberg ex Britton) Rydberg (Yellow ladies'-tresses Orchid verified in North 

Carolina. Castanea 48: 48-49. 

–––. 1983b. Spiranthes ovalis var. erostellata (Orchidaceae), a new autogamous variety from the eastern United States. 

Brittonia 35: 120-125. 

–––. 1991. Systematics of Malaxis bayardii and M. unifolia. Lindleyana 6: 3-23. 

–––. 1998. A synopsis of the genus Proserpinaca in the southeastern United States. Castanea 63: 408-414. 

–––. 2004. A synopsis of the genus Hexalectris in the United States and a new variety of Hexalectris revoluta. Native Orchid 

Conference Journal 1: 5-25. 

–––, L. Dumouchel, and V.R. Brownell. 1998. Pollination of the Miccosukee gooseberry (Ribes echinellum). Castanea 63: 402- 

407. 

–––, and V.S. Engel. 1993. Systematics and distribution of Hexalectris spicata var. arizonica (Orchidaceae). Lindleyana 8: 

119-125. 

–––, and K.B. Gregg. 1992. Systematics of the genus Cleistes in North America. Lindleyana 7: 57-73. 

–––, S.M. McKay-Kuja, and G. Mitrow. 1999. Rank and typification in North American dwarf cherries, and a key to taxa. 

Taxon 48: 483-488. 

Caulkins, D.B., and R. Wyatt. 1990. Variation and taxonomy of Phytolacca americana and P. rigida in the southeastern United 

States. Bull. Torrey Bot. Club 117: 357-367. 

Chafin. L.G. 2000. Field guide to the rare plants of Florida. Florida Natural Areas Inventory, Tallahassee, FL. 

Chafin, L.G. 2006 {in prep.}. Field guide to the rare plants of Georgia. State Botanical Garden of Georgia, Athens. 

Chamberlain, D.F. 1982. A revision of Rhododendron. II. Subgenus Hymenanthes. Notes R.B.G. Edinb. 39: 209-486. 

Chambers, H. 1993. [add Pycnanthemum reference] 

–––, and J. Hamer. 1992. More about picky Pycnanthemums; can taxonomy be practical after all? Tipularia 7: 19-24. 

Chambers, K.L. 1989. The taxonomic relationships of Allocarya corallicarpa (Boraginaceae). Madroño 36: 280-281. 

–––. 2004. Taxonomic notes on Krigia (Asteraceae). Sida 21: 225-236. 

Channell, R.B. 1957. A revisional study of the genus Marshallia (Compositae). Contr. Gray Herbarium Harv. Univ. 181: 41- 

130. 

–––, and C.W. James. 1964. Nomenclatural and taxonomic corrections in Warea (Cruciferae). Rhodora 66: 18-26. 

–––, and C.E. Wood, Jr. 1959. The genera of the Primulales of the southeastern United States. J. Arnold Arb. 40: 268-288. 

–––, and C.E. Wood, Jr. 1962. The Leitneriaceae in the southeastern United States. J. Arnold Arb. 43: 435-438. 

–––, and C.E. Wood, Jr. 1987. The Buxaceae in the southeastern United States. J. Arnold Arb. 68: 241-257. 

Chapman, A.W. 1863. Flora of the southern United States: containing an abridged description of the flowering plants and ferns 

of Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the natural 

system, first edition. American Book Company, NY.


–––. 1878. An enumeration of some plants – chiefly from the semi-tropical regions of Florida – which are either new, or which 

have not hitherto been recorded as belonging to the flora of the southern states. Botanical Gazette 3: 2-6. 

–––. . 1883. Flora of the southern United States: containing an abridged description of the flowering plants and ferns of 

Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the natural system, 

second edition. American Book Company, NY. 

–––. 1897. Flora of the southern United States: containing an abridged description of the flowering plants and ferns of 

Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the natural system, 

third edition. American Book Company, NY. 

Chase, M.W., and 41 other authors. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid 

gene rbcL. Ann Mo. Bot. Gard. 80: 528-580. 

–––, D.E. Soltis, P.S. Soltis, P.J. Rudall, M.F. Fay, W.H. Hahn, S. Sullivan, J. Joseph, M. Molvray, P.J. Kores, T.J. Givnish, K.J. 

Sytsma, and J.C. Pires. 2000. Higher-level systematics of the monocotyledons: an assessment of current knowledge and a 

new classification. In: K.L. Wilson & D. A. Morrison, eds., Monocots: systematics and evolution. CSIRO, Melbourne. 

–––, S. Zmarzty, M.D. Lledó, K.J. Wurdack, S.M. Swensen, and M.F. Fay. 2002. When in doubt, put it in the Flacourtiaceae: a 

molecular phylogenetic analysis based on rbcL DNA sequences. Kew Bulletin 57: 141-181. 

Chaudhri, M.N. 1968. A revision of the Paronychiinae. Drukkerij H. Gianotten N.V., Tilburg. 440 pp. 

Cheek, M. 1994. The correct names for the subspecies of Sarracenia purpurea L. Carnivorous Plant Newletter 23: 69-73. 

–––. 2001. Good news: Drosera longifolia L. rejected, Sarracenia purpurea L. conserved with a new type. Carnivorous Plant 

Newsletter 30: 29-30. 

Chen, Chia Jui, M.G. Mendenhall, and B.L. Turner. 1994. Taxonomy of Thermopsis (Fabaceae) in North America. Ann. 

Missouri Bot. Gard. 81: 714-742. 

Cheplick, G.P. 1988. Influence of environment and population origin on survivorship and reproduction in reciprocal transplants 

of amphicarpic peanutgrass (Amphicarpum purshii). Am. J. Bot. 75: 1048-1056. 

–––. 1989. Nutrient availability, dimorphic seed production, and reproductive allocation in the annual grass Amphicarpum 

purshii. Can J. Bot. 67: 2514-2521. 

–––, and J.A. Quinn. 1982. Amphicarpum purshii and the "pessimistic strategy" in amphicarpic annuals with subterranean fruit. 

Oecologia 52: 327-332. 

–––, and J.A. Quinn. 1983. The shift in aerial/subterranean fruit ratio in Amphicarpum purshii: causes and significance. 

Oecologia 57: 374-379. 

–––, and J.A. Quinn. 1986. Self-fertilization in Amphicarpum purshii: its influence on fitness and variation in progeny from 

aerial panicles. Am. Midl. Nat. 116: 394-402. 

–––, and J.A. Quinn. 1987. The role of seed depth, litter, and fire in the seedling establishment of amphicarpic peanutgrass 

(Amphicarpum purshii). Oecologia 73: 459-464. 

–––, and J.A. Quinn. 1988a. Quantitative variation of life history traits in amphicarpic peanutgrass (Amphicarpum purshii) and 

its evolutionary significance. Am J. Bot. 75: 123-131. 

–––, and J.A. Quinn. 1988b. Subterranean seed production and population responses to fire in Amphicarpum purshii 

(Gramineae). J. Ecol. 76: 263-273. 

Chester, E.W., B.E. Wofford, R. Kral. 1997. Atlas of Tennessee vascular plants. Vol. 2. Angiosperms: dicots. Misc. Publ. No. 

13, Center for Field Biology, Austin Peay State Univ., Clarksville, TN. 242 pp. 

–––, B.E. Wofford, R. Kral, H.R. DeSelm, and A.M. Evans. 1993. Atlas of Tennessee vascular plants. Vol. 1. Pteridophytes, 

gymnosperms, angiosperms: monocots. Misc. Publ. No. 9, Center for Field Biology, Austin Peay State Univ., Clarksville, 

TN. 

Choi, H.-K., and J. Wen. 2000. A phylogenetic analysis of Panax (Araliaceae): integrating cp DNA restriction site and nuclear 

rDNA ITS sequence data. Plant Syst. Evol. 224: 109-120. 

Chuang, T.I., and L. Constance. 1977. Cytogeography of Phacelia ranunculacea (Hydrophyllaceae). Rhodora 79: 115-122. 

Church, G.L. 1967. Taxonomic and genetic relationships of eastern North American species of Elymus with setaceous glumes. 

Rhodora 69: 121-162. 

Church, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneuploidy and breeding system evolution 

in the radiation of the lineage across North America. Molecular Phylogenetics and Evolution 27: 223-238. 

–––, and D.R. Taylor. 2005. Speciation and hybridization among Houstonia (Rubiaceae) species: the influence of polyploidy on 

reticulate evolution. Amer. J. Botany 92: 1372-1380. 

Churchill, J.A., and E. Schell. 1992. Noteworthy collections: Tennessee. Castanea 57: 293. 

–––, N. Churchill, M.J. Waterway, S. de Blois, and C. Schell. 1992. Noteworthy collections: Tennessee. Castanea 57: 151. 

Cialdella, A.M., and L.M. Giussani. 2002. Phylogenetic relationships of the genus Piptochaetium (Poaceae, Pooideae, Stipeae): 

evedence from morphological data. Ann. Missouri Bot. Gard. 89: 305-336 

Clancy, K., and M. Sullivan. 1990. Distribution of the needle palm, Rhapidophyllum hystrix. Castanea 55: 31-39. 

Clark, R.B. 1942. A revision of the genus Bumelia in the United States. Ann. Missouri Bot. Gard. 29:155-182. 

Clark, R.C. 1971. The woody plants of Alabama. Annals Missouri Bot. Garden 58: 99-242. 

–––, C.G. Hewins, J.D. Husband, C.T. Kirk, and R.W. Long. 1997. Noteworthy collections: Kentucky. Castanea 62: 288. 

–––, R.L. Jones, T.J. Weckman, R.L. Thompson, J.W. Thieret, Kentucky Nature Preserves Commission, and K. Freeman. 2005. 

State records and other noteworthy collections for Kentucky. Sida 21: 1909-1916. 

Clausen, R.T. 1939. Silene caroliniana. Rhodora 41: 575-584.


–––. 1975. Sedum of North America north of the Mexican plateau. Cornell Univ. Press, Ithaca, NY. 

Clay, K. 1983. Myrmechory in the trailing arbutus (Epigaea repens L.). Bull. Torrey Bot. Club 110: 166-169. 

–––. 1995. Noteworthy collections: North Carolina. Castanea 60: 84-85. 

Clayton, W.D., and S.A. Renvoize. 1986. Genera graminum; grasses of the world. Kew Bulletin Additional Series 13. Her 

Majesty's Stationery Office, London. 

Clemants, S.E. 1990. Juncaceae (Rush Family) of New York State. New York State Museum Bulletin 475: 1-67. 

Clements, R.K., J.M. Baskin, and C.C. Baskin. 1998. The comparative biology of the two closely-related species Penstemon 

tenuiflorus Pennell and P. hirsutus (L.) Willd. (Scrophulariaceae, section Graciles): I. Taxonomy and geographical 

distribution. Castanea 63: 138-153. 

Clevinger, J.A. 2004. New combinations in Silphium (Asteraceae; Heliantheae). Novon 14: 275-277. 

Clewell, A.F. 1966a. Native North American species of Lespedeza (Leguminosae). Rhodora 68: 359-405. 

–––. 1966. Identification of the Lespedezas in North America. Bull. Tall Timbers Research Station 7. 

–––. 1985. Guide to the vascular plants of the Florida Panhandle. University Presses of Florida, Tallahassee, FL. 605 pp. 

–––. 1990. Establishment of Lespedeza virgata (Leguminosae) in the southeastern United States. J. Elisha Mitchell Sci. Soc. 

106: 32-37. 

–––, and J.W. Wooten. 1971. A revision of Ageratina (Compositae: Eupatorieae) from eastern North America. Brittonia 23: 

123-143. 

Cochrane, T.S. 1976. Taxonomic status of the Onosmodium molle complex (Boraginaceae) in Wisconsin. Michigan Botanist 

15: 103-110. 

Coffey, V.J., and S.B. Jones, Jr. 1980. Biosystematics of Lysimachia section Seleucia (Primulaaceae). Brittonia 32: 309-322. 

Coile, N.C. 1988. Taxonomic studies on the deciduous species of Ceanothus L. (Rhamnaceae). Ph.D. dissertation, Univ. of 

Georgia, Athens. 

Coker, W.C. 1919. The distribution of Rhododendron catawbiense, with remarks on a new form. J. Elisha Mitchell Sci. Soc. 

25: 76-82. 

–––. 1943. Magnolia cordata Michaux. J. Elisha Mitchell Sci. Soc. 59: 81-88. 

–––, and H.R. Totten. 1945. Trees of the southeastern United States, including Virginia, North Carolina, South Carolina, 

Tennessee, Georgia, and northern Florida. Univ. of North Carolina Press, Chapel Hill, NC. 

Coleman, J.R. 1966. A taxonomic revision of section Ximenesia of the genus Verbesina (Compositae). Amer. Midl. Naturalist 

76: 475-481. 

Collins, J.L. 1976. A revision of the annulate Scutellaria (Labiatae). Ph.D. dissertation, Vanderbilt Univ. 

–––, and T.F. Wieboldt. 1992. Trifolium calcaricum (Fabaceae), a new clover from limestone barrens of eastern United States. 

Castanea 57: 282-286. 

Collins, S.L., and W.H. Blackwell, Jr. 1979. Bassia (Chenopodiaceae) in North America. Sida 8: 57-64. 

Columbus, J.T. 1999. An expanded circumscription of Bouteloua (Gramineae: Chloridoideae): new combinations and names. 

Aliso 18: 61-65. 

Compton, J.A., A. Culham, and S.L. Jury. 1998. Reclassification of Actaea to include Cimicifuga and Souliea (Ranunculaceae): 

phylogeny inferred from morphology, nrDNA, ITS, and cpDNA trnL-F sequence variation. Taxon 47: 593-634. 

Constance, L. 1940. The genus Ellisia. Rhodora 42: 33-39. 

–––. 1941. The genus Nemophila Nutt. Univ. of Cal. Publ. in Bot. 19: 341-398. 

–––. 1942. The genus Hydrophyllum L. Amer. Midl. Nat. 27: 710-731. 

–––. 1949. A revision of Phacelia subgenus Cosmanthus (Hydrophyllaceae). Contr. Gray Herb. 168: 1-48. 

–––. 1963. Chromosome number and classification in Hydrophyllaceae. Brittonia 15: 273-285. 

Cook, R.E., and J.C. Semple. 2004. A new name and a new combination in Solidago subsect. Glomeruliflorae (Asteraceae: 

Astereae). Sida 21: 221-244. 

Cooper, A.W., and E.P. Mercer. 1977. Morphological variation in Fagus grandifolia Ehrh. in North Carolina. J. Elisha Mitch. 

Sci. Soc. 93: 136-149. 

Cooperrider, T.S. 1985. Thaspium and Zizia in Ohio. Castanea 50: 116-119. 

–––. 1995. The Dicotyledonae of Ohio. Part 2. Linaceae through Campanulaceae. Ohio State University Press, Columbus. 656 

pp. 

–––, and G.A. McCready. 1975. On separating Ohio specimens of Lindernia dubia and L. anagallidea (Scrophulariaceae). 

Castanea 40: 191-197. 

Core, E.L. 1936. The American species of Scleria. Brittonia 2: 1-105. 

Correa, M.D., and R.L. Wilbur. 1969. A revision of the genus Carphephorus (Compositae-Eupatoriae). J. Elisha Mitch. Sci. 

Soc. 85: 79-91. 

Correll, D.S. 1936. Epidendrum conopseum in North Carolina. J. Elisha Mitchell Sci. Soc. 52: 91-92. 

–––. 1937. The orchids of North Carolina. J. Elisha Mitchell Sci. Soc. 53: 139-172. 

–––. 1950. Native orchids of North America north of Mexico. Chronica Botanica Cp., Waltham, MA. 399 pp. 

–––, and H.B. Correll. 1982. Flora of the Bahama Archipelago (including the Torks and Caicos Islands). J. Cramer, Vaduz. 

1692 pp. 

Costea, M., G.L. Nesom, and S. Stefanović. 2006a. Taxonomy of the Cuscuta pentagona complex (Convolvulaceae) in North 

America. Sida 22: 151-176.


–––, G.L. Nesom, and S. Stefanović. 2006b. Taxonomy of the Cuscuta gronovii and Cuscuta umbrosa (Convolvulaceae). Sida 

22: 197-208. 

–––, G.L. Nesom, and S. Stefanović. 2006c. Taxonomy of the Cuscuta indecora (Convolvulaceae) complex in North America. 

Sida 22: 209-226. 

–––, A. Sanders, and G. Waines. 2001a. Preliminary results toward a revision of the Amaranthus hybridus species complex 

(Amaranthaceae). Sida 19: 931-974. 

–––, A. Sanders, and G. Waines. 2001b. Notes on some little known Amaranthus taxa (Amaranthaceae) in the United States. 

Sida 19: 975-992. 

–––, and F.J. Tardif. 2003a. Nomenclatural changes in the genus Polygonum section Polygonum (Polygonaceae). Sida 20: 987- 

997. 

–––, and F.J. Tardif. 2003b. Conspectus and notes on the genus Amaranthus in Canada. Rhodora 105: 260-281. 

–––, and F.J. Tardif. 2003c. Polygonum aviculare subsp. rurivagum (Polygonaceae) in North America. Sida 20: 1709-1711. 

Coulter, J.M., and J.N. Rose. 1900. Monograph of the North American Umbelliferae. Contr. U.S. Nat. Herb. 7: 1-256. 

Crane, E.H. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic Bot. 22: 509-517. 

Cranfill, R. 1983. The distribution of Woodwardia areolata. Amer. Fern J. 73: 46-52. 

Crawford, D.J., and E.B. Smith. 1984. Allozyme divergence and intraspecific variation in Coreopsis grandifolia (Compositae). 


–––, and M.E. Mort. 2005. Phylogeny of eastern North American Coreopsis (Asteraceae – Coreopsideae): insights from nuclear 

and plastid sequences, and comments on character evolution. Amer. J. Bot. 92: 330-336. 

Crins, W.J. 1989a. Status of the few-flowered club-rush, Scirpus verecundus (Cyperaceae), in Canada. Can. Field-Naturalist 

103: 57-60. 

–––. 1989b. The Tamaricaceae in the southeastern United States. J. Arnold Arb. 70: 403-425. 

–––. 1991. The genera of Paniceae (Gramineae: Panicoideae) in the southeastern United States. J. Arnold Arb., Supplementary 

Series 1: 171-312. 

–––, and P.W. Ball. 1983. The taxonomy of the Carex pensylvanica complex (Cyperaceae) in North America. Can. J. Bot. 61: 

1692-1717. 

Cronquist, A. 1945. Studies in the Sapotaceae, III. Dipholis and Bumelia. J. Arnold Arbor. 26:435-471. 

–––. 1947. Notes on the Compositae of the northeastern United States – V. Astereae. Bull. Torrey Bot. Club 74: 142-150. 

–––. 1980. Asteraceae, Volume I, Vascular flora of the Southeastern United States. University of North Carolina Press, Chapel 

Hill, N.C. 

–––. 1981. An integrated system of classification of flowering plants. New York Botanical Garden, N.Y. 1262 pp. 

–––. 1982. Reduction of Pseudotaenidia to Taenidia (Apiaceae). Brittonia 34: 365-367. 

–––. 1985. Eupatorium godfreyanum (Asteraceae), a "new" species from eastern United States. Brittonia 37: 237-242. 

Croom, H.B. 1837. A catalogue of plants, native or naturalized, in the vicinity of New Bern, North Carolina, with remarks and 

synonyms. G.P. Scott, New York, NY. 

Crow, G.E. 1978. A taxonomic revision of Sagina (Caryophyllaceae) in North America. Rhodora 80: 1-91. 

–––, and C.B. Hellquist. 2000a. Aquatic and wetland plants of northeastern North America: a revised and enlarged edition of 

Norman C. Fassett's A Manual of Aquatic Plants. Vol. 1. Pteridophytes, gymnosperms, and angiosperms: dicotyledons. 

Univ. of Wisconsin Press, Madison, WI. 

–––, and C.B. Hellquist. 2000b. Aquatic and wetland plants of northeastern North America: a revised and enlarged edition of 

Norman C. Fassett's A Manual of Aquatic Plants. Vol. 2. Angiosperms: monocotyledons. Univ. of Wisconsin Press, 

Madison, WI. 

Cruden, R.W. 1991. A revision of Isidrogalvia (Liliaceae): recognition of Ruíz and Pavón's genus. Systematic Bot. 16: 270- 

282. 

Cullen, J. 1980. A revision of Rhododendron. I. Subgenus Rhododendron sections Rhododendron & Pogonanthum. Notes 

R.B.G. Edinb. 39: 1-207. 

Cullings, K.W., and L. Hileman. 1997. The Monotropoideae is a monophyletic sister group to the Arbutoideae (Ericaceae): a 

molecular test of Copeland's hypothesis. Madroño 44: 297-304. 

Culwell, D.E. 1970. A taxonomic study of the section Hypericum in the eastern United States. Ph.D. dissertation, University of 

North Carolina at Chapel Hill. 

Curtis, M.A. 1843. An account of some new and rare plants of North Carolina. Amer J. Sci. 44: 80-84. 

–––. 1860. The woody plants of North Carolina. Holden, Raleigh NC 

Cusick, A.W. 1985. Lithospermum (Boraginaceae) in Ohio, with a new taxonomic rank for Lithospermum croceum Fernald. 

Mich. Botanist 24: 63-69. 

–––. 1987. A binomial for a common hybrid Lycopodium. Amer. Fern J. 77: {}. 

–––. 1992. Carex section Acrocystis (Cyperaceae) in Ohio. Michigan Botanist 31: 99-108. 

–––. 1994. Noteworthy collections: West Virginia. Castanea 59: 79-80. 

–––. 1996. Notes on the genus Carex (Cyperaceae) in West Virginia. Castanea 61: 161-167. 

–––. 2002. A binomial for the hybrid Polypodium of eastern North America. Amer. Fern J. 92: 240-241. 

D'Arcy, W.G., and E.H. Eshbaugh. 1974. New World peppers [Capsicum - Solanaceae] north of Colombia: a résumé. Baileya 

19: 93-105. 

Dahlgren, R.M.T., and H.T. Clifford. 1982. The monocotyledons: a comparative study. Academic Press, London. 378 pp.


–––, H.T. Clifford, and P.F. Yeo. 1985. The families of the monocotyledons: structure, evolution, and taxonomy. Springer- 

Verlag, Berlin. 520 pp. 

Dane, F., and P. Lang. 2004. Sequence variation at cpDNA regions of watermelon and related wild species: implications for the 

evolution of Citrullus haplotypes. Amer. J. Bot. 91: 1922-1929. 

Danin, A., and L.C. Anderson. 1986. Distribution of Portulaca oleracea L. (Portulacaceae) subspecies in Florida. Sida 11: 318- 

324. 

Daoud, H.S., and R.L. Wilbur. 1965. A revision of the North American species of Helianthemum (Cistaceae). Rhodora 67: 63- 

312 (pagination interrupted). 

Darbyshire, S.J. 1993. Realignment of Festuca subgenus Schedonardus with the genus Lolium (Poaceae). Novon 3: 239-243. 

–––, and L.E. Pavlick. 1997. Nomenclatural notes on North American grasses. Phytologia 82: 73-78. 

Davidian, H.H. 1982. The Rhododendron species. Volume I: Lepidotes. Timber Press, Portland, OR. 

Davidson, J.F. 1950. The genus Polemonium (Tournefort) L. Univ. California Publ. Bot. 23: 209-282. 

Davison, P.G. 1997. Noteworthy collections: Georgia and South Carolina. Castanea 62: 129. 

Davies, P.A. 1952. Geographical variation in Shortia galacifolia. Rhodora 54: 121-124. 

Davis, R.J. The North American perennial species of Claytonia. Brittonia 18: 285-303. 

de Wet, J.M.J. 1978. Systematics and evolution of Sorghum sect. Sorghum (Gramineae). Amer. J. Bot. 65: 477-484. 

–––, J.R. Harlan, and D.E. Brink. 1982. Systematics of Tripsacum dactyloides (Gramineae). Amer. J. Botany 69: 1251-1257. 

Decker-Walters, D.S., S.-M. Chung, J.E. Staub, H.D. Quemada, and A.I. López-Sesé. 2002. The origin and genetic affinities of 

wild populations of melon (Cucumis melo, Cucurbitaceae) in North America. Plant. Syst. Evol. 233: 183-197. 

Degtjareva, G.V., T.E. Kramina, D.D. Sokoloff, T.H. Samigullin, C.M. Valiejo-Roman, and A.S. Antonov. 2006. Phylogeny of 

the genus Lotus (Leguminosae, Loteae): evidence from nrITS sequaences and morphology. Can. J. Bot. 84: 813-830. 

DeJong, D.C.D. 1965. A systematic study of the genus Astranthium (Compositae, Astereae). Publ. Mus. Michigan State Univ. 

Biol. Ser. 2: 429-528. 

del Castillo, R. F. 1994. Factors influencing the genetic structure of Phacelia dubia, a species with a seed bank and large 

fluctuations in population size. Heredity 72: 446-458. 

–––. 1998. Fitness consequences of maternal and nonmaternal components of inbreeding in the gynodioecious Phacelia dubia. 

Evolution 52: 44-60. 

Delahoussaye, A.J., and J.W. Thieret. 1967. Cyperus subgenus Kyllinga (Cyperaceae) in the continental United States. Sida 3: 

128-136. 

DeLaney, K.R., N. Bissett, and J.D. Weidenhamer. 1999. A new species of Carphephorus (Asteraceae; Eupatorieae) from 

peninsular Florida. The Botanical Explorer 1: 1-15. 

–––, R.P. Wunderlin, and J.C. Semple. 2003. Chrysopsis delaneyi (Asteraceae, Astereae), another new species from peninsular 

Florida. Botanical Explorer 3: 1-37. 

Dellinger, B. 1989. Noteworthy collections: North Carolina: Trientalis borealis. Castanea 54: 127. 

Dempster, L.T. 1978. The genus Galium (Rubiaceae) in Mexico and Central America. Univ. of Calif. Publ. in Botany 73: 1-33 

–––. 1981. The genus Galium (Rubiaceae) in South America. II. Allertonia 2: 393-426. 

Dennis, W.M. 1980. Sarracenia oreophila (Kearney) Wherry in the Blue Ridge Province of northeastern Georgia. Castanea 45: 

101-103. 

–––, and D.H. Webb. 1981. The distribution of Pilularia americana A. Br. (Marsileaceae) in North America, north of Mexico. 

Sida 9: 19-24. 

DePoe, C.E., and E.O. Beal. 1969. Origin and maintenance of clinal variation in Nuphar (Nymphaeaceae). Brittonia 21: 15-28. 

Des Marais, D.L., A. R. Smith, D.M. Britton, and K.M. Pryer. 2003. Phylogenetic relationships and evolution of extant 

horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F). Int. J. Plant Sci. 164: 737-751. 

Detling, L.E. 1939. A revision of the North American species of Descurainia. Amer. Midland Nat. 22: 481-520. 

DeVore, M.L. 1991. The occurrence of Acicarpha tribuloides (Calyceraceae) in eastern North America. Rhodora 93: 26-35. 

Dhillion, S.S., and R.C. Anderson. 1999. Growth and photosynthetic response of first-year garlic mustard (Alliaria petiolata) to 

varied irradiance. J. Torrey Bot. Soc. 126: 9-14. 

Diamond, A.R., Jr., and R.S. Boyd. 2004. Distribution, habitat characteristics amd population trends of the rare southeastern 

endemic Rudbeckia auriculata (Perdue) Kral (Asteraceae). Castanea 69: 249-264. 

Diamond, P. 1999. Paederia foetida (Rubiaceae), new to the flora of North Carolina. Sida 18: 1273-1276. 

Diane, N., H. Förther, & H.H. Hilger. 2002. A systematic analysis of Heliotropium, Tournefortia, and allied taxa of the 

Heliotropiaceae (Boraginales) based on ITS1 sequences and morphological data. Amer. J. Botany 89: 287-295 

Dibble, A.C., and C.S. Campbell. 1995. Distribution and conservation of Nantucket shadbush, Amelanchier nantucketensis 

(Rosaceae). Rhodora 97: 339-349. 

Dietrich, W., and W.L. Wagner. 1988. Systematics of Oenothera section Oenothera subsection Raimannia and subsection 

Nutantigemma (Onagraceae). Systematic Bot. Monographs 24: 1-91. 

Dietrich, W., W.L. Wagner, and P.H. Raven. 1997. Systematics of Oenothera section Oenothera subsection Oenothera 

(Onagraceae). Systematic Bot. Monographs 50: 1-234. 

Digital Flora of Texas. 2005. Texas vascular plant image gallery. http://www.csdl.tamu.edu/FLORA/gallery.htm . Accessed 5 

December 2005. 

Dirr, M.A. 2004. Hydrangeas for American gardens. Timber Press, Portland. 236 pp. 

Dore, W.G. 1964. Two kinds of blue cohosh. Ontario Naturalist.


Dorn, R.D. 1984. Vascular plants of Wyoming. Mountain West Publishing, Cheyenne, WY. 

–––. 1988. Vascular plants of Wyoming, second edition. Mountain West Publishing, Cheyenne, WY. 

–––. 1995. A taxonomic study of Salix section Cordatae subsection Luteae (Salicaceae). Brittonia 47: 160-174. 

–––. 2001. Vascular plants of Wyoming, third edition. Mountain West Publishing, Cheyenne, WY. 

Dorr, L.J. 1990. A revision of the North American genus Callirhoe (Malvaceae). Mem. New York Bot. Garden 56: 1-76. 

–––, and F.R. Barrie. 1993. Typification of the Linnaean names in Pyrola (Ericaceae, Pyroloideae). Brittonia 45: 177-180. 

Douglass, C.C. 1980. Waldsteinia lobata (Baldw.) T. & G. (Rosaceae) verified for South Carolina. Castanea 45: 228-232. 

Downer, R.G., and P.E. Hyatt. 2003. Recommendations concerning the identification of Carex retroflexa and Carex texensis 

(Cyperaceae; section Phaestoglochin Dumort.). Castanea 68: 245-253. 

Downie, S.R., and J.D. Palmer. 1992. Restriction site mapping of the chloroplast DNA inverted repeat: a molecular phylogeny 

of the Asteridae. Ann. Missouri Bot. Gard. 79: 266-283. 

–––, S. Ramanath, D.S. Katz-Downie, and E. Llanas. 1998. Molecular systematics of Apiaceae subfamily Apioideae: 

phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer and plastid RPOC1 intron sequences. Am. J. 

Bot. 85: 563-591. 

Doyle, J.D. 1990. Systematics of the Opuntia humifusa complex. Ph.D. dissertation, University of North Carolina at Chapel 

Hill. 

Drábková, L., J. Kirschner, O. Seberg, G. Petersen, and Č. Vlček. 2003. Phylogeny of the Juncaceae based on rbcL sequences, 

with special emphasis on Luzula DC. and Juncus L. Plant Syst. Evol. 240: 133-147. 

Drapalik, D.J. 1969. A biosystematic study of the genus Matelea in the southeastern United States. Ph.D. dissertation, 

University of North Carolina, Chapel Hill. 225 pp. 

Dubuisson, J.-Y., S. Hennequin, E.J.P. Douzery, R.B. Cranfill, A.R. Smith, and K.M. Pryer. 2003. rbcL phylogeny of the fern 

genus Trichomanes (Hymenophyllaceae), with special reference to neotropical taxa. Int. J. Plant Sci. 164: 753-761. 

Dudley, T.R. 1974. The correct authority for Cardamine clematitis (Cruciferae). Rhodora 76: 53-57. 

Duistermaat, H. 1996. Monograph of Arctium L. (Asteraceae): generic delimitation (including Cousinia Cass. p.p.), revision of 

the species, pollen morphology, and hybrids. Gorteria Spplement 3, Rijksherbarium, Leiden. 

Duke, J.A. 1955. Distribution and speciation of the genus Ludwigia in North Carolina. J. Elisha Mitchell Sci. Soc. 71: 255-269. 

–––. 1961. Preliminary revision of the genus Drymaria. Ann. Mo. Bot. Gard. 48: 173-268. 

Duley, M.L., and M.A. Vincent. 2003. A synopsis of the genus Cladrastis (Leguminosae). Rhodora 105: 205-239. 

Duncan, T. 1980. A taxonomic study of the Ranunculus hispidus Michaux complex in the Western Hemisphere. Univ. of 

California Publications in Botany, vol. 77. 

Duncan, W.H. 1967. Woody vines of the southeastern states. Sida 3: 1-76. 

–––. 1969. Celastrus (Celastraceae) in the southeastern states. Sida 3: 309-310. 

–––. 1979. Changes in Galactia (Fabaceae) of the southeastern United States. Sida 8: 170-180. 

–––. 1985. Ten additions to the vascular flora of Georgia. Castanea 50: 52-55. 

–––, and N.E. Brittain. 1966. The genus Gaylussacia (Ericaceae) in Georgia. Bull. Georgia Academy of Sci. 24: 13-26. 

–––, and D.W. Dejong. 1964. Taxonomy and heterostyly of North American Gelsemium (Loganiaceae). Sida 1: 346-357. 

–––, and M.B. Duncan. 1988. Trees of the southeastern United States. University of Georgia Press, Athens. 

–––, and M.B. Duncan. [in prep.]. Shrubs of the southeastern United States. 

–––, and R.B. McCartney. 1992. About Lupinus cumulicola (Fabaceae). Sida 15: 346-347. 

–––, and T.M. Pullen. 1962. Lepidote Rhododendrons of the southeastern United States. Brittonia 14: 290-298. 

Duvall, M.R., and 10 other authors. 1993. Phylogenetic hypotheses for the monocotyledons constructed from rbcL sequence 

data. Ann. Mo. Bot. Gard. 80: 607-619. 

Easterly, N.W. 1957. A morphological study of Ptilimnium. Brittonia 9: 136-145. 

Ebinger, J.E. 1974. A systematic study of the genus Kalmia (Ericaceae). Rhodora 76: 315-398. 

–––, D.S. Seigler, and H.D. Clarke. 2002. Notes on the segregates of Acacia farnesiana (L.) Willd. (Fabaceae: Mimoisoideae) 

and related species in North America. Southwestern Naturalist 47: 86-91. 

EckenwalderJ.E. 1977. North American cottonwoods (Populus, Salicaceae) of sections Abaso and Aigeiros. J. Arnold Arb. 58: 

193-208. 

–––. 1984. Natural intersectional hybrids between North American species of Populus (Salicaceae) in sections Aigeiros and 

Tacamahaca. II. Taxonomy. Can. J. Bot. 62: 325-335. 

–––. 1996. Systematics and evolution of Populus. In Stettler, R.F., H.D. Bradshaw, Jr., P.E. Heilman, and T.M. Hinckley, eds. 

Biology of Populus and its implications for management and conservation. NRC Research Press, Ottawa. 

Eddie, W.M.M., T. Shulkina, J. Gaskin, R.C. Haberle, and R.K. Jansen. 2003. Phylogeny of Campanulaceae s. str. inferred from 

ITS sequences of nuclear ribosomal DNA. Ann. Missouri Bot. Gard. 554-575. 

Edmondson, J.R. 2005. A new combination in Oxypolis Rafinesque (Apiaceae). Novon 15: 109. 

Edwards, J.M., J.A. Churchill, and U. Weiss. 1970. A chemical contribution to the taxonomic status of Lophiola americana. 

Phytochem. 9: 1563-1564. 

Ehdaie, M., and S.D. Russell. 1984. Megagametophyte development of Nandina domestica and its taxonomic implications. 

Phytomorphology 34: 221-225. 

Eigsti, O.J. 1942. A cytological investigation of Polygonatum using the colchicine-pollen tube technique. Am. J. Bot. 29: 626- 

636.


Eiten, G. 1963. Taxonomy and regional variation of Oxalis section Corniculatae. I. Introduction, keys and synopsis of species. 

Amer. Midl. Nat. 69: 257-309. 

Eleuterius, L.N. 1977?. A revised description of the salt-marsh rush, Juncus roemerianus. Sida 7: 355-360. 

Elias, T.S. 1971a. The genera of Fagaceae in the southeastern United States. J. Arnold Arb. 52: 159-195. 

–––. 1971b. The genera of Myricaceae in the southeastern United States. J. Arnold Arb. 52: 305-318. 

–––. 1972. The genera of Juglandaceae in the southeastern United States. J. Arnold Arb. 53: 26-51. 

Ellison, A.M., H.L. Buckley, T.E. Miller, and N.J. Gotelli. 2004. Morphological variation in Sarracenia purpurea 

(Sarraceniaceae): geographic, environmental, and taxonomic correlates. Amer. J. Bot. 91: 1930-1935. 

Epling, C. 1942. The American species of Scutellaria. Univ. Calif. Publ. in Botany 20: 1-146. 

Eriksson, T., and M.J. Donoghue. 1997. Phylogenetic relationships of Sambucus and Adoxa (Adoxoideae, Adoxacaeae) based 

on nuclear ribosomal ITS sequences and preliminary morphological data. Systematic Bot. 22: 555-573. 

–––, M.J. Donoghue, and M.S. Hibbs. 1998. Phylogenetic analysis of Potentilla using DNA sequences of nuclear ribosomal 

internal transcribed spacers (ITS), and implications for the classification of Rosoideae (Rosaceae). Pl. Syst. Evol. 211: 155- 

179. 

–––, M.S. Hibbs, A.D. Yoder, C.F. Delwiche, and M.J. Donoghue. 2003. The phylogeny of Rosoideae (Rosaceae) based on 

sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA and the trnL/F region of chloroplast DNA. 

Int. J. Plant Sci. 164: 197-211. 

Ertter, B. 2000. Floristic surprises in North America north of Mexico. Ann. Missouri Bot. Garden. 87: 81--109. 

Esselman, E.J., and D.J. Crawford. 1997. Molecular and morphological evidence for the origin of Solidago albopilosa 

(Asteraceae), a rare endemic of Kentucky. Systematic Bot. 22: 245-257. 

Esser, H.-J. 2002. A revision of Triadica Lour. (Euphorbiaceae). Harvard Papers in Botany 7: 17-21. 

Essig, F.B. 1990. The Clematis virginiana (Ranunculaceae) complex in the Southeastern United States. Sida 14: 49-68. 

Estes, D. 2004. Noteworthy records: middle Tennessee. Castanea 69: 69-74. 

–––, and J. Beck. 2005. Sporobolus heterolepis (Poaceae) new to Tennessee. Sida 21: 1923-1926. 

–––, and C. Fleming. 2006. Clematis morefieldii (Ranunculaceae) new to Tennessee. Sida 22: 821-824. 

Evert, D.S. 1957. Dionaea transplants in the New Jersey Pine Barrens. Bartonia 29: 3-4. 

Evrard, C., and C. Van Hove. 2004. Taxonomy of the American Azolla species (Azollaceae): a critical review. Syst. & Geogr. 

Pl. 74: 301-318. 

Eyde, R.H. 1966. The Nyssaceae in the Southeastern United States. J. Arnold Arb. 47: 117-125. 

–––. 1977. Reproductive structures and evolution in Ludwigia (Onagraceae). I. Androecium, placentation, merism. Ann. Mo. 

Bot. Gard. 64: 644-655. 

–––. 1978. Reproductive structures and evolution in Ludwigia (Onagraceae). II. Fruit and seed. Ann. Mo. Bot. Gard. 65: 656- 

675. 

–––. 1981. Reproductive structures and evolution in Ludwigia (Onagraceae). III. Vasculature, nectaries, conclusions. Ann. 

Mo. Bot. Gard. 68: 379-412. 

–––. 1987. The case for keeping Cornus in the broad Linnaean sense. Systematic Botany 12: 505-518. 

Faber-Langendoen, D., and D. Tart. 2001. Proposal for revisions to the national standards for physiognomic levels of vegetation 

classification in the United States. Federal Geographic Data Committee, Vegetation Subcommitee, Washington, DC. 

Fairbrothers, D.E., and J.R. Gray. 1972. Microstegium vimineum (Trin.) A. Camus (Gramineae) in the United States. Torreya 

99: 97-100. 

Fairey, J.E, III. 1967. The genus Scleria in the southeastern United States. Castanea 32: 37-71. 

–––, & A.T. Whittemore. 1999. Proposal to conserve the name Scleria pauciflora (Cyperaceae) with a conserrved type. Taxon 

48: 575-576. 

Fan, C., and Q.-Y. Xiang. 2001. Phylogenetic relationships within Cornus (Cornaceae) based on 26S rDNA sequences. Amer. 

J. Bot. 88: 1131-1138. 

Fantz, P.R. 2000. Nomenclatural notes on the genus Clitoria for the Flora North American project. Castanea 65: 89-92. 

–––. 2002a. Distribution of Centrosema (Leguminosae: Phaseoleae: Clitoriinae) for the Flora of North America project. Vulpia 

1: 41-81. 

–––. 2002b. Distribution of Clitoria (Leguminosae: Phaseoleae: Clitoriinae) for the Flora of North America project. Vulpia 1: 

82-132. 

Farjon, A. 1998. World checklist and bibliography of conifers. Royal Botanic Gardens, Kew, England. 

–––. 2005. A monograph of Cupressaceae and Sciadopitys. Royal Botanic Gardens, Kew. 643 pp. 

Farmer, S.B., and E.E. Schilling. 2002. Phylogenetic analyses of Trilliaceae based on morphological and molecular data. 

Systematic Botany 27: 674-692. 

Farrar, D.R. 1974. Gemmiferous fern gametophytes – Vittariaceae. Am. J. Bot. 61: 146-155. 

Farrar, D.R. 1978. Problems in the identity and origin of the Appalachian Vittaria gametophyte, a sporophyteless fern of the 

eastern United States. Am. J. Bot. 65: 1-12. 

–––. 1992. Trichomanes intricatum: the independent Trichomanes gametophyte in the eastern United States. Amer. Fern J. 82: 

68-74. 

–––, and J.T. Mickel. 1991. Vittaria appalachiana: a name for the "Appalachian Gametophyte." Amer. Fern J. 81: 69-75. 

–––, J.C. Parks, and B.W. McAlpin. 1983. The fern genera Vittaria and Trichomanes in the northeastern United States. 

Rhodora 85: 83-92.


–––, and J.F. Wendel. 1996. Eastern moonworts: genetics and relationships [abstract]. Am. J. Bot. 83: 124. 

Fassett, N.C. 1935. A study of Streptopus. Rhodora 37: 88-113. 

–––. 1944. Dodecatheon in eastern North America. Amer. Midland Naturalist 31: 455-486. 

Fay, M.F., and M.W. Chase. 1996. Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, 

Amaryllidaceae and Agapanthoideae. Taxon 45: 441-451. 

–––, P.J. Rudall, S. Sullivan, K.L. Stobart, A.Y. de Bruijn, G. Reeves, F. Qamaruz-Zaman, W.-P. Hong, J. Joseph, W.J. Hahn, 

J.G. Conran, and M.W. Chase. 2000. Phylogenetic studies of Asparagales based on four plastid DNA regions. In: K.L. 

Wilson & D. A. Morrison, eds., Monocots: systematics and evolution. CSIRO, Melbourne. 

Ferguson, C.J., F. Krämer, and R.K. Jansen. 1999. Relationships of eastern North American Phlox (Polemoniaceae) based on 

ITS sequence data. Systematic Bot. 24: 616-631. 

Ferguson, D.M. 1998. Phylogenetic analysis and relationships in Hydrophyllaceae based on ndhF sequence data. Systematic 

Botany 23: 253-268. 

Ferguson, I.K. 1965. The genera of Valerianaceae and Dipsacaceae in the southeastern United States. J. Arnold Arb. 46: 218- 

231. 

–––. 1966a. The genera of Caprifoliaceae in the southeastern United States. J. Arnold Arb. 47: 33-59. 

–––. 1966b. The genera of Sterculiaceae in the southeastern United States. J. Arnold Arb. 47: 60-74. 

–––. 1966c. Notes on the nomenclature of Cornus. J. Arnold. Arb. 47: 100-105. 

–––. 1966d. The Cornaceae in the southeastern United States. J. Arnold Arb. 47: 106-116. 

–––, and G.K. Brizicky. 1965. Nomenclatural notes on Dipsacus fullonum and Dipsacus sativus. J. Arnold Arb. 46: 362-365. 

Ferguson, R.L., J.A. Rivera, and L.L. Wood. 1989. Submerged aquatic vegetation in the Albemarle-Pamlico estuarine system. 

Albemarle-Pamlico Estuarine Study Project No. 88-10. 

Fernald, M.L. 1911. The northern variety of Gaylussacia dumosa. Rhodora 13: 95-99. 

–––. 1943. Virginian botanizing under restrictions. Rhodora 45: 357-511 (pagination interrupted) 

–––. 1950. Gray's manual of botany, eighth (centennial) edition. Corrected printing, 1970. D. Van Nostrand Co., New York, 

N.Y. 

–––. 1950b. The North American variety of Milium effusum. Rhodora 52:218–222. 

–––, and B.G. Schubert. 1949. Some identities in Breweria. Rhodora 51: 35-43. 

Ferry, R.J., Sr., and R.J. Ferry, Jr. 1987. Calycanthus brockiana (Calycanthaceae), a new spicebush from north central Georgia. 

Sida 12: 339-341. 

Figlar, R.B., and H.P. Nooteboom. 2004. Notes on Magnoliaceae IV. Blumea 49: 87-100. 

Fishbein, M., and W.D. Stevens. 2005. Resurrection of Seutera Reichenbach (Apocynaceae, Asclepiadoideae). Novon 15: 531- 

533. 

Fisher, D.D., H.J. Schenk, J.A. Thorsch, and W.R. Ferren, Jr. 1997. Leaf anatomy and subgeneric affiliations of C3 and C4 

species of Suaeda (Chemopodiaceae) in North America. Am. J. Bot. 84: 1198-1210. 

Fisher, T.R. 1957. Taxonomy of the genus Heliopsis (Compositae). Ohio J. of Sci. 57: 171-191. 

Fleming, G.P., and J.C. Ludwig. 1996. Noteworthy collections: Virginia. Castanea 61: 89-94. 

Flora of North America Editorial Committee. 1993a. Flora of North America north of Mexico. Volume 1, introduction. Oxford 

Univ. Press, New York, NY. 372 pp. 

–––. 1993b. Flora of North America north of Mexico. Volume 2, pteridophytes and gymnosperms. Oxford Univ. Press, New 

York, NY. 475 pp. 

–––. 1997. Flora of North America north of Mexico. Volume 3, Magnoliophyta: Magnoliidae and Hamamelidae. Oxford 


–––. 2000. Flora of North America north of Mexico. Volume 22, Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in 

part), and Zingiberidae. Oxford Univ. Press, New York, NY. 352 pp. 

–––. 2002a. Flora of North America north of Mexico. Volume 26, Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford 


–––. 2002b. Flora of North America north of Mexico. Volume 23, Magnoliophyta: Commelinidae (in part): Cyperaceae. 

Oxford Univ. Press, New York, NY. 608 pp. 

–––. 2003a. Flora of North America north of Mexico. Volume 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. 


–––. 2003b. Flora of North America north of Mexico. Volume 4, Magnoliophyta: Caryophyllidae, part 1. Oxford Univ. Press, 

New York, NY. 559 pp. 

–––. 2005. Flora of North America north of Mexico. Volume 5, Magnoliophyta: Caryophyllidae, part 2. Oxford Univ. Press, 

New York, NY. 656 pp. 

–––. 2006a. Flora of North America north of Mexico. Volume 19, Magnoliophyta: Asteridae, part 6: Asteraceae, part 1. 


–––. 2006b. Flora of North America north of Mexico. Volume 20, Magnoliophyta: Asteridae, part 6: Asteraceae, part 2. 


–––. 2006c. Flora of North America north of Mexico. Volume 19, Magnoliophyta: Asteridae, part 6: Asteraceae, part 1. 


Flores-Cruz, M., H.D. Santana-Lira, S.D. Koch, and R. Grether. 2004. Taxonomic significance of leaflet anatomy in Mimosa 

series Quadrivalvis (Leguminosae, Mimosoideae). Systematic Botany 29: 892-902.


Folsom, J.P. 1984. Una reinterpretacion del estatus y relaciones de las taxa del complejo de Platanthera ciliaris [a 

reinterpretation of the status and relationships of taxa of the yellow-fringed orchid complex]. Orquidea (Méx.) 9: 321-345. 

Ford, B.A., D.A.R. McQueen, R.F.C. Naczi, and A.A. Reznicek. 1998. Allozyme variation and genetic relationships among 

species in the Carex willdenowii complex (Cyperaceae). Amer. J. Bot. 85: 546-552 

Forest, F., and A. Bruneau. 2000. Phylogenetic analysis, organization, and molecular evolution of the nontranscribed spacer of 

5S ribosomal RNA genes in Corylus (Betulaceae). Int. J. Plant Sci. 161: 793-806. 

Fosberg, F.R., and L. Artz. 1953. The varieties of Monarda fistulosa L. Castanea 18: 128-130. 

Foster, S. 1991. Echinacea: nature's immune enhancer. Healing Arts Press, Rochester, VT. 150 pp. 

Fox, W.B., R.K. Godfrey, and H.L. Blomquist. 1950. Notes on distribution of North Carolina plants – II. Rhodora 52: 253-271. 

–––, R.K. Godfrey, and H.L. Blomquist. 1952. Notes on distribution of North Carolina plants – III. Rhodora 54: 165-182. 

Franklin, M.A. 2001. Factors affecting seed production in natural populations of Lysimachia asperulifolia Poir. (Primulaceae), a 

rare, self-incompatible plant species. M.S. thesis, Dept. of Botany, N.C. State Univ., Raleigh, NC. 

Franklin, M.A. 2004. Natural Heritage Program list of the rare plants of North Carolina. North Carolina Natural Heritage 

Program, Raleigh, NC. 

Franz, N.M, A.S. Weakley, and R.K. Peet. [in prep.] On the use of taxonomic concepts in support of biodoiversity research and 

taxonomy. New Taxonomy Proceedings (Systematics Association). 

Franzke, A., K. Pollmann, W. Bleeker, R. Kohrt, and H. Hurka. 1998. Molecular systematics of Cardamine and allied genera 

(Brassicaceae): ITS and non-coding chloroplast DNA. Folia Geobotanica 33: 225-240. 

Freckmann, R.W. 1981. Realignments in the Dichanthelium acuminatum complex (Poaceae). Phytologia 48: 99-110. 

–––, and M.G. Lelong. 2002. Nomenclatural changes and innovations in Panicum and Dichanthelium (Poaceae, Paniceae). Sida 

20: 161-174. 

Freeman, C.C. 2004. A new combination in Persicaria (Polygonaceae). Sida 21: 291-292. 

Freeman, J.D. 1975. Revision of Trillium subgenus Phyllantherum (Liliaceae). Brittonia 27: 1-62. 

Freudenstein, J.V. 1992. Systematics of Corallorhiza and the Corallorhizinae (Orchidaceae). Ph.D. dissertation, Cornell 

University, Ithaca, NY. 

–––. 1997. A monograph of Corallorhiza (Orchidaceae). Harvard Papers in Botany 10: 5-51. 

–––. 1999a. Relationships and character transformation in Pyroloideae (Ericaceae) based on ITS sequences, morphology, and 

development. Systematic Botany 24: 398-408. 

–––. 1999b. A new species of Corallorhiza (Orchidaceae) from West Virginia, U.S.A. Novon 9: 511-513. 

Freytag, G.F., and D.G. Debouck. 2002. Taxonomy, distribution, and ecology of the genus Phaseolus (Leguminosae – 

Papilionoidae) in North America, Mexico and Central America. Sida, Bot. Misc. 23: 1-300. 

Fritsch, P.W., and S.D. Lucas. 2000. Clinal variation in the Halesia carolina complex (Styracaceae). Systematic Botany 25: 

197-210. 

Frodin, D.G., and R. Govaerts. 1996. World checklist and bibliography of Magnoliaceae. Kew Botanic Gardens, Kew, England. 

–––, and R. Govaerts. 2003. World checklist and bibliography of Araliaceae. Kew Botanic Gardens, Kew, England. 

Fross, D., and D. Wilken. 2006. Ceanothus. Timber Press, Portland, OR. 272 pp. 

Frye, C.T., and C. Lea. 2002. Atlas and annotated list of Carex (Cyperaceae) of Maryland and the District of Columbia. The 

Maryland Naturalist 44: 41-108. 

Fryxell, P.A. 2002. An Abutilon nomenclator. Lundellia 5: 79-118. 

Fu, C., H. Kong, Y. Qiu, and K.M. Cameron. 2005. Molecular phylogeny of the east Asian - North American disjunct Smilax 

sect. Nemexia (Smilacaceae). Int. J. Plant Sci. 166: 301-309. 

Fuertes Aguilar, J., P.A. Fryxell, and R.K. Jansen. 2003. Phylogenetic relationships and classification of the Sida generic 

alliance (Malvaceae) based on nrDNA ITS evidence. Systematic Botany 28: 352-364. 

Furlow, J.J. 1987a. The Carpinus caroliniana complex in North America. I. A multivariate analysis of geographical variation. 


–––. 1987a. The Carpinus caroliniana complex in North America. II. Systematics. Systematic Bot. 12: 416-434. 

–––. 1990. The genera of Betulaceae in the southeastern United States. J. Arnold Arb. 71: 1-67. 

Fusiak, F., and E.E. Schilling. 1984. Systematics of the Prenanthes roanensis complex (Asteraceae: Lactuceae). Bull. Torrey 

Bot. Club 111: 338-348. 

Gaddy, L.L. 1981. Two carices new to South Carolina. Castanea 46: 237-238. 

–––. 1986. A new heartleaf (Hexastylis) from Transylvania County, North Carolina. Brittonia 38: 82-85. 

–––. 1987a. A review of the taxonomy and biogeography of Hexastylis (Aristolochiaceae). Castanea 52: 186-196. 

–––. 1987b. Hexastylis shuttleworthii var. harperi (Aristolochiaceae), a new variety of heartleaf from Alabama and Georgia. 

Sida 12: 51-56. 

–––. 1990. [Echinacea] 

–––. 1995. Carex radfordii (section Laxiflorae: Cyperaceae), a new species from the Southern Appalachians. Novon 5: 259- 

261. 

–––, and D.A. Rayner. 1980. Rare or overlooked? Recent plant collections from the Coastal Plain of South Carolina. Castanea 

45: 181-184. 

Gadek, P.A., D.L. Alpers, M.M. Heslewood, and C.J. Quinn. 2000. Relationships within Cupressaceae sensu lato: a combined 

morphological and molecular approach. Am. J. Bot. 87: 1044-1057. 

Gale, S. 1944. Rhynchospora, section Eurhynchospora, in Canada, the United States and the West Indies. Rhodora 46: 89-278.


Ganders, F.R., M. Berbee, and M. Pirseyedi. 2000. ITS base sequence phylogeny in Bidens (Asteraceae): evidence for the 

continental relatives of Hawaiian and Marquesan Bidens. Systematic Bot. 25: 122-133. 

Gandhi, K.N. 1989. A biosystematic study of the Schizachyrium scoparium complex. Ph. D. dissertation, Texas A. & M. Univ. 

–––. 1999. Nomenclatural novelties for the Western Hemisphere plants. II. Harvard papers in Botany 4: 295-299. 

–––, R.D. Thomas, and S.L. Hatch. 1987. Cuscutaceae of Louisiana. Sida 12: 361-379. 

–––, and M.E. Barkworth. 2003. Nomenclatural and taxonomic review of knotroot bristle grass (Setaria parviflora, Gramineae). 

Rhodora 105: 197-204. 

–––, and B.E. Dutton. 1993. Palisot de Beauvois, the correct combining author of Erianthus giganteus (Poaceae). Taxon 42: 

855-856. 

–––, and R.D. Thomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4: 1-202. 

–––, and R.D. Thomas. 1991. Additional notes on the Asteraceae of Louisiana. Sida 14: 514-517. 

Garrison, J. 1992. The other side of Lygodium palmatum. Fiddlehead Forum 19: 10. 

Gaskin, J.F., F. Ghahremani-nejad, D.-y. Zhang, and J.P. Londo. 2004. A systematic overview of Frankeniaceae and 

Tamaricaceae from nuclear rDNA and plastid sequence data. Ann. Missouri Bot. Gard. 401-409. 

Gastony, G.J. 1977. Chromosomes of the independently reproducing Appalachian gametophyte – a new source of taxonomic 

evidence. Systematic Bot. 2: 43-48. 

–––. 1988. The Pellaea glabella complex: electrophoretic evidence for the deivations of the agamosporous taxa and a revised 

taxonomy. Amer. Fern. J. 78: 44-67. 

–––, and D.R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern United States and adjacent 

Mexico – a molecular phylogenetic reassessment of generic lines. Aliso 17: 131-144. 

–––, and D.E. Soltis. 1977. Chromosome studies of Parnassia and Lepuropetalon (Saxifragaceae) from the eastern United 

States. A new base number for Parnassia. Rhodora 79: 573-578. 

–––, and M.C. Ungerer. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: 

Onocleeae). Am. J. Bot. 84: 840-849. 

–––, G. Yatskievych, and C.K. Dixon. 1992. Chloroplast DNA restriction site variation in the fern genus Pellaea: phytogenetic 

relationships of the Pellaea glabella complex. Am. J. Bot. 79: 1072-1080. 

Gattinger, A. 1901. The flora of Tennessee and a philosophy of botany respectfully dedicated to the citizens of Tennessee. 

Gospel Advocate Publishing Company, Nashville, TN. 

Gensel, W.H. 1988. Rhododendron subsection Caroliniana. Rhododendron Notes & Records vol. 2. Proc. of 3 rd 

Rhododendron Conference, April 29-30, 1985. Rhododendron Species Foundation, Washington, DC. 

Geoffroy, M., and W.G. Berendson. 2003. The concept problem in taxonomy: importance, components, approaches. In: W.G. 

Berendsohn (ed.). MoReTax: handling factual information linked to taxonomic concepts in biology. Schriftenreihe für 

Vegetationskunde 39: 5-14. 

Gernandt, D.S., G. Geada López, S. Ortiz García, and A. Liston. 2005. Phylogeny and classification of Pinus. Taxon 54: 29-42. 

Gibson, T.C. 1991. Differential escape of insects from carnivorous plant traps. Am. Midl. Nat. 125: 55-62. 

Gil-ad, N.L. 1998. The micromorphologies of seed coats and petal trichomes of the taxa of Viola subsect. Boreali-Americanae 

(Violaceae) and their utility in discerning orthospecies from hybrids. Brittonia 50: 91-121. 

Gill, L.S. 1977. A cytosystematics study of the genus Monarda L. (Labiatae) in Canada. Caryologia 30: 381-394. 

Gillespie, J.P. 1962. A theory of relationships in the Lycopodium inundatum complex. Amer. Fern J. 52: 19-26. 

Gillett, G.W. 1964. Genetic barriers in the Cosmanthus Phacelias (Hydrophyllaceae). Rhodora 66: 359-368. 

–––. 1968. Systematic relationships in the Cosmanthus Phacelias (Hydrophyllaceae). Brittonia 20: 368-374. 

Gillett, J.M. 1957. A revision of the North American species of Gentianella Moench. Ann. Mo. Bot. Garden 44: 195-269. 

–––. 1959. A revision of Bartonia and Obolaria (Gentianaceae). Rhodora 61: 43-63. 

Gillis, W.T. 1971. The systematics and ecology of poison-ivy and the poison-oaks (Toxicodendron, Anacardiaceae). Rhodora 

73: 72-159, 161-237, 370-443, 465-540. 

Ginzbarg, S. 1992. A new disjunct variety of Croton alabamensis (Euphorbiceae) from Texas. Sida 15: 41-52. 

Gleason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. New 

York Botanical Garden and Hafner Press, New York, N.Y. 

Gleason, H.A., and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, second 

edition. New York Botanical Garden, Bronx, NY. 

Godfrey, R.K. 1948. Studies in the Compositae of North Carolina. I. Liatris. J. Elisha Mitchell Scientific Society 64: 241-249. 

–––. 1949. Studies in the Compositae of North Carolina. II. The Compositae of Wake, Durham, and Orange counties. J. Elisha 

Mitchell Scientific Society 65: 276-305. 

–––––-. 1969. Pieris phillyreifolia (Hook.) DC. (Ericaceae) in South Carolina. Sida 3: 447-448. 

–––. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. University of Georgia Press, 

Athens. 

–––, and P. Adams. 1964. The identity of Sagittaria isoetiformis (Alismataceae). Sida 1: 269-273. 

–––, and R. Kral. 1958. Observations on the Florida flora. Brittonia 10: 166-177. 

–––, and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States, monocotyledons. Univerity of Georgia 

Press, Athens, Georgia. 

–––, and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United States, dicotyledons. Univerity of Georgia 

Press, Athens, Georgia.


Godt, M.J.W., and J.L. Hamrick. 1995. Low levels of allozyme differentiation between Pyxidanthera (pyxie-moss) taxa 

(Diapensiaceae). Pl. Syst. Evol. 195: 159-168. 

–––, and J.L. Hamrick. 1999. Genetic divergence among infraspecific taxa of Sarracenia purpurea. Systematic Botany 23: 427- 

438. 

Goetsch, L., A.J. Eckert, and B.D. Hall. 2005. The molecular systematics of Rhododendron (Ericaceae): a phylogeny based 

upon RPB2 gene sequences. Systematic Botany 30: 616-626. 

Goldblatt, P. 1976. Chromosome number and its significance in Batis maritima (Bataceae). J. Arnold Arb. 57: 526-530. 

–––, and D.J. Mabberley. 2005. Belamcanda included in Iris, and the new combination I. domestica (Iridaceae: Irideae). Novon 

15: 128-132. 

–––, J. Manning, and G. Dunlop. 2004. Crocosmia and Chasmanthe. Royal Horticultural Society Plant Collector Guide. 

Timber Press, Portland, OR. 219 pp. 

Goldman, D.H. 1998. Hovenia dulcis (Rhamnaceae) naturalized in central Texas. Sida 18: 350-352. 

–––. 1999. Distribution update: Sabal minor in Mexico. Palms 43: 40-44. 

–––, C. van den Berg, and M.P. Griffith. 2004. Morphometric circumscription of species and infraspecific taxa in Calopogon R. 

Br. (Orchidaceae). Plant Syst. Evol. 247: 37-60 

Gonsoulin, G.J. 1974. A revision of Styrax (Styracaceae) in North America, Central America, and the Caribbean. Sida 5: 191- 

258. 

Goodspeed, T. H. 1954. The genus Nicotiana: origins, relationships and evolution of its species in the light of their distribution, 

morphology and cytogenetics. Chronica Botanica Co., Waltham, Mass. 536 pp. 

Gordon, J.E. 1981. Arachniodes simplicior new to South Carolina and the United States. Amer. Fern J. 71: 65-68. 

Gottlieb, J.E. 2002. Lycopodium lagopus new in West Virginia. Amer. Fern J. 92: 241-242. 

Gould, F.W. 1967. The grass genus Andropogon in the United States. Brittonia 19: 70-76. 

–––. 1975. The grasses of Texas. Texas A. & M. University Press, College Station, Texas. 

–––. 1979. The genus Bouteloua (Poaceae). Ann. Missouri Bot. Gard. 66: 348-416. 

–––, and C.A. Clark. 1978. Dichanthelium (Poaceae) in the United States and Canada. Ann. Missouri Bot. Gard. 65: 1088- 

1132. 

Gould, K.R. 1996. A new, disjunct variety of Spigelia gentianoides (Loganiaceae) from Bibb County, Alabama. Sida 17: 417- 

421. 

Gould, K.R., and M.J. Donoghue. 2000. Phylogeny and biogeography of Triosteum (Caprifoliaceae). Harvard Papers in Botany 

5: 157-166. 

Govaerts, R. 1998. World checklist and bibliography of Fagales (Betulaceae, Corylaceae, Fagaceae and Ticodendraceae). 

Royal Botanic Gardens, Kew, England. 

–––, D.G. Frodin, and T.D. Pennington. 2001. World checklist and bibliography of Sapotaceae. Royal Botanical Gardens, Kew, 

England. 

–––, D.G. Frodin, and A. Radcliffe-Smith. 2000. World checklist and bibliography of Euphorbiaceae (with Pandaceae). 

Volumes 1-4. Royal Botanic Gardens, Kew, England. 

Govus, T.E. 1987. The occurrence of Sarracenia oreophila (Kearney) Wherry in the Blue Ridge Province of southwestern 

North Carolina. Castanea 52: 310-311. 

Graetz, K.E. 1973. Seacoast plants of the Carolinas for conservation and beautification. U.S. Dept. of Agriculture and Soil 

Conservation Service, Raleigh, NC and Columbia SC. 

Graham, S.A. 1966. The genera of Araliaceae in the southeastern United States. J. Arnold Arb. 47: 126-136. 

–––. 1975. Taxonomy of the Lythraceae in the southeastern United States. Sida 6: 80-103. 

–––. 1985. A revision of Ammannia (Lythraceae) in the western hemisphere. J. Arnold Arb. 66: 395-420. 

–––, and C.E. Wood, Jr. 1965. The genera of Polygonaceae in the southeastern United States. J. Arnold Arb. 46: 91-121. 

–––, and C.E. Wood, Jr. 1975. The Podostemaceae of the southeastern United States. J. Arnold Arb. 56: 456-465. 

Grant, A.L. 1924. A monograph of the genus Mimulus. Ann. Mo. Bot. Garden 11: 99-389. 

Grant, E., and C. Epling. 1943. A study of Pycnanthemum (Labiatae). Univ. of Calif. Publ. in Botany 20: 195-240. 

Grant, V. 1956. A synopsis of Ipomopsis. Aliso 3: 351-362. 

–––. 1997. Nomenclature of subfamilies and tribes in the Polemoniaceae. Phytologia 83: 385-389. 

–––. 1998. Primary classification and phylogeny of the Polemoniaceae, with comments on molecular cladistics. Amer. J. Bot. 

85: 741-752. 

–––. 2003. Taxonomy of the Polemoniaceae: the subfamilies and tribes. Sida 20: 1371-1385. 

Grant, W.F., and B.K. Thompson. 1975. Observations on Canadian birches, Betula cordifolia, B. neoalaskana, B. populifolia, B. 

papyrifera, and B. × caerulea. Can J. Bot. 53: 1478-1490. 

Gray, J.R., and D.E. Fairbrothers. 1971. A clarification of some misconceptions about Amphicarpum purshii (Gramineae). Bull. 

Torrey Bot. Club 98: 174-175. 

Grayum, M.H. 1987. A summary of evidence and arguments supporting the removal of Acorus from the Araceae. Taxon 36: 

723-729. 

Grear, J.W. 1978. A revision of the New World species of Rhynchosia (Leguminosae-Faboideae). Mem. New York Bot. Gard. 

31: 1-168. 

Green, E.P., and F. T. Short. 2003. World Atlas of seagrasses. Prepared by the UNEP World Consservation Monitoring Centre. 

Univ. of Calif. Press, Berkeley, Calif., US.


Green, P.S. 1962. Watercress in the New World. Rhodora 64: 32-43. 

–––. 1966. Identification of the species and hybrids in the Lonicera tatarica complex. J. Arnold Arb. 47: 75-88. 

Greene, C.W. 1980. The systematics of Calamagrostis (Gramineae) in eastern North America. Ph. D. thesis, Harvard 

University, Cambridge, Mass. 

Greene, E.L. 1892. On certain Spiraeaceae. Pittonia 2: 219-222. 

Gregg, K.B. 1991. Defrauding the deceitful orchid: pollen collection by pollinators of Cleistes divaricata and C. bifaria. 

Lindleyana 6: 214-220. 

Greuter, W., J. McNeill, F.R. Barrie, H.-M. Burdet, V. Demoulin, T. S. Filggueiras, D.H. Nicolson, P.C. Silva, J.E. Skog, P. 

Trehane, N.J. Turland, and D.L. Hawksworth. 2000. International Code of Botanical Nomenclature (St. Louis Code) 

adopted by the Sixteenth International Botanical Congress, St. Louis, Missouri, July-August 1999. Regnum Vegetabile 131. 

Koeltz Scientific Books, Königstein. 

Grimes, J.W. 1988. Systematics of New World Psoraleae (Leguminosae-Faboideae). Ph.D. dissertation, Univ. of Texas at 

Austin. 

–––. 1990. A revision of the New World species of Psoraleeae (Leguminosae: Papilionoideae). Memoirs N.Y. Bot. Gard. 61: 

1-114. 

Groves, C.R. 2003. Drafting a conservation blueprint: a practitioner’s guide to planning for biodiversity. Island Press. 457 pp. 

Guillon, J.-M. 2004. Phylogeny of horsetails (Equisetum) based on the chloroplast rps4 gene and adjacent noncoding sequences. 


Gusman, G., and L. Gusman. 2002. The genus Arisaema: a monograph for botanists and nature lovers. Gantner, Ruggell, 

Lichtenstein. 438 pp. 

Gustafson, D.J., G. Romano, R.E. Latham, and J.K. Morton. 2003. Amplified fragment length polymorphism analysis of genetic 

relationships among the serpentine barrens endemic Cerastium velutinum Rafinesque var. villosissimum Pennell 

(Caryophyllaceae) and closely related Cerastium species. J. Torrey Bot. Soc. 130: 218-223. 

Gustafsson, M.H.G., V. Bittrich, and P.F. Stevens. 2002. Phylogeny of Clusiaceae based on rbcL sequences. Int. J. Plant Sci. 

163: 1045-1054. 

Guthrie, W. 1820. A universal geography; or, a view of the present state of the known world. Benj. Warner, Philadelphia, PA. 

Hágsater, E. 2000. New names for Florida orchids. North American Native Orchid Journal 6: 299-309. 

Haines, A.A. 2002. A new combination in Lycopodiella (Lycopodiaceae). Rhodora 104: 296-298. 

–––. 2003a. The families Huperziaceae and Lycopodiaceae of New England: a taxonomic and ecological reference. V.F. 

Thomas Co., Bowdoin, ME. 100 pp. 

–––. 2003b. Lycopodiella ×gilmanii (Lycopodiaceae), a new hybrid bog clubmoss from northeastern North America. Amer. 

Fern J. 93: 196-202. 

–––. 2004. New combination in Poa. Botanical Notes 10: 1-5. 

Halda, J.J. 1996. The genus Gentiana. SEN, Dobré. 

Hall, D.W. 1982. Sorghastrum (Poaceae) in Florida. Sida 9: 302-308. 

–––. 1998. Is Cogon Grass really an exotic? Wildland Weeds 1: 14-15. 

Hall, J.C., K.J. Sytsma, and H.H. Iltis. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. 

Amer. J. Botany 89: 1826-1842. 

Hämet-Ahti, L. 1980. Juncus trifidus L. subsp. carolinianus Hämet-Ahti, n. subsp., in eastern North America. Veröff. Geobot. 

Inst. ETH Stiftung Rübel, Zurich 69: 7-13. 

Hamilton, C.W., and S.H. Reichard. 1992. Current practice in the use of subspecies, variety, and forma in the classification of 

wild plants. Taxon 41: 485-498. 

Hamzeh, M., and S. Dayanandan. 2004. Phylogeny of Populus (Salicaceae) based on nucleotide sequences of chloroplast trnTtrnF 

region and nuclear rDNA. Amer. J. Botany 91: 1398-1408. 

Hancock, J.F. 2004. Plant evolution and the origin of crop species, second edition. CABI Publishing, Oxon, UK. 313 pp. 

Hancock, T.E., and P.E. Hosier. 2003. Ecology of the threatened species Amaranthus pumilus Rafinesque. Castanea 68: 236- 

244. 

Hanks, G.R., ed. 2002. Narcissus and daffodil: the genus Narcissus. Taylor & Francis, London. 428 pp. 

Hansen, B.F., and R.P. Wunderlin. 1988. Synopsis of Dichanthelium (Poaceae) in Florida. Ann. Missouri Bot. Gard. 75: 1637- 

1657. 

Hao, G., Y.-M. Yuan, C.-M. Hu, X.-J. Ge, and N.-X. Zhao. 2004. Molecular phylogeny of Lysimachia (Myrsinaceae) based on 

chlorplast trnL-F and nuclear ribosomal ITS sequences. Molecular Phylogenetics and Evolustion 31: 323-339. 

Hardin, J.W. 1952. The Juglandaceae and Corylaceae of Tennessee. Castanea 17: 78-89. 

–––. 1957a. A revision of the American Hippocastanaceae – I. Brittonia 9: 145-171. 

–––. 1957b. A revision of the American Hippocastanaceae – II. Brittonia 9: 173-195. 

–––. 1961. A hybrid population of Habenaria and variation in H. blephariglottis. Castanea 26: 120-123. 

–––. 1963. Pachystima canbyi in North Carolina. Castanea 28: 177-178. 

–––. 1964a. A comparison of Phytolacca americana and P. rigida. Castanea 29: 155-164. 

–––. 1964b. Variation in Aconitum of eastern United States. Brittonia 16: 80-94. 

–––. 1968. Diervilla (Caprifoliaceae) of the southeastern United States. Castanea 33: 31-36. 

–––. 1971a. Studies of the southeastern United States flora. I. Betulaceae. J. Elisha Mitch. Sci. Soc. 87: 39-41. 

–––. 1971b. Studies of the southeastern United States flora. II. The gymnosperms. J. Elisha Mitchell Sci. Soc. 87: 43-50.


–––. 1972. Studies of the southeastern United States flora. III. Magnoliaceae and Illiciaceae. J. Elisha Mitchell Sci. Soc. 88: 

30-32. 

–––. 1973. The enigmatic chokeberries (Aronia, Rosaceae). Bull. Torrey Bot. Club 100: 178-184. 

–––. 1974. Studies of the southeastern United States flora. IV. Oleaceae. Sida 5: 274-285. 

–––. 1975. Hybridization and introgression in Quercus alba. J. Arnold Arb. 56: 336-363. 

–––. 1976. Terminology and classification of Quercus trichomes. J. Elisha Mitch. Sci. Soc. 92: 151-161. 

–––. 1979. Stellate and "stellate" trichomes and stellate vestiture. ASB Bulletin 26: 74. 

–––. 1985. Foliar trichomes in american beech. ASB Bulletin 32: 46. 

–––. 1990. Variation patterns and recognition of varieties in Tilia americana s.l. Systematic Bot. 15: 33-48. 

–––. 1992. Foliar morphology of the common trees of North Carolina and adjacent states. N.C. Agricultural Research Service 

Tech. Bull. 298. 135 pp. 

–––, and R.L. Beckmann. 1982. Atlas of foliar surface features in woody plants. V. Fraxinus (Oleaceae) of eastern North 

America. Brittonia 34: 129-140. 

–––, and G.P. Johnson. 1985. Atlas of foliar surface features in woody plants, VIII. Fagus and Castanea (Fagaceae) of eastern 

North America. Bull. Torrey Bot. Club 112: 11-20. 

–––, and K.A. Jones. 1989. Atlas of foliar surface features in woody plants, X. Magnoliaceae of the United States. Bull. Torrey 

Bot. Club 116: 164-173. 

–––, and L.L. Phillips. 1985a. Atlas of foliar surface features in woody plants, VII. Rhus subg. Rhus (Anacardiaceae) of North 

America. Bull. Torrey Bot. Club 112: 1-10. 

–––, and L.L. Phillips. 1985b. Hybridization in eastern North American Rhus (Anacardiaceae). ASB Bulletin 32: 99-106. 

–––, and D.E. Stone. 1984. Atlas of foliar surface features in woody plants, VI. Carya (Juglandaceae) of North America. 

Brittonia 36: 140-153. 

–––, R.L. Kologiski, J.R. Massey, J.F. Matthews, J.D. Pittillo, and A.E. Radford. 1977. Vascular plants. In J.E. Cooper, S.S. 

Robinson, and J.B. Funderburg (eds.). Endangered and threatened plants and animals of North Carolina: proceedings of a 

symposium on endangered and threatened biota of North Carolina, Meredith College, Raleigh, November 7-8, 1975. 

Harms, V.L. 1974. A preliminary conspectus of Heterotheca section Chrysopsis (Compositae). Castanea 39: 155-165. 

Harper, R.M. 1905. Mesadenia lanceolata and its allies. Torreya 5: 182-185. 

–––. 1906. Some new or otherwise noteworthy plants from the coastal plain of Georgia. Bull. Torrey Bot. Club 33: 229-233. 

–––. 1944. Notes on Plantago, with special reference to P. cordata. Castanea 9: 121-130. 

Hart, J.A., and R.A. Price. 1990. The genera of Cupressaceae (including Taxodiaceae) in the southeastern United States. J. 

Arnold Arb. 71: 275-322. 

Hartman, R.L., and B.E. Nelson. 1998. Taxonomic novelties from North America north of Mexico: a 20-year vascular plant 

diversity baseline. Monographs in Systematic Botany from the Missouri Botanical Garden 67: 1-59. 

Harvill, A.M., Jr., T.R. Bradley, C.E. Stevens, T.F. Wieboldt, D.M.E. Ware, D.W. Ogle, G.W. Ramsey, and G.P. Fleming. 1992. 

Atlas of the Virginia flora, third edition. Virginia Botanical Associates, Burkeville, VA. 

Haskins, M.L., and W.J. Hayden. 1987. Anatomy and affinities of Penthorum. Am. J. Bot. 74: 164-177. 

Hatley, J.R. 1977. An analysis of variation in Shortia galacifolia. M.S. thesis, Dept. of Botany, North Carolina State University. 

Hauber, D.P., and L. Legé. 1999. A survey of allozymic variation among three members of the Sagittaria graminea complex 

(Alismataceae) from the southeastern United States. J. Torrey Bot. Soc. 126: 181-187. 

Haufler, C.M., D.E. Soltis, and P.S. Soltis. 1995. Phylogeny of the Polypodium vulgare complex: insights from chloroplast 

DNA restriction site data. Systematic Bot. 20: 110-119. 

–––, and M.D. Windham. 1991. New species of North American Cystopteris and Polypodium, with comments on their reticulate 

relationships. Amer. Fern J. 81: 7-23. 

–––, M.D. Windham, and E.W. Rabe. 1995. Reticulate evolution in the Polypodium vulgare complex. Systematic Bot. 20: 89- 

109. 

–––, M.D. Windham, and T.A. Ranker. 1990. Biosystematic analysis of the Cystopteris tennesseensis (Dryopteridaceae) 

Complex. Ann. Missouri Bot. Gard. 77: 314-329. 

Hauk, W.D. 1996. Phylogenetics of Ophioglossaceae: the evolutionary consequences of morphological reduction. Ph.D. 

dissertation, University of North Carolina at Chapel Hill, Biology dept. 

–––, C.R. Parks, and M.W. Chase. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL-F plastid DNA 

sequences and morphology. Molecular Phylogenetics and Evolution 28: 131-151. 

Hauke, R.L. 1979. Equisetum ramosissimum in North America. Amer. Fern J. 69: 1-5. 

–––. 1984. Equisetum ramosissimum in Louisiana. Amer. Fern J. 74: 61. 

–––. 1992. Revisiting Equisetum ramosissimum. Amer. Fern J. 82: 83-84. 

Hayden, W.J., and S.M. Hayden. 1984. Wood anatomy and relationships of Betula uber. Castanea 49: 26-30. 

Hayes, D.W. 1946. Two remarkable range extensions. Castanea 11: 61-62. 

Haynes, R.R. 1971. A monograph of the genus Conopholis (Orobanchaceae). Sida 4: 246-264. 

–––. 1977. The Najadaceae in the southeastern United States. J. Arnold Arb. 58: 161-170. 

–––. 1978. The Potamogetonaceae in the southeastern United States. J. Arnold Arb. 59: 170-191. 

–––. 1979. Revision of North and Central American Najas (Najadaceae). Sida 8: 34-56. 

–––. 1987. The Zannichelliaceae in the southeastern United States. J. Arnold Arb. 68: 259-268. 

–––. 1998. Noteworthy collections: Alabama. Castanea 63: 81-82.


–––, and J.R. Burkhalter. 1998. A new species of Echinodorus (Alismataceae) from the United States of America. Castanea 63: 

180-182 

–––, and C.B. Hellquist. 1996. New combinations in North American Alismatidae. Novon 6: 370-371. 

–––, D.H. Les, and M. Král. 1998. Two new combinations in Stuckenia, the correct name for Coleogeton (Potamogetonaceae). 

Novon 8: 241. 

Hays, J.F. 1998a. Priority of the name Agalinis harperi (Scrophulariaceae) over the names Agalinis delicatula and Agalinis 

pinetorum. Sida 18: 369-370. 

–––. 1998b. Agalinis (Scrophulariaceae) in the Ozark highlands. Sida 18: 555-577. 

Heafner, K.D. 2001. Pellaea wrightiana Hooker (Pteridaceae) in North Carolina revisted with a new record for eastern North 

America and a key to Pellaea species in the Carolinas. Castanea 66: 319-326. 

Heard, S.B., and J.C. Semple. 1988. The Solidago rigida complex (Compositae: Astereae): a multivariate morphometric 

analysis and chromosome numbers. Can. J. Bot. 66: 1800-1807. 

Heiser, C.B., Jr., and B. Pickersgill. 1975. Names for the bird peppers [Capsicum - Solanaceae]. Baileya 19: 151-156. 

–––, Jr, D.M. Smith, S.B. Clevenger, and W.C. Martin, Jr. 1969. The North American sunflowers (Helianthus). Mem. Torrey 

Bot. Club 22: 1-218. 

Helfgott, D.M., and R.J. Mason-Gamer. 2004. The evolution of North American Elymus (Triticeae, Poaceae) allotetraploids: 

evidence from phosphoenolpyruvate carboxylase gene sequences. Systematic Botany 29: 850-861. 

Henderson, A., G. Galeano, and R. Bernal. 1995. Field guide to the palms of the Americas. Princeton Univ. Press, Princeton, 

NJ. 352 pp. 

Henrard, J.T. 1929. A monograph of the genus Aristida. Mededeelingen Rijks-Herb. 58: 1-325. 

Henrickson, J. 1987. A taxonomic reevaluation of Gossypianthus and Guilleminea (Amaranthaceae). Sida 12: 307-337. 

–––. 1999. Studies in New World Amaranthus (Amaranthaceae). Sida 18: 783-807. 

Henry, M.G. 1946. A new lily from southern Alabama and northern Florida. Bartonia 24: 1-4. 

Hermann, F.J. 1947. A new species of Carex from Tennessee. Castanea 12: 113-115 

Herndon, A. 1993. Notes on Chamaesyce (Euphorbiaceae) in Florida. Rhodora 95: 352-368. 

Herrera Arrieta, Y., P.M. Peterson, and M. de la Cerda Lemus. 2004. Revisión de Bouteloua Lag. Comisión Nacional para el 

Conocimiento y Uso de la Biodiversidad y Instituto Politécnico Nacional, Durango, Mexico. 187 pp. 

Hershkovitz, M.A., and E.A. Zimmer. 2000. Ribosomal DNA evidence and disjunctions of western American Portulacaceae. 

Molecular Phylogenetics and Evolution 15: 419-439. 

Hess, W.J., and N.A. Stoynoff. 1998. Taxonomic status of Quercus acerifolia (Fagaceae) and a morphological comparison of 

four members of the Quercus shumardii complex. Systematic Bot. 23: 89-100. 

Hickey, R.J. 1977. The Lycopodium obscurum complex in North America. Amer. Fern J. 67: 45-48. 

–––, and J.M. Beitel. 1979. A name change for Lycopodium flabelliforme. Rhodora 81: 137-140. 

Hilger, H.H., and N. Diane. 2003. A systematic analysis of Heliotropiaceae (Boraginales) based on trnL and ITS1 sequence 

data. Bot. Jahrb. Syst. 125: 19-51. 

Hill, L.M. 1992. A floristic and chromosomal study of the Fumariaceae in Virginia. Castanea 57: 273-281. 

Hill, S.R. 1992. Calciphiles and calcareous habitats of South Carolina. Castanea 57: 25-33. 

–––––-. 1999. The relict flora of ice ponds in South Carolina. Castanea 64: 14-22. 

–––, and C.N. Horn. 1997. Additions to the flora of South Carolina. Castanea 62: 194-208. 

Hillig, K.W., and P.G. Mahlberg. 2004. A chemotaxonomic analysis of cannabinoid variation in Cannabis (Cannabaceae). 

Amer. J. Bot. 91: 966-975. 

Hilton, J.L., and R.S. Boyd. 1996. Microhabitat requirements and seed/microsite limitation of the rare granite outcrop endemic 

Amphianthus pusillus (Scrophulariaceae). Bull. Torrey Bot. Club 123: 189-196. 

Hilu, K.W. 1980. Noteworthy collections: Eleusine tristachya. Madroño 27: 177-178. 

Hinton, B.D. 1968. Parietaria praetermissa (Urticaceae), a new species from the southeastern United States. Sida 3: 191-194. 

Hitchcock, A.S., and A. Chase. 1910. The North American species of Panicum. Contr. U.S. Natl. Herb. 15: 1–396. 

–––, and A. Chase. 1950. Manual of the grasses of the United States, second edition. U.S. Dept. of Agriculture Miscellaneous 

Publication No. 200 (reprinted in 1971 by Dover Publications, New York). 

Hitchcock, C.L. 1944. The Tofieldia glutinosa complex of western North America. Amer. Midl. Naturalist 31: 487-498. 

Ho, T.-N., and S.-W. Liu. 1990. The infrageneric classification of Gentiana (Gentianaceae). Bull. Br. Mus. nat. Hist. (Bot.) 20: 

169-192. 

–––, and S.W. Liu. 2001. A worldwide monograph of Gentiana. Science Press, Beijing. 

Hodgdon, A.R. 1938. A taxonomic study of Lechea. Rhodora 40: 29-69, 87-131. 

Hodkinson, T.R., M.W. Chase, M.D. Lledó, N. Salamin, and S.A. Renvoize. 2002. Phylogenetics of Miscanthus, Saccharum, 

and related genera (Saccharinae, Andropogoneae, Poaceae) based on DNA sequences from ITS nuclear ribosomal DNA and 

plastid trnL intron and trnL-F intergenic spacers. J. Plant. Res. 115: 381-392. 

Holm, T. 1896. A study of some anatomical characters of North American Gramineae. VII. The genus Amphicarpum. Bot. 

Gazette {}: 403-406. 

Holmes, W.C. 1995. Review preparatory to an infrageneric classification of Mikania (Tribe: Eupatoriae). In Hind, D.J.N., C. 

Jeffrey, and G.V. Pope (eds.). Advances in Compositae systematics, pp. 239-254. Royal Botanic Gardens, Kew. 

Holmgren, N.H. 1994. Redefinition of Dodecatheon dentatum (Primulaceae) and rationale for use of varietal rank. Brittonia 46: 

87-94.


Holub, J. 1975a. Diphasiastrum, a new genus in Lycopodiaceae. Preslia (Praha) 47: 97-110. 

–––. 1975b. Notes on some species of Diphasiastrum. Preslia (Praha) 47: 232-240. 

Homoya, M.A. 1993. Orchids of Indiana. Indiana Academy of Science, Bloomington, IN. 276 pp. 

Hoot, S.B., S. Magallón, and P.R. Crane. 1999. Phylogeny of basal eudicots based on three molecular data sets: atpB, rbcL, and 

18S nuclear ribosomal DNA sequences. Ann. Missouri Bot. Garden 86: 1-32. 

–––, N.S. Napier, and W.C. Taylor. 2004. Revealing unknown or extinct lineages within Isoëtes (Isoëtaceae) using DNA 

sequences from hybrids. Amer. J. Bot. 91: 899-204. 

–––, A.A. Reznicek, and J.D. Palmer. 1994. Phylogenetic relationships in Anemone (Ranunculaceae) based on morphology and 

chloroplast DNA. Systematic Bot. 19: 169-200. 

Hopkins, C.O., and W.H. Blackwell, Jr. 1977. Synopsis of Suaeda (Chenopodiaceae) in North America. Sida 7: 147-173. 

Hopkins, M. 1937. Arabis in eastern and central North America. Rhodora 39: 63-186. 

Horn, C.N. 1997. An ecological study of Frasera caroliniensis in South Carolina. Castanea 62: 185-193. 

–––. 1998. Noteworthy collections: North Carolina and Virginia. Castanea 63: 495. 

Hornberger, K.L. 1991. The blue-eyed-grasses (Sisyrinchium: Iridaceae) of Arkansas. Sida 14: 597-604. 

Horton, J.H. 1963. A taxonomic revision of Polygonella (Polygonaceae). Brittonia 15: 177-203. 

–––. 1972. Studies of the southeastern United States flora. IV. Polygonaceae. J. Elisha Mitchell Sci. Soc. 88: 92-102. 

Hoshizaki, B.J., and K.A. Wilson. 1999. The cultivated species of the fern genus Dryopteris in the United States. American 

Fern Journal 89: 1-98. 

Howard, R.A., and G.W. Staples. 1983. The modern names for Catesby's plants. J. Arnold Arb. 64: 511-546. 

Hsiao, J.Y., and M.L. Lin. 1995. A chemotaxonomic study of essential oils from the leaves of genus Clerodendrum 

(Verbenaceae) native to Taiwan. Bot. Bull. Acad. Sin. 36: 247-251. 

Hu, Shiu-ying. 1954-56. A monograph of the genus Philadelphus. J. Arnold Arb. 35: 276-333; 36: 52-109; 37: 15-90. 

–––. 1979. Ailanthus. Arnoldia 39: 29-50. 

Huck, R.B. 1984. Systematics and evolution of Dicerandra (Labiatae). Ph.D. dissertation, Univ. of North Carolina at Chapel 

Hill, Dept. of Biology. 

–––. 1987. Systematics and evolution of Dicerandra (Labiatae). Phanerogamarum Monographiae Tomus XIX. J. Cramer, 

Berlin. 343 pp. 

–––, and H.L. Chambers. 1997. Polyploidy: a factor in the evolution of Dicerandra Benth. (Labiatae). Edinb. J. Bot. 54: 217- 

229. 

Huft, M.J. 1979. A monograph of Euphorbia section Tithymalopsis. Ph.D. dissertation, Univ. of Michigan. 

Hughes, C. 1998. Monograph of Leucaena (Leguminosae-Mimosoideae). Systematic Botany Monographs 55: 1-244. 

Hunt, D., ed. 1998. Magnolias and their allies. Proceedings of an international symposium, Royal Holloway, University of 

London, Egham, Surrey, U.K., 12-13 April 1996. International Dendrological Society and the Magnolia Society. 

Hunt, D.M. 1994. Morphology and ecology of Quercus series Laurifoliae, Marilandicae and Nigrae. Pp. 99-188 in Ed. A. 

Miyawaki, K. Iwatsuki, and M. M. Grandtner (eds). Vegetation in eastern North America. Vegetation system and dynamics 

under human activity in the eastern North American cultural region in comparison with Japan. University of Tokyo Press, 

Tokyo, Japan. 515 pp. 

–––. 1990. A systematic review of Quercus series Laurifoliae, Marilandicae and Nigrae. Unpublished Ph. D. dissertation, 

University of Georgia, Athens, GA. 

–––, and R.E. Zaremba. 1992. The northeastward spread of Microstegium vimineum (Poaceae) into New York and adjacent 

states. Rhodora 94: 167-170. 

Hunt, D.R. 1983. New names in Commelinaceae. American Commelinaceae: XI. Kew Bull. 38: 131-133. 

–––. 1986. Campelia, Rhoeo, and Zebrina united with Tradescantia. American Commelinaceae: XIII. Kew Bull. 41: 401-412. 

Hunziker, A.T. 2001. Genera Solanacearum: the genera of Solanaceae illustrated, arranged according to a new system. A.R.G. 

Gantner, Ruggell. 500 pp. 

Huttleston, D.G. 1949. The three subspecies of Arisaema triphyllum. Bull. Torrey Bot. Club 76: 407-413. 

–––. 1981. The four subspecies of Arisaema triphyllum. Bull. Torrey Bot. Club 108: 479-481. 

Iltis, H.H. 1960. Studies in the Capparidaceae – VII. Old World Cleomes adventive in the New World. Brittonia 12: 279-294. 

–––. 1965. The genus Gentianopsis (Gentianaceae): transfers and phytogeographic comments. Sida 2: 129-154. 

Irving, R.S. 1980. The systematics of Hedeoma (Labiatae). Sida 8: 218-295. 

Irwin, H.S., and R.C. Barneby. 1982. The American Cassiinae: a synoptical revision of Leguminosae tribe Cassieae subtribe 

Cassiinae in the New World. Memoirs N.Y. Bot. Gard. 35: 1-918. 

Isely, D. 1973. Leguminosae of the United States. I. Subfamily Mimosoideae. Memoirs N.Y. Bot. Gard. 25: 1-152. 

–––––-. 1975. Leguminosae of the United States. II. Subfamily Caesalpinioideae. Memoirs N.Y. Bot. Gard. 25: 1-228. 

–––. 1981. Leguminosae of the United States. III. Subfamily Papilionoidae: Tribes Sophoreae, Podalyriaeae, Loteae. 

Memoirs N.Y. Bot. Gard. 25: 1-264. 

–––. 1986a. Notes about Psoralea sensu auct., Amorpha, Baptisia, Sesbania and Chamaecrista (Leguminosae) in the 

southeastern United States. Sida 11: 429-440. 

–––. 1986b. Notes on Leguminosae: Papilionoideae of the southeastern United States. Brittonia 38: 352-359. 

–––. 1990. Leguminosae (Fabaceae), volume 3, part 2, Vascular flora of the southeastern United States. University of North 

Carolina Press, Chapel Hill, NC.


–––. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and Hawaii). Monte L. 

Bean Life Science Museum, Brigham Young Univ., Provo, UT. 

–––, and F.J. Peabody. 1984. Robinia (Leguminosae: Papilionoideae). Castanea 49: 187-202. 

Jacono, C.C. 1999. Salvinia molesta (Salviniaceae), new to Texas and Louisiana. Sida 18: 927-928. 

Jansen, R.K. 1985. The systematics of Acmella (Asteraceae-Heliantheae). Systematic Bot. Monographs 8. 

Järvinen, P., A. Palmé, L. Orlando Morales, M. Lännenpää, M. Keinänen, T. Sopanen, and M. Lascoux. 2004. Phylogenetic 

relationships of Betula species (Betulaceae) based on nuclear ADH and chloroplast matK sequences. Amer. J. Bot. 91: 

1834-1845. 

Jefferson-Brown, M.J. 1969. Daffodils and Narcissi: a complete guide to the Narcissus family. Faber and Faber, London. 224 

pp. 

–––. 1991. Narcissus. Timber Press, Portland, OR. 224 pp. 

Jenne, G.E. 1966. A study of variation in North American Hamamelis L. (Hamamelidaceae). Master's thesis, Vanderbilt 

University, Biology Dept. 204 pp. 

Jensen, R.J. 1977. Numerical analysis of the scarlet oak complex (Quercus subgen. Erythrobalanus) in the eastern United 

States: relationships above the species level. Systematic Bot. 2: 122-133. 

Johansson, J.T. 1998. Chloroplast DNA restriction site mapping and the phylogeny of Ranunculus (Ranunculaceae). Pl. Syst. 

Evol. 213: 1-19. 

Johnson, A.F. 1982. Some demographic characteristics of the Florida Rosemary Ceratiola ericoides Michx. Amer. Midland 

Nat. 108: 170-174. 

Johnson, G.P. 1988. Revision of Castanea section Balanocastanon (Fagaceae). J. Arnold Arb. 69: 25-49. 

–––. 1992. Noteworthy collections: Arkansas. Castanea 57: 150-151. 

–––. 1994. Noteworthy collections: Arkansas. Castanea 59: 78. 

Johnson. M.F. 1980. The genus Prenanthes L. (Cichorieae – Asteraceae) in Virginia. Castanea 45: 24-30. 

Johnson, R.G. 1968. {Disporum}. Castanea 33: 262-266. 

Johnson, S.R. 1992. Observations on populations of Chamaesyce polygonifolia Small and C. ingallsii Small (Euphorbia 

ammannioides HBK) on barrier islands of Virginia and North Carolina. Castanea 57: 291-292. 

Johnston, M.C. 1957. Synopsis of the United States species of Forestiera (Oleaceae). Southwestern Naturalist 2: 140-151. 

Jones, A.G. 1979. A study of wild leek, and the recognition of Allium burdickii (Liliaceae). Systematic Bot. 4: 29-43. 

–––. 1980a. A classification of the New World species of Aster (Asteraceae). Brittonia 32: 230-239. 

–––. 1980b. Data on chromosome numbers in Aster (Asteraceae), with comments on the status and relationships of certain 

North American species. Brittonia 32: 240-261. 

–––. 1984. Nomenclatural notes on Aster (Asteraceae) – II. New combinations and some transfers. Phytologia 55: 373-388. 

–––. 1992. Aster and Brachyactis (Asteraceae) in Oklahoma. Sida, Bot. Misc. 8: 1-46. 

–––, and D.A. Young. 1983. Generic concepts of Aster (Asteraceae): a comparison of cladistic, phenetic, and cytological 

approaches. Systematic Bot. 8: 71-84. 

Jones, D.L. 1993. Cycads of the world. Smithsonian Institution Press, Washington, DC. 312 pp. 

Jones, G.N. 1939. A synopsis of the North American species of Sorbus. Jour. Arnold Arb. 20: 1-43. 

–––. 1940. A monograph of the genus Symphoricarpos. Jour. Arnold Arb. 21: 201-252. 

Jones, Q. 1951. A cytotaxonomic study of the genus Disporum in North America. Contr. Gray Herb. 173: 1-40. 

Jones, R.L. 1983. A systematic study of Aster section Patentes (Asteraceae). Sida 10: 41-81. 

–––. 1992. Additional studies of Aster georgianus, A. patens, and A. phlogifolius (Asteraceae). Sida 15: 305-315. 

–––. 2005. Plant life of Kentucky: an illustrated guide to the vascular flora. Univ. Press of Kentucky. 834 pp. 

Jones, S.B., Jr. 1982. The genera of Vernonieae (Compositae) in the southeastern United States. J. Arnold Arb. 63: 489-507. 

–––, and N.C. Coile. 1988. The distribution of the vascular flora of Georgia. Dept. of Botany, Univ. of Georgia, Athens, 

Georgia. 

Jones, S.D., and G.D. Jones. 1993. A new species of Carex (Cyperaceae: Triquetrae) from the Chisos Mountains, Texas, and a 

key to species of section Triquetrae. Sida 15: 509-518. 

–––, and A.A. Reznicek. 1995. Carex conjuncta (Cyperaceae) verified from Arkansas, and notes on the range of Carex 

oklahomensis. Sida 16: 772-774. 

Joseph, and Heimburger. 1966. [Anemone]. Canad. J. Bot. 44: 899-928. 

Judd, W.S. 1979. Generic relationships in the Andromedeae (Ericaceae). J. Arnold Arb. 60: 477-503. 

––––––. 1981. A monograph of Lyonia (Ericaceae). J. Arnold Arbor. 62: 63-209, 315-436. 

–––. 1982. A taxonomic revision of Pieris (Ericaceae). J. Arnold Arb. 63: 103-144. 

–––. 1983. The taxonomic status of Stipulicida filiformis (Caryophyllaceae). Sida 10: 33-36. 

–––. 1984. A taxonomic revision of the American species of Agarista (Ericaceae). J. Arnold Arb. 65: 255-342. 

–––. 1996. The Pittosporaceae in the southeastern United States. Harvard Papers in Botany 8: 15-26. 

–––. 1998. The Smilacaceae in the southeastern United States. Harvard Papers in Botany 3: 147-169. 

–––. 2000. The Hypoxidaceae in the southeastern United States. Harvard Papers in Botany 5: 79-98. 

–––. 2003. The genera of Ruscaceae in the southeastern United States. Harvard Papers in Botany 7: 93-149. 

–––, and I.K. Ferguson. 1999. The genera of Chenopodiaceae in the southeastern United States. Harvard Papers in Botany 4: 

365-416.


–––, and K.A. Kron. 1993. Circumscription of Ericaceae (Ericales) as determined by preliminary cladistic analyses based on 

morphological, anatomical, and embryological features. Brittonia 45: 99-114. 

–––, and K.A. Kron. 1995. A revision of Rhododendron VI. Subgenus Pentanthera (sections Sciadorhodion, Rhodora and 

Viscidula). Edinb. J. Bot. 52: 1-54. 

–––, and K.A. Kron. 1996. Phylogenetic relationships of the Lyonia-group of Andromedeae (Ericaceae): evidence from 

morphology and matK sequence data. Am. J. Bot. 83: 165. [abstract] 

–––, R.W. Sanders, and M.J. Donoghue. 1994. Angiosperm family pairs: preliminary phylogenetic analyses. Harvard Papers in 

Botany 5: 1-51. 

Judziewicz, E.J., R.J. Soreng, G. Davidse, P.M. Peterson, T.S. Filgueiras, and F.O. Zuloaga. 2000. Catalogue of New World 

grasses: I. Subfamilies Anomochlooideae, Bambusoideae, Ehrhartoideae, and Pharoideae. Contributions from the U.S. 

National Herbarium 39: 1-128. 

Kadereit, J.W. 2004. The families and genera of vascular plants. VII. Flowering plants - Dicotyledons - Lamiales (except 

Acanthaceae including Avicenniaceae). Springer, Berlin. 478 pp. 

Källersjö, M., G. Bergqvist, and A. Anderberg. 2000. Generic realignment in primuloid families of the Ericales s.l.: a 

phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology. Am. J. Bot. 87: 1325-1341. 

Kartesz, J.T. 1994. A synonymized checklist of the vascular plants of the United States, Canada, and Greenland. Timber Press, 

Portland, Oregon. 

–––. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and 

Greenland. First Edition. In: Kartesz, J.T., and C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North 

Carolina Botanical Garden, Chapel Hill, NC. 

–––. In prep. Common names for the North American flora. Timber Press, Portland, Oregon. 

–––, and K.N. Gandhi. 1991. Cymophyllus fraserianus (Ker-Gawler) Kartesz & Gandhi (Cyperaceae), the correct name for 

Fraser's sedge. Rhodora 93: 136-140. 

–––, and K.N. Gandhi. 1992. Nomenclatural notes for the North American flora. X. Phytologia 72: 80-92. 

–––, and K.N. Gandhi. 1994. Nomenclatural notes for the North American flora. XIII. Phytologia 76: 441-457. 

–––, and K.N. Gandhi. 1995. Nomenclatural notes for the North American flora. XIV. Phytologia 78: 1-17. 

Kato, H., S. Kawano, R. Terauchi, M. Ohara, and F.H. Utech. 1995. Evolutionary biology of Trillium and related genera 

(Trilliaceae). I. Restriction site mapping and variation of chloroplast DNA and its systematic implications. Plant Species 

Biol. 10: 17-29. 

Kauffman, G.E., G.L. Nesom, A.S. Weakley, T.E. Govus, and L.M. Cotterman. 2004. A new species of Symphyotrichum 

(Asteraceae: Astereae) from a serpentine barren in western North Carolina. Sida 21: 827-839. 

Kawano, S., and H.H. Iltis. 1963. Cytotaxonomy of the genus Polygonatum (Liliaceae). I. Karyotype analysis of some eastern 

North American species. Cytologia 28: 321-330. 

–––, and H. Kato. 1995. Evolutionary biology of Trillium and related genera (Trilliaceae). II. Cladistic analyses on gross 

morphological characters, and phylogeny and evolution of the genus Trillium. Plant Species Biol. 10: 169-183. 

Kazempour Osaloo, S., F.H. Utech, M. Ohara, and S. Kawano. 1999. Molecular systematics of Trilliaceae. I. Phylogenetic 

analyses of Trillium using matK gene sequences. J. Plant Res. 112: 35-49. 

Keating, R.C. 2004. Vegetative anatomical data and its relationship to a revised classification of the genera of the Araceae. 

Ann. Missouri Bot. Gard. 91: 485-494. 

Keener, B.R., and R. Kral. 2003. A new species of Solidago (Asteraceae: Astereae) from north central Alabama. Sida 20: 1589- 

1593. 

Keener, C.S. 1967. A biosystematic study of Clematis section Integrifoliae (Ranunculaceae). J. Elisha Mitchell Sci. Soc. 83: 1- 

41. 

–––. 1975. Studies in the Ranunculaceae of the southeastern United States. III. Clematis L. Sida 6: 33-47. 

–––. 1976. Studies in the Ranunculaceae of the southeastern United States. V. Ranunculus L. Sida 6: 266-283. 

–––. 1977. Studies in the Ranunculaceae of the southeastern United States. VI. Miscellaneous genera. Sida 7: 1-12. 

–––. 1981. The status of Thalictrum hepaticum Greene (Ranunculaceae). Castanea 46: 43-49. 

–––, E.T. Dix, and B.E. Dutton. 1996. The identity of Anemone riparia (Ranunculaceae). Bartonia 59: 37-47. 

–––, and S.B. Hoot. 1987. Ranunculus section Echinella (Ranunculaceae) in the southeastern United States. Sida 12: 57-68. 

Kelloff, C.L., and C.R. Werth. 1998. Allozyme evidence for genetic divergence between two eastern North American varieties 

(angustum and asplenioides) of the Athyrium filix-femina complex [abstract]. Am. J. Bot. 85 [supplement]: 101. 

–––, J. Skog, L. Adamkewicz, and C.R. Werth. 2002. Differentiation of eastern North American Athyrium filix-femina taxa: 

evidence from allozymes and spores. Amer. Fern J. 92: 185-213. 

Kelly, L.M. 1997. A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy. Am. J. 

Bot. 84: 1752-1765. 

–––. 1998. Phylogenetic relationships in Asarum (Aristolochiaceae) based on morphology and ITS sequences. Am. J. Bot. 85: 

1454-1467. 

–––, and F. González. 2003. Phylogenetic relationships in Aristolochiaceae. Systematic Botany 28: 236-249. 

Kesler, T.R., L.C. Anderson, and S.M. Hermann. 2003. A taxonomic reevaluation of Aristida stricta (Poaceae) using anatomy 

and morphology. Southeastern Naturalist 2: 1-10. 

Kessler, J.W. 1987. A treatment of Scleria (Cyperaceae) for North America north of Mexico. Sida 12: 391-407.


Kiers, A.M., T.H.M. Mes, R. van der Meijden, and K. Bachmann. 1999. Morphologically defined Cichorium (Asteraceae) 

species reflect lineages based on chloroplast and nuclear (ITS) DNA data. Systematic Bot. 24: 645-659. 

Kiger, R.W. 1971. Arthraxon hispidus (Gramineae) in the United States: taxonomy and floristic status. Rhodora 73: 39-46. 

Kilpatrick, E.S., and P.D. McMillan. 2003. Noteworthy collections: South Carolina. Castanea 68: 182. 

Kim, K.-J., and B.L. Turner. 1992. Systematic overview of Krigia (Asteraceae - Lactuceae). Brittonia 44: 173-198. 

Kim, S.-C., D.C. Crawford, M. Tadesse, M. Berbee, F.R. Ganders, M. Pirseyedi, and E.J. Esselman. 1999. ITS sequences and 

phylogenetic relationships in Bidens and Coreopsis (Asteraceae). Systematic Botany 24: 480-493. 

–––, C.-W. Park, Y.-D. Kim, and Y. Suh. 2001. Phylogenetic relationships in family Magnoliaceae inferred from ndhF 

sequences. Am. J. Bot. 88: 717-728. 

Kim, Y.-D. 1998. Chloroplast DNA restriction site variation and phylogeny of the Berberidaceae. Amer. J. Bot. 85: 1766-1778. 

–––, and R.K. Jansen. 1996. Phylogenetic implications of rbcL and ITS sequence variation in the Berberidaceae. Systematic 

Botany 21: 381-396. 

–––, S.-H. Kim, and L.R. Landrum. 2004. Taxonomic and phytogeographic implications from ITS phylogeny in Berberis 

(Berberidaceae). J. Plant Res. 117: 175-182. 

Kimball, R.T., D.J. Crawford, J.R. Page, and P.J. Harmon. 2002. Inter-simple sequence repeat (ISSR) diversity within Monarda 

fistulosa var. brevis (Lamiaceae) and divergence between var. brevis and var. fistulosa. Brittonia 53: 511-518. 

King, R.M., and H. Robinson. 1987. The genera of the Eupatoriae (Asteraceae). Monographs in Systematic Botany 22: 1-581. 

Kintsch, J.A., and D.L. Urban. 2002. Focal species, community representation, and physical proxies as conservation strategies: a 

case study in the Amphibolite Mountains, North Carolina, U.S.A. Conservation Biology 16: 936-947. 

Kirkman, W.B., and J.R. Ballington. 1990. Creeping blueberries (Ericaceae: Vaccinium sect. Herpothamnus) – a new look at V. 

crassifolium including V. sempervirens. Systematic Bot. 15: 679-699. 

–––, T.R. Wentworth, and J.R. Ballington. 1989. The ecology and phytosociology of the creeping blueberries, Vaccinium 

section Herpothamnus. Bull. Torrey Bot. Club 116: 114-133. 

Kirschner, J., et al. 2002a. Juncaceae 1: Rostkovia to Luzula. Species Plantarum: Flora of the World 6: 1-237. 

–––, et al. 2002b. Juncaceae 2: Juncus subg. Juncus. Species Plantarum: Flora of the World 7: 1-336. 

–––, et al. 2002c. Juncaceae 3: Juncus subg. Agathryon. Species Plantarum: Flora of the World 8: 1-192. 

Knapp, S., M.W. Chase, and J.J. Clarkson. 2004. Nomenclatural changes and a new sectional classification in Nicotiana 

(Solanaceae). Taxon 53: 73-82. 

Knapp, W.M. 2004. Taxonomic status of Juncus longii, a putative taxon within the Juncus marginatus complex (Juncaceae 

sect. Graminifolii) [abstract]. Southeastern Biology 51: 134. 

–––, and D. Estes. 2006. Gratiola brevifolia (Plantaginaceae) new to the flora of Delaware, the Delmarva Peninsula, and the 

mid-Atlantic. Sida 22: 825-829. 

Knepper, D.A., D.M. Johnson, and L.J. Musselman. 2002. Marsilea mutica in Virginia. Amer. Fern J. 92: 243-244. 

Knobloch, I.W., and D.B. Lellinger. Cheilanthes castanea and its allies in Virginia and West Virginia. Castanea 34: 59-61. 

Knox, J.S. 1987. An experimental garden test of characters used to distinguish Helenium virginicum Blake from H. autumnale 

L. Castanea 52: 52-58. 

–––, M.J. Gutowski, D.C. Marshall, and O.G. Rand. 1995. Tests of the genetic bases of character differences between Helenium 

virginicum and H. autumnale (Asteraceae) using common gardens and transplant studies. Systematic Bot. 20: 120-131. 

–––. 1997. A nine year demographic study of Helenium virginicum (Asteraceae), a narrow endemic seasonal wetland plant. J. 

Torrey Bot. Soc. 124: 236-245. 

Koch, E.W., and R.J. Orth. 2003. The seagrasses of the mid-Atlantic coast of the United States. Pp. 216-223 in E.P. Green and 

F. T. Short. 2003. World Atlas of seagrasses. Prepared by the UNEP World Consservation Monitoring Centre. Univ. of 

Calif. Press, Berkeley, Calif., US. 

Koch, M., and I.A. Al-Shehbaz 2002. Molecular data indicate complex intra- and intercontinental differentiation of American 

Draba (Brassicaceae). Ann. Missouri Bot. Gard. 89: 88-109. 

–––, and I. A. Al-Shehbaz. 2004. Taxonomic and phylogenetic evaluation of the American "Thlaspi" species: identity and 

relationship to the Eurasian genus Noccaea (Brassicaceae). Systematic Botany 29: 375-384 

–––, J. Bishop, and T. Mitchell-Olds. 1999. Molecular systematics and evolution of Arabidopsis and Arabis. Plant Biol. 1: 529- 

537. 

Koch, S.D. 1978. Notes on the genus Eragrostis (Gramineae) in the southeastern United States. Rhodora 80: 390-403. 

Koperski, M., M. Sauer, W. Braun, and S.R. Gradstein. 2000. Referenzliste der Moose Deutschlands. Schriftenreihe für 

Vegetationskunde 34: 1-519. 

Korall, P., P. Kenrick, and J.P. Therrien. 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships 

based on rbcL gene sequences. J. Plant Sci. 160: 585-594. 

Kott, L.S., and D.M. Britton. 1982. A comparative study of sporophyte morphology of three cytotypes of Polypodium 

virginianum in Ontario. Can. J. Bot. 60: 1360-1370. 

–––, and D.M. Britton. 1983. Spore morphology and taxonomy of Isoetes in northeastern North America. Can. J. Bot. 61: 3140- 

3163. 

Koyama, T. 1987. Grasses of Japan and its neighboring regions: an identification manual. Kodansha Ltd., Tokyo. 

Krakow, G.A. 1989. A systematic study of Ilex ambigua, Ilex decidua and related taxa. M.S. thesis, Univ. of Georgia, Athens. 

Král, M. 1966. Die Begrenzung der Gattung Parageum Nakai et Hara. Preslia (Praha) 38: 151-153. 

Kral, R. 1960. A revision of Asimina and Deeringothamnus (Annonaceae). Brittonia 12: 233-278.


–––. 1966a. Xyris (Xyridaceae) of the continental United States and Canada. Sida 2: 177-260. 

–––. 1966b. Observations on the flora of the southeastern United States with special reference to northern Louisiana. Sida 2: 

395-408. 

–––. 1966c. {Eriocaulaceae} Sida 2: 285-332. 

–––. 1971. A treatment of Abildgaardia, Bulbostylis, and Fimbristylis (Cyperaceae) for North America. Sida 4: 57-227. 

–––. 1973. Some notes on the flora of the southern states, particularly Alabama and middle Tennessee. Rhodora 75: 366-410. 

–––. 1976. A treatment of Delphinium for Alabama and Tennessee. Sida 6: 243-265. 

–––. 1978a. A synopsis of Fuirena (Cyperaceae) for the Americas north of South America. Sida 7: 309-354. 

–––. 1978b. A new species of Xyris (sect. Xyris) from Tennessee and northwestern Georgia. Rhodora 80: 444-447. 

–––. 1981a. Notes on some "quill"-leaved umbellifers. Sida 9:124-134. 

–––. 1981b. Some distributional reports of weedy or naturalized foreign species of vascular plants for the southern states, 

particularly Alabama and middle Tennessee. Castanea 46: 334-339. 

–––. 1982. A new phyllodial-leaved Sagittaria (Alismaceae) from Alabama. Brittonia 34: 12-17. 

–––. 1983a. A report on some rare, threatened, or endangered forest-related vascular plants of the South. Vol. I and II. USDA 

Forest Service Tech. Publ. R8-TP2. Atlanta, GA. 

–––. 1983b. The Xyridaceae in the southeastern United States. J. Arnold Arb. 64: 421-429. 

–––. 1992. A new species of Fimbristylis (Cyperaceae) from the sandstone and granitic outcrops of Alabama and Georgia. Sida 

15: 317-321. 

–––. 1996. Supplemental notes on Rhynchospora crinipes and related species in section Fuscae (Cyperaceae). Sida 17: 385- 

411. 

–––. 1999. A revised taxonomy for two North American Rhynchospora (Cyperaceae) and for two North American Xyris 

(Xyridaceae). Novon 9: 205-219. 

–––. 2004. An evaluation of Anthenantia. Sida 21: 293-310. 

–––, and P.E. Bostick. 1969. The genus Rhexia (Melastomataceae). Sida 3: 387-440. 

–––, and R.K. Godfrey. 1958. Synopsis of the Florida species of Cacalia. Quart. J. Florida Acad. Sci. 21: 193-206. 

–––, and G.L. Nesom. 2003. Two new species of Liatris series Graminifoliae (Asteraceae: Eupatorieae) from the southeastern 

United States. Sida 20: 1573-1583. 

–––, and B.A. Sorrie. 1998. Proposal to conserve the name Eriocaulon lineare (Erocaulaceae) with a conserved type. Taxon 47: 

741-742. 

Kramer, K.U., and P.S. Green. 1990. Pteridophytes and Gymnosperms. In K. Kubitzki, ed., The families and genera of vascular 

plants. Springer-Verlag, Berlin. 

Kress, W.J., G.D. Maddox, and C.S. Roesel. 1994. Genetic variation and protection priorities in Ptilimnium nodosum 

(Apiaceae), an endangered plant of the eastern United States. Conservation Biology 8: 271-276. 

Krings, A. 2002. Floral variation and disgnosis of Richardia (Rubiaceae) in the Carolinas. Castanea 67: 329-330. 

–––. 2003. Typification and nomenclatural history of Trachelospermum difforme (Apocynaceae). Sida 20: 1641-1644. 

–––. 2005. Neotypification of Ceropegia palustris and Lyonia maritima (Apocynaceae: Asclepiadoideae). Sida 21: 1507-1513. 

–––, and J.C. Neal. 2001a. A Scutellaria (Lamiaceae) new to North Carolina and a key to the small-flowered Carolina 

congeners. Sida 19: 735-739. 

–––, and J.C. Neal. 2001b. South American skullcap (Scutellaria racemosa: Lamiaceae) in the southeastern United States. Sida 

19: 1171-1179. 

–––, and R.J. Richardson. 2006. Cayratia japonica (Vitaceae) new to North Carolina and an updated key to the genera of 

Vitaceae in the Carolinas. Sida 22: 813-815. 

–––, A.S. Weakley, J.C. Neal, and E.C. Swab. 2005. Ranunculus ficaria (Ranunculaceae) new to North carolina and an updated 

key to Carolina congeners. Sida 21: 2429-2437. 

–––, R. Westbrooks, and J. Lloyd. 2002. Cirsium nuttallii (Asteraceae: Cynareae) new to North Carolina and an illustrated key 

to southeastern congeners. Sida 20: 845-848. 

–––, and Q.-Y. (Jenny) Xiang, 2004. The Gonolobus complex (Apocynaceae: Asclepiadoideae) in the southeastern United 

States. Sida 21: 103-116. 

–––, and Q.-Y. (Jenny) Xiang. 2005. Taxonomy of the Gonolobus complex (Apocynaceae, Asclepiadoideae) in the southeastern 

United States: ISSR evidence and parsimony analysis. Harvard Papers in Botany 10: 147-159. 

Kron, K.A. 1993. A revision of Rhododendron section Pentanthera. Edinb. J. Bot. 50: 249-364. 

–––, and M.W. Chase. 1993. Systematics of the Ericaceae, Empetraceae, Epacridaceae and related taxa based upon rbcL 

sequence data. Ann. Mo. Bot. Gard. 80: 735-741. 

–––, and M. Creel. 1999. A new species of deciduous azalea (Rhododendron section Pentanthera; Ericaceae) from South 

Carolina. Novon 9: 377-380. 

–––, and J.M. King. 1996. Cladistic relationships of Kalmia, Leiophyllum, and Loiseleuria (Phyllodoceae, Ericaceae) based on 

rbcL and nrITS data. Systematic Bot. 21: 17-30. 

–––, W.S. Judd, P.F. Stevens, D.M. Crayn, A.A. Anderberg, P.A. Gadek, C.J. Quinn, and J.L. Luteyn. 2002. Phylogenetic 

classification of Ericaceae: molecular and morphological evidence. The Botanical Review 68: 335-423. 

Kruijt, R.C. 1996. A taxonomic monograph of Sapium Jacq., Anomostachys (Baill.) Hurus., Duvigneaudia J. Léonard and 

Sclerocroton Hochst. (Euphorbiaceae tribe Hippomaneae). E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart.


Kubitzki, K., J.G. Rohwer, and V. Bittrich, eds. 1993. The families and genera of vascular plants. II. Flowering plants - 

Dicotyledons - Magnoliid, Hamamelid and Caryophyllid families. Springer, Berlin. 653 pp. 

–––, ed. 1998a. The families and genera of vascular plants. III. Flowering plants - Monocotyledons - Lilianae (except 

Orchidaceae). Springer, Berlin. 478 pp. 

–––, ed. 1998b. The families and genera of vascular plants. IV. Flowering plants - Monocotyledons - Alismatanae and 

Commelinanae (except Gramineae). Springer, Berlin. 511 pp. 

–––, ed. 2004. The families and genera of vascular plants. VI. Flowering plants - Dicotyledons - Celastrales, Oxalidales, 

Rosales, Cornales, Ericales. Springer, Berlin. 489 pp. 

–––, and C. Bayer, eds. 2003. The families and genera of vascular plants. V. Flowering plants - Dicotyledons - Malvales, 

Capparales, and non-betalain Caryophyllales. Springer, Berlin. 418 pp. 

Kuijt, J. 1982. The Viscaceae in the southeastern United States. J. Arnold Arb. 63: 401-410. 

–––. 2003. Monograph of Phoradendron (Viscaceae). Systematic Botany Monographs 66: 1-643. 

Kurashige, Y., J.I. Etoh, T. Handa, K. Takayanagi, and T. Yukawa. 2001. Sectional relationships in the genus Rhododendron 

(Ericaceae): evidence from matK and trnK intron sequences. Plant Systematics and Evolution 228: 1-14. 

Kurz, H., and R.K. Godfrey. 1962. Trees of northern Florida. University Press of Florida, Gainesville, FL. 311 pp. 

LaFrankie, J.V., Jr. 1986. Transfer of the species of Smilacina to Maianthemum (Liliaceae). Taxon 35: 584-589. 

Lakela, O. 1937. A monograph of the genus Tiarella L. in North America. Amer. J. Bot. 24: 344-351. 

Lamb Frye, A.S., and K.A. Kron. 2003. rbcL phylogeny and character evolution in Polygonaceae. Systematic Botany 28: 326- 

332. 

Lamboy, W.F. 1987. Aster section Biotia (Asteraceae) in New England and the status of Aster glomeratus. Rhodora 89: 299- 

318. 

–––. 1988. The status of Aster commixtus and a new species of Aster from the southeastern United States. Systematic Botany 

13: 187-195. 

–––. 1992. The taxonomic status and probable origin of Aster chlorolepis, a Southern Appalachian endemic. Castanea 57: 52- 

65. 

Lamont, E.E. 1990. A new combination in Eupatorium section Verticillata (Asteraceae). Phytologia 69: 467-468. 

–––. 1995. Taxonomy of Eupatorium section Verticillata (Asteraceae). Memoirs New York Botanical Garden 72: 1-68. 

Lance, R. 1995. The hawthorns of the southeastern United States. Published by the author, Fletcher, NC. 136 pp. 

Landolt, E. 1980. Key to the determination of taxa within the family of Lemnaceae. Veröffentlichungen des Geobotanischen 

Institutes der Eidg. Techn. Hochschule, Stiftung Rübel, Zürich 70: 13-21. 

–––. 1986. The family of Lemnaceae – a monographic study. Veröffentlichungen des Geobotanischen Institutes der Eidg. 

Techn. Hochschule, Stiftung Rübel, Zürich 71: 1-566. 

Lane, C. 2005. Witch hazels. Timber Press & Royal Horticulatural Society Plant Collectors Guide. 

Lane, M.A., D.R. Morgan, Y. Suh, B.B. Simpson, and R.K. Jansen. 1996. Relationships of North American genera of Astereae, 

based on chloroplast DNA restriction site data. Pp. 49-77 in D.J.N. Hind and H.J. Beentje (eds.) Compositae: Systematics. 

Proceedings of the International Compositae Conference, Kew, 1994, vol. 1. 

Larisey. M.M. 1940a. A monograph of the genus Baptisia. Ann. Mo. Bot. Garden 27: 119-244. 

–––. 1940b. A revision of the North American species of the genus Thermopsis. Ann. Mo. Bot. Garden 27: 245-258. 

Le Duc, A. 1995. A revision of Mirabilis section Mirabilis (Nyctaginaceae). Sida 16: 613-648. 

Leadlay, S.A., and V.H. Heywood. 1990. The biology and systematics of the genus Coincya Porta & Rigo ex Rouy (Cruciferae). 

Bot. J. Linn. Soc. 102: 313-398. 

LeBlond, R.J. 1997. Distribution of Rhynchospora harperi. Castanea 62: 278-280. 

–––. 2000. Solidago villosicarpa (Asteraceae: Astereae), a rare new southeastern Coastal Plain endemic. Sida 19: 291-300. 

–––. 2001a. Taxonomy of the Dichotoma group of Dichanthelium (Poaceae). Sida 19: 821-837. 

–––. 2001b. Endemic plants of the Cape Fear Arch region. Castanea 66: 83-97. 

–––, and A.S. Weakley. 2002. Schizaea pusilla Pursh (Schizaeaceae) in North Carolina. Rhodora 104: 86-91. 

–––, A.S. Weakley, A.A. Reznicek, and W.J. Crins. 1994. Carex lutea (Cyperaceae), a rare new Coastal Plain endemic from 

North Carolina. Sida 16: 153-161. 

Lee, N.S., T. Sang, D.J. Crawford, S.H. Yeau, S.-C. Kim. 1996. Molecular divergence between disjunct taxa in eastern Asia and 

eastern North America. Am. J. Bot. 83: 1373-1378. 

Lee, Yin-Tse. 1976. The genus Gymnocladus and its tropical affinity. J. Arnold Arb. 57: 91-112. 

Lellinger, D.B. 1985. A field manual of the ferns and fern allies of the United States and Canada. Smithsonian Institution 

Press, Washington, D.C. 

Lelong, M.G. 1984. New combinations for Panicum subgenus Panicum and subgenus Dichanthelium (Poaceae) of the 

southeastern United States. Brittonia 36: 262-273. 

–––. 1986. A taxonomic treatment of the genus Panicum (Poaceae) in Mississippi. Phytologia 61: 251-269. 

Leonard, E.C. 1927. The North American species of Scutellaria. Contr. U.S. National Herbarium 22: 703-748. 

Leonard, M.R., R.E. Cook, and J.C. Semple. 2005. A multivariate morphometric study of the aster genus Sericocarpus 

(Asteraceae: Astereae). Sida 21: 1472-1505. 

Leonard, S.W. 1971a. The distribution of Thelypteris torresiana in the southeastern United States. Amer. Fern J. 62: 97-99. 

–––. 1971b. Additions to the flora of the Carolinas. J. Elisha Mitchell Sci. Soc. 87: 97-100. 

–––. 1981a. Fimbristylis perpusilla Harper in South Carolina. Castanea 46: 235-236.


–––. 1981b. Fimbristylis perpusilla Harper in South Carolina. Castanea 46: 341-343. 

–––. 1987. Fimbristylis perpusilla in North Carolina. Castanea 52: 150. 

–––. 2006. A new species of witch-hazel (Hamamelis: Hamamelidaceae) apparently endemic to southern Mississippi. Sida 22: 

849-856. 

Les, D.H. 1985. The taxonomic significance of plumule morphology in Ceratophyllum (Ceratophyllaceae). Systematic Bot. 10: 

338-346. 

–––. 1986. The evolution of achene morphology in Ceratophyllum (Ceratophyllaceae), I. Fruit-spine variation and 

relationships of C. demersum, C. submersum, and C. apiculatum. Systematic Bot. 11: 549-558. 

–––. 1988a. The evolution of achene morphology in Ceratophyllum (Ceratophyllaceae), II. Fruit variation and systematics of 

the "spiny-margined" group. Systematic Bot. 13: 73-86. 

–––. 1988b. The evolution of achene morphology in Ceratophyllum (Ceratophyllaceae), III. Relationships of the "faciallyspined" 

group. Systematic Bot. 13: 509-518. 

–––. 1988c. The origin and affinities of the Ceratophyllaceae. Taxon 37: 326-345. 

–––. 1989. The evolution of achene morphology in Ceratophyllum (Ceratophyllaceae), IV. Summary of proposed relationships 

and evolutionary trends. Systematic Bot. 14: 254-262. 

–––, G.J. Anderson, and M.A. Cleland. 1995. Sterility in the North American lake cress Neobeckia aquatica (Brassicaceae): 

Inferences from chromosome number. Rhodora 97: 185-200. 

–––, and D.J. Crawford. 1999. Landoltia (Lemnaceae), a new genus of duckweeds. Novon 9: 530-533. 

–––, and R.P. Wunderlin. 1981. Hygrophila polysperma (Acanthaceae) in Florida. Florida Sci. 44: 189-192. 

–––, R.S. Capers, and N.P. Tippery. 2006. Introduction of Glossostigma (Phrymaceae) to North America: a taxonomic and 

ecological overview. Amer. J. Bot. 93: 927-939. 

Levin, G.A. 1999a. Evolution in the Acalypha gracilens/monococca complex (Euphorbiaceae): morphological analysis. 


–––. 1999b. Notes on Acalypha (Euphorbiaceae) in North America. Rhodora 101: 217-233. 

Levy, F. 1991a. Morphological differentiation in Phacelia dubia and P. maculata. Rhodora 93: 11-25. 

Levy, F. 1991b. A genetic analysis of reproductive barriers in Phacelia dubia. Heredity 67:331-345. 

Levy, F. 1997. Non-homeotic meristic flower mutants in Phacelia dubia. J. Heredity 88:31-37. 

Levy, F. and K. A. Malone. 2001. Phacelia dubia in South Carolina: the interface of morphology, genetics and taxonomy. 

Castanea 66:134-144. 

Levy F. and C. L. Neal. 1999. Spatial and temporal genetic structure in chloroplast and allozyme markers in Phacelia dubia 

implicate genetic drift. Heredity 82:422-431. 

Levy, F., Antonovics, J., Boynton, J. E. and N. W. Gillham. 1996. A population genetic analysis of chloroplast DNA in 

Phacelia. Heredity 76:143-155. 

Lewis, D.Q. 2000. A revision of the New World species of Lindernia (Scrophulariaceae). Castanea 65: 93-122. 

Lewis, W.H. 2006. Hedyotis australis (Rubiaceae) new to Missouri and Florida and related species in the south-central United 

States. Sida 22: 831-836. 

–––, and R.L. Oliver. 1965. Realignment of Calystegia and Convolvulus (Convolvulaceae). Ann. Missouri Bot. Gard. 52: 217- 

222 

–––, and R.L. Oliver. 1974. Revision of Richardia (Rubiaceae). Brittonia 26: 271-301. 

Li, J., and M.J. Donoghue. 1999. More molecular evidence for interspecific relationships in Liquidambar (Hamamelidaceae). 

Rhodora 101: 87-91. 

–––, J. Alexander III, T. Ward, P. del Tredici, and R. Nicholson. Phylogenetic relationships of Empetraceae inferred from 

sequences of chloroplast gene matK and nuclear ribosomal DNA ITS region. Molecular Phylogenetics and Evolution 25: 

306-315. 

–––, P. del Tredici, S. Yang, and M.J. Donoghue. 2002. Phylogenetic relationships and biogeography of Stewartia 

(Camellioideae, Theaceae) inferred from nuclear ribosomal DNA ITS sequences. Rhodora 104: 117-133. 

–––, J. Ledger, T. Ward, and P. del Tredici. 2004. Phylogenetics of Calycanthaceae based on molecular and morphological data, 

with a special reference to divergent paralogues of the nrDNA ITS region. Harvard Papers in Bot. 9: 69-82. 

Libby, G.W., and C.T. Bloom. 1998. Nestronia umbellula Rafinesque (Santalaceae) from the Highland Rim of Kentucky. 

Castanea 63: 161-164. 

Lidén, M. 1981. Proposal to change the typification of Corydalis nomen conservandum. Taxon 30: 323-325. 

–––. 1986. Synopsis of Fumarioideae (Papaveraceae) with a monograph of the tribe Fumarieae. Opera Botanica 88: 1-133. 

–––, T. Fukuhara, J. Rylander, and B. Oxelman. 1997. Phylogeny and classification of Fumariaceae, with emphasis on Dicentra 

s.l., based on the plastid gene rps16 intron. Pl. Syst. Evol. 206: 411-420. 

Liede, S. 1997a. Subtribes and genera of the tribe Asclepiadaceae (Apocynaceae, Asclepiadoideae) – a synopsis. Taxon 46: 

233-247. 

–––. 1997b. American Cynanchum (Asclepiadaceae) – a preliminary infrageneric classification. Novon 7: 172-181. 

–––, and U. Meve. 1997. Some clarifications, new species, and new combinations in American Cynanchinae (Asclepiadaceae). 

Novon 7: 38-45. 

–––, and U. Meve. 2003. Dissolution of Cynanchum sect. Macbridea (Apocynaceae – Asclepiadoideae). Nordic J. of Bot. 22: 

579–591


–––, and A. Täuber. 2002. Circumscription of the genus Cynanchum (Apocynaceae – Asclepiadoideae). Systematic Botany 27: 

789-800. 

Lihová, J., K. Marhold, H. Kudoh, and M.A. Koch. 2006. Worldwide phylogeny and biogeography of Cardamine flexuosa 

(Brassicaceae) and its relatives. Amer. J. Botany 93: 1206-1221. 

Lint, H., and C. Epling. 1945. A revision of Agastache. Amer. Midl. Nat. 33: 207-230. 

Lipow, S.R., and R. Wyatt. 1998. Reproductive biology and breeding system of Gonolobus suberosus (Asclepiadaceae). J. 

Torrey Bot. Soc. 125: 183-193. 

Little, D.P., A.E. Schwarzbach, R.P. Adams, and C.-F. Hsieh. 2004. The circumscription and phylogenetic relationships of 

Callitropsis and the newly described genus Xanthocyparis (Cupressaceae). Amer. J. Bot. 91: 1872-1881. 

Little, E.L., Jr. 1969. Two varietal transfers in Carya (Hickory). Phytologia 19: 186-190. 

Liu, T.-S. 1971. A monograph of the genus Abies. Dept. of Forestry, National Taiwan University, Taipei. 

Lledó, M.D., M.B. Crespo, K.M. Cameron, M.F. Fay, and M.W. Chase. 1998. Systematics of Plumbaginaceae based upon 

cladistic analysis of rbcL sequence data. Systematic Bot. 23: 21-29. 

Loconte, H., and W.H. Blackwell. 1981. A new species of blue cohosh (Caulophyllum, Berberidaceae) in eastern North 

America. Phytologia 49: 483. 

––––––, and W.H. Blackwell. 1984. Berberidaceae in Ohio. Castanea 49: 39-43. 

–––, and W.H. Blackwell. 1985. Intrageneric taxonomy of Caulophyllum (Berberidaceae). Rhodora 87: 463-469. 

–––, and J.R. Estes. 1989a. Generic relationships within Leonticeae (Berberidaceae). Can. J. Bot. 67: 2310-2316 

–––, and J.R. Estes. 1989b. Phylogenetic systematics of Berberidaceae and Ranunculales (Magnoliidae). Systematic Bot. 14: 

565-579. 

Long, R.W. 1970. The genera of Acanthaceae in the southeastern United States. J. Arnold Arb. 51: 257-309. 

Longhi-Wagner, H.M., and S.A. Renvoize. 2004. The genus Ctenium (Poaceae – Cynodonteae) in Bolivia. Kew Bull. 59: 305- 

309. 

Lourteig, A. 1979. Oxalidaceae extra-Austroamericanae. II. Oxalis L. Sectio Corniculatae DC. Phytologia 42: 57-198. 

Lowden, R.M. 1973. Revision of the genus Pontederia L. Rhodora 75: 426-487. 

Lowry, P.P., II, and A.G. Jones. 1979. Biosystematic investigations and taxonomy of Osmorhiza Section Osmorhiza (Apiaceae) 

in North America. Amer. Midl. Naturalist 101: 21-27. 

Ludwig, J.C. 1999. The flora of dolomite and limestone barrens in southwestern Virginia. Castanea 64: 209-230. 

Luebke, N.T., and J.M. Budke. 2003. Isoëtes tennesseensis (Isoëtaceae), an octoploid quillwort from Tennessee. American Fern 

J. 93: 184-190. 

Luer, C.A. 1972. The native orchids of Florida. New York Botanical Garden, Bronx, New York. 

–––. 1975. The native orchids of the United States and Canada, excluding Florida. New York Botanical Garden, Bronx, New 

York. 

Luken, J.O., J.W. Thieret, and J.R. Kartesz. 1993. Erucastrum gallicum (Brassicaceae): invasion and spread in North America. 

Sida 15: 569-582. 

Luteyn, J.L. 1976. Revision of Limonium (Plumbaginaceae) in eastern North America. Brittonia 28: 303-317. 

–––, W.S. Judd, S.P. Vander Kloet, L.J. Dorr, G.D. Wallace, K.A. Kron, P.F. Stevens, and S.E. Clemants. 1996. Ericaceae of 

the southeastern United States. Castanea 61: 101-144. 

Ma, Yu-Chuan. 1951. Gentianopsis: a new genus of Chinese Gentianaceae. Acta Phytotax. Sinica 1: 5-19. 

Mabberley, D.J. 1997a. The plant-book: a portable dictionary of the vascular plants. Cambridge Univ. Press, Cambridge, U.K. 

–––. 1997b. A classification for edible Citrus (Rutaceae). Telopea 7: 167-172. 

Mackenzie, K.K. 1931-1935. Poales, Cyperaceae, Cypereae (pars). North American Flora, vol. 18, Parts 1-7. 

MacRoberts, M.H., and B.R. MacRoberts. 1992. Observations on toothache grass (Ctenium aromaticum [Poaceae: Chlorideae]) 

with particular reference to fire. Phytologia 73: 439-444. 

–––, and B.R. MacRoberts. 2004. Sarracenia purpurea (Sarraceniaceae) in Louisiana, Sida 21: 1149-1152. 

–––, B.R. MacRoberts, and L.S. Jackson. 2004. Observations on Parnassia grandifolia DC. (Saxifragaceae) in the west Gulf 

Coastal Plain. Phytologia 86: 98-103. 

Maddox, D., and R. Bartgis. 1990. Harperella (Ptilimnium nodosum) recovery plan. U.S. Fish and Wildlife Service, Newton 

Corner, MA. 55 pp. 

Maguire, B. 1950. Studies in the Caryophyllaceae – IV. a synopsis of the North American species of the subfamily Silenoideae. 

Rhodora 52: 233-245 

Mahler, W.F. 1975. Typification and distribution of the varieties of Gnaphalium helleri Britton (Compositae-Inuleae). Sida 6: 

30-32. 

Mahoney, A. M. 1998. Packera paupercula – predatory compilo-species or mare's nest of convergent species-in-progress 

[abstract]. Am. J. Bot. 85 [supplement]: 109-110. 

–––, and R.R. Kowal. Four new varieties and one new combination in the Packera paupercula complex in eastern North 

America. Sida [in press]. 

Manen, J.-F., M.C. Boulter, and Y. Naciri-Graven. 2002. The complex history of the genus Ilex L. (Aquifoliaceae): evidence 

from the comparison of plastid and nuclear DNA sequences and from fossil data. Plant Syst. Evol. 235: 79-98. 

–––, C. Habashi, D. Jeanmonod, J.-M. Park, and G.M. Schneeweiss. 2004. Phylogeny and infraspecific variability of 

holoparasitic Orobanche (Orobanchaceae) inferred from plastid rbcL sequences. Molec. Phylo. and Evol. 33: 482-500.


Mangaly, J.K. 1968. A cytotaxonomic study of the herbaceous species of Smilax: section Coprosmanthus. Rhodora 70: 55-82, 

247-273. 

Manhart, J.R. 1984. A biosystematic study of Carex section Laxiflorae. Ph.D. dissertation, University of Georgia. 

Manitz, H. 1983. Zur Nomenclatur einiger Convolvulaceae und Cuscutaceae. I. Feddes Repert. 94: 173-182. 

Manning, W.E. 1950. A key to the hickories north of Virginia with notes on the two pignuts, Carya glabra and C. ovalis. 

Rhodora 52: 188-199. 

Manning, S.D. 2000. The genera of Bignoniaceae in the southeastern United States. Harvard Papers in Botany 5: 1-77. 

Marazzi, B., P.K. Endress, L. P. de Queiroz, and E. Conti. 2006. Phylogenetic relationships within Senna (Leguminosae, 

Cassiinae) based on three chloroplast DNA regions: patterns in the evolution of flora symmetry and extrafloral nectaries. 

Amer. J. Bot. 93: 288-303. 

Maréchal, R., J.-M. Mascherpa, and F. Stainier. 1978. Étude taxonomique d'un groupe complexe d'espèces des genres Phaseolus 

et Vigna (Papilionaceae) sur le base de données morphologiques et polliniques, traitées par l'analyse informatique. Boissiera 

28: 1-273. 

Martínez, M. 1993. The correct application of Physalis pruinosa L. (Solanaceae). Taxon 42: 103-104. 

Martins, L., C. Oberprieler, and F.H. Hellwig. 2003. A phylogenetic analysis of Primulaceae s.l. based on internal transcribed 

spacer (ITS) DNA sequence data. Plant. Syst. Evol. 237: 75-85. 

Maskas, S.D., and M.B. Cruzan. 2000. Patterns of infraspecific diversification in the Piriqueta caroliniana complex in 

southeastern North America and the Bahamas. Evolution 54: 815-827. 

Maslin, B.R., J.T. Miller, and D.S. Seigler. 2003. Overview of the generic status of Acacia (Leguminosae: Mimosoideae). 

Australian Systematic Botany 16: 1-18. 

Massey, A.B. 1944. The ferns and fern allies of Virginia. Bull. Va. Polytechnic Institute 37: 1-110 

Massey, J.R. 1975. Fatoua villosa (Moraceae): additional notes on distribution in the southeastern United States. Sida 6: 116. 

–––––, D.K.S. Otte, T.A. Atkinson, and R.D. Whetstone. 1983. An atlas and illustrated guide to the threatened and endangered 

vascular plants of the mountains of North Carolina and Virginia. Southeastern Forest Experiment Station General Technical 

Report SE-20, Asheville, N.C. 

Mast, A.R., D.M.S. Feller, S. Kelso, and E. Conti. 2004. Buzz-pollinated Dodecatheon originated fro within the heterostylous 

Primula subgenus Auriculastrum (Primulaceae): a seven-region cpDNA phylogeny and its implications for floral evolution. 

Amer. J. Bot. 91: 926-942. 

Mathew, B. 1992. A taxonomic and horticultural review of Erythronium L. (Liliaceae). Bot. J. Linn. Soc. 109: 453-471. 

–––. 1996. A review of Allium sect. Allium. Royal Botanic Gardens, Kew, England. 

Mathias, M.E., and L. Constance. 1945. Umbelliferae. North American Flora, vol. 28B: 43-397. N.Y. Botanical Garden, New 

York. 

Matthews, C.R., and J.H. Howard. 1999. Genetic variation in the federally endangered Schweinitz's sunflower, Helianthus 

schweinitzii T. & G. (Asteraceae). Castanea 64: 231-242. 

Matthews, J.F., L.S. Barden, and C.R. Matthews. 1997. Corrections of the chromosome number, distribution and 

misidentifications of the federally endangered sunflower, Helianthus schweinitzii T. & G. J. Torrey Botanical Society 124: 

198-209. 

–––, J.R. Allison, R.T. Ware, Sr., and C. Nordman. 2002. Helianthus verticillatus Small (Asteraceae) rediscovered and 

redescribed. Castanea 67: 13-24. 

–––, W.R. Faircloth, and J.R. Allison. 1991. Portulaca biloba Urban (Portulacaceae), a species new to the United States. 


–––, and D.W. Ketron. 1991. Two new combinations in Portulaca (Portulacaceae). Castanea 56: 304-305. 

–––, D.W. Ketron, and S.F. Zane. 1992a. The reevaluation of Portulaca pilosa and P. mundula (Portulacaceae). Sida 15: 71-89. 

–––, D.W. Ketron, and S.F. Zane. 1992b. Portulaca umbraticola Kunth (Portulacaceae) in the United States. Castanea 57: 202- 

208. 

–––, D.W. Ketron, and S.F. Zane. 1993. The biology and taxonomy of the Portulaca oleracea L. (Portulaceae) complex in 

North America. Rhodora 95: 166-183. 

–––, and P.A. Levins. 1985a. The genus Portulaca in the southeastern United States. Castanea 50: 96-104. 

–––, and P.A. Levins. 1985b. Portulaca pilosa L., P. mundula I.M. Johnst. and P. parvula Gray in the Southwest. Sida 11: 45- 

61. 

–––, and P.A. Levins. 1986. The systematic significance of seed morphology in Portulaca (Portulacaceae) under scanning 

electron microscopy. Systematic Bot. 11: 302-308. 

Matthews, J.F., and A.E. Radford. 1985. New reports of Calamagrostis porteri A. Gray from North Carolina. Castanea 50: 202. 

Mayfield, M.H. 2002. The varieties of Liatris elegans (Asteraceae). Sida 20: 597-603. 

Mazzeo, P.M. 1974. Betula uber – what is it and where is it? Castanea 39: 273-278. 

McAllister, H., and K. Ashburner. 2004. Plate 487. Betula lenta forma uber; Betulaceae. Curtis's Botanical Magazine 21: 54- 

60. 

McAtee, W.L. 1956. A review of the nearctic Viburnum. Privately published by the author, Chapel Hill, NC. 

McAvoy, W.A. 2002. Amaranthus pumilus Raf. (seabeach amaranth, Amaranthaceae) rediscovered in Sussex County, 

Delaware. Bartonia 61: 147-148. 

McCartney, R.B., K. Wurdack, and J.H. Moore. 1989. The genus Lindera in Florida. The Palmetto, Summer 1989: 3-8. 

McClintock, E., and C. Epling. 1942. A review of the genus Monarda (Labiatae). Univ. of Calif. Publ. in Botany 20: 147-194.


McClintock, E. 1957. A monograph of the genus Hydrangea. Proc. Calif. Acad. Sci. 29: 147-256. 

McClintock, K.A., and M.J. Waterway. 1994. Genetic differentiation between Carex lasiocarpa and C. pellita (Cyperaceae) in 

North America. Am. J. Bot. 81: 224-231. 

McClure, F.A. 1963. A new feature in bamboo rhizome anatomy. Rhodora 65: 134-136. 

–––. 1973. Genera of bamboos native to the New World. Smithsonian Contr. Bot. 9: 1-148. 

McCormac, J.S., J.K. Bissell, and S.J. Stine, Jr. 1995. The status of Fraxinus tomentosa (Oleaceae) in Ohio with notes on its 

occurrence in Michigan and Pennsylvania. Castanea 60: 70-78. 

McCormick, J.F., and R.B. Platt. 1964. Ecotypic differentiation in Diamorpha cymosa. Bot. Gazette 125: 271-279. 

McCoy, J.-A. 2004. Noteworthy collections: North Carolina. Castanea 69: 329. 

McDade, L.A., and M.L. Moody. 1999. Phylogenetic relationships among Acanthaceae: evidence from noncoding trnL-trnF 

chloroplast DNA sequences. Amer. J. Bot. 86: 70-80. 

–––, S.E. Masta, M.L. Moody, and E. Waters. 2000. Phylogenetic relationships among Acanthaceae: evidence from two 

genomes. Systematic Bot. 25: 106-121. 

McDaniel, J.C. 1966. Variations in the sweet bay magnolias. Morris Arboretum Bull. 17: 7-12. 

McDaniel, S. 1968. Harperocallis, a new genus of the Liliaceae from Florida. J. Arnold Arb. 49: 35-40. 

–––. 1971. The genus Sarracenia (Sarraceniaceae). Tall Timbers Research Station Bull. 9: 1-36. 

–––. 1986. Taxonomic study of three Sarracenia subspecies (S. rubra ssp. alabamensis, S. rubra ssp. wherryi, and S. rubra ssp. 

jonesii). Report to the U.S. Fish and Wildlife Service. 

McDougal, K.M., and C.R. Parks. 1984. Elevational variation in foliar flavonoids of Quercus rubra L. (Fagaceae). Am. J. Bot. 

71: 301-308. 

McDowell, G.W. 1969. American Yew in North Carolina. J. Elisha Mitchell Sci. Soc. 85: 16-17. 

–––. 1984. Bogbean and shinleaf in North Carolina. Castanea 49: 203. 

McGregor, R.L. 1968. The taxonomy of the genus Echinacea (Compositae). Univ. Kansas Sci. Bull. 48: 113-142. 

McKenney, T.C. 1967. Differentiation of sterile specimens of Nyssa sylvatica Marsh. and Diospyros virginiana L. Castanea 32: 

186-189. 

McKenzie, P.M., B. Jacobs, C.T. Bryson, G.C. Tucker, and R. Carter. 1998. Cyperus fuscus (Cyperaceae), new to Missouri and 

Nevada, with comments on its occurrence in North America. Sida 18: 325-333. 

–––, and D. Ladd. 1995. Status of Bromus nottowayanus (Poaceae) in Missouri. Missouriensis 16: 57-68. 

–––, L.E. Urbatsch, and C. Aulbach-Smith. 1987. Eustachys caribaea (Poaceae), a species new to the United States and a key to 

Eustachys in the United States. Sida 12: 227-232. 

McKenzie, R.J., E.M. Muller, A.K.W. Skinner, P.O. Karis, and N.P. Barker. 2006. Phylogenetic relationships and generic 

delimitation in subtribe Arctotidineae (Asteraceae: Arctotideae) inferred by DNA sequence data from ITS and five 

chloroplast regions. Amer. J. Botany 93: 1222-1235. 

McKeown, K. 1999. A review of the taxonomy of the genus Echinacea, in J. Janick (ed.). Perspectives on new crops and new 

uses. ASHS Press, Alexandria, VA. http://www.hort.purdue.edu/newcrop/proceedings1999/pdf/v4-482.pdf. Accessed 8 

December 2005. 

McKinney, L.E. 1992. A taxonomic revision of the acaulescent blue violets (Viola) of North America. Sida Bot. Miscellany 7: 

1-60. 

–––, and N.H. Russell. 2002. Violaceae of the southeastern United States. Castanea 67: 369-379. 

McMillan, P.D. 2003. Noteworthy collections: South Carolina. Castanea 68: 345-347. 

–––, R.K. Peet, R.D. Porcher, and B.A. Sorrie. 2002. Noteworthy botanical collections from the fire-maintained pineland and 

wetland communities of the coastal plain of the Carolinas and Georgia. Castanea 67: 61-83. 

McNamara, J., and J.A. Quinn. 1977. Resource allocation and reproduction in populations of Amphicarpum purshii 

(Gramineae). Am. J. Bot. 64: 17-23. 

McNeill, J. 1976. Nomenclature of four perennial species of Bromus in eastern North America, with a proposal for the listing of 

B. purgans L. as a rejected name under Article 69. Taxon 25: 611-616. 

–––. 1979. Diplachne and Leptochloa (Poaceae) in North America. Brittonia 31: 399-404. 

McRoy, C.P., and C. Helfferich, eds. 1977. Seagrass ecosystems: a scientific perspective. Marcel Dekker, Inc., New York, NY. 

314 pp. 

McVaugh, R. 1936. Studies in the taxonomy and distribution of the eastern North American species of Lobelia. Rhodora 38: 

241-263, 276-298, 305-329. 

–––. 1944. The genus Cnidoscolus: generic limits and intergeneric groups. Bull. Torrey Bot. Club 71: 457-474. 

–––. 1945. The genus Triodanis Rafinesque, and its relationships to Specularia and Campanula. Wrightia 1: 13-52. 

–––. 1948. Generic status of Triodanis and Specularia. Rhodora 50: 38-49. 

–––. 1951. A revision of the North American black cherries (Prunus serotina Ehrh., and relatives). Brittonia 7: 279-315. 

–––. 1972. Compositarum Mexicanarum pugillus. Contr. Univ. Michigan Herb. 9: 359-484. 

–––, M.R. McVaugh, and M. Ayers. 1996. Chapel Hill and Elisha Mitchell the botanist. Occasional Publication No. 1 of the 

Chapel Hill Historical Society and Contribution No. 1 of the N.C. Botanical Garden. Botanical Garden Foundation, Chapel 

Hill, NC. 122 pp. 

Meacham, C.A. 1980. Phylogeny of the Berberidaceae with an evaluation of classifications. Systematic Bot. 5: 149-172. 

Mears, J.A. 1975. The taxonomy of Parthenium Section Partheniastrum DC. (Asteraceae--Ambrosiinae). Phytologia 31: 463- 

482.


Medley, M.E. 1989. Silphium wasiotensis (Asteraceae), a new species from the Appalachian plateaus in eastern Kentucky. Sida 

13: 285-291. 

–––. 1993. An annotated catalog of the known or reported vascular flora of Kentucky. Ph.D. dissertation, Dept. of Biology, 

University of Louisville, KY. 

–––, H. Bryan, J. MacGregor, and J.W. Thieret. 1985. Achyranthes japonica (Miq.) Nakai (Amaranthaceae) in Kentucky and 

West Virginia: new to North America. Sida 11: 92-95. 

Mellichamp, T.L., J.F. Matthews, and P.J. Smithka. 1987. New state and regional records of vascular plants in the Carolinas. 

Castanea 52: 95-111. 

–––, J.F. Matthews, and P.J. Smithka. 1988. It's Anthriscus sylvestris, not Conioselinum chinensis, new to North Carolina- 

Tennessee. Castanea 53: 81-82. 

Mellinger, A.C. 1972. Ecological life cycle of Viguiera porteri and factors responsible for its endemism to granite outcrops of 

Georgia and Alabama. Ph.D. thesis, University of North Carolina at Chapel Hill, Botany department. 

Menapace, F.J., P.G. Davison, and D.H. Webb. 1998. Noteworthy collections: Mississippi. Castanea 63: 80-81. 

Mendenhall, M.G. 1994a. New combinations in Thermopsis and Baptisia. Phytologia 76: 383-384. 

Mendenhall, M.G. 1994b. Phylogeny of Baptisia and Thermopsis (Leguminosae) as inferred from chloroplast DNA and nuclear 

ribosomal DNA sequences, secondary chemistry, and morphology. Ph.D. dissertation, Univ. of Texas at Austin. 

Meng, S.-W., A.W. Douglas, D.-Z. Li, Z.-D. Chen, H.-X. Liang, and J.-B. Yang. 2003. Phylogeny of Saururaceae based on 

morphology and five regions from three plant genomes. Ann. Missouri Bot. Gard. 90: 592-602. 

Mennema, J. 1989. A taxonomic revision of Lamium (Lamiaceae). Leiden Botanical Series 11: 1-196. 

Meyer, F.G. 1976. A revision of the genus Koelreuteria (Sapindaceae). J. Arnold Arb. 57: 129-166. 

–––, and J.W. Hardin. 1987. Status of the name Aesculus flava Solander (Hippocastanaceae). J. Arnold Arb. 68: 335-341. 

Meyer, F.K. 1973. Conspectus der "Thlaspi" Arten Europas, Afrikas und Vorderasiens. Feddes Repertorium 84: 449-470. 

–––. 1979. Kritische Revision der "Thlaspi" Arten Europas, Afrikas und Vorderasiens. Feddes Repertorium 90: 129-154. 

Mickel, J.T. 1979. How to know the ferns and fern allies. Wm. C. Brown Company, Dubuque, Iowa. 

Miller, A.J., D.A. Young, and J. Wen. 2001. Phylogeny and biogeography of Rhus (Anacardiaceae) based on ITS sequence data. 

Int. J. Plant Sci. 162: 1401-1407. 

Miller, G.N. 1955. The genus Fraxinus, the ashes, in North America, north of Mexico. Memoir 335, Cornell Univ. Agri. 

Experiment Station, Ithaca, NY. 

Miller, K.I., and G.L. Webster. 1967. A preliminary revision of Tragia (Euphorbiaceae) in the United States. Rhodora 69: 241- 

305. 

Miller, J.N., and K.L. Chambers. 2006. Systematics of Claytonia (Portulacaceae). Systematic Botany Monographs 78: 1-236. 

Miller, N.G. 1971a. The genera of the Urticaceae in the southeastern United States. J. Arnold Arb. 52: 40-68. 

–––. 1971b. The Polygalaceae in the southeastern United States. J. Arnold Arb. 52: 267-284. 

–––. 1991. The genera of Meliaceae in the southeastern United States. J. Arnold Arb. 71: 453-486. 

–––, and C.E. Wood, Jr. 2003. The Asian weed Fatoua villosa (Moraceae) in New York state and Massachusetts. Rhodora 105: 

286-291. 

Milne, R.I. 2004. Phylogeny and biogeography of Rhododendron subsection Pontica, a group with a Tertiary relict distribution. 

Molecular Phylogenetics and Evolution 33: 389-401. 

Mitchell, R.J. 1990. Trillium. Part 4 – the pedicellate species of eastern North America. The Plantsman 12: 44-60. 

Mitchell, R.S. 1970. A re-evaluation of Polygonum meisnerianum in North America. Rhodora 72: 182-188. 

–––, and J.K. Dean. n.d. Polygonaceae (Buckwheat Family) of New York state. Bulletin Numb. 431. N.Y. State Museum, 

Albany, NY. 

Moldenke, H.N. 1980. A sixth summary of the Verbenaceae, Avicenniaceae, Stilbaceae, Chloanthaceae, Symphoremaceae, 

Nyctanthaceae, and Eriocaulaceae of the world as to valid taxa, geographic distribution and synonymy. Phytologia Memoirs 

II. Privately published, Plainfield, NJ. 

Montgomery, J.D. 1982. Dryopteris in North America. Part II: the hybrids. Fiddlehead Forum 9: 23-30. 

–––, and E.M. Paulton. 1981. Dryopteris in North America. Fiddlehead Forum 8: 25-31. 

Moore, M.O. 1991. Classification and systematics of eastern North American Vitis L. (Vitaceae) north of Mexico. Sida 14: 

339-367. 

Moran, R.C. 1982. The Asplenium trichomanes complex in the United States and adjacent Canada. Am. Fern J. 72: 5-11. 

–––. 1983. Cystopteris tenuis (Michx.) Desv.: a poorly understood species. Castanea 48: 218-223. 

–––. 1998. Ferns, flashlights, and Tertiary forests. Fiddlehead Forum 25: 1-7. 

–––, and A.R. Smith. 1999. Salvinia adnata Desv. versus S. molesta D.S. Mitch. Am. Fern J. 89: 268-269. 

Morden, C.W. and S.L. Hatch. 1989. An analysis of morphological variation in Muhlenbergia capillaris (Poaceae) and its allies 

in the southeastern United States. Sida 13: 303-314. 

Morgan, D.R., and D.E. Soltis. 1993. Phylogenetic relationships among members of Saxifragaceae sensu lato based on rbcL 

sequence data. Ann. Mo. Bot. Gard. {}: 631-660. 

Morgan, J.T. 1966. A taxonomic study of the genus Boltonia (Asteraceae). Ph.D. dissertation, University of North Carolina at 

Chapel Hill. 

Morse, L.E. 1979. Systematics and ecological biogeography of the genus Hudsonia (Cistaceae), the sand heathers. Ph.D. 

dissertation, Harvard University.


Mort, M.E., and D.E. Soltis. 1999. Phylogenetic relationships and the evolution of ovary position in Saxifraga section 

Micranthes. Systematic Botany 24: 139-147. 

Morton, C.M., B. Isaac, J. Kartesz, and R. Coxe. 2004. Additions to and noteworthy records for the vascular flora of West 

Virginia. Sida 21: 481-485. 

Morton, C.V. 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U.S. National Herbarium 38: 153- 

214. 

Morton, G.H. 1973. [dissertation] 

Morton, G.H. 1974. A new subspecies and other nomenclatural changes in the Solidago arguta complex. Phytologia 28: 1-3. 

Mosquin, T. 1966. A new taxonomy for Epilobium angustifolium L. (Onagraceae). Brittonia 18: 167-188. 

Mosyakin, S.L., and S.E. Clemants. 1996. New infrageneric taxa and combinations in Chenopodium L. (Chenopodiaceae). 

Novon 6: 398-403. 

Muasya, A.M., J.J. Bruhl, D.A. Simpson, A. Culham, and M.W. Chase. 2000. Suprageneric phylogeny of Cyperaceae: a 

combined analysis. In: K.L. Wilson & D. A. Morrison, eds., Monocots: systematics and evolution. CSIRO, Melbourne. 

Mullahey, J.J. Tropical soda apple (Solanum viarum Dunal), a biological pollutant threatening Florida. Castanea 61: 255-260. 

–––, M. Nee, R.P. Wunderlin, and K.R. Delaney. 1993. Tropical soda apple (Solanum viarum): a new weed threat in 

subtropical regions. Weed Technology 7: 783-786. 

Mulligan, G.A. 1980. The genus Cicuta in North America. Can. J. Bot. 58: 1755-1767. 

Müller, K., and T. Borsch. 2005. Phylogenetics of Utricularia (Lentibulariaceae) and molecular evolutuion of the trnK intron in 

a lineage with high substitutional rates. Plant Syst. Evol. 250: 39-67. 

Müller-Doblies, D., and U. Müller-Doblies. 1996. Tribes and subtribes and some species combinations in Amaryllidaceae J. St.- 

Hil. emend. R. Dahlgren & al. 1985. Feddes Repertorium 107: 5-6, S.c. 1-S.c. 9. 

Mummenhoff, K., and M. Koch. 1994. Chloroplast DNA restriction site variation and phylogenetic relationships in the genus 

Thlaspi sensu lato (Brassicaceae). Systematic Bot. 19: 73-88. 

Munz, P.A. 1937. Studies in Onagraceae. X. The subgenus Kneiffia (genus Oenothera) and miscellaneous new species of 

Oenothera. Bull. Torrey Bot. Club 64: 287-306. 

–––. 1942. Studies in Onagraceae. XII. A revision of the New World species of Jussiaea. Darwiniana 4: 179-284. 

–––. 1944. Studies in Onagraceae. XIII. The American species of Ludwigia. Bull. Torrey Bot. Club 71: 152-165. 

–––. 1946. Aquilegia: the cultivated and wild columbines. Gentes Herb. 7: 1-150. 

–––. 1965. Onagraceae. N. Amer. Fl. II. 5: 1-278. 

Murakami, N., S. Nogami, M. Watanabe, and K. Iwatsuki. 1999. Phylogeny of Aspleniaceae inferred from rbcL nucleotide 

sequences. Am. Fern J. 89: 232-243. 

Murdy, W.H. 1966. The systematics of Phacelia maculata and P. dubia var. georgiana, both endemic to granite outcrop 

communities. Am. J. Bot. 53: 1028-1036. 

–––. 1968. Plant speciation associated with granite outcrop communities of the southeastern Piedmont. Rhodora 70: 394-407. 

Murray, A.E., Jr. 1970. A monograph of the Aceraceae. Ph.D. thesis, Pennsylvania State University. 

Murray, E. 1982. Notae Spermatophytae (Spermatophyta notes). Kalmia 12: 18-28. 

Murrell, Z.E. 1993. Phylogenetic relationships in Cornus (Cornaceae). Systematic Botany 18: 469-495. 

–––, P.E. Carroll, S.A. Myers, and P.J. Lawless. 1998. Examination of species boundaries in Hexastylis contracta Blomquist 

and H. rhombiformis Gaddy [abstract]. Amer. J. Bot. 85 [supplement]: 146-147. 

Musselman, L.J. 1982. The Orobanchaceae of Virginia. Castanea 47: 266-275. 

–––. 1984. An unusual specimen of Orobanche from North Carolina collected by John Ball in 1884. Castanea 49: 91-93. 

–––. 2001. Georgia quillworts. Tipularia 16: 2-19, 40. 

–––, R.D. Bray, and D.A. Knepper. 1996. Isoetes ×bruntonii (Isoetes engelmannii × I. hyemalis), a new hybrid quillwort from 

Virginia. Amer. Fern J. 86: 8-15. 

–––, R.D. Bray, and D.A. Knepper. 1997. Isoetes ×carltaylorii (Isoetes acadiensis × I. engelmannii), a new interspecific 

quillwort hybrid from the Chesapeake Bay. Can. J. Bot. 75: 301-309. 

–––, W.C. Taylor, and R.D. Bray. 2001. Isoetes massaponica (Isoetaceae), a new diploid quillwort from freshwater tridal 

marshes of Virginia. Novon 11: 200-204. 

–––, and D.A. Knepper. 1994. Quillworts of Virginia. Amer. Fern J. 84: 48-68. 

–––, D.A. Knepper, R.D. Bray, C.A. Caplen, and C. Ballou. 1995. A new Isoetes hybrid from Virginia. Castanea 60: 245-254. 

Myint, T. 1966. Revision of the genus Stylisma (Convolvulaceae). Brittonia 18: 97-117. 

Mymudes, M.S., and D.H. Les. Morphological and genetic variability in Plantago cordata (Plantaginaceae), a threatened aquatic 

plant. Am. J. Bot. 80: 351-359. 

Naczi, R.F.C. 1989. Carex asynchrona, a new species of section Griseae (Cyperaceae) from Tamaulipas, Mexico. Sida 13: 

487-492. 

–––. 1990. The taxonomy of Carex bromoides (Cyperaceae). Contr. Univ. Mich. Herb. 17: 215-222. 

–––. 1993. Carex brysonii and Carex godfreyi, new species of Carex section Griseae (Cyperaceae) from the southeastern United 

States. Contr. Univ. Michigan Herb. 19: 195-205. 

–––. 1997. Carex pigra, a new species of Carex section Griseae (Cyperaceae) from the southeastern United States of America. 

Novon 7: 67-71. 

–––. 1999a. Carex planispicata, a widespread and frequent new species of Carex section Griseae (Cyperaceae) from the eastern 

United States of America. J. Ky. Acad. Sci. 60: 37-44.


–––. 1999b. Chromosome numbers of some eastern North American species of Carex and Eleocharis (Cyperaceae). Contr. 

Univ. Michigan Herb. 22: 105-119. 

–––, and B.A. Ford. 1998. Systematics of the Carex jamesii complex (section Phyllostachys, Cyperaceae) [abstract]. Am. J. 

Bot. 85 [supplement]: 147. 

–––, and J.W. Thieret. 1996. Invasion and spread of Coincya monensis (Brassicaceae) in North America. Sida 17: 43-53. 

–––, C.T. Bryson, and T.S. Cochrane. 2002. Seven new species and one new combination in Carex (Cyperaceae) in North 

America. Novon 12: 508-532. 

–––, A.A. Reznicek, and B.A. Ford. 1998. Morphological, geographical, and ecological differentiation in the Carex willdenowii 

complex (Cyperaceae). Am. J. Bot. 85: 434-447. 

–––, E.M. Soper, F.W. Case, Jr., and R.B. Case. 1999. Sarracenia rosea (Sarraceniaceae), a new species of pitcher plant from 

the southeastern United States. Sida 18: 1183-1206. 

Nanda, J.S., and S.D. Sharma, eds. 2003. Monograph on genus Oryza. Science Publishers, Inc., Enfield, NH. 400 pp. 

Nash, G.V. 1900. Some new grasses from the southern states. N.Y. Bot. Garden Bull. 1: 429-436. 

NatureServe. 2003. A working classification of terrestrial ecological systems in the coterminous United States. International 

terrestrial ecological systems classification. NatureServe, Arlington, VA. 61 pp. plus appendices. 

NatureServe. 2005a. International ecological classification standard: terrestrial ecological classifications. NatureServe Central 

Databases. Arlington, VA. Data current as of March 2005. 

NatureServe. 2005b. NatureServe explorer, an online encyclopedia of life. http://www.natureserve.org/explorer/. Accessed 28 

November 2005. 

Navaro, A.M., and W.H. Blackwell. 1990. A revision of Paxistima (Celastraceae). Sida 14: 231-249. 

Neinhuis, C., S. Wanke, K.W. Hilu, K. Müller, and T. Borsch. 2005. Phylogeny of Aristolochiaceae based on parsimony, 

likelihood, and Bayesian analyses of trnL-trnF sequences. Plant Syst. Evol. 250: 7-26. 

Nelson, A.D., W.J. Elisens, and D. Benish. 1998. Notes on chromosome numbers in Chelone (Scrophulariaceae). Castanea 63: 

183-187. 

Nelson, G. 1994. The trees of Florida: a reference and field guide. Pineapple Press, Sarasota, FL. 338 pp. 

–––. 1996. The shrubs and woody vines of Florida: a reference and field guide. Pineapple Press, Sarasota, FL. 391 pp. 

–––. 2000. The ferns of Florida: a reference and field guide. Pineapple Press, Sarasota, FL. 208 pp. 

Nelson, J.B. 1980. Mitreola vs. Cynoctonum, and a new combination. Phytologia 46: 338-340. 

–––. 1981. Stachys (Labiatae) in Southeastern United States. Sida 9: 104-123. 

–––. 1993. Noteworthy collections: South Carolina. Castanea 58: 59-63. 

–––, ed. 2003. South Carolina Plant Atlas. http://cricket.biol.sc.edu/herb/ 

–––, and J.E. Fairey III. 1979. Misapplication of the name Stachys nuttallii to a new southeastern species. Brittonia 31: 491- 

494. 

–––, and K.B. Kelly. 1997. Noteworthy collections: South Carolina. Castanea 62: 283-288. 

–––, and D.A. Rayner. 1988. Isanthus brachiatus and Helianthus schweinitzii in York County, South Carolina. Castanea 53: 

82-83. 

Nesom, G.L. 1980. Erigeron tenuis T. & G. (Asteraceae) distantly disjunct in North Carolina. Castanea 45: 70-71. 

–––. 1983. Galax (Diapensiaceae): geographic variation in chromosome number. Systematic Bot. 8: 1-14. 

–––. 1989. New species, new sections, and a taxonomic overview of American Pluchea (Compositae: Inuleae). Phytologia 67: 

158-167. 

–––. 1990. Taxonomic status of Gamochaeta (Asteraceae: Inuleae) and the species of the United States. Phytologia 68: 186- 

198. 

–––. 1993a. Taxonomy of Sericocarpus (Asteraceae: Astereae). Phytologia 75: 45-54. 

–––. 1993b. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae: Astereae) and observations on their 

phylogenetic position. Phytologia 75: 1-44. 

–––. 1993c. Sageretia mexicana (Rhamnaceae), a new species from southwestern Mexico. Phytologia 75: 369-376. 

–––. 1993d. Taxonomy of Doellingeria (Asteraceae: Astereae). Phytologia 75: 452-462. 

–––. 1994a. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 193-274. 

–––. 1994b. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. 

Phytologia 77: 141-297. 

–––. 1995. Revision of Chaptalia (Asteraceae: Mutisieae) from North America and continental Central America. Phytologia 

78: 153-188. 

–––. 1997. Taxonomic adjustments in North American Aster sensu latissimo (Asteraceae: Astereae). Phytologia 82: 281-288. 

–––. 1999. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in four southeastern states and new for North America. Sida 

18: 1259-1264. 

–––. 2000a. Noteworthy collections from Herbarium NCU. Castanea 65: 80-83. 

–––. 2000b. Generic conspectus of the tribe Astereae (Asteraceae) in North America and Central America, the Antilles, and 

Hawaii. Sida Botanical Miscellany 20: 1-100. 

–––. 2000c. Callery pear (Pyrus calleryana – Rosaceae) naturalized in North Carolina. Rhodora 102: 361-364. 

–––. 2000d. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in Georgia. Sida 19: 413. 

–––. 2000e. Noteworthy collections: North Carolina. Castanea 65: 170. 

–––. 2001a. Notes on variation in Pseudognaphalium obtusifolium (Asteraceae: Gnaphalieae). Sida 19: 615-619.


–––. 2001b. Taxonomic review of Chrysogonum (Asteraceae: Heliantheae). Sida 19: 811-820. 

–––. 2002. New combination in Salix (Salicaceae). Sida 20: 523-524. 

–––. 2004a. Notes on typification in Pluchea (Asteraceae: Plucheae). Sida 21: 59-64. 

–––. 2004b. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United States and comments on similar 

species. Sida 21: 717-741. 

–––. 2004c. New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States. Sida 21: 1175-1185. 

–––. 2004d. Asteraceae from wool mill sites in South Carolina, including new records for North America. Sida 21: 1215-1223. 

–––. 2005a. Taxonomic review of Astranthium integrifolium (Asteraceae: Astereae). Sida 21: 2015-2021. 

–––. 2005b. Taxonomy of the Symphyotrichum (Aster) subulatum group and Symphyotrichum (Aster) tenuifolium (Asteraceae: 

Astereae). Sida 21: 2125-2140. 

–––, and V.M. Bates. 1984. Reevaluations of infraspecific taxonomy in Polygonella (Polygonaceae). Brittonia 36: 37-44. 

–––, and J.T. Kartesz. 2000. Observations on the Ludwigia uruguayensis complex (Onagraceae) in North America. Castanea 

65: 123-125. 

–––, and P.J.Leary. 1992. A new species of Ionactis (Asteraceae: Astereae) from southern Nevada and an overview of the 

genus. Brittonia 44: 247-252. 

–––, and B.L. Turner. 1998. Variation in the Berlandiera pumila (Asteraceae) complex. Sida 18: 493-502. 

Neufeld, H.S. 1986. Ecophysiological implications of tree architecture for two cypress taxa, Taxodium distichum (L.) Rich. and 

T. ascendens Brongn. Bull. Torrey Bot. Club 113: 118-124. 

Newell, R.E., and R.B. Newell. 1994. Juncus caesariensis Coville (New Jersey Rush) in Nova Scotia, Canada. Bartonia 58: 

121-124. 

Nevling, L.I., Jr. 1962. The Thymelaeaceae in the southeastern United States. J. Arnold Arb. 43: 428-434. 

Neyland, R. 2001. A phylogeny inferred from large ribosomal subunit (26S) rDNA sequences suggests that Cuscuta is a derived 

member of Convolvulaceae. Brittonia 53: 108-115. 

–––, and M.K. Hennigan. 2004. A cladistic analysis of Monotropa uniflora (Ericaceae) inferred from large ribosomal subunit 

(26S) rRNA gene sequences. Castanea 69: 265-271. 

Nicely, K.A. 1965. A monographic study of the Calycanthaceae. Castanea 30: 38-81. 

Nickrent, D.L., and V. Malécot. 2001. A molecular phylogeny of the Santalales. Presented at the 7th International Parasitic 

Weed Symposium, Nantes, France, June 5-8, 2001. http://www.parasiticplants.siu.edu/Santalales.IPWC/Sants.IPWC.html. 

Accessed 10 December 2005. 

Nixon, K.C., and J.M. Poole. 2003. Revision of the Mexican and Guatemalan species of Platanus (Platanaceae). Lundellia 6: 

103-137. 

Nordborg, G. 1966. Sanguisorba L., Sarcopoterium Spach, and Bencomia Webb et Berth. Opera Botanica 11: 2. C. Blum, 

Lund, Sweden. 

–––. 1967. The genus Sanguisorba section Poterium. Opera Botanica No. 16. C.W.K. Gleerup, Lund, Sweden. 

Noss, R.F., and A.Y. Cooperrider. 1994. Saving nature’s legacy: protecting and restoring biodiversity. Island Press, 

Washington, DC. 

Noyes, R.D. 2000. Biogeographical and evolutionary insights on Erigeron and allies (Asteraceae) from ITS sequence data. Pl. 

Syst. Evol. 220: 93-114. 

–––, and L.H. Rieseberg. 1999. ITS sequence data support a single origin for North American Astereae (Asteraceae) and reflect 

deep geographic divisions in Aster s.l. Amer. J. Bot. 86: 398-412. 

O'Kane, S.L., Jr., and I.A. Al-Shehbaz. 1997. A synopsis of Arabidopsis. Novon 7: 323-327. 

–––, and I.A. Al-Shehbaz. 2002. Paysonia, a new genus segregated from Lesquerella (Brassicaceae). Novon 12: 379-381. 

–––, and I.A. Al-Shehbaz. 2003. Phylogenetic position and generic limits of Arabidopsis (Brassicaceae) based on sequences of 

nuclear ribosomal DNA. Ann. Missouri Bot. Gard. 90: 603-612. 

Ogden, E.C. 1974. Potamogeton in New York. N.Y. State Museum Bull. 423. 

Ogle, D.W. 1991a. Spiraea virginiana Britton: I. Delineation and distribution. Castanea 56: 287-296. 

–––. 1991b. Spiraea virginiana Britton: II. Ecology and species biology. Castanea 56: 297-303. 

–––, and P. W. Mazzeo. 1976. Betula uber, the Virginia round-leaf birch, rediscovered in Southwest Virginia. Castanea 41: 

248-256. 

Oh, S.-H., and D. Potter. 2005. Molecular phylogenetic systematics and biogeography of tribe Neillieae (Rosaceae) using DNA 

sequences of cpDNA, rDNA, and LEAFY. Amer. J. Botany 92: 179-192. 

Ohi-Toma, T., T. Sugawara, H. Murata, S. Wanke, C. Neinhuis, and J. Murata. 2006. Molecular phylogeny of Aristolochia 

sensu lato (Aristolochiaceae) based on sequences of rbcL, matK, and phyA genes, with special reference to differentiation of 

chromosome numbers. Systematic Bot. 31: 481-492 

Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153-178. 

Olmstead, R.G., and P.A. Reeves. 1995. Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and 

ndhF sequences. Ann. Missouri Bot. Gard. 82: 176-193. 

–––, B. Bremer, K.M. Scott, and J.D. Palmer. 1993. A parsimony analysis of the Asteridae sensu lato based on rbcL sequences. 


–––, and J.D. Palmer. 1997. Implications for the phylogeny, classification, and biogeography of Solanum from cpDNA 

restriction site variation. Systematic Bot. 22: 19-30.


–––, C.W. DePamphilis, A.D. Wolfe, N.D. Young, W.J. Elisons, and P.A. Reeves. 2001. Disintegration of the Scrophulariaceae. 

Amer. J. Bot. 88: 348-361. 

Olsen, J. 1979. Taxonomy of the Verbesina virginica complex (Asteraceae). Sida 8: 128-134. 

Ortiz-Diaz, J.-J., and A. Culham. 2000. Phylogenetic relationships of the genus Sporobolus (Poaceae: Eragrostideae) based on 

nuclear ribosomal DNA ITS sequences. Pp. 184-188 in S.W.L. Jacobs and J. Everett (eds.) Grasses: systematics and 

evolution. CSIRO, Melbourne. 

Orzell, S.L., and E.L. Bridges. 2002. Notes on Carphephorus odoratissimus (Asteraceae) in peninsular Florida, U.S.A. Sida 20: 

559-569. 

Ownbey, G.B. 1947. Monograph of the North American species of Corydalis. Annals. Mo. Bot. Garden 34: 187-251. 

Ownbey, R.P. 1944. The liliaceous genus Polygonatum in North America. Ann. Mo. Bot. Garden 31: 373-413. 

Oxelman, B., M. Backlund, and B. Bremer. 1999. Relationships of the Buddlejaceae s.l. investigated using parsimony jackknife 

and branch support analysis of chloroplast ndhF and rbcL sequence data. Systematic Botany 24: 164-182. 

Packer, J.G. 1993. Two new combinations in Triantha (Liliaceae). Novon 3: 278-279. 

Padgett, D.J. 1999. Nomenclatural novelties in Nuphar (Nymphaeaceae). Sida 18: 823-826. 

Paler, M.H., and D.S. Barrington. 1995. The hybrid Cystopteris fragilis × C. tenuis (Dryopteridaceae) and the relationship 

between its tetraploid progenitors. Systematic Bot. 20: 528-545. 

Palmer, M.W., G.L. Wade, and P. Neal. 1995. Standards for the writing of floras. BioScience 45: 339-345. 

Palmer, P.G. 1975. A biosystematic study of the Panicum amarum--P. amarulum complex (Gramineae). Brittonia 27: 142-150. 

Palomino, G., P. Martínez, C. Bernal, and M. Sousa S. 1993. Diferencias cromosómicas entre algunas especies de los géneros 

Sophora L. y Styphnolobium Schott. Ann. Missouri Bot. Gard. 80: 284-290. 

Park, Chong-Wook. 1988. Taxonomy of Polygonum section Echinocaulon (Polygonaceae). Mem. N.Y. Bot. Garden 47: 1-82. 

–––, and Hyun-Woo Lee. 1996. Taxonomic notes on Cimicifuga purpurea, stat. nov. (Ranunculaceae). Novon 6: 93-95. 

Park, K. 1998. Monograph of Euphorbia sect. Tithymalopsis (Euphorbiaceae). Edinb. J. Bot. 55: 161-208. 

Park, M.M. 1992. A biosystematic study of Thalictrum section Leucocoma (Ranunculaceae). Ph.D. dissertation, Pennsylvania 

State University. 

Parker, E.S., and S.B. Jones. 1975. A systematic study of the genus Balduina (Compositae, Heliantheae). Brittonia 27: 355-361. 

Parker, M.A. 1996. Cryptic species within Amphicarpaea bracteata (Leguminosae): evidence from isozymes, morphology, and 

pathogen specificity. Can J. Bot. 74: 1640-1650. 

Parkinson, P.G. 1988. Adansonian nomina rejicienda et nomina conservanda proposita, 1983-1986. Taxon 37: 148-151. 

Parks, C.R., and J.W. Hardin. 1963. Yellow Erythroniums of the eastern United States. Brittonia 15: 245-259. 

–––, and J.F. Wendel. 1990. Molecular divergence between Asian and North American species of Liriodendron (Magnoliaceae) 

with implications for interpretation of fossil floras. Amer. J. Bot. 77: 1243-1256. 

–––, J.F. Wendel, M.M. Sewell, and Y.-L. Qiu. 1994. The significance of allozyme variation and introgression in the 

Liriodendron tulipifera complex (Magnoliaceae). Amer. J. Bot. 81: 878-889. 

Parrish, J.D., D.P. Braun, and R.S. Unnasch. 2003. Are we conserving what we say we are? Measuring ecological integrity 

within protected areas. BioScience 53: 851-860. 

Patrick, T.S. 1986. The trilliums of eastern North America. Published privately by the author, Social Circle, GA. 

–––, J.R. Allison, G.A. Krakow. 1995. Protected plants of Georgia: an information manual on plants designated by the state of 

Georgia as endangered, threatened, rare, or unusual. Georgia Dept. of Natural Resources, Social Circle, GA. 246 pp. 

Paun, O., C. Lehnebach, J.T. Johansson, P. Lockhart, and E. Hörandl. 2005. Phylogenetic relationships and biogeography of 

Ranunculus and allied genera (Ranunculaceae) in the Mediterranean region and in the European alpine system. Taxon 54: 

911-930. 

Pavlick, L.E. 1995. Bromus L. of North America. Royal British Columbia Museum, Victoria, BC. 160 pp. 

Peck, J.H. 2003. Arkansas flora: additions, reinstatements, exclusions, and re-exclusions. Sida 20: 1737-1757. 

Peet, R.K. 1993. A taxonomic study of Aristida stricta and A. beyrichiana. Rhodora 95: 25-37. 

Peirson, J.A., P.D. Cantino, and H.E. Ballard, Jr. 2006. A taxonomic revision of Collinsonia (Lamiaceae) based on phonetic 

analyses of morphological variation. Syst. Bot. 31: 398-409. 

Pelotto, J.P., and M.A. Del Pero Martínez. 1998. Flavonoids in Strophostyles species and the related genus Dolichopsis 

(Phaseolinae, Fabaceae): distribution and phylogenetic significance. Sida 18: 213-222. 

Peng, Ching-I. 1984. Ludwigia ravenii (Onagraceae), a new species from the Coastal Plain of the southeastern United States. 


–––. 1986. A new combination in Ludwigia sect. Microcarpium (Onagraceae). Ann. Mo. Bot. Gard. 73: 490. 

–––. 1988. The biosystematics of Ludwigia sect. Microcarpium (Onagraceae). Ann. Mo. Bot. Gard. 75: 970-1009. 

–––. 1989. The systematics and evolution of Ludwigia sect. Microcarpium (Onagraceae). Ann. Mo. Bot. Gard. 76: 221-302. 

–––, and H. Tobe. 1987. Capsule wall anatomy in relation to capsular dehiscence in Ludwigia sect. Microcarpium (Onagraceae). 

Am. J. Bot. 74: 1102-1110. 

Pennell, F.W. 1916. Notes on plants of the southern United States – II. Bull. Torrey Bot. Club 43: 407-421. 

–––. 1935. The Scrophulariaceae of eastern temperate North America. Academy of Natural Sciences of Philadelphia 

Monograph No. 1. 

Pennington, T.D. 1991. The genera of Sapotaceae. Royal Botanic Gardens, Kew & N.Y. Botanical Gardens, Bronx, New York. 

Perdue, R.E., Jr. 1957. Synopsis of Rudbeckia subgenus Rudbeckia. Rhodora 59: 293-299.


Perry, J.E., D.M.E. Ware, and A. McKenney-Mueller. 1998. Aeschynomene indica L. (Fabaceae) in Virginia. Castanea 63: 191- 

194. 

Perry, J.P., III, and L.J. Musselman. 1994. Psilotum nudum new to North Carolina. Amer. Fern J. 84: 102-104. 

Perry, L.M. 1937. Notes on Silphium. Rhodora 39: 281-297. 

Petersen, G., and O. Seberg. 2003. Phylogenetic analyses of the diploid species of Hordeum (Poaceae) and a revised 

classification of the genus. Syst. Bot. 28: 293-306. 

Peterson, P.M., S.L. Hatch, and A.S. Weakley. [in press]. Sporobolus in M.E. Barkworth, K.M. Capels, and L.A. Vorobik 

(eds.), Manual of grasses for the Continental United States and Canada, Department of Agriculture Miscellaneous 

Publication. 

–––-, E.E. Terrell, E.C. Uebel, C.A. Davis, H. Scholz, and R.J. Soreng. 1999. Oplismenus hirtellus subspecies undulatifolius, a 

new record for North America. Castanea 64: 201-202. 

Pfeil, B.E., and M.D. Crisp. 2005. What to do with Hibiscus? A proposed nomenclatural resolution for a large and well known 

genus of Malvaceae and comments on paraphyly. Australian Systyematic Botany 18: 49-60 

Pfosser, M., W. Wetschnig, S. Ungar, and G. Prenner. 2003. Phylogenetic relationships among genera of Massonieae 

(Hyacinthaceae) inferred from plastid DNA and seed morphology. J. Plant Res. 116: 115-132. 

Philbrick, C.T., and G.E. Crow. 1983. Distribution of Podostemum ceratophyllum Michx. (Podostemaceae). Rhodora 85: 325- 

341. 

–––, R.A. Aakjar, Jr., and R.L. Stuckey. 1998. Invasion and spread of Callitriche stagnalis (Callitrichaceae) in North America. 

Rhodora 100: 25-38. 

Phipps, J.B. 1988. Crataegus (Maloideae, Rosaceae) of the southeastern United States, I. Introduction and series Aestivales. J. 

Arnold Arb. 69: 401-431. 

–––. 1998. Synopsis of Crataegus series Apiifoliae, Cordatae, Microcarpae, and Brevispinae (Rosaceae subfam. Maloideae). 


–––, R.J. O'Kennon, and R.W. Lance. 2003. Hawthorns and medlars. Royal Horticultural Society Plant Collector Guide. 

Timber Press, Portland OR. 139 pp. 

Pigniotti, L., and L.M. Mariotti. 2004. Micromorphology of Scirpus (Cyperaceae) and related genera in south-west Europe. 

Bot. J. Linn. Soc. 145: 45-58. 

Pilatowski, R.E. 1982. A taxonomic study of the Hydrangea arborescens complex. Castanea 47: 84-98. 

Pinson, J.N., Jr., and W.T. Batson. 1971. The status of Muhlenbergia filipes Curtis (Poaceae). J. Elisha Mitchell Sci. Soc. 87: 

188-191. 

Pippen, R.W. 1978. Cacalia. N. Amer. Fl. II 10: 151-159. 

Pittillo, J.D., and A.E. Brown. 1988. Additions to the vascular flora of the Carolinas, III. J. Elisha Mitchell Sci. Soc. 104: 1-18. 

–––, J.H. Horton, and K.E. Herman. 1972. Additions to the vascular flora of the Carolinas. II. J. Elisha Mitchell Sci. Soc. 88: 

144-152 

–––, W.H. Wagner, Jr., D.R. Farrar, and S.W. Leonard. 1975. New pteridophyte records in the Highlands Biological Station 

area, Southern Appalachians. Castanea 40: 263-272. 

Pittman, A.B. 1988. Systematic studies in Scutellaria sect. Mixtae (Labiatae). Ph.D. dissertation, Vanderbilt Univ. 

Plant Information Center. 2005. Virtual herbarium. http://www.ibiblio.org/pic/herbarium.htm. Accessed 27 November 2005. 

Platt, S.G., and J.F. Townsend. 1996. Noteworthy collections: South Carolina. Castanea 61: 397-398. 

Pohl, R.W. 1969. Muhlenbergia, subgenus Muhlenbergia (Gramineae) in North America. Amer. Midl. Naturalist 82: 512-542. 

Poiani, K.P., B.D. Richter, M.G. Anderson, and H.E. Richter. 2000. Biodiversity conservation at multiple scales: functional 

sites, landscapes, and networks. BioScience 50:133-146. 

Poindexter, D.B. 2006. Eight new plant distributional records to Alleghany County, North Carolina. J. North Carolina Academy 

Sci. 122: 101-105. 

Porter, D.M. 1969. The genus Kallstroemia (Zygophyllaceae). Contr. Gray Herb. 198: 41-153. 

–––. 1976. Zanthoxylum (Rutaceae) in North America north of Mexico. Brittonia 28: 443-447. 

–––, and T.F. Wieboldt. 1991. Vascular plants. In K. Terwilliger, coord., Virginia's endangered species: proceedings of a 

symposium. McDonald and Woodward Publ. Co., Blacksburg VA. 

Possingham, H.P., S.J. Andelman, M.A. Burgman, R.A. Medelin, L.L. Master, and D.A. Keith. 2002. Limits to the use of 

threatened species lists. Trends in Ecology and Evolution 17: 503-507. 

Powell, M., V. Savolainen, P. Cuénoud, J.-F. Manen, and S. Andrews. 2000. The mountain holly (Nemopanthus mucronatus: 

Aquifoliaceae) revisited with molecular data. Kew Bulletin 55: 341-347. 

Prance, G.T. 1970. The genera of Chrysobalanaceae in the southeastern United States. J. Arnold Arb. 51: 521-528. 

–––. 1972. Chrysobalanaceae. Flora Neotropica Monograph No. 9. Hafner Press, New York, NY. 410 pp. 

–––, and C.A. Sothers. 2003. Chrysobalanaceae 1: Chrysobalanus to Parinari. Species Plantarum: Flora of the World 9: 1-319. 

Prather, L.A., and J.A. Keith. 2003. Monarda humilis (Lamiaceae), a new combination for a species from New Mexico, and a 

key to the species of section Cheilyctis. Novon 13: 104-109. 

–––, C.J. Ferguson, and R.K. Jansen. 2000. Polemoniaceae phylogeny and classification: implications of sequence data from the 

chloroplast gene ndhF. Am. J. Bot. 87: 1300-1308. 

Preston, C.D., M.G. Telfer, H.R. Arnold, and P. Rothery. 2002. The changing flora of Britain, 1930-1999. In C.D. Preston, 

D.A. Pearman, and T.D. Dines (eds.). New Atlas of the British and Irish flora: an atlas of the vascular plants of Britain, 

Ireland, the Isle of Man and the Channel Islands. Oxford University Press, Oxford.


Price, R.A. 1989. The genera of Pinaceae in the southeastern United States. J. Arnold Arb. 70: 247-305. 

–––. 1990. The genera of Taxaceae in the southeastern United States. J. Arnold Arb. 71: 69-91. 

–––, A. Liston, and S.H. Strauss. 1998. Phylogeny and systematics of Pinus. Pp. 49-68 in D.M. Richardson, ed., Ecology and 

biogeography of Pinus. Cambridge Univ. Press. 527 pp. 

Pridgeon, A.M., R.M. Bateman, A.V. Cox, J.R. Hapeman, and M.W. Chase. 1997. Phylogenetics of subtribe Orchidinae 

(Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 1. Intergeneric relationships and polyphyly of Orchis sensu 

lato. Lindleyana 12: 89-109. 

–––, P.J. Cribb, M.W. Chase, and F.N. Rasmussen. 1999a. Genera orchidacearum. Volume 1: General introduction, 

Apostasioideae, Cypripedioideae. Oxford Univ. Press. 

–––, P.J. Cribb, M.W. Chase, and F.N. Rasmussen. 1999b. Genera orchidacearum. Volume 3: Orchidioideae (part 1). Oxford 

Univ. Press. 

–––, P.J. Cribb, M.W. Chase, and F.N. Rasmussen. 1999c. Genera orchidacearum. Volume 3: Orchidoideae (part 2), 

Vanilloideae. Oxford Univ. Press. 

–––, P.J. Cribb, M.W. Chase, and F.N. Rasmussen. 2005. Genera orchidacearum. Volume 4: Epidendroideae (part 1). Oxford 

Univ. Press. 

Prince, L.M. 2002. Circumscription and biogeographic patterns in the eastern North American – east Asian genus Stewartia 

(Theaceae: Stewartieae): insight from chloroplast and nuclear DNA sequence data. Castanea 67: 290-301. 

–––, and C.R. Parks. 2001. Phylogenetic relationships of Theaceae inferred from chloroplast DNA sequence data. Amer. J. Bot. 

88: 2309-2320. 

Pringle, J.S. 1967. Taxonomy of Gentiana, section Pneumonanthe, in eastern North America. Brittonia 19: 1-32. 

–––. 1971. Taxonomy and distribution of Clematis, sect. Atragene (Ranunculaceae), in North America. Brittonia 23: 361-393. 

–––. 1977. Gentiana linearis (Gentianaceae) in the Southern Appalachians. Castanea 42: 1-8. 

–––. 2002. Nomenclature of the heart-leaved hedge-nettle, Stachys cordata (Lamiaceae). Sida 20: 583-584. 

–––. 2004. Nomenclature of the Virginia bluebell, Mertensia virginica (Boraginaceae). Sida 21: 771–775. 

–––, and A.J. Sharp. 1964. Gentiana austromontana, a new species from the Southern Appalachians. Rhodora 66: 402-404. 

Pruski, J.F. 1998. Helianthus porteri (A. Gray) Pruski (Compositae), a new combination validated for the Confederate Daisy. 

Castanea 63: 74-75. 

–––. 2004. Panphalea heterophylla (Compositae: Mutisioideae: Nassauvieae), a genus and species new for the flora of North 

America. Sida 21: 1225-1228. 

–––, and G.L. Nesom. 2004. Gamochaeta coarctata, the correct name for Gamochaeta spicata (Asteraceae: Gnaphalieae). Sida 

21: 711-714. 

Pryer, K.M., and C.H. Haufler. 1993. Isozymic and chromosomal evidence for the allotetraploid origin of Gymnocarpium 

dryopteris (Dryopteridaceae). Systematic Bot. 18: 150-172. 

––––, and L.R. Phillippe. 1989. A synopsis of the genus Sanicula (Apiaceae) in eastern Canada. Can. J. Bot. 67: 694-707. 

–––, H. Schneider, A.R. Smith, R. Cranfill, P.G. Wolf, J.S. Hunt., and S.D. Sipes. 2001. Horsetails and ferns are a monophyletic 

group and the closest living relatives to seed plants. Nature 409: 618-622. 

–––, E. Schuettpelz, P.G. Wolf, H. Schneider, A.R. Smith, and R. Cranfill. 2004. Phylogeny and evolution of ferns 

(monilophytes) with a focus on the early leptosporangiate divergences. Amer. J. Botany 91: 1582-1598. 

Puff, C. 1976. The Galium trifidum group (Galium sect. Aparinoides, Rubiaceae). Can. J. Bot. 54: 1911-1925. 

–––. 1977. The Galium obtusum group (Galium sect. Aparinoides, Rubiaceae). Bull. Torrey Bot. Club 104: 202-208. 

Pyck, N., A. Van Lysebetten, J. Stessens, and E. Smets. 2002. The phylogeny of Patrineae sensu Graebner (Valerianaceae) 

revisited: additional evidence from ndhF sequence data. Plant Syst. Evol. 233: 29-46. 

Pyšek, P., D.M. Richardson, M. Rejmánek, G.L. Webster, M. Williamson, and J. Kirschner. 2004. Alien plants in checklists and 

floras: towards better communication between taxonomists and ecologists. Taxon 53: 131-143. 

Qualls, D.A. 1984. A revision of the New World species of Lindernia Allioni (Scrophulariaceae). M.A. Thesis, Dept. of 

Biology, Univ. of North Carolina at Chapel Hill, Chapel Hill, NC. 

Quinn, J.A., and D.E. Fairbrothers. 1971. Habitat ecology and chromosome numbers of natural populations of the Danthonia 

sericea complex. Amer. Midland Natur. 85: 531-536. 

Rabeler, R.K. 1985. Petrorhagia (Caryophyllaceae) of North America. Sida 11: 6-44. 

–––. 1991. Moenchia erecta (Caryophyllaceae) in eastern North America. Castanea 56: 150-151. 

–––, and J.W. Thieret. 1988. Comments on the Caryophyllaceae of the southeastern United States. Sida 13: 149-156. 

Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina 

Press, Chapel Hill, N.C. 1183 pp. 

Ramsey, G.W. 1987. Morphological considerations in the North American Cimicifuga (Ranunculaceae). Castanea 52: 129-141. 

–––. 1988. A comparison of vegetative characteristics of several genera with those of the genus Cimicifuga (Ranunculaceae). 

Sida 13: 57-63. 

Randall, J.L., and K.W. Hilu. 1986. Biosystematic studies of North American Trisetum spicatum (Poaceae). Systematic Bot. 

11: 567-578. 

Raven, P.H. 1963. The Old World species of Ludwigia (including Jussiaea), with a synopsis of the genus (Onagraceae). 

Reinwardtia 6: 327-427. 

–––, and D.P. Gregory. 1972. A revision of the genus Gaura (Onagraceae) Mem. Torrey Bot. Club 23: 1-96. 

–––, and W. Tai. 1979. Observations of chromosomes in Ludwigia (Onagraceae). Ann. Mo. Bot. Gard. 66: 862-879.


Rayner, D.A., and J. Henderson. 1980. Vaccinium sempervirens (Ericaceae), a new species from Atlantic White-cedar bogs in 

the sandhills of South Carolina. Rhodora 82: 503-507. 

Reddoch, A.H., and J.M. Reddoch. 1993. The species pair Platanthera orbiculata and P. macrophylla (Orchidaceae): 

taxonomy, morphology, distributions and habitats. Lindleyana 8: 171-188. 

Redman, D.E. 1995. Distribution and habitat types for Nepal Microstegium [Microstegium vimineum (Trin.) Camus] in 

maryland and the District of Columbia. Castanea 60: 270-275. 

Reed, C.F. 1953. The ferns and fern allies of Maryland and Delaware including District of Columbia. Reed Herbarium, 

Baltimore, MD. 

–––. 1961a. Andrographis, a genus of Acanthaceae, new to eastern United States. Castanea 26: 128. 

–––. 1961b. Amaranthaceae new to eastern United States. Castanea 26: 123-127. 

–––. 1964. A flora of the chrome and manganese ore piles at Canton, in the Port of Baltimore, Maryland and at Newport News, 

Virginia, with descriptions of genera and species new to the flora of eastern United States. Phytologia 10: 324-406. 

Reeder, J.R., and M.A. Ellington. 1960. Calamovilfa, a misplaced genus of Gramineae. Brittonia 12: 71-77. 

Rehder, A. 1903. Synopsis of the genus Lonicera. Missouri Botanical Garden Annual Report 1903: 27-232. 

–––. 1945. Carya alba proposed as nomen ambiguum. J. Arnold Arb. 26: 482-483. 

Renner, S.S., L.-B. Zhang, and J. Murata. 2004. A chloroplast phylogeny of Arisaema (Araceae) illustrates Tertiary floristic 

links between Asia, North America, and East Africa. Amer. J. Bot. 91: 881-888. 

Rettig, J.H. 1988. A biosystematic study of the Carex pensylvanica group (section Acrocystis) in North America. Ph.D. 

dissertation. Univ. of Georgia, Athens. 

–––. 1989. Nomenclatural changes in the Carex pensylvanica group (section Acrocystis, Cyperaceae) of North America. Sida 

13: 449-452. 

–––. 1990. Achene micromorphology of the Carex nigromarginata complex (section Acrocystis, Cyperaceae). Rhodora 92: 70- 

79. 

Reveal, J.L. 1989. A checklist of the Eriogonoideae (Polygonaceae). Phytologia 66: 266-294. 

–––. 1993a. A splitter's guide to the higher taxa of the flowering plants (Magnoliophyta) generally arranged to follow the 

sequence proposed by Thorne (1992) with certain modifications. Phytologia 74: 203-263. 

–––. 1993b. The correct name of the northern expression of Sarracenia purpurea L. (Sarraceniaceae). Phytologia 74: 180-184. 

–––. 1993c. Streptopus lanceolatus (Aiton) Reveal, a new name for Streptopus roseus Michx. (Convallariaceae). Phytologia 74: 

185-189. 

–––. 2004. Nomenclatural summary of Polygonaceae subfamily Eriogonoideae. Harvard Papers in Bot. 9: 143-230. 

–––, P.H. Raven, P. Hoch, R.R. Haynes, and C.B. Hellquist. 2003. (1603-1605) Proposals to conserve the name Ludwigia 

repens (Onagraceae) with a conserved type, and to reject the names Potamogeton oblongifolium and P. rotundifolium 

(Potamogetonaceae), all published in Forster's Flora Americae Septentrionalis. Taxon 52: 864-866. –––, and F.R. Barrie. 

1992. Matelea suberosa (L.) Shinners (Asclepiadaceae) – once again. Bartonia 57: 36-38. 

–––, and C.S. Keener. 1981. Virgulus Raf. (1837), an earlier name for Lasallea Greene (1903) (Asteraceae). Taxon 30: 648- 

651. 

–––, and M.J. Seldin. 1976. On the identity of Halesia carolina L. (Styracaceae). Taxon 25: 123-140. 

–––, and W.B. Zomlefer. 1998. Two new orders for monocotyledonous plants. Novon 8: 176-177. 

Reznicek, A.A. 1993. Carex pumila (Cyperaceae) in North America. Castanea 58: 220-224. 

–––, and P.W. Ball. 1980. The taxonomy of Carex section Stellulatae in North America north of Mexico. Contr. Univ. Mich. 

Herb. 14: 153-204. 

–––, and P.M. Catling. 1986. Carex striata, the correct name for C. walteriana (Cyperaceae). Rhodora 88: 405-406. 

Rhoads, A.F., and W.M. Klein, Jr. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American 

Philosophical Society, Philadelphia, PA. 

Richards, E.L. 1968. A monograph of the genus Ratibida. Rhodora 70: 348-393. 

Richardson, D.M., ed. 1998. Ecology and biogeography of Pinus. Cambridge Univ. Press. 527 pp. 

Richardson, J.E., M.F. Fay, Q.C.B. Cronk, D. Bowman, and M.W. Chase. 2000a. A phylogenetic analysis of Rhamnaceae using 

rbcL and trnL-F plastid DNA sequences. Am. J. Bot. 87: 1309-1324. 

–––, M.F. Fay, Q.C.B. Cronk, and M.W. Chase. 2000b. A revision of the tribal classification of Rhamnaceae. Kew Bulletin 55: 

311-340. 

Ridsdale, C.E. 1976. A revision of the tribe Cephalantheae (Rubiaceae). Blumea 23: 177-188. 

Riggins, R. 1977. A biosystematic study of the Sporobolus asper complex (Gramineae). Iowa State J. of Research 51: 287-321. 

Ringius, G.S. 1985. [Solidago] 

Risk, A.C., and D.L. Wyrick. 1996. Silphium wasiotense Medley in Tennessee. Castanea 61: 194-196. 

Roalson, E.H., and E.A. Friar. 2000. Infrageneric classification of Eleocharis (Cyperaceae) revisited: evidence from the internal 

transcribed spacer (ITS) region of nuclear ribosomal DNA. Syst. Bot. 25: 323-336. 

Robbins, H.C. 1968. The genus Pachysandra. Sida 3: 211-248. 

Roberts, P.R., and H.J. Oosting. 1958. Responses of Venus fly trap (Dionaea muscipula) to factors involved in its endemism. 

Ecol. Monographs 28: 193-218. 

Robertson, K.R. 1971. The Linaceae in the southeastern United States. J. Arnold Arb. 52: 649-665. 

–––. 1973. The Krameriaceae in the southeastern United States. J. Arnold Arb. 54: 322-327. 

–––. 1974. The genera of Rosaceae in the southeastern United States. J. Arnold Arb. 55: 303-332, 344-401, 611-662.


–––. 1975. The Oxalidaceae in the southeastern United States. J. Arnold Arb. 56: 223-239. 

–––. 1976. The genera of Haemodoraceae in the southeastern United States. J. Arnold Arb. 57: 205-216. 

–––. 1981. The genera of Amaranthaceae in the southeastern United States. J. Arnold Arb. 62: 267-314. 

–––, and Yin-Tse Lee. 1976. The genera of Caesalpinioideae (Leguminosae) in the southeastern United States. J. Arnold Arb. 

57: 1-53. 

–––, J.B. Phipps, J.R. Rohrer, and P.G. Smith. 1991. A synopsis of genera in Maloideae (Rosaceae). Systematic Botany 16: 

376-394. 

Robinson, H. 1978. Studies in the Heliantheae (Asteraceae). XII. Re-establishment of the genus Smallanthus. Phytologia 39: 

47-53. 

Robson, N.K.B. 1977. Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. Bull. Nat. Hist. Mus. 

Lond. (Bot.) 5: 291-355. 

–––. 1981. Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bull. Nat. Hist. Mus. Lond. (Bot.) 8: 

55-226. 

–––. 1987. Studies in the genus Hypericum L. (Guttiferae). 7. Section 29. Brathys (part 1). Bull. Nat. Hist. Mus. Lond. (Bot.) 

16: 1-106. 

–––. 1990. Studies in the genus Hypericum L. (Guttiferae). 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. Bull. Nat. 

Hist. Mus. Lond. (Bot.) 20: 1-151. 

–––. 1996. Studies in the genus Hypericum L. (Guttiferae). 6. Sections 20. Myriandra to 28. Elodes. Bull. Nat. Hist. Mus. 

Lond. (Bot.) 26: 75-217. 

–––. 2000. Studies in the genus Hypericum L. (Guttiferae). 4(1). Sections 7. Roscyna to 9. Hypericum sensu lato (part 1). Bull. 

Nat. Hist. Mus. Lond. (Bot.) 31: 37-88. 

–––. 2002. Studies in the genus Hypericum L.( Guttiferae). 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. 

Hypericum series 1. Hypericum. Bull. Nat. Hist. Mus. Lond. (Bot.) 32: 61-123. 

–––. 2006. Studies in the genus Hypericum L.( Clusiaceae). Section 9. Hypericum sensu lato (part 3): subsection 1. Hypericum 

series 2. Senanensia, subsection 2. Erecta and section 9b. Graveolentia. Systematics and Biodoiversity4: 19-98. 

–––, and P. Adams. 1968. Chromosome numbers in Hypericum and related genera. Brittonia 20: 95-106. 

Rock, H.F.L. 1957. A revision of the vernal species of Helenium (Compositae). Rhodora 59: 101-116, 128-159, 168-178, 203- 

216. 

Rodgers, C.L. 1950. The Umbelliferae of North Carolina and their distribution in the Southeast. J. of the Elisha Mitchell 

Scientific Socity 66: 195-266. 

Rodman, J.E. 1974. Systematics and evolution of the genus Cakile (Cruciferae). Contr. Gray Herb. 205: 3-146. 

Roecker, R., and T. Socha. 2004. Hawaiian plant threatens South Carolina dunes. Wildland Weeds 7: 19–20. 

Rogers, C.M. 1963. Yellow flowered species of Linum in eastern North America. Brittonia 15: 97-122. 

–––. 1984. Linaceae. North American Flora, Series II, Part 12, New York Botanical Garden, Bronx, NY. 

Rogers, G.K. 1982a. The Stemonaceae in the southeastern United States. J. Arnold Arb. 63: 327-336. 

–––. 1982b. The Bataceae in the southeastern United States. J. Arnold Arb. 63: 375-386. 

–––. 1983. The genera of Alismataceae in the southeastern United States. J. Arnold Arb. 64: 383-420. 

–––. 1985. The genera of Phytolaccaceae in the southeastern United States. J. Arnold Arb. 66: 1-37. 

–––. 1986. The genera of Loganiaceae in the southeastern United States. J. Arnold Arb. 67: 143-185. 

–––. 1987. The genera of Cinchonoideae (Rubiaceae) in the southeastern United States. J. Arnold Arb. 68: 137-183. 

–––. 2005. The genera of Rubiaceae in the southeastern United States, part II. Subfamily Rubioideae, and subfamily 

Cinchonoideae revisited (Chiococca, Erithalis, and Guettarda). Harvard Papers in Botany 10: 1-45. 

Rollins, R.C. 1941. A monographic study of Arabis in western North America. Rhodora 43: 313-325. 

–––. 1961. A weedy crucifer again reaches North America. Rhodora 63: 345-346. 

–––. 1993. The Cruciferae of continental North America: systematics of the mustard family from the Arctic to Panama. 

Stanford Univ. Press, Stanford, CA. 976 pp. 

–––, and E.A. Shaw. The genus Lesquerella (Cruciferae) in North America. Harvard Univ. Press, Cambridge. 288 pp. 

Romanowski, N. 2002. Gardening with carnivores: Sarracenia pitcher plants in cultivation and in the wild. University Press of 

Florida, Gainesville, FL. 106 pp. 

Ronse De Craene, L. P., and J.R. Akeroyd. 1988. Generic limits in Polygonum and related genera (Polygonaceae) on the basis 

of floral characters. Bot. J. Linn. Soc. 98: 321-371. 

–––, S.P. Hong, and E.F. Smets. 2004. What is the taxonomic status of Polygonella? Evidence of floral morphology. Ann. 


Rosatti, T.J. 1984. The Plantaginaceae in the southeastern United States. J. Arnold Arb. 65: 533-562. 

–––. 1986. The genera of Sphenocleaceae and Campanulaceae in the southeastern United States. J. Arnold Arb. 67: 1-64. 

–––. 1987a. The genera of Pontederiaceae in the southeastern United States. J. Arnold Arb. 68: 35-71. 

–––. 1987b. Field and garden studies of Arctostaphylos uva-ursi (Ericaceae) in North America. Systematic Bot. 12: 61-77. 

–––. 1989. The genera of the suborder Apocynineae (Apocynaceae and Asclepiadaceae) in the Southeastern United States. J. 

Arnold Arb. 70: 443-514. 

Rose, J.N. 1911. Two new species of Harperella. Contr. U.S. Nat. Herb. 13: 289-290. 

–––, and P.C. Standley. 1911. The genus Talinum in Mexico. Contr. U.S. Nat. Herb. 13: 281-288.


Rosendahl, C.O., F.K. Butters, and O. Lakela. 1936. A monograph on the genus Heuchera. Minnesota Studies in Plant Science 

2: 1-180. 

Rossetto, M., B.R. Jackes, K.D. Scott, and R.J. Henry. 2002. Is the genus Cissus (Vitaceae) monophyletic? Evidence from 

plastid and nuclear ribosomal DNA. Systematic Botany 27: 522-533. 

Rossignol, L., M. Rossignol, and R. Haicour. 1987. A systematic revision of Phyllanthus subsection Urinaria (Euphorbiaceae). 

Am. J. Bot. 74: 1853-1862. 

Rothrock, P.E., A.A. Reznicek, and L.R. Ganion. 1997. Taxonomy of the Carex straminea complex (Cyperaceae). Can. J. Bot. 

75: 2177-2195. 

Rudall, P.J., K.L. Stobart, W.-P. Hong, J.G. Conran, C.A. Furness, G.C. Kite, and M.W. Chase. 2000. Consider the lilies: 

systematics of Liliales. In: K.L. Wilson & D. A. Morrison, eds., Monocots: systematics and evolution. CSIRO, Melbourne. 

Rudd, V.E. 1955. The American species of Aeschynomene. Contr. U.S. National Herbarium 32: 1-172. 

–––. 1972. Leguminosae-Faboideae-Sophoreae. North American Flora Series II, Part 7. 

Russell, N.H. 1965. Violets (Viola) of central and eastern United States: an introductory survey. Sida 2: 1-113. 

Rydberg, P.A. 1915. Cardulaes. Carduaceae. Heleniae, Tageteae. North American Flora 34, Part 2: 81-180. 

Saarela, J.M., P.M. Peterson, R.J. Soreng, and R.E. Chapman. 2003. A taxonomic revision of the eastern North American and 

eastern Asian disjunct genus Brachyelytrum (Poaceae): evidence from morphology, phytogeography, and AFLPs. 


Saltonstall, K. 2002. Cryptic invasion by a non-native genotype of the common reed, Phragmites australis, into North America. 

Proceedings of the National Academy of Sciences, USA 99(4): 2445-2449. 

–––, P.M. Peterson, and R.J. Soreng. 2004. Recognition of Phragmites australis subsp. americanus (Poaceae: Arundinoideae) in 

North America: evidence from morphological and genetic anlyses. Sida 21: 683-692. 

Samuel, R., W. Gutermann, T.F. Stuessy, C.F. Ruas, H.-W. Lack, K. Tremetsberger, S. Talavera, B. Hermanowski, and F. 

Ehrendorfer. 2006. Molecular phylogenetics reveals Leontodon (Asteraceae, Lactuceae) to be diphyletic. Amer. J. Botany 

93: 1193-1205. 

Sanders, R.W. 1987. Identity of Lantana depressa and L. ovatifolia (Verbenaceae) of Florida and the Bahamas. Syst. Botany 

12: 44-60. 

–––. 2006. Taxonomy of Lantana sect. Lantana (Verbenaceae): I. Correct application of Lantana camara and associated names. 

Sida 22: 381-421. 

–––, and W.S. Judd. 2000. Incorporating phylogenetic results into floristic treatments. Sida 18: 97-112. 

Sargent, C.S. 1918. Notes on North American trees. II. Carya. Bot. Gazette 66: 229-258. 

–––. 1921. Notes on North American trees. VIII. J. Arnold Arb. 2: 164-174. 

Sauer, J.D. 1955. Revision of the dioecious amaranths. Madroño 13: 5-46. 

–––. 1972. Revision of Stenotaphrum (Gramineae: Paniceae) with attention to its historical geography. Brittonia 24: 202-222. 

Saunders, R.M.K. 2001. Schisandraceae. Species Plantarum: Flora of the World 4: 1-62. 

Savolainen, V., M.F. Fay, D.C. Albach, A. Backlund, M. van der Bank, K.M. Cameron, S.A. Johnson, M.D. Lledó, J.-C. Pintaud, 

M. Powell, M.C. Sheahan, D.E. Soltis, P.S. Soltis, P. Weston, W.M. Whitten, K..J. Wurdack, and M.W. Chase. 2000. 

Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bulletin 55: 257-309. 

Schafale, M.P., and A.S. Weakley. 1990. Classification of the natural communities of North Carolina, third approximation. 

North Carolina Natural Heritage Program, Raleigh, N.C. 

Schanzer, I.A. 1994. Taxonomic revision of the genus Filipendula Mill. (Rosaceae). J. Jpn. Bot. 69: 290-319. 

Scheen, A-C., C. Brochmann, A.K. Brysting, R. Elvar, A. Morris, D.E. Soltis, P.S. Soltis, and V.A. Albert. 2004. Northern 

hemisphere biogeography of Cerastium (Caryophyllaceae): insights from phylogenetic analysis of noncoding plastid 

nucleotide sequences. Amer. J. Bot. 91: 943-952. 

Schell, C.M., and M.J. Waterway. 1992. Allozyme variation and the genetic structure of populations of the rare sedge Carex 

misera (Cyperaceae). Plant Species Biol. 7: 141-150. 

Schilling, E.E. 1981. Systematics of Solanum sect. Solanum (Solanaceae) in North America. Systematic Bot. 6: 172-185. 

–––, C.R. Linder, R.D. Noyes, and L.H. Rieseberg. 1998. Phylogenetic relationships in Helianthus (Asteraceae) based on 

nuclear ribosomal DNA internal transcribed spacer region sequence data. Systematic Bot. 23: 177-187. 

Schlessman, M.A. 1985. Floral biology of American ginseng (Panax quinquefolium). Bull. Torrey Bot. Club 112: 129-133. 

Schmidt, G.J., and E.E. Schilling. 2000. Phylogeny and biogeography of Eupatorium (Asteraceae: Eupatorieae) based on 

nuclear ITS sequence data. Amer. J. Botany 87: 716-726. 

Schnabel, A., and J.F. Wendel. 1998. Cladistic biogeography of Gleditsia (Leguminosae) based on ndhF and rpl16 chloroplast 

gene sequences. Amer. J. Bot. 85: 1753-1765. 

Schneider, H., S.J. Russell, C.J. Cox, F. Bakker, S. Henderson, F. Rumsey, J. Barrett, M. Gibby, and J.C. Vogel. 2004. 

Chloroplast phylogeny of Asplenioid ferns based on rbcL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its 

implication for biogeography. Systematic Botany 29: 260-274. 

Schnell, D.E. 1976. Carnivorous plants of the United States and Canada. John F. Blair, Winston-Salem, NC. 125 pp. 

–––. 1977. Infraspecific variation in Sarracenia rubra Walt.: some observations. Castanea 42: 142-170. 

–––. 1979. A critical review of published variants of Sarracenia purpurea L. Castanea 44: 47-59. 

–––. 1980a. Pinguicula caerulea Walt. forma leucantha: a new form. Castanea 45: 56-60. 

–––. 1980b. Notes on the biology of Sarracenia oreophila (Kearney) Wherry. Castanea 45: 166-170. 

–––. 1981. Sarracenia purpurea L. ssp. venosa (Raf.) Wherry: variations in the Carolinas Coastal Plain. Castanea 46: 225-234.


–––. 1993. Sarracenia purpurea L. ssp. venosa (Raf.) Wherry var. burkii Schnell (Sarraceniaceae) – a new variety of the Gulf 

Coastal Plain. Rhodora 95: 6-10. 

–––. 1995. Drosera filiformis Raf.: one species or two? Carnivorous Plant Newsletter 24: 11-15. 

–––. 1998. A pitcher key to the genus Sarracenia L. (Sarraceniaceae). Castanea 63: 489-492. 

–––. 2002a. Sarracenia minor Walt. var. okeefenokeensis Schnell: a new variety. Carnivorous Plant Newsletter 31: 36-39. 

–––. 2002b. Carnivorous plants of the United States and Canada. 2nd edition. Timber Press, Portland, OR. 468 pp. 

–––, and R.O. Determann. 1997. Sarracenia purpurea L. ssp. venosa (Raf.) Wherry var. montana Schnell & Determann 

(Sarraceniaceae): a new variety. Castanea 62: 60-62. 

Scholz, U. 1981. Monographie der Gattung Oplismenus (Gramineae). Phanerog. Monog. 13. J. Cramer, Vaduz, Germany. 217 

pp. 

Schrader, J.A., and W.R. Graves. 2002. Infraspecific systematics of Alnus maritima (Betulaceae) from three widely disjunct 

populations. Castanea 67: 380-401. 

–––, and W.R. Graves. 2004. Systematics of Dirca (Thymelaeaceae) based on ITS sequences and ISSR polymorphisms. Sida 

21: 511-524. 

Schultheis, L.M., and M.J. Donoghue. 2004. Molecular phylogeny and biogeography of Ribes (Grossulariaceae), with an 

emphasis on gooseberries (subg. Grossularia). Systematic Botany 29: 77-96. 

Schumacher, A. 1947. Die Moorlilien (Narthecium)-Arten Europas. Archiv für Hydrobiologie 41:112-195. 

Schuyler, A.E. 1962. A new species of Scirpus in the northeastern United States. Rhodora 64: 43-49. 

–––. 1967. A taxonomic revision of North American leafy species of Scirpus. Proc. Acad. Nat. Sci. Phila. 119: 295-323. 

–––. 1974. Typification and application of the names Scirpus americanus Pers., S. olneyi Gray, and S. pungens Vahl. Rhodora 

76: 51-52. 

–––. 1975. Scirpus cylindricus: an ecologically restricted eastern North American tuberous bulrush. Bartonia 43: 29-37. 

–––. 1989. Intertidal variants of Bacopa rotundifolia and B. innominata in the Chesapeake Bay drainage. Bartonia 55: 18-22. 

–––. 1996. Taxonomic status of Panicum hirstii Swallen. Bartonia 59: 95-96. 

Schweitzer, J.A., and K.C. Larson. 1999. Greater morphological plasticity of exotic honeysuckle species may make them better 

invaders than native species. J. Torrey Bot. Soc. 126: 15-23. 

Scora, R.W. 1967. Interspecific relationships in the genus Monarda (Labiatae). Univ. of Calif. Publ. in Botany 41: 1-69. 

Scott, P.J., and R.T. Day. 1983. Diapensiaceae: a review of the taxonomy. Taxon 32: 417-423. 

Seigler, D.S., and J.E. Ebinger. 2005. New combinations in the genus Vachellia (Fabaceae: Mimosaoideae) from the New 

World. Phytologia 87: 139-178 

Seine, R., and W. Barthlott. 1994. Some proposals on the infrageneric classification of Drosera L. Taxon 43: 583-589. 

Semple, J.C. 1981. A revision of the goldenaster genus Chrysopsis (Nutt.) Ell. nom. cons. (Compositae-Astereae). Rhodora 83: 

323-384. 

–––. 1983. Range expansion of Heterotheca camporum (Compositae: Astereae) in the southeastern United States. Brittonia 35: 

140-146. 

–––. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae): the prairie and montane 

goldenasters of North America. Univ. of Waterloo Biological Series 37. 

–––. 2003. New names and combinations in goldenrods, Solidago (Asteraceae: Astereae). Sida 20: 1605-1616. 

–––. 2004. Miscellaneous nomenclatural changes in Astereae (Asteraceae). Sida 21: 759-765. 

–––, and L. Brouillet. 1980a. A synopsis of North American asters: the subgenera, sections, and subsections of Aster and 

Lasallea. Am. J. Bot. 67: 1010-1026. 

–––, and L. Brouillet. 1980b. Chromosome numbers and satellite morphology in Aster and Lasallea. Am. J. Bot. 67: 1027- 

1039. 

–––, and F.D. Bowers. 1985. A revision of the goldenaster genus Pityopsis Nutt. (Compositae: Astereae). Univ. of Waterloo 

Biological Series 29: 1-34. 

–––, J.G. Chmielewski, and M.A. Lane. 1989. Chromosome number determinations in fam. Compositae, tribe Astereae. III. 

Additional counts and comments on generic limits and ancestral base numbers. Rhodora 91: 296-314. 

–––, S.B. Heard, and ChunSheng Xiang. 1996. The asters of Ontario (Compositae: Astereae): Diplactis Raf., Oclemena E.L. 

Greene, Doellingeria Nees and Aster L. (including Canadanthus Nesom, Symphyotrichum Nees, and Virgulus Raf.). Univ. 

of Waterloo Biology Series 38. 

–––, G.S. Ringius, and J.J. Zhang. 1999. The goldenrods of Ontario: Solidago L. and Euthamia Nutt. 3rd Edition. Univ. 

Waterloo Biol. Ser. 39: 1-90. 

Sennblad, B., and B. Bremer. 1996. The familial and subfamilial relationships of Apocynaceae and Asclepiadaceae evaluated 

with rbcL data. Pl. Syst. Evol. 202: 153-175. 

Senters, A.E., and D.E. Soltis. 2003. Phylogenetic relationships in Ribes (Grossulariaceae) inferred from ITS sequence data. 

Taxon 52: 51-66. 

Serviss, B.E., S.T. McDaniel, and C.T. Bryson. 2000. Occurrence, distribution, and ecology of Alocasia, Caladium, Colocasia, 

and Xanthosoma (Araceae) in the southeastern United States. Sida 19: 149-174. 

Shaw, J., and R.L. Small. 2004. Addressing the "hardest puzzle in American pomology:" phylogeny of Prunus sect. 

Prunocerasus (Rosaceae) based on seven noncoding chloroplast DNA regions. Amer. J. Bot. 91: 985-996. 

Shaw, J.M.H. 2000. A taxonomic revision of Podophyllum in the wild and in cultivation. The New Plantsman 7: 30-41, 103- 

113, 142-159, 220-235.


–––. 2002. Podophyllum. In Stearn, W.T. 2002. The genus Epimedium and other herbaceous Berberidaceae, including the 

genus Podophyllum. Timber Press, Portland, OR. 

Shaw, R.B., and R.D. Webster. 1987. The genus Eriochloa (Poaceae: Paniceae) in North and Central America. Sida 12: 165- 

207. 

Shen, Chung-Fu. 1992. A monograph of the genus Fagus Tourn. ex L. (Fagaceae). Ph.D. dissertation, Biology dept., City 

University of New York. 390 pp. 

Sherff, E.E., and E.J. Alexander. 1955. Compositae – Heliantheae – Coreopsidinae. North American Flora, series II, part 2. 

New York Botanical Garden. 

Shetler, S.G. 1982. Variation and evolution of the Nearctic harebells (Campanula subsect. Heterophylla). Phanerogamarum 

Monographiae XI. J. Cramer, Vaduz. 516 pp. 

–––, and N.R. Morin. 1986. Seed morphology in North American Campanulaceae. Ann. Mo. Bot. Gard. 73: 653-688. 

–––, and S.S. Orli. 2000. Annotated checklist of the vascular plants of the Washington-Baltimore area. Part I: ferns, fern allies, 

gymnosperms, and dicotyledons. Dept. of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, DC. 

–––, and S.S. Orli. 2002. Annotated checklist of the vascular plants of the Washington-Baltimore area. Part I: monocotyledons. 

Dept. of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC. 

Sheviak, C.J. 1991. Morphological variation in the compilospecies Spiranthes cernua (L.) L.C. Rich.: ecologically-limited 

effects of gene flow. Lindleyana 6: 228-234. 

–––. 1994. Cypripedium parviflorum Salisb. I. The small-flowered varieties. Amer. Orchid Soc. Bull. 63: 664-669. 

–––, and P.M. Catling. 1980. The identity and status of Spiranthes ochroleuca (Rydberg) Rydberg. Rhodora 82: 525-562. 

Shinners, L.H. 1946. Revision of the genus Kuhnia L. Wrightia 1: 122-144. 

–––. 1957. Synopsis of the genus Eustoma (Gentianaceae). Southwestern Naturalist 2: 38-43. 

–––. 1962a. Calamintha (Labiatae) in the southern United States. Sida 1: 69-75. 

–––. 1962b. Synopsis of Collinsonia (Labiatae). Sida 1: 76-83. 

–––. 1962c. Drosera (Droseraceae) in the southeastern United States: an interim report. Sida 1: 53-59. 

–––. 1962d. Synopsis of United States Bonamia, including Breweria and Stylisma (Convolvulaceae). Castanea 27: 65-77. 

–––. 1962e. Vegetative key to woody Labiatae of the southeastern Coastal Plain. Sida 1: 92-93. 

–––. 1962f. Micromeria brownei and its allies. Sida 1: 94-97. 

–––. 1962g. Synopsis of Conradina. Sida 1: 84-88. 

–––. 1971. Kuhnia L. transferred to Brickellia Ell. (Compositae). Sida 4: 274. 

Shinwari, Z.K., R. Terauchi, F.H. Utech, and S. Kawano. 1994. Recognition of the New World Disporum section Prosartes as 

Prosartes (Liliaceae) based on the sequence data of the rbcL gene. Taxon 43: 353-366. 

Shulkina, T.V., J.F. Gaskin, and W.M.M. Eddie. 2003. Morphological studies toward an improved classification of 

Campanulaceae s. str. Ann. Missouri Bot. Gard. 90: 576-591. 

Siedo, S.J. 1999. A taxonomic treatment of Sida sect. Ellipticifoliae (Malvaceae). Lundellia 2: 100-127. 

Sieren, D.J. 1981. The taxonomy of the genus Euthamia. Rhodora 83: 551-579. 

Silberhorn, G.M. 1998. Invasion of Cuscuta indecora Choisy (Convolvulaceae) in a tidal brackish marsh in Virginia. Castanea 

63: 190-191. 

Silveus, W.A. 1942. Grasses: classification and description of species of Paspalum and Panicum in the United States. 

Published by the author, San Antonio, Texas. 

Simmers, R.W., and R. Kral. 1992. A new species of Blephilia (Lamiaceae) from northern Alabama. Rhodora 94: 1-14. 

Simpson, B.B,, A. Weeks, D.M. Helfgott, and L.L. Larkin. 2004. Species relationships in Krameria (Krameriaceae) based on 

ITS sequences and morphology: implications for character utility and biogeography. Systematic Botany 29: 97-108. 

Simpson, M.G. 1983. Pollen ultrastructure of the Haemodoraceae and its taxonomic significance. Grana 22: 79-103. 

–––, and W.C. Dickison. 1981. Comparative anatomy of Lachnanthes and Lophiola (Haemodoraceae). Flora 171: 95-113. 

Singhurst, J.R., and W.C. Holmes. 2004. Comments on the rediscovery and distribution of Cunila origanoides (Lamiaceae) in 

Texas. Sida 21: 1161-1163. 

–––, E.L. Keith, and W.C. Holmes. 2005. Three species of vascular plants new to Texas. Phytologia 87: 124-128. 

Siripun, K.C., and E.E. Schilling. 2006. Molecular confirmation of the hybrid origin of Eupatorium godfreyanum (Asteraceae). 

Amer. J. Bot. 93: 319-325. 

Sipple, W.S. 2002. Pine-barren golden-heather (Hudsonia ericoides L.) reported for the first time in Maryland. Bartonia 61: 

149-150. 

Skeen, J.N., P.D. Doerr, and D.H. Van Lear. 1993. Oak-hickory-pine forests. In W.H. Martin, S.G. Boyce, and A.C. 

Echternacht, eds. Biodiversity of the southeastern United States. John Wiley & Sons, New York, NY. 

Skinner, M.W., and B.A. Sorrie. 2002. Conservation and ecology of Lilium pyrophilum, a new species of Liliaceae from the 

Sandhills region of the Carolinas and Virginia, U.S.A. Novon 12: 94-105. 

Škoda, B. 1997. Taxonomic comments on the "Flora of North America north of Mexico," vol. 2, with some nomenclatural 

combinations for Pteridophyta. Preslia, Praha 68: 341-359. 

Skog, J.T., and N.H. Nickerson. 1972. Variation and speciation in the genus Hudsonia. Ann Mo. Bot. Gard. 59: 454-464. 

Skvortsov, A.K. 1979. Taxonomy and distribution of Circaea (Onagraceae) in the U.S.S.R. Ann Mo. Bot. Club 66: 880-892. 

Sleumer, H. 1967a. Die Gattung Gaylussacia H.B.K. Botanische Jahrbücher Syst. 86: 309-384. 

–––. 1967b. Monographia Clethracearum. Botanische Jahrbücher Syst. 87: 36-175.


Small, E. 1978. A numerical and nomenclatural analysis of morpho-geographic taxa of Humulus. Systematic Bot. 3: 37-76. 

Small, J.K. 1903. Flora of the southeastern United States, being descriptions of the seed-plants, ferns and fern-allies growing 

naturally in North Carolina, South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, Louisiana, and in 

Oklahoma and Texas east of the one hundredth meridian. Published by the author, New York, NY. 

–––. 1913. Flora of the southeastern United States, being descriptions of the seed-plants, ferns and fern-allies growing naturally 

in North Carolina, South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, Louisiana, and in 

Oklahoma and Texas east of the one hundredth meridian, second edition. Published by the author, New York, NY. 1394 

pp. 

–––. 1924. A new bog-asphodel from the mountains. Torreya 24: 86-87. 

–––. 1933. Manual of the southeastern flora, being descriptions of the seed plants growing naturally in Florida, Alabama, 

Mississippi, eastern Louisiana, Tennessee, North Carolina, South Carolina, and Georgia. University of North Carolina 

Press, Chapel Hill, N.C. 

–––. 1938. Ferns of the southeastern states. The Science Press, Lancaster, Pa. 

Smedmark, J.E.E. 2006. Recircumscription of Geum L. (Coluriae: Rosaceae). Bot. Jahrb. Syst. 126: 1-9. 

–––, and T. Eriksson. 2002. Phylogenetic relationships of Geum (Rosaceae) and relatives inferred from the nrITS and trnL-trnF 

regions. Syst. Bot. 27: 303-317. 

Smith, A.C. 1944. Araliaceae. North American Flora 28B: 3-41. N.Y. Botanical Garden, New York. 

Smith, A.R. 1992. A review of the fern genus Micropolypodium (Grammitidaceae). Novon 2: 419-425. 

–––, and R.B. Cranfill. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae). Amer. Fern J. 92: 131- 

149. 

–––, K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, & P.G. Wolf. 2006. A classification of extant ferns. Taxon 55: 705- 

731. 

Smith, B.D., J.B. beck, A.T. Denham, and P.J. Calie. 2004. High resolution GIS mapping and current status of the ten viable 

populations of Short's goldenrod (Solidago shortii – Asteraceae) in Kentucky. Sida 21: 1121-1130. 

Smith, E.B. 1976. A biosystematic survey of Coreopsis in eastern United States and Canada. Sida 6: 123-215. 

–––. 1981. New combinations in Croptilon (Compositae - Asteraceae). Sida 9: 59-63. 

–––. 1982a. Juvenile and adult leaflet phases in Aralia spinosa (Araliaceae). Sida 9: 330-332. 

–––. 1982b. A new variety of Cardamine angustata (Cruciferae) from the Ouachita Mountains of Arkansas. Brittonia 34: 376- 

380. 

–––. 1988. An atlas and annotated list of the vascular plants of Arkansas, second edition. Published by the author. Fayetteville, 

AR. 489 pp. 

Smith, G.L., and W.S. Flory. 1990. Studies on Hymenocallis henryae (Amaryllidaceae). Brittonia 42: 212-220. 

–––, and M.A. Garland. 1996. Taxonomic status of Hymenocallis choctawensis and Hymenocallis puntagordensis 

(Amaryllidaceae). Sida 17: 305-319. 

–––, and M.A. Garland. 2003. Nomenclature of Hymenocallis taxa (Amaryllidaceae) in the southeastern United States. Taxon 

52: 805-817 

Smith, L.B., and C.E. Wood, Jr. 1975. The genera of Bromeliaceae in the southeastern United States. J. Arnold Arb. 56: 375- 

397. 

Smith, M., and N. Parker. 2005. Quercus montana (Fagaceae), new to Missouri. Sida 21: 1921-1922. 

Smith, R.R., and D.B. Ward. 1976. Taxonomy of the genus Polygala series Decurrentes (Polygalaceae). Sida 6: 284-310. 

Smith, S.D., R.S. Cowan, K.B. Gregg, M.W. Chase, N. Maxted, and M.F. Fay. 2004. Genetic discontinuities among populations 

of Cleistes (Orchidaceae, Vanilloideae) in North America. Bot. J. Linn. Soc. 145: 87-95. 

Smith, S.G. 1995. New combinations in North American Schoenoplectus, Bolboschoenus, Isolepis, and Trichophorum 

(Cyperaceae). Novon 5: 97-102. 

–––, and E. Hayasaka. 2002. New combinations within North American Schoenoplectus smithii and S. purshianus (sect. 

Actaeogeton, Cyperaceae) and comparsion with eastern Asian allies. Novon 12: 106-111. 

Smith, T.W., J.T. Donaldson, T.F. Wieboldt, G.L. Kauffman, and M.J. Waterway. 2006. The geographic and ecological 

distribution of the Roan Mountain sedge, Carex roanensis (Cyperaceae). Castanea 71: 45-53. 

Snoeijer, W. 1996. Catharanthus roseus, the Madagascar Periwinkle, a review of its cultivars. Wageningen Agricultural 

University Papers 96-3: 47-120. 

Snow, N. 1998. Caryopsis morphology of Leptochloa sensu lato (Poaceae, Chloridioideae). Sida 18: 271-282. 

Snyder, D. 1996. The genus Rhexia in New Jersey. Bartonia 59: 55-70. 

Snyder, L.H., Jr., and J.G. Bruce. 1986. Field guide to the ferns and other pteridophytes of Georgia. Univ. of Georgia Press, 

Athens, GA. 270 pp. 

Socorro González-Elizondo, M., and P.M. Peterson. 1997. A classification of and key to the supraspecific taxa in Eleocharis 

(Cyperaceae). Taxon 46: 433-449. 

Soejima, A., and J. Wen. 2006. Phylogenetic analysis of the grape family (Vitaceae) based on three chloroplast markers. Amer. 

J. Bot. 93: 278-287. 

Soják, J. 1992. Generische problematik der Selaginellaceae. Preslia, Praha 64: 151-158. 

–––. 2004. Potentilla L. (Rosaceae) and related genera in the former USSR (identification key, checklist and figures). Notes on 

Potentilla XVI. Bot. Jahrb. Syst. 125: 253-340.


Soltis, D.E. 1980. A biosystematic study of Sullivantia and related studies in the Saxifragaceae. Ph.D. thesis, Indiana 

University. 236 pp. 

–––. 1985. Allozymic differentiation among Heuchera americana, H. parviflora, H. pubescens, and H. villosa (Saxifragaceae). 


–––, B.A. Bohm, and G.L. Nesom. 1983. Flavonoid chemistry of cytotypes in Galax (Diapensiaceae). Systematic Bot. 8: 15-23. 

–––, P.S. Soltis, M.W. Chase, M.E. Mort, D.C. Albach, M. Zanis, V. Savolainen, W.H. Hahn, S.B. Hoot, M.F. Fay, M. Axtell, 

S.M. Swensen, L.M. Prince, W.J. Kress, K.C. Nixon, and J.S. Farris. 2000. Angiosperm phylogeny inferred from 18S 

rDNA, rbcL, and atpB sequences. Bot. J. Linn. Soc. 133: 381-461. 

–––, Qiu-Yun Xiang, and L. Hurford. 1995. Relationships and evolution of Hydrangeaceae based on rbcL sequence data. Am. 

J. Bot. 82: 504-514. 

–––, R.K. Kuzoff, E. Conti, R. Gornall, and K. Ferguson. 1996. matK and rbcL gene sequence data indicate that Saxifraga 

(Saxifragaceae) is polyphyletic. Am. J. Bot. 83: 371-382. 

Somers, P., and W.R. Buck. 1975. Selaginella ludoviciana, S. apoda, and their hybrids in the southeastern United States. Am. 

Fern J. 65: 76-82. 

Soreng, R.J. 1998. An infrageneric classification for Poa in North America, and other notes on sections, species, and subspecies 

of Poa, Puccinellia, and Dissanthelium (Poaceae). Novon 8: 187-202. 

–––, and E.E. Terrell. 1997. Taxonomic notes on Schedonorus, a segregate genus from Festuca or Lolium, with a new 

nothogenus, ×Schedololium, and new combinations. Phytologia 83: 85-88. 

–––, P.M. Peterson, G. Davidse, E.J. Judziewicz, F.O. Zuloaga, T.S. Filgueiras, and O. Morrone. 2003. Catalogue of New 

World grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. National Herbarium 48: 1-730. 

Sorrie, B.A. 1997. Notes on Lycopus cokeri (Lamiaceae). Castanea 62: 119-126. 

–––. 1998a. Distribution of Drosera filiformis and D. tracyi (Droseraceae): phytogeographic implications. Rhodora 100: 239- 

260. 

–––. 1998b. Noteworthy collections: Georgia. Castanea 63: 496-500. 

–––. 2000. Rhynchospora leptocarpa (Cyperaceae), an overlooked species of the southeastern United States. Sida 19: 139-147. 

–––, and R.J. LeBlond. 1997. Vascular plants new to the Bahamas and Andros Island. Bahamas J. of Science 4: 14-18. 

–––, and S.W. Leonard. 1999. Noteworthy records of Mississippi vascular plants. Sida 18: 889-908. 

–––, and P. Somers. 1999. The vascular plants of Massachusetts: a county checklist. Massachusetts Division of Fisheries and 

Wildlife, Natural Heritage & Endangered Species Program, Westborough, MA. 187 pp. 

–––, and A.S. Weakley. 2001. Coastal plain vascular plant endemics: phytogeographic patterns. Castanea 66: 50-82. 

–––, and A.S. Weakley. [in press]. Developing a blueprint for conservation of the endangered longleaf pine ecosystem based on 

centers of Coastal Plain endemism. Applied Vegetation Science. 

–––, M.H. MacRoberts, B.R. MacRoberts, and S.B. Walker. 2003. Oxypolis ternata (Apiaceae) deleted from the Texas flora. 

Sida 20: 1323-1324. 

–––, B. van Eerden, M.J. Russo. 1997. Noteworthy plants from Fort Bragg and Camp MacKall, North Carolina. Castanea 62: 

239-259. 

Sousa S., M., and V.E. Rudd. 1993. Revisión del género Styphnolobium (Leguminosae: Papilionoideae: Sophoreae). Ann. 


South Carolina Heritage Trust. 1993. Rare, threatened, and endangered species of South Carolina. S.C. Heritage Trust, 

Columbia, SC. 

Southall, R.M., and J.W. Hardin. 1974. A taxonomic revision of Kalmia (Ericaceae). J. Elisha Mitchell Sci. Soc. 90: 1-23. 

Spalik, K. 1996. Species boundaries, phylogenetic relationships, and ecological differentiation in Anthriscus (Apiaceae). Pl. 

Syst. Evol. 199: 17-32. 

Spangler, R.E., and R.G. Olmstead. 1999. Phylogenetic analysis of Bignoniaceae based on the cpDNA gene sequences rbcL and 

ndhF. Ann. Missouri Bot. Gard. 86: 33-46. 

Speer, W.D., and K.W. Hilu. 1999. Relationships between two infraspecific taxa of Pteridium aquilinum (Dennstaedtiaceae). I. 

Morphological evidence. Systematic Bot. 23: 305-312. 

–––, C.R. Werth, and K.W. Hilu. 1999. Relationships between two infraspecific taxa of Pteridium aquilinum 

(Dennstaedtiaceae). II. Isozyme evidence. Systematic Bot. 23: 313-325. 

Spongberg, S.A. 1971. The Staphyleaceae in the southeastern United States. J. Arnold Arb. 52: 196-203. 

–––. 1972. The genera of Saxifragaceae in the southeastern United States. J. Arnold Arb. 53: 409-498. 

–––. 1974. A review of deciduous-leaved species of Stewartia (Theaceae). J. Arnold Arb. 55: 182-214. 

–––. 1977. Ebenaceae hardy in temperate North America. J. Arnold Arb. 58: 146-160. 

–––. 1998. Magnoliaceae hardy in cooler temperate regions. In D. Hunt, ed. Magnolias and their allies. Proceedings of an 

international symposium, Royal Holloway, University of London, Egham, Surrey, U.K., 12-13 April 1996. International 

Dendrological Society and the Magnolia Society. 

Spooner, D.M. 1984. Infraspecific variation in Gratiola viscidula Pennell (Scrophulariaceae). Rhodora 86: 79-87. 

–––, and J.S. Shelly. 1983. The national historical distribution of Platanthera peramoena (A. Gray) A. Gray (Orchidaceae) and 

its status in Ohio. Rhodora 85: 55-64. 

–––, A.W. Cusick, G.F. Hall, and J.M. Baskin. 1985. Observations on the distribution and ecology of Sida hermaphrodita (L.) 

Rusby (Malvaceae). Sida 11: 215-225. 

Stace, C. 1997. New flora of the British Isles, second edition. Univ. of Cambridge Press, Cambridge. 1130 pp.


Staff of the Bailey Hortorium. 1976. Hortus third: a concise dictionary of plants cultivated in the United States and Canada. 

MacMillan, NY. 

Stalter, R., and E. Lamont. 1996. Noteworthy collections: Virginia. Castanea 61: 396-397. 

Standley, L.A. 1983. A clarification of the status of Carex crinita and C. gynandra (Cyperaceae). Rhodora 85: 229-241. 

–––, J.L. Dudley, and L.P. Bruederle. 1991. Electrophoretic variability in the rare sedge, Carex polymorpha (Cyperaceae). Bull. 

Torrey Bot. Club 118: 444-450. 

Stanford, A.M. 1998. The biogeography and phylogeny of Castanea, Fagus, and Juglans based on MATK and ITS sequence 

data. Ph.D. dissertation, Biology Departement, University of North Carolina at Chapel Hill. 

Stanford, A.M., R. Harden, and C.R. Parks. 2000. Phylogeny and biogeography of Juglans (Juglandaceae) based on matK and 

ITS sequence data. Am. J. Bot. 87: 872-882. 

Staples, G.W., J.H. Wiersema, N.A. Chambers, and D.F. Austin. 2005. The restoration of Ipomoea muricata (L.) Jacq. 

(Convolvulaceae). Taxon 54: 1075-1079. 

Steane, D.A., R.P.J. de Kok, and R.G. Olmstead. 2004. Phylogenetic relationships betrween Clerodendrum (Lamiaceae) and 

other Ajugoid genera inferred from nuclear and chloroplast DNA sequence data. Molecular Phylogenetics and Evolution 

32: 39-45. 

–––, R.W. Scotland, D.J. Mabberley, and R.G. Olmstead. 1999. Molecular systematics of Clerodendrum (Lamiaceae): ITS 

sequences and total evidence. Amer. J. Bot. 86: 98-107. 

Stearn, W.T. 2002. The genus Epimedium and other herbaceous Berberidaceae, including the genus Podophyllum. Timber 

Press, Portland, OR. 

Stefanović, S., L. Krueger, and R.G. Olmstead. 2002. Monophyly of the Convolvulaceae and circumscription of their major 

lineages based on DNA sequences of multiple chloroplast loci. Amer. J. Bot. 89: 1510-1522. 

–––, D.F. Austin, and R.G. Olmstead. 2003. Classification of Convolvulaceae: a phylogenetic approach. Systematic Botany 28: 

791-806. 

Stein, J., D. Binion, and R. Acciavatti. 2003. Field guide to native oak species of eastern North America. Forest Health 

Technology Enterprise Team Publ. 2003-01. 

Stephenson, S.N. 1971. The biosystematics and ecology of the genus Brachyelytrum in Michigan. Mich. Bot. 10: 19-33. 

Steury, B.W. 1999. Noteworthy collections: Maryland. Castanea 64: 271-272. 

–––. 2004a. Noteworthy collections: District of Columbia and Maryland. Castanea 69: 154-157 

–––. 2004b. Noteworthy collections: Virginia. Castanea 69: 241-242. 

–––, R.W. Tyndall, and G. Cooley. 1996. Noteworthy collections: Maryland. Castanea 61: 392-396. 

Stevens, P.F. 2006. Angiosperm Phylogeny Website. Version 7, May 2006 [and more or less continuously updated since]. 

http://www.mobot.org/MOBOT/research/APweb/ 

Stevenson, D.W. 1991. The Zamiaceae in the southeastern United States. J. Arnold Arboretum, Supp. Series 1: 367-384. 

Steyermark, J.A. 1949. Lindera melissaefolia. Rhodora 51: 153-162. 

–––. 1951. A glabrous variety of Silphium terebinthinaceum. Rhodora 53: 133-135. 

–––, and C.S. Steyermark. 1960. Hepatica in North America. Rhodora 62: 223-232. 

Stiles, B.J., and C.L. Howel. 1998. Floristic survey of Rabun County, Georgia, part II. Castanea 63: 154-160. 

Stone, D.E. 1961. Ploidal level and stomatal size in the American hickories. Brittonia 13: 293-302. 

–––. 1968. Cytological and morphological notes on the southeastern endemic Schisandra glabra (Schisandraceae). J. Elisha 

Mitchell Sci. Soc. 84: 351-356. 

–––, and J.L. Freeman. 1968. Cytotaxonomy of Illicium floridanum and I. parviflorum (Illiciaceae). J. Arnold Arb. 49: 41-51. 

–––, G.A. Adrouny, and R.H. Flake. 1969. New World Juglandaceae. II. Hickory nut oils, phenetic similarities, and 

evolutionary implications in the genus Carya. Am. J. Bot. 56: 928-935. 

Stoynoff, N., and W.J. Hess. 1990. A new status for Quercus shumardii var. acerifolia (Fagaceae). Sida 14: 267-271. 

Straley, G.B. 1977. Systematics of Oenothera sect. Kneiffia (Onagraceae). Ann. Missouri Bot. Gard. 64: 381-424. 

Strand, A.E., and R. Wyatt. 1991. Geographical variation and biosystematics of sand myrtle, Leiophyllum buxifolium 

(Ericaceae). Systematic Bot. 16: 529-545. 

Strausbaugh, P.D., and E.L. Core. 1978. Flora of West Virginia, second edition, Seneca Books, Grantsville, WV. 

Stritch, L.R. 1984. Nomenclatural contributions to a revision of the genus Wisteria. Phytologia 56: 183-184. 

Strong, M.T. 1994. Taxonomy of Scirpus, Trichophorum, and Schoenoplectus (Cyperaceae) in Virginia. Bartonia 58: 29-68. 

Struwe, L., and V.A. Albert, eds. 2002. Gentianaceae: systematics and natural history. Cambridge Univ. Press, Cambridge. 652 

pp. 

–––, V.A. Albert, and B. Bremer. 1994. Cladistics and family level classification of the Gentianales. Cladistics 10: 175-206. 

–––, J.W. Kadereit, J. Klackenberg, S. Nilsson, M. Thiv, B. von Hagen, and V.A. Albert. 2002. Systematics, character 

evolution, and biogeography of Gentianaceae, including a new tribal and tribal classification. In L. Struwe and V.A. Albert, 

eds. 2002. Gentianaceae: systematics and natural history. Cambridge Univ. Press, Cambridge. 652 pp. 

Sugawara, T. 1987. Cytotaxonomic study of Asarum senso lato. Proc. Sino-Jpn. Symposium Pl. Chromos. {}: 147-150. 

Sullivan, J.R. 1978. Putative hybridization in the genus Eupatorium (Compositae). Rhodora 80: 513-527. 

–––. 1985. Systematics of the Physalis viscosa complex (Solanaceae). Systematic Bot. 10: 426-444. 

–––. 2004. The genus Physalis (Solanaceae) in the southeastern United States. Rhodora 106: 305-326. 

Sundberg, S.D. 2004. New combinations in North American Symphyotrichum subgenus Astropolium (Asteraceae: Astereae). 

Sida 21: 903-910.


Sundell, E., R.D. Thomas, C. Amason, R.L. Stuckey, and J. Logan. 1999. Noteworthy vascular plants from Arkansas. Sida 18: 

877-887. 

Sutter, R., L. Mansberg, and J. Moore. 1983. Endangered, threatened, and rare plant species of North Carolina: a revised list. 

Association of Southeastern Biologists Bulletin 30:153-163. 

Sutton, D.A. 1988. A revision of the tribe Antirrhineae. British Museum (Natural History), Oxford Univ. Press, London. 

Swallen, J.R. 1961. A new species of Panicum from New Jersey. Rhodora 63: 235-236. 

Sweeney, C.R. 1970. Monograph of the genus Silphium. I. Silphium compositum Michaux (Compositae). Ohio J. of Sci. 70: 

226-233. 

Sweeney, P.W., and R.A. Price. 2000. Polyphyly of the genus Dentaria (Brassicaceae): evidence from trnL intron and ndhF 

sequence data. Systematic Botany 25: 468-478. 

–––, and R.A. Price. 2001. A multivariate morphological analysis of the Cardamine concatenata alliance (Brassicaceae). 

Brittonia 53: 82-95. 

Sytsma, K.J., J. Morawetz, J.C. Pires, M. Nepokroeff, E. Conti, M. Zjhra, J.C. Hall, and M.W. Chase. 2002. Urticalean rosids: 

circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. Amer. J. Botany 89: 1531- 

1546. 

Takahashi, M., and S. Kawano. 1989. Pollen morphology of the Melanthiaceae and its systematic implications. Ann. Mo. Bot. 

Gard. 76: 863-876. 

Takhtajan, A. 1986. Floristic regions of the world. Univ. of Calif. Press, Berkeley, CA. 522 pp. 

Takhtajan, A. 1997. Diversity and classification of flowering plants. Columbia Univ. Press, NY. 643 pp. 

Tamura , M.N., J. Yamashita, S. Fuse, and M. Haraguchi. 2004. Molecular phylogeny of monocotyledons inferred from 

combined analysis of plastid matK and rbcL gene sequences. J. Plant Res. 117: 109-120. 

Tank, D.C., P.M. Beardsley, S.A. Kelchner, and R.G. Olmstead. 2006. review of the systematics of Scrophulariaceae s.l. and 

their current disposition. Australian Systematic Botany: 19: 289-307.. 

Taylor, C.E.S., and R.J. Taylor. 1983. New species, new combinations and notes on the goldenrods (Euthamia and Solidago – 

Asteraceae). Sida 10: 176-183. 

Taylor, C.M. 1994. Revision of Tetragonia (Aizoaceae) in South America. Systematic Bot. 19: 575-589. 

Taylor, P. 1989. The genus Utricularia – a taxonomic monograph. Her Majesty's Stationery Office, London. 

Taylor, S. I. and F. Levy. 2002. Responses to soils and a test for preadaptation to serpentine in Phacelia dubia 

(Hydrophyllaceae). New Phytologist 155:437-447. 

Terrell, E.E. 1959. A revision of the Houstonia purpurea group (Rubiaceae). Rhodora 61: 157-181, 188-207. 

–––. 1978. Taxonomic notes on Houstonia purpurea var. montana (Rubiaceae). Castanea 43: 25-29. 

–––. 1986. Taxonomic and nomenclatural notes on Houstonia nigricans (Rubiaceae). Sida 11: 471-481. 

–––. 1991. Overview and annotated list of North American species of Hedyotis, Houstonia, Oldenlandia, and related genera. 

Phytologia 71: 212-243. 

–––. 1996. Revision of Houstonia (Rubiaceae-Hedyotidae). Systematic Bot. Monographs 48: 1-118. 

–––. 2001. Taxonomy of Stenaria (Rubiaceae: Hedyotitideae), a new genus including Hedyotis nigricans. Sida 19: 591-614. 

–––, and J.L. Reveal. 1996. Noteworthy collections: Maryland. Castanea 61: 95-96. 

–––, and H. Robinson. 2006. Taxonomy of North American species of Oldenlandia (Rubiaceae). Sida 22: 305-329. 

–––, P.M. Peterson, J.L. Reveal, and M.R. Duvall. 1997. Taxonomy of North American species of Zizania (Poaceae). Sida 17: 

533-549. 

Tharp, B.C., and M.C. Johnston. 1961. Recharacterization of Dichondra (Convolvulaceae) and a revision of the North American 

species. Brittonia 13: 346-360. 

Therman, E. 1950. Chromosome numbers in American Polygonatum species. Am. J. Bot. 37: 407-413. 

–––. 1953. Chromosomal evolution in the genus Polygonatum. Hereditas 39: 277-288. 

–––. 1956. Cytotaxonomy of the tribe Polygonatae. Am. J. Bot. 43: 134-142. 

Thien, L.B., E.G. Ellgaard, M.S. Devall, S.E. Ellgaard, and P.F. Ramp. 1994. Population structure and reproductive biology of 

Saururus cernuus L. (Saururaceae). Plant Species Biol. 9: 47-55. 

Thieret, J.W. 1971. The genera of Orobanchaceae in the southeastern United States. J. Arnold Arb. 52: 404-434. 

–––. 1972. The Phrymaceae in the southeastern United States. J. Arnold Arb. 53: 226-233. 

–––. 1975. The Mayacaceae in the southeastern United States. J. Arnold Arb. 56: 248-255. 

–––. 1977. The Martyniaceae in the southeastern United States. J. Arnold Arb. 58: 25-39. 

–––. 1982. The Sparganiaceae in the southeastern United States. J. Arnold Arb. 63: 341-355. 

–––. 1988. The Juncaginaceae in the southeastern United States. J. Arnold Arb. 69: 1-23. 

–––, and J.R. Baird. 1985. Thlaspi alliaceum (Cruciferae) in Kentucky and Indiana. Trans. Kentucky Academy of Science 46: 

145-146. 

–––, and J.O. Luken. 1996. The Typhaceae in the southeastern United States. Harvard Papers in Botany 8: 27-56. 

Thomas, J.L. 1960. A monographic study of the Cyrillaceae. Contr. Gray Herb. Harvard Univ. 186: 1-114. 

Thomas, R.D., and C.M. Allen. 1993. Atlas of the vascular flora of Louisiana. Volume I: Ferns & fern allies, conifers, & 

monocotyledons. Louisiana Dept. of Wildlife and Fisheries, Natural Heritage Program, Baton Rouge, LA. 

–––, and C.M. Allen. 1996. Atlas of the vascular flora of Louisiana. Volume II: Dicotyledons, Acanthaceae-Euphorbiaceae. 

Louisiana Dept. of Wildlife and Fisheries, Natural Heritage Program, Baton Rouge, LA.


–––, and C.M. Allen. 1998. Atlas of the vascular flora of Louisiana. Volume III: Dicotyledons, Fabaceae-Zygophyllaceae. 

Louisiana Dept. of Wildlife and Fisheries, Natural Heritage Program, Baton Rouge, LA. 

–––, and P.S. Marx. 1979. Notes on three species of Ophioglossum from North Carolina. Sida 8: 113. 

Thomas, W.W. 1984. The systematics of Rhynchospora section Dichromena. Mem. New York Bot. Garden 37. 

Thomson, P.M., and R.H. Mohlenbrock. 1979. Foliar trichomes of Quercus subgenus Quercus in the eastern United States. J. 

Arnold Arb. 60: 350-366. 

Thompson, S.W., and T.G. Lammers. 1997. Phenetic analysisof morphological variation in the Lobelia cardinalis complex 

(Campanulaceae: Lobelioideae). Systematic Botany 22: 315-331. 

Thorne, R.F. 1992. Classification and geography of the flowering plants. Bot. Review 58: 225-348. 

Threadgill, P.F., and J.M. Baskin. 1978. Swertia caroliniensis or Frasera caroliniensis? Castanea 43: 20-22. 

Threlkeld, S.J., and E.D. Soehren. 2003. Noteworthy collections: Alabama. Castanea 68: 182-183. 

Timmerman-Erskine, M., and R.S. Boyd. 1999. Reproductive biology of the endangered plant Clematis socialis 

(Ranunculaceae). J. Torrey Bot. Soc. 126: 107-116. 

–––, R.R. Dute, and R.S. Boyd. 2002. Morphometric analysis of the Trillium pusillum Michaux complex (Trilliaceae) of the 

southeastern United States. Castanea 67: 109-119. 

Tobe, H., and R.C. Keating. 1985. The morphology and anatomy of Hydrastis (Ranunculales): systematic reevaluation of the 

genus. Bot Mag. Tokyo 98: 291-316. 

Tobe, J.D. 1998. The phylogeny of Magnolia in eastern North America. In D. Hunt, ed. Magnolias and their allies. 

Proceedings of an international symposium, Royal Holloway, University of London, Egham, Surrey, U.K., 12-13 April 

1996. International Dendrological Society and the Magnolia Society. 

Tomb, A.S. 1980. Taxonomy of Lygodesmia (Asteraceae). Systematic Botany Monographs 1: 1-51. 

Tomlinson, P.B. 1986. The biology of trees native to tropical Florida. Published by the author. 480 pp. 

Towe, L.C. 2004. American azaleas. Timber Press, Portland, OR. 146 pp. 

Townsend, J.F., and V. Karaman-Castro. 2006. A new species of Boltonia (Asteraceae) from the Ridge and Valley 

physiographic province, U.S.A. Sida 22: 873-886. 

–––, R. Carter, R.D. Porcher, and P.D. McMillan. 2000. Noteworthy Collections: Georgia and South Carolina. Castanea 65: 

231-232. 

Trapnell D.W., J.L. Hamrick, and D.E. Giannasi. 2004. Genetic variation and species boundaries in Calopogon (Orchidaceae). 


Trauth-Nare, A.E., and R.F.C. Naczi. 1998. Taxonomic status of the varieties of Seneca snakeroot, Polygala senega L. 

(Polygalaceae) [abstract]. Am. J. Bot 85 [supplement]: 163. 

Treiber, M. 1980. Biosystematics of the Arisaema triphyllum complex. Ph.D. dissertation, Univ. of North Carolina-Chapel Hill, 

Department of Botany. 

Triplett, J.K., L.G., Clark, and A.S. Weakley. 2006. Hill cane (Arundinaria appalachiana), a new species of bamboo (Poaceae: 

Bambusoideae) from the Southern Appalachian Mountains. Sida 22: 79-95. 

Tripp, K.E. 1995. Cephalotaxus: the plum yew. Arnoldia 55: 24-39. 

Tryon, R.M. 1955. Selaginella rupestris and its allies. Annals Mo. Bot. Garden 42: 1-99. 

Tucker, A.O., N.H. Dill, T.D. Pizzolato, and R.D. Kral. 1983. Nomenclature, distribution, chromosome numbers, and fruit 

morphology of Oxypolis canbyi and O. filiformis (Apiaceae). Systematic Bot. 8: 299-304. 

Tucker, G.C. 1984. A revision of the genus Kyllinga Rottb. (Cyperaceae) in Mexico and Central America. Rhodora 86: 507- 

538. 

–––. 1986. The genera of the Elatinaceae in the southeastern United States. J. Arnold Arb. 67: 471-483. 

–––. 1987. The genera of Cyperaceae in the southeastern United States. J. Arnold Arb. 68: 361-445. 

–––. 1988. The genera of Bambusoideae (Gramineae) in the southeastern United States. J. Arnold Arb. 69: 239-273. 

–––. 1989. The genera of Commelinaceae in the southeastern United States. J. Arnold Arb. 70: 97-130. 

–––. 1990. The genera of Arundinoideae (Gramineae) in the southeastern United States. J. Arnold Arb. 71: 145-177. 

–––. 1996. The genera of Poöideae (Gramineae) in the southeastern United States. Harvard Papers in Botany 9: 11-90. 

Turner, B.L. 1995a. Synopsis of the genus Onosmodium (Boraginaceae). Phytologia 78: 39-60. 

–––. 1995b. Taxonomic overview of Hedyotis nigricans (Rubiaceae) and closely allied taxa. Phytologia 79: 12-21. [with 

corrected map: Phytologia 80: 142-143] 

–––. 2006. Overview of the genus Baptisia (Leguminosae). Phytologia 88: 253-268. 

–––, and D. Dawson. 1980. Taxonomy of Tetragonotheca (Asteraceae-Heliantheae). Sida 8: 296-303. 

–––, and M.G.Mendenhall. 1993. A revision of Malvaviscus (Malvaceae). Ann. Missouri Bot. Gard. 80: 439-457. 

–––, and M.I. Morris. 1976. Systematics of Palafoxia (Asteraceae: Heleniae). Rhodora 78: 567-628. 

–––, and M. Whalen. 1975. Taxonomic study of Gaillardia pulchella (Asteraceae – Heliantheae). Wrightia 5: 189-192. 

–––, H. Nichols, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, Volume 1. Sida, Botanical Miscellany 

24. 

Turrill, N.L., D.K. Evans, and F.S. Gilliam. 1994. Identification of West Virginia members of the Dentaria complex [D. 

diphylla Michx., D. heterophylla Nutt., and D. laciniata Muhl. ex Willd. (Brassicaceae)] using above-ground vegetative 

characters. Castanea 59: 22-30. 

Tyndall, R.W., B.J. Holt, and G. Lam. 1996. Aeschynomene virginica (L.) BSP. in Maryland. Castanea 61: 86-89. 

Ulmer, T., and J.M. MacDougal. 2004. Passiflora: passionflowers of the world. Timber Press, Portland, OR. 430 pp.


Umber, R.E. 1979. The genus Glandularia (Verbenaceae) in North America. Systematic Bot. 4: 72-102. 

USDA NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, 

LA 70874-4490 USA. Accessed November 2005. 

Urbatsch, L.E. 1972. Systematic study of the Altissimae and Giganteae species groups of the genus Vernonia (Compositae). 

Brittonia 24: 229-238 

–––, R.P. Roberts, and V. Karaman. 2003. Phylogenetic evaluation of Xylothamia, Gundlachia, and related genera (Asteraceae, 

Astereae) based on ETS and ITS NRDNA sequence data. Amer. J. Botany 90: 634-649. 

Uttal, L.J. 1974. The varieties of Spiraea betulifolia. Bull. Torrey Bot. Club 101: 35-36. 

–––. 1985. Virginia's two kinds of blue cohosh. Jeffersonia 16: 20-27. 

–––. 1986a. Once and for all it is Paxistima. Castanea 51: 67-68. 

–––. 1986b. Taxonomic and nomenclatural notes on Vaccinium L. section Cyanococcus (Ericaceae). Sida 11: 397-399. 

–––. 1986c. Updating the genus Vaccinium L. (Ericaceae) in West Virginia. Castanea 51: 197-201. 

–––. 1987. The genus Vaccinium L. (Ericaceae) in Virginia. Castanea 52: 231-255. 

–––. 1991. Notes on Uvularia puberula Michaux (Liliaceae). Castanea 56: 70. 

Valder, P. 1995. Wisterias: a comprehensive guide. Timber Press, Portland, OR. 160 pp. 

van Alstine, N.E., W.H. Moorhead III, A. Belden, Jr., T.J. Rawinski, and J.C. Ludwig. 1996. Recently discovered populations of 

small whorled pogonia (Isotria medeoloides) in Virginia. Banisteria 7: 3-7. 

van Bergen, M.A. 1996. Revision of Catharanthus G. Don; series of revisions of Apocynaceae XLI. Wageningen Agricultural 

University Papers 96-3: 1-46. 

van de Wouw, M., N. Maxted, and B.V. Ford-Lloyd. 2003. A multivariate and cladistic study of Vicia L. ser. Vicia (Fabaceae) 

based on analysis of morphological characters. Plant Syst. Evol. 237: 19-39. 

Van der Bank, M., M.F. Fay, and M.W. Chase. 2002. Molecular phylogenetics of Thymelaeaceae, with particular reference to 

African and Australian genera. Taxon 51: 329-339. 

van der Werff, H., and H.G. Richter. 1996. Toward an improved classification of Lauraceae. Ann. Mo. Bot. Garden 83: 409- 

418. 

van Gelderen, C.J., and D.M. van Gelderen. 2004. Encyclopedia of Hydrangeas. Timber Press, Portland, OR. 279 pp. 

van Gelderen, D.M., P.C. de Jong, H.J. Oterdoom. 1994. Maples of the world. Timber Press, Portland OR. 458 pp. 

van Welzen, P.C. 1981. A taxonomic revision of the genus Arthraxon Beauv. Blumea 27: 255-300. 

Vander Kloet, S.P. 1977. Potential and actual gene exchange among three sympatric species of Vaccinium § Cyanococcus in 

Highlands County, Florida. Can. J. Bot 55: 2668-2672. 

–––––. 1978a. Systematics, distribution, and nomenclature of the polymorphic Vaccinium angustifolium. Rhodora 80: 358-376. 

–––. 1978b. The taxonomic status of Vaccinium pallidum, the hillside blueberries including Vaccinium vacillans. Can J. Bot. 

56: 1559-1574. 

–––. 1980. The taxonomy of the highbush blueberry, Vaccinium corymbosum. Can. J. Bot. 58: 1187-1201. 

–––. 1982. A note on the occurrence of root-shoots in Vaccinium corymbosum L. Rhodora 84: 447-450. 

–––. 1983a. The taxonomy of Vaccinium § Oxycoccus. Rhodora 85: 1-43. 

–––. 1983b. The taxonomy of Vaccinium § Cyanococcus: a summation. Can J. Bot. 61: 256-266. 

–––. 1988. The genus Vaccinium in North America. Publication 1828, Research Branch, Agriculture Canada, Ottawa. 

–––, and I.V. Hall. 1981. The biological flora of Canada. 2. Vaccinium myrtilloides Michx., velvet-leaf blueberry. Can. Field- 

Nat. 95: 329-345. 

Vega, A.S. 2000. Revisión taxonómica de las especies americanas del género Bothriochloa (Poaceae: Panicoideae: 

Andropogoneae). Darwiniana 38: 127-186. 

Veldkamp, J.F. 1999. A revision of Chrysopogon Trin. including Vetiveria Bory (Poaceae) in Thailand and Malesia with notes 

on some other species from Africa and Australia. Austrobaileya 5: 503-533. 

–––, R. de Koning, and M.S.M. Sosef. 1986. Generic delimitation of Rottboellia and related genera (Gramineae). Blumea 31: 

281-307. 

Verdcourt, B. 2004. The variation of Sida rhombifolia L. (Malvaceae) in East Africa. Kew Bull.59: 233-239. 

Vincent, M.A. 2004. Spread of Fatoua villosa (mulberry weed; Moraceae) in North America. J. Ky. Acad. Sci. 65: 67-75. 

Virginia Botanical Associates. 2006. Digital Atlas of the Virginia Flora. http://www.biol.vt.edu/digital_atlas/. Accessed 9 

February 2006. 

Vitt, P., and C.S. Campbell. 1997. Reproductive biology of Isotria medeoloides (Orchidaceae). Rhodora 99: 56-63. 

Vogelmann, J.E. 1985. Crossing relationships among North American and eastern Asian populations of Agastache sect. 

Agastache (Labiatae). Systematic Bot. 10: 445-452. 

von Balthazar, M., P.K. Endress, and Y.-L. Qiu. 2000. Phylogenetic relationships in Buxaceae based on internal transcribed 

spacers and plastid ndhF sequences. Int. J. Plant Sci. 161: 785-792. 

Voss, E.G. 1972. Michigan flora: a guide to the identification and occurrence of the native and naturalized seed-plants of the 

state. Part I, gymnosperms and monocots. Cranbrook Institute of Science Bulletin No. 55 and Univ. of Mich. Herbarium, 

Bloomfield Hills, MI. 488 pp. 

–––. 1985. Michigan flora: a guide to the identification and occurrence of the native and naturalized seed-plants of the state. 

Part II, dicots (Saururaceae-Cornaceae). Cranbrook Institute of Science Bulletin No. 59 and Univ. of Mich. Herbarium, Ann 

Arbor, MI. 724 pp.


–––. 1996. Michigan flora: a guide to the identification and occurrence of the native and naturalized seed-plants of the state. 

Part III, dicots (Pyrolaceae-Compositae). Cranbrook Institute of Science Bulletin No. 61 and Univ. of Mich. Herbarium, 

Ann Arbor, MI. 622 pp. 

Vuilleumier, B.S. 1969. The tribe Mutisieae (Compositae) in the southeastern United States. J. Arnold Arb. 50: 620-625. 

Wagenknecht, B.L. 1960. Revision of Heterotheca, section Heterotheca (Compositae). Rhodora 62: 61-76, 97-107. 

Wagner, W.H., Jr. 1977. Systematic implications of the Psilotaceae. Brittonia 29: 54-63. 

–––. 1992. Hiemobotrychium, a new section of Botrychium subgenus Sceptridium from the southeastern United States. Novon 

2: 267-268. 

–––, J.M. Beitel, and R.C. Moran. 1989. Lycopodium hickeyi: a new species of North American clubmoss. Amer. Fern J. 79: 

119-121. 

–––, and J.M. Beitel. 1992. Generic classification of modern North American Lycopodiaceae. Ann. Mo. Bot. Gard. 79: 676- 

686. 

–––, E.M. Bush, C.R. Werth, and R.L. Bartgis. 1991. First records of alternate-leaved spleenwort, Asplenium ×alternifolium, in 

the New World. Castanea 56: 128-134, 

–––, F.S. Wagner, C.N. Miller, Jr., and D.H. Wagner. 1978. New observations on the royal fern hybrid Osmunda ×ruggii. 

Rhodora 80: 92-106. 

Wagner, W.L., and H. Robinson. 2001. Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing 

their natural limits and a synopsis. Brittonia 53: 539-561. 

–––, and P. Hoch. 2000. Proposal to reject the name Gaura mollis (Onagraceae). Taxon 49: 101-102. 

Wagstaff, S.J., and R.G. Olmstead. 1997. Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Systematic 

Bot. 22: 165-179. 

Wahl, H.A. 1954. A preliminary study of the genus Chenopodium in North America. Bartonia 27: 1-46. 

Walker, J.B., K.J. Sytsma, J. Treutlein, and M. Wink. 2004. Salvia (Lamiaceae) is not monophyletic: implications for the 

systematics, radiation, and ecological specializations of Salvia and tribe Mentheae. Amer. J. Bot. 91: 1115-1125. 

Wallace, L.A., and M.A. Case. 2000. Contrasting allozyme diversity between northern and southern populations of 

Cypripedium parviflorum (Orchidaceae): implications for Pleistocene refugia and taxonomic boundaries. Syst. Bot. 25: 

281-296. 

Walters, S.M., and D.A. Webb. 1972. Veronica. In T.G. Tutin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H. Valentine, 

S.M. Walters, and D.A. Webb. Flora Europaea. Volume 3. Cambridge, England. 

Wang, Y., P.W. Fritsch, S. Shi, F. Almeda, B.C. Cruz, and L.M. Kelly. 2004. Phylogeny and infrageneric classification of 

Symplocos (Symplocaceae) inferred from DNA sequence data. Amer. J. Bot. 91: 1901-1914. 

Ward, A.B., and C.N. Horn. 1998. A status survey of Dirca palustris L. (Leatherwood, Thymelaeaceae) in South Carolina. 

Castanea 63: 165-173. 

Ward, D.B. 1968. Contributions to the flora of Florida – 3, Evolvulus (Convolvulaceae). Castanea 33: 76-79. 

–––. 1974. Contributions to the flora of Florida – 6, Vaccinium (Ericaceae). Castanea 39: 191-205. 

–––. 1977a. Keys to the flora of Florida – 2, Paronychia (Caryophyllaceae. Phytologia 35: 414-418. 

–––. 1977b. Corrections in Paronychia (Caryophyllaceae). Phytologia 37: 449-450. 

–––. 1977c. Keys to the flora of Florida – 5, Dioscoreaceae. Phytologia 38: 151-154. 

–––. 1998. Pueraria montana: the correct scientific name of the kudzu. Castanea 63: 76-77. 

–––. 2001. New combinations in the Florida flora. Novon 11: 360-365. 

–––. 2004a. Keys to the flora of Florida – 9, Oxalis (Oxalidaceae). Phytologia 86: 32-41. 

–––. 2004b. Acer floridanum: the correct scientific name of the Florida maple. Castanea 69: 230-233. 

–––. 2004c. New combinations in the Florida flora II. Novon 14: 365-371. 

–––. 2004d. Keys to the flora of Florida – 11, Elytraria (Acanthaceae). Phytologia 86: 200-202. 

–––. 2005a. Rediscovery of Sisyrinchium corymbosum Bicknell (Iridaceae), lost for one hundred years. Castanea 70: 155-157. 

–––. 2005b. A case of disputed orthography: is it Echinchloa colona; or is it Echinochloa colonum (Gramineae). Sida 21: 2171- 

2183. 

–––. 2006a. A nomenclatural history of southeastern filiferous Yucca, with selection of a neotype for Y. flaccida. Castanea 71: 

80-84. 

–––. 2006b. Keys to the flora of Florida – 13, Vitis (Vitaceae). Phytologia 88: 216-223. 

–––, and A.K. Gholson. 1987. The hidden abundance of Lepuropetalon spathulatum (Saxifragaceae) and its first reported 

occurrence in Florida. Castanea 52: 59-68. 

–––, and D.W. Hall. 2004. Keys to the flora of Florida – 10, Galactia (Leguminosae). Phytologia 86: 65-74. 

–––, and W.K. Taylor. 1999. Discovery of tree-form gopher apple (Licania michauxii Prance), with implication of an arboreous 

ancestor. Castanea 64: 263-265. 

Ware, D.M.E. 1973. Floristic survey of the Thompson River watershed. Castanea 38: 349-378. 

–––. 1983. Genetic fruit polymorphism in North American Valerianella (Valerianaceae) and its taxonomic implications. 


Ware, S. 1967. A new Talinum (Portulaceae) from the cedar glades of Middle Tennessee. Rhodora 69: 466-475. 

–––. 1992. Where are all the hickories in the Piedmont oak-hickory forest? Castanea 57: 4-12. 

–––, and G. Pinion. 1990. Substrate adaptation in rock outcrop plants: eastern United States Talinum (Portulacaceae). Bull. 

Torrey Bot. Club 117: 284-290.


Warners, D.P., and D.C. Laughlin. 1999. Evidence for a species-level distinction of two co-occurring asters: Aster puniceus L. 

and Aster firmus. 

Warnock, M.J. 1995. A taxonomic conspectus of North American Delphinium. Phytologia 78: 73-101. 

Wasshausen, D.C. 1998. Acanthaceae of the southeastern United States. Castanea 63: 99-116. 

Waterway, M.J. 1986. A reevaluation of Lycopodium porophilum and its relationship to L. lucidulum (Lycopodiaceae). 


Watkins, J.E., Jr., and D.R. Farrar. 2002. A new name for an old fern from north Alabama. Amer. Fern J. 92: 171-178. 

–––, and D.R. Farrar. 2005. Origin and taxonomic affinities of Thelypteris (subgen. Stegnogramma) burksiorum 

(Thelypteridaceae). Brittonia 57: 183-201. 

Watson, L.E., W.J. Elisens, and J.R. Estes. 1991. Electrophoretic and cytogenetic evidence for allopolyploid origin of 

Marshallia mohrii (Asteraceae). Am. J. Bot. 78: 408-416. 

–––, and J.R. Estes. 1990. Biosystematic and phenetic analysis of Marshallia (Asteraceae). Systematic Bot. 15: 403-414. 

–––, R.K. Jansen, and J.R. Estes. 1991. Tribal placement of Marshallia (Asteraceae) using chloroplast DNA restriction site 

mapping. Am. J. Bot. 78: 1028-1035. 

–––, A.B. Kornkven, C.R. Miller, J.R. Allison, N.B. McCarty, and M.M. Unwin. 2002. Morphometric and genetic variation in 

Eriocaulon koernickianum Van Heurck & Muller-Argoviensis (Eriocaulaceae): a disjunct plant species of the southeastern 

United States. Castanea 67: 416-426. 

Weakley, A.E. 2002. Evolutionary relationships within the genus Philadelphus L. (Hydrangeaceae). Master's thesis, Biology 

Dept., Univ. of North Carolina, Chapel Hill, NC. 

Weakley, A.S. 1990. Natural Heritage Program list of the rare plants of North Carolina. North Carolina Natural Heritage 

Program, Raleigh, NC. 

–––. 2005. Flora of the Carolinas, Virginia, and Georgia, working draft of June 2005. University of North Carolina Herbarium, 

N.C. Botanical Garden, Chapel Hill, NC. http://www.herbarium.unc.edu/. Accessed 10 December 2005. 

–––, and G.L. Nesom. 2004. A new species of Ptilimnium (Apiaceae) from the Atlantic coast. Sida 21: 743-752. 

–––, and P.M. Peterson. 1998. Taxonomy of the Sporobolus floridanus complex (Poaceae: Sporobolinae). Sida 18: 247-270. 

Weatherwax, P. 1934. Flowering and seed production in Amphicarpon floridanum. Bull. Torrey Bot. Club 61: 211-215. 

Weaver, R.E., Jr., and L. Rüdenberg. 1975. Cytytaxonomic notes on some Gentianaceae. J. Arnold Arb. 56: 211-222. 

Webb. 1980. {Hypericum} 

Webber, J.M. and P.W. Ball. 1980. Introgression in Canadian populations of Lycopus americanus Muhl. and L. europaeus L. 

(Labiatae). Rhodora 82: 281-304. 

–––, and P.W. Ball. 1984. The taxonomy of the Carex rosea group (section Phaestoglochin) in Canada. Can. J. Bot. 62: 2058- 

2073. 

Weber, W.A. 1995. New names and combinations, principally in the Rocky Mountain flora – IX. Phytologia 79: 65-67. 

Webster, G.L. 1967. The genera of Euphorbiaceae in the southeastern United States. J. Arnold Arb. 48: 303-430. 

–––. 1970. A revision of Phyllanthus (Euphorbiaceae) in the continental United States. Brittonia 22: 44-76. 

–––. 1992. Realignments in American Croton (Euphorbiaceae). Novon 2: 269-273. 

–––. 1993. A provisional synopsis of the sections of the genus Croton (Euphorbiaceae). Taxon 42: 793-823. 

–––. 1994. Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Ann. Missouri Bot. Gard. 81: 33-144. 

Webster, R.D. 1980. Distribution records for Digitaria bicornis in eastern United States. Sida 8: 352-353. 

–––. 1987. Taxonomy of Digitaria section Digitaria in North America (Poaceae: Paniceae). Sida 12: 209-222. 

–––. 1988. Genera of the North American Paniceae (Poaceae: Panicoideae). Systematic Bot. 13: 576-609. 

–––. 1980. Distribution records for Digitaria bicornis in eastern United States. Sida 8: 352-353. 

–––. 1992. Character significance and generic similarities in the Paniceae (Poaceae: Panicoideae). Sida 15: 185-213. 

–––. 1993. Nomenclature of Setaria (Poaceae: Paniceae). Sida 15: 447-489. 

–––. 1995. Nomenclatural changes in Setaria and Paspalidium (Poaceae: Paniceae). Sida 16: 439-446. 

–––, J.H. Kirkbride, and J.V. Reyna. 1989. New World genera of the Paniceae (Poaceae: Panicoideae). Sida 13: 393-417. 

–––, and R.B. Shaw. 1995. Taxonomy of the native North American species of Saccharum (Poaceae: Andropogoneae). Sida 

16: 551-580. 

Weckman, T.J., J.E. Weckman, R.L. Thompson, and D.L. White. 2002. Noteworthy Collections: Kentucky. New records and a 

summary of naturalized Viburnum taxa in Kentucky. Castanea 67: 104-106. 

Weigant, P.L. 2002. Distribution of Aletris in North America. J. North Carolina Academy of Science 118: 44-49. 

Weigend, M., O. Mohr, and T.J. Motley. 2002. Phylogeny and classification of the genus Ribes (Grossulariaceae) based on 5S- 

NTS sequences and morphological and anatomical data. Bot. Jahrb. Syst. 124: 163-182. 

Weldy, T.W., H.T. Mlodozeniec, L.E. Wallace, and M.A. Case. 1996. The current status of Cypripedium kentuckiense 

(Orchidaceae) including a morphological analysis of a newly discovered population in eastern Virginia. Sida 17: 423-435. 

Weller, S.G. 1970. A preliminary report on the varieties of Maianthemum canadense in northern Michgan. Michigan Botanist 

9: 48-52. 

Wells, E.F. 1984. A revision of the genus Heuchera (Saxifragaceae) in eastern North America. Systematic Bot. Monographs 3: 

45-121. 

Wen, J. 1993. Generic delimitation of Aralia (Araliaceae). Brittonia 45: 47-55. 

–––. 1998. Systematics and biogeography of the Aralia -- Panax complex (Araliaceae) [abstract]. Am. J. Bot. 85 [supplement]: 

166.


–––, and R.K. Jansen. 1995. Morphological and molecular comparisons of Campsis grandiflora and C. radicans 

(Bignoniaceae), and eastern Asian and eastern North American vicariad pair. Pl. Syst. Evol. 196: 173-183. 

–––, and S. Shi. 1999. A phylogenetic and biogeographic study of Hamamelis (Hamamelidaceae), an eastern Asian and eastern 

North American disjunct genus. Biochemical Systematics and Ecology 27: 55-66. 

–––, S. Shi, R.K. Jansen, and E.A. Zimmer. 1998. Phylogeny and biogeography of Aralia sect. Aralia (Araliaceae). Am. J. Bot. 

85: 866-875. 

–––, and T.F. Stuessy. 1993. The phylogeny and biogeography of Nyssa (Cornaceae). Systematic Bot. 18: 68-79. 

–––, and E.A. Zimmer. 1996. Phylogeny and biogeography of Panax L. (the ginseng genus, Araliaceae): inferences from ITS 

sequences of nuclear ribosomal DNA. Molecular Phylogenetics and Evolution 6: 167-177. 

–––, P.P. Lowry II, J.L. Walck, and K.-O. Yoo. 2002. Phylogenetic and biogeographic diversification in Osmorhiza (Apiaceae). 

Ann. Missouri Bot. Gard. 89: 414–428. 

Werth, C.R. 1991. Isozyme studies on the Dryopteris "spinulosa" complex, I: The origin of the log fern Dryopteris celsa. 


–––, Linghe Zeng, and W.V. Baird. 1997. An enigmatic tetraploid Eleusine (Gramineae) discovered in South Carolina. ASB 

Bulletin 44: 97. 

Wherry, E.T. 1929. Acidity relations of the Sarracenias. J. Washington Acad. Sci. 19: 379-390. 

–––. 1933. The Appalachian relative of Sarracenia flava. Bartonia 15: 7-8. 

–––. 1940. A novelty in the genus Tiarella (Saxifragaceae). Notulae Naturae (Academy of Natural Sciences of Philadelphia) 

42: 1-4. 

–––. 1946. A key to the eastern North American lilies. Bartonia 24: 5-8. 

–––. 1949. Further observations on eastern Tiarellas. Bartonia 25: 70. 

–––. 1955. The genus Phlox. Morris Arboretum Monographs III. Philadelphia, PA. 

–––. 1972. Notes on the Sarracenia subspecies. Castanea 37: 146-147. 

Whetstone, R.D. 1983. The Sterculiaceae in the flora of the southeastern United States. Sida 10: 15-23. 

Whitcher, I.N., and Jun Wen. 2001. Phylogeny and biogeography of Corylus (Betulaceae): inferences from ITS sequences. 


Whitehead, D.R. 1963. "Northern" elements in the Pleistocene flora of the Southeast. Ecology 44: 403-406. 

Whittemore, A.T. 2003. Noteworthy collections: District of Columbia. Castanea 68: 261. 

–––. 2004. Sawtooth oak (Quercus acutissima, Fagaceae) in North America. Sida 21: 447-454. 

–––, and K.C. Nixon. 2005. Proposal to reject the name Quercus prinus (Fagaceae). Taxon 54: 213-214. 

Widrlechner, M.P. 1998. The genus Rubus L. in Iowa. Castanea 63: 415-465. 

Wieboldt, T.F. 1987. The shale barren endemic, Arabis serotina (Brassicaceae). Sida 12: 381-389. 

–––. 1992. Cardamine micranthera Rollins, small-anthered bittercress in Patrick County: new to the Virginia flora. Banisteria 

1: 16-17. 

–––. 1995. A new station for smooth cliffbrake, Pellaea glabella, (Pteridaceae) on masonry walls. Banisteria 6: 23-24. 

–––, and S. Bentley. 1982. Cheilanthes feei new to Virginia. Amer. Fern J. 72: 76-78. 

–––, and J.S. Semple. 2003. Solidago faucibus (Asteraceae: Astereae), a new mesic forest goldenrod from the Appalachian 

mountains. Sida 20: 1595-1603. 

–––, G.P. Fleming, J.C. Ludwig, and F.C. Huber. 1998. Noteworthy collections: Virginia. Castanea 63: 82-91. 

Wiegand, K.M. 1928. Aster lateriflorus and some of its relatives. Rhodora 30: 161-179. 

Wiegleb, G., and Z. Kaplan. 1998. An account of the species of Potamogeton L. (Potamogetonaceae). Folia Geobotanica 33: 

241-316. 

Wiegrefe, S.J., K.J. Sytsma, and R.P. Guries. 1994. Phylogeny of elms (Ulmus, Ulmaceae): molecular evidence for a sectional 

classification. Systematic Bot. 19: 590-612. 

Wiggins, I.L. 1932. Plants recently established in the San Francisco bay region. Torreya 32: 3-4. 

Wikström, N., and P. Kenrick. 2000. Relationships of Lycopodium and Lycopodiella based on combined plastid rbcL gene and 

trnL intron sequence data. Systematic Bot. 25: 495-510. 

–––, and P. Kenrick. 2001. Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by 

use of nonparametric smoothing. Molecular Phylogenetics and Evolution 19: 177-186. 

Wilbur, R.L. 1963. The leguminous plants of North Carolina. North Carolina Agricultural Experiment Station Tech. Bull. No. 

151, Raleigh, N.C. 294 pp. 

–––. 1964. A revision of the dwarf species of Amorpha (Leguminosae). J. Elisha Mitchell Sci. Soc. 80: 51-65. 

–––. 1968. The status of Hedyotis procumbens var. hirsuta (Rubiaceae). Rhodora 70: 306-311. 

–––. 1970a. Taxonomic and nomenclatural observations on the eastern North American genus Asimina (Annonaceae). J. Elisha 

Mitchell Sci. Soc. 86: 88-96. 

–––. 1970b. Infraspecific classification in the Carolina flora. Rhodora 72: 51-65. 

–––. 1975. A revision of the North American genus Amorpha (Leguminosae-Psoraleae). Rhodora 77: 337-409. 

–––. 1988a. What do we know about Diamorpha smallii (Crassulaceae), "one of the better-known taxa in the Southeastern 

flora?" Sida 13: 1-16. 

–––. 1988b. The authority for Lepuropetalon spathulatum (Saxifragaceae). Castanea 53: 306-308. 

–––. 1988c. The correct scientific name of the pale, yellow, or white gentian of the eastern United States. Sida 13: 161-165. 

–––. 1994. The Myricaceae of the United States and Canada: genera, subgenera, and series. Sida 16: 93-107.


–––. 2002. The identity and history of Myrica caroliniensis (Myricaceae). Rhodora 104: 31-41. 

–––. 2003. What is the correct name for the bristly greenbrier? Rhodora 105: 250-259. 

–––. 2004. The subgeneric nomenclature for the herbaceous-stemmed Smilax species (Smilacaceae) of North America. 

Brittonia 56: 166-168. 

–––, and S. Bloodworth. 2004. Notes on the box huckleberry, Gaylussacia brachycera (Ericaceae), and its unexpected presence 

in North Carolina. Rhodora 106: 371-377. 

–––, and M.K. Whitson. 2005. The identity of Riddell’s seven validly published but over-looked pteridophytic binomials. 

Amer. Fern J. 95: 160-169. 

–––, and H.S. Daoud. 1961. The genus Lechea (Cistaceae) in the southeastern United States. Rhodora 63: 103-118. 

–––, and H.S. Daoud. 1964. The genus Helianthemum (Cistaceae) in the southeastern United States. J. Elisha Mitchell Sci. Soc. 

70: 38-43. 

–––, and C.H. Racine. 1971. The genus Leiophyllum (Ericaceae): morphological variation, distribution, and classification. J. 

Elisha Mitchell Sci. Soc. 87: 222-226 

Wilce, J.H. 1972. Lycopod spores, I. General spore patterns and the generic segregates of Lycopodium. Amer. Fern J. 62: 65- 

79. 

Williams, C. 1999. André Michaux and the discovery of Magnolia macrophylla in North Carolina. Castanea 64: 1-13. 

Williams, J.G., and A.E. Williams. 1983. Field guide to orchids of North America from Alaska, Greenland, and the Arctic south 

to the Mexican border. Universe Books, New York. 

Williges, K.A., and C.S. Loftin. 1995. Noteworthy plant species from the Okefenokee Swamp, Georgia. Sida 16: 775-780. 

Wilson, C.A. 2004. Phylogeny of Iris based on chloroplast matK gene and trnK intron sequence data. Molecular Phylogenetics 

and Evolution 33: 402-412. 

Wilson, J.S. 1965. Variation of three taxonomic complexes of the genus Cornus in eastern United States. Trans. Kans. Acad. 

Sci. 67: 747-817. 

Wilson, K.A. 1960a. The genera of Hydrophyllaceae and Polemoniaceae in the southeastern United States. J. Arnold Arb. 41: 

197-212. 

–––. 1960b. The genera of Convolvulaceae in the southeastern United States. J. Arnold Arb. 41: 298-317. 

Wilson, P. 1932. Talinum. In P.A. Rydberg, Portulacaceae. North American Flora, volume 21, part 4. New York Botanical 

garden, New York, NY. 

Windham, M.D. 1987. Argyrochosma, a new genus of Cheilanthoid ferns. Amer. Fern J. 77: 37-41. 

Windler, D.R. 1974. A systematic treatment of the native unifoliolate Crotalarias of North America (Leguminosae). Rhodora 

76: 151-204. 

Winkworth, R.C., and M.J. Donoghue. 2005. Viburnum phylogeny based on combined molecular data: implications for 

taxonomy and biogeography. Amer. J. Botany 92: 653-666. 

Winter, K., M.R. Schmitt, and G.E. Edwards. 1982. Microstegium vimineum, a shade adapted C4 grass. Plant Science Letters 

24: 311-318. 

Wipff, J.K. 1996a. Nomenclatural combinations in Schizachyrium (Poaceae: Androponeae). Phytologia 80: 35-39. 

–––. 1996b. Nomenclatural combinations in Digitaria (Poaceae: Paniceae). Phytologia 80: 348-349. 

–––. 1996c. Nomenclatural combinations in the Andropogon gerardii complex (Poaceae: Andropogoneae). Phytologia 80: 

343-347. 

–––, and S.L. Hatch. 1994. A systematic sutudy of Digitaria sect. Pennatae (Poaceae: Paniceae) in the New World. Systematic 

Bot. 19: 613-627. 

–––, and S.D. Jones. 1995. Nomenclatural combination in Poaceae. Phytologia 78: 244-245. 

–––, R.I. Lonard, S.D. Jones, and S.L. Hatch. 1993. The genus Urochloa (Poaceae: Paniceae) in Texas, including one 

previously unreported species for the state. Sida 15: 405-414. 

–––, and B.S. Rector. 1993. Rottboellia cochinchinensis (Poaceae: Andropogoneae) new to Texas. Sida 15: 419-424. 

Wise, D.A., and M.Y. Menzel. 1971. Genetic affinities of the North American species of Hibiscus sect. Trionum. Brittonia 23: 

425-437. 

Wiser, S.K. 1991. Two North Carolina locations for Calamagrostis cainii Hitch., previously considered endemic to Mt. 

LeConte, Tennessee. Castanea 56:147-149. 

Wofford, B.E. 1976. The taxonomic status of Ageratina luciae-brauniae (Fern.) King & H. Robins. Phytologia 33: 369-371. 

–––. 1983. A new Lindera (Lauraceae) from North America. J. Arnold Arb. 64: 325-331. 

–––. 1989. Guide to the vascular plants of the Blue Ridge. Univ. of Georgia Press, Athens, GA. 

–––. 2006. A new species of Stenanthium (Melianthaceae) from Tennessee, U.S.A. Sida 22: 447-459. 

–––, and E.W. Chester. 2002. Guide to the trees, shrubs, and woody vines of Tennessee. Univ. of Tennessee Press, Knoxville. 

–––, and R.L. Jones. 1988. Fimbristylis perpusilla Harper (Cyperaceae) from the Cumberland Plateau of Tennessee. Castanea 

53: 299-302. 

–––, and R. Kral. 1993. Checklist of the vascular plants of Tennessee. Sida, Bot. Misc. 10: 1-66. 

–––, J. de Paula-Souza, A.S. Weakley, and T.E. Govus. 2004. The rediscovery of the South American Hybanthus parviflorus 

(Violaceae) in North America. Sida 21: 1209-1214. 

Wojciechowski, M.F., M. Lavin, and M.J. Sanderson. 2004. A phylogeny of legumes (Leguminosae) based on analysis of the 

plastid matK gene resolves many well-supported subclades within the family. Amer. J. Bot. 91: 1846-1862. 

Wood, C.E., Jr. 1949. The American barbistyled species of Tephrosia (Leguminosae). Contr. Gray Herbarium 170: 193-384.


–––. 1958. The genera of the woody Ranales in the southeastern United States. J. Arnold Arb. 39: 296-346. 

–––. 1960. The genera of Sarraceniaceae and Droseraceae in the southeastern United States. J. Arnold Arb. 41: 152-163. 

–––. 1961. The genera of Ericaceae in the southeastern United States. J. Arnold Arb. 42: 10-80. 

–––. 1966. On the identity of Drosera brevifolia. J. Arnold Arb. 47: 89-99. 

–––. 1971. The Saururaceae in the southeastern United States. J. Arnold Arb. 52: 479-485. 

–––. 1975. The Balsaminaceae in the southeastern United States. J. Arnold Arb. 56: 413-426. 

–––. 1983a. The genera of Menyanthaceae in the southeastern United States. J. Arnold Arb. 64: 431-445. 

–––. 1983b. The genera of Burmanniaceae in the southeastern United States. J. Arnold Arb. 64: 293-307. 

–––, and W.P. Adams. 1976. The genera of Guttiferae (Clusiaceae) in the southeastern United States. J. Arnold Arb. 57: 74-90. 

–––, and R.K. Godfrey. 1957. Pinguicula (Lentibulariaceae) in the southeastern United States. Rhodora 59: 217-230. 

–––, and R.E. Weaver, Jr. 1982. The genera of Gentianaceae in the southeastern United States. J. Arnold Arb. 63: 441-487. 

Woodland, D.W. 1982. Biosystematics of the perennial North American taxa of Urtica. II. Taxonomy. Systematic Bot. 7: 282- 

290. 

–––, I.J. Bassett, C. Crompton, and S. Forget. 1982. Biosystematics of the perennial North American taxa of Urtica. I. 

Chromosome number, hybridization, and palynology. Systematic Bot. 7: 269-281. 

Woods, K., K.W. Hilu, J.H. Wiersema, and T. Borsch. 2005a. Pattern of variation and systematics of Nymphaea odorata: I. 

Evidence from morphology and inter-simple sequence repeats (ISSRs). Systematic Botany 30: 471-480. 

–––, K.W. Hilu, T. Borsch, and J.H. Wiersema. 2005b. Pattern of variation and systematics of Nymphaea odorata: II. Sequence 

information from ITS and trnL-trnF. Systematic Botany 30: 481-493. 

Woods, M. 2005. A revision of the North American species of Apios (Fabaceae). Castanea 70: 85-100. 

–––, A.R. Diamond, Jr., and D.N. Searcy. 2003. Noteworthy collections: Alabama. Castanea 68: 91-92. 

Woodson, R.E., Jr. 1928. Studies in the Apocynaceae. III. A monograph of the genus Amsonia. Ann. Missouri Bot. Garden 

15: 379-434. 

–––. 1930. Studies in the Apocynaceae. I. A critical study of the Apocynoideae (with special reference to the genus 

Apocynum). Ann. Missouri Bot. Garden 17: 1-212. 

–––. 1954. The North American species of Asclepias L. Ann. Missouri Bot. Garden 41: 1-211. 

Wooten, J.W., and A.F. Clewell. 1971. Fleischmannia and Conoclinium (Compositae, Eupatorieae) in eastern North America. 

Rhodora 73: 566-574. 

Wujek, D.E., and F.J. Menapace. 1986. Taxonomy of Carex section Folliculatae using achene morphology. Rhodora 88: 399- 

403. 

Wunderlin, R.P. 1981. Polygonella polygama (Polygonaceae) in Florida. Florida Sci. 44: 78-80. 

–––. 1982. Guide to the vascular plants of central Florida. University Presses of Florida, Tampa, FL. 472 pp. 

–––, and B.F. Hansen. 2003. Guide to the vascular plants of Florida, second edition. University Press of Florida, Gainesville, 

FL. 

–––, and B.F. Hansen. 2005. Atlas of Florida vascular plants . [S.M. Landry and K.N. 

Campbell (application development), Florida Center for Community Design and Research.] Institute for Systematic Botany, 

University of South Florida, Tampa. Accessed 28 November 2005. 

–––, B.F. Hansen, and D.W. Hall. 1985. The vascular flora of central Florida: taxonomic and nomenclatural changes, additional 

taxa. Sida 11: 232-244. 

–––, B.F. Hansen, K.R. Delaney, M. Nee, and J.J. Mullahey. 1993. Solanum viarum and S. tampicense (Solanaceae): two 

weedy species new to Florida and the United States. Sida 15: 605-611. 

–––, and J.E. Poppleton. 1977. The Florida species of Ilex (Aquifoliaceae). Florida Sci. 40: 7-21. 

Wurdack, J.J., and R. Kral. 1982. The genera of Melastomataceae in the southeastern United States. J. Arnold Arb. 63: 429- 

439. 

Wurdack, K.J., P. Hoffmann, and M.J. Chase. 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae 

(Euphorbiaceae senso stricto) using plastid rbcL and trnL-F DNA sequences. Amer. J. Bot. 92: 1397-1420. 

–––, P. Hoffmann, R. Samuel, A. de Bruijn, M. van der Bank, and M.W. Chase. 2004. Molecular phylogenetic analysis of of 

Phyllanthaceae (Phyllanthoideae pro parte, Euphorbiaceae sensu lato) using plastid rbcL DNA sequences. Amer. J. Bot. 91: 

1882-1900. 

Wyatt, R. 1983. Reproductive biology of the granite outcrop endemic Sedum pusillum (Crassulaceae). Systematic Bot. 8: 24- 

28. 

–––. 1985. Aesculus parviflora in South Carolina: phytogeographical implications. Bull. Torrey Bot. Club 112: 194-195. 

–––. 1996. More on the southward spread of common milkweed, Asclepias syriaca L. Bull. Torrey Bot. Club 123: 68-69. 

–––, S.B. Broyles, J.L. Hamrick, A. Stoneburner. 1993. Systematic relationships within Gelsemium (Loganiaceae): evidence 

from isozymes and cladistics. Systematic Bot. 18: 345-355. 

–––, A. Stoneburner, S.B. Broyles, and J.R. Allison. 1993. Range extension southward in the common milkweed, Asclepias 

syriaca L. Bull. Torrey Bot. Club 120: 177-179. 

Wynne, F.E. 1944. Drosera in eastern North America. Bull. Torrey Bot. Club 71: 166-174. 

Xiang, Chunsheng, and J.C. Semple. 1996. Molecular systematic study of Aster sensu lato and related genera (Asteraceae: 

Astereae) based on chloroplast DNA restriction site analyses and mainly North American taxa. Pp. 393-423 in D.J.N. Hind 

and H.J. Beentje (eds.) Compositae: Systematics. Proceedings of the International Compositae Conference, Kew, 1994, 

vol. 1.


Xiang, Q.-Y. (Jenny), D.E. Soltis, and P.S. Soltis. 1998. Phylogenetic relationships of Cornaceae and close relatives inferred 

from matK and rbcL sequences. Amer. J. Bot. 85: 285-297. 

Xiang, Q.-Y. (Jenny), M.L. Moody, D.E. Soltis, C.z. Fan, and P.S. Soltis. 2002. Relationships within Cornales and 

circumscription of Cornaceae – matK and rbcL sequence data and effects of outgroups and long branches. Molecular 

Phylogenetics and Evolution 24: 35-57. 

Xiang, Q.-Y. (Jenny), D.T. Thomas, W. Zhang, S.R. Manchester, and Z. Murrell. 2006. Species level phylogeny of the genus 

Cornus (Cornaceae) based on molecular and morphological evidence – implications for taxonomy and Tertiary 

intercontinental migration. Taxon 55: 9-30. 

Yamashita, J., and M.N. Tamura. 2000. Molecular phylogeny of the Convallariaceae (Asparagales). In: K.L. Wilson & D. A. 

Morrison, eds., Monocots: systematics and evolution. CSIRO, Melbourne. 

Yang, S.-X., J.-B. Yang, L.-G. Lei, D.-Z. Li, H. Yoshino, and T. Ikeda. 2004. Reassessing the relationships between Gordonia 

and Polyspora (Theaceae) based on the combined analyses of molecular data from the nuclear, plastid, and mitochondrial 

genomes. Plant Syst. Evol. 248: 45-55. 

Yatabe, Y., H. Nishida, and N. Murakami. 1999. Phylogeny of Osmundaceae inferred from rbcL nucleotide sequences and 

comparison to the fossil evidences. J. Plant Res. 112: 397-404. 

Yates, H.O. 1966a. Morphology and cytology of Uniola (Gramineae). Southwestern Naturalist 11: 145-189. 

–––. 1966b. Revision of grasses traditionally referred to Uniola, I. Uniola and Leptochloöpsis. Southwestern Naturalist 11: 372- 

394. 

–––. 1966c. Revision of grasses traditionally referred to Uniola, II. Chasmanthium. Southwestern Naturalist 11: 415-455. 

Yelton, J.D. 1974. Houstonia montana, a species, not an ecological variety. Castanea 39: 149-155. 

Yeo, P.F. 1984. Fruit-discharge type in Geranium (Geraniaceae): its use in classification and its evolutionary implications. J. 

Linn. Soc. Bot. 89: 1-36. 

Yeou-Ruenn, Ling. 1995. The New World Artemisia L. In Hind, D.J.N., C. Jeffrey, and G.V. Pope (eds.). Advances in 

Compositae systematics, pp. 239-254. Royal Botanic Gardens, Kew. 

Ying, Tsuen-Shen, Susumu Terabayashi, and D.E. Boufford. 1984. A monograph of Diphylleia (Berberidaceae). J. Arnold Arb. 

65: 57-94. 

Yuan, Y.-M., and P. Küpfer. 1995. Molecular phylogenetics of the subtribe Gentianinae (Gentianaceae) inferred from the 

sequences of internal transcribed spacers (ITS) of nuclear ribosomal DNA. Plant Systematics and Evolution 196: 207-226. 

–––, P. Küpfer, & J.J. Doyle. 1996. Infrageneric phylogeny of the genus Gentiana (Gentianaceae) inferred from nucleotide 

sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA. Amer. J. Bot. 83: 641-652. 

Yuncker, T.G. 1921. Revision of the North American and West Indian species of Cuscuta. Illinois Biol. Monogr. 6: 91-231. 

–––. 1965. Cuscuta. N. Am. Fl. II (4). 51 pp. 

Zahner, R., and S.M. Jones. 1983. Resolving the type locality for Shortia galacifolia T. & G. Castanea 48: 163-173. 

Zardini, E.M., H. Gu, & P.H. Raven. 1991. On the separation of two species within the Ludwigia uruguayensis complex 

(Onagraceae). Systematic Bot. 16: 242-244. 

Zavada, M.S. 1983. Comparative morphology of monocot pollen and evolutionary trends of apertures and wall structures. Bot. 

Rev. 49: 331-379. 

–––, and M. Kim. 1996. Phylogenetic analysis of Ulmaceae. Plant Systematics and Evolution 200: 13-20. 

–––, Xue-Lin Xu, and J.M. Edwards. 1983. On the taxonomic status of Lophiola aurea Ker-Gawler. Rhodora 85: 73-81. 

Zettler, L.W., N.S. Ahuja, and T.M. McInnis, Jr. 1996. Insect pollination of the endangered Monkey-face Orchid (Platanthera 

integrilabia) in McMinn County, Tennessee – one last glimpse of a once common spectacle. Castanea 61: 14-24. 

Zhang, J. 1996. A molecular biosystematic study on North American Solidago and related genera (Asteraceae: Astereae) based 

on chloroplast DNA RFLP analysis. Ph.D. dissertation, Univ. of Waterloo, Ontario, Canada. 

Zhang, W.-H., Z.-D. Chen, J.-H. Li, H.-B. Chen, and Y.-C. Tang. 2003. Phylogeny of the Dipsacales s.l. based on chloroplast 

trnL-F and ndhF sequences. Molecular Phylogenetics and Evolution 26: 176-189. 

Zika, P.F. 2003. The native subspecies of Juncus effusus (Juncaceae) in western North America. Brittonia 55: 150-156. 

–––, and A.L. Jacobson. 2003. An overlooked hybrid Japanese knotweed (Polygonum cuspidatum × sachalinense; 

Polygonaceae) in North America. Rhodora 105: 143-152. 

Ziman, S., C.S. Keener, Y. Kadota, E. Bulakh, O. Tsarenko, and B.E. Dutton. 2004. A taxonomic revision of Anemone L. 

subgenus Anemonanthea (DC.) Juz. sensu lato (Ranunculaceae). J. Japn. Bot. 79: 43-71, 196-206, 281-310. 

Zohary, D., and M. Hopf. 1994. Domestication of plants in the Old World. The origin and spread of cultivated plants in west 

Asia, Europe, and the Nile Valley. Second edition. Clarendon Press, Oxford. 279 pp. 

Zomlefer, W.B. 1996. The Trilliaceae in the southeastern United States. Harvard Papers in Botany 1: 91-120. 

–––. 1997a. The genera of Melanthiaceae in the southeastern United States. Harvard Papers in Botany 2: 133-177. 

–––. 1997b. The genera of Nartheciaceae in the southeastern United States. Harvard Papers in Botany 2: 195-211. 

–––. 1997c. The genera of Tofieldiaceae in the southeastern United States. Harvard Papers in Botany 2: 179-194. 

–––. 1998. The genera of Hemerocallidaceae in the southeastern United States. Harvard Papers in Botany 3: 113-145. 

–––. 1999. Advances in angiosperm systematics: examples from the Liliales and Asparagales. J. Torrey Bot. Soc. 126: 58-62. 

–––. 2003. Documented chromosome numbers 2003: 1. Chromosome number of Toxicoscordion nuttallii (Liliales: 

Melanthiaceae) and clarification of the genus. Sida 20: 1085-1092. 

–––, and W.S. Judd. 2002. Resurrection of segregates of the polyphyletic genus Zigadenus s.l. (Liliales: Melanthiaceae) and 

resulting new combinations. Novon 12: 299-308.


–––, and G.L. Smith. 2002. Documented chromosome numbers 2002: 1. Chromosome number of Stenanthium (Liliales: 

Melanthiaceae) and its significance in the taxonomy of tribe Melanthieae. Sida 20: 221–226. 

–––, N.H. Williams, W.M. Whitten, and W.S. Judd. 2001. Generic circumscription and relationships in the tribe Melanthieae 

(Liliales, Melanthiaceae), with emphasis on Zigadenus: evidence from ITS and trnL-F sequence data. Amer. J. Bot. 88: 

1657-1669. 

–––, W.M. Whitten, N.H. Williams, and W.S. Judd. 2003. An overview of Veratrum s.l. (Liliales: Melanthiaceae) and an 

infrageneric phylogeny based on ITS sequence data. Systematic Botany 28: 250-269. 

–––, W.M. Whitten, N.H. Williams, and W.S. Judd. 2006. Infrageneric phylogeny of Schoenocaulon (Liliales: Melanthiaceae) 

with clarification of cryptic species based on ITS sequence data and geographical distribution. Amer. J. Botany 93: 1178- 

1192. 

Zona, S. 1997. The genera of Palmae (Arecaceae) in the southeastern United States. Harvard Papers in Botany 27: 1-107. 

Zuloaga, F.O., O. Morrone, A.S. Vega, and L.M. Giussani. 1998. Revisión y análisis cladístico de Steinchisma (Poaceae: 

Panicoideae: Paniceae). Ann. Missouri Bot. Gard. 631-656.


INDEX of FAMILIES and GENERA 

Abelia ........................................481 

Abelmoschus .............................488 

Abies............................................60 

Abrus .........................................355 

Abutilon.....................................488 

Acacia........................................355 

Acalypha....................................344 

ACANTHACEAE........................68 

Acanthospermum......................119 

Acer ...........................................639 

ACERACEAE .............................71 

Achillea .....................................119 

Achyranthes ................................77 

Acicarpha ..................................260 

Acinos........................................446 

Acmella......................................119 

Aconitum...................................576 

ACORACEAE ...........................687 

Acorus .......................................687 

Acroptilon..................................120 

Actaea........................................577 

Actinidia......................................71 

ACTINIDIACEAE......................71 

Adiantum.....................................43 

Adlumia.....................................408 

Adonis........................................578 

ADOXACEAE.............................71 

Aegilops.....................................837 

Aegopodium ................................85 

Aeschynomene ..........................355 

Aesculus ....................................642 

Aethusa........................................85 

Agalinis .....................................526 

Agarista .....................................325 

Agastache ..................................446 

AGAVACEAE ...........................687 

Ageratina...................................120 

Ageratum...................................121 

Agrimonia .................................596 

Agropyron .................................837 

Agrostemma ..............................270 

Agrostis......................................837 

Ailanthus...................................655 

Aira............................................838 

AIZOACEAE ..............................75 

Ajuga .........................................446 

Akebia........................................472 

Albizia........................................356 

Alcea..........................................488 

Alchemilla .................................596 

Aletris ........................................818 

Aleurites ....................................345 

Alisma........................................689 

ALISMATACEAE ....................689 

ALLIACEAE.............................693 

Alliaria ......................................237 

Allium........................................694 

Alnus .........................................228 

Alopecurus ................................838 

Alophia......................................794 

Aloysia .......................................675 

Alstroemeria ..............................696 

ALSTROEMERIACEAE ..........696 

Alternanthera ..............................77 

Althaea.......................................488 

ALTINGIACEAE........................76 

Alysicarpus ................................356 

Alyssum .....................................238 

AMARANTHACEAE..................76 

Amaranthus.................................77 

AMARYLLIDACEAE ...............696 

Amblyolepis ...............................121 

Ambrosia ...................................121 

Amelanchier ..............................596 

Amianthium...............................813 

Ammannia .................................483 

Ammi ...........................................85 

Ammophila ................................839 

Ammoselinum..............................85 

Amorpha....................................356 

Ampelaster.................................122 

Ampelopsis.................................683 

Amphiachyris ............................122 

Amphianthus .............................542 

Amphicarpaea ...........................357 

Amphicarpum............................839 

Amsinckia..................................233 

Amsonia.....................................100 

ANACARDIACEAE....................81 

Anagallis....................................503 

Anaphalis...................................122 

Anchusa.....................................233 

Andrographis...............................68 

Andromeda ................................325 

Andropogon...............................840 

Anemone....................................578 

Anemonella................................580 

Anethum ......................................86 

Angadenia .................................101 

Angelica.......................................86 

ANNONACEAE ..........................84 

Anoda.........................................488 

Antennaria.................................122 

Anthemis....................................123 

Anthenantia...............................844 

Anthoxanthum ..........................845 

Anthriscus ...................................86 

Anticlea......................................814 

Antigonon..................................563 

Antirrhinum ..............................542 

Apera .........................................845 

Aphanes .....................................597 

APIACEAE..................................85 

Apios ..........................................358 

Apium ..........................................87 

Aplectrum ..................................821 

APOCYNACEAE ........................99 

Apocynum..................................101 

Apteria .......................................708 

AQUIFOLIACEAE...................108 

Aquilegia................................... 580 

Arabidopsis ............................... 238 

Arabis........................................ 238 

ARACEAE ................................ 700 

Arachis...................................... 358 

Arachniodes................................ 23 

Aralia ........................................ 112 

ARALIACEAE.......................... 112 

Arctium ..................................... 124 

Arctostaphylos .......................... 325 

Arctotis...................................... 124 

ARECACEAE........................... 705 

Arenaria.................................... 271 

Arethusa.................................... 821 

Argemone.................................. 534 

Argentina .................................. 597 

Argyrochosma............................. 44 

Arisaema................................... 701 

Aristida...................................... 845 

Aristolochia............................... 115 

ARISTOLOCHIACEAE........... 115 

Armoracia ................................. 239 

Arnica ....................................... 124 

Arnoglossum............................. 125 

Aronia ....................................... 597 

Arrhenatherum......................... 848 

Artemisia................................... 126 

Arthraxon ................................. 848 

Arum ......................................... 701 

Aruncus..................................... 598 

Arundinaria .............................. 848 

Arundo ...................................... 849 

Asarum...................................... 115 

ASCLEPIADACEAE ............... 118 

Asclepias ................................... 101 

Asimina....................................... 84 

ASPARAGACEAE ................... 707 

Asparagus ................................. 707 

Asperula.................................... 623 

ASPLENIACEAE....................... 17 

Asplenium ................................... 17 

Aster.......................................... 127 

ASTERACEAE......................... 118 

Astilbe ....................................... 649 

Astragalus ................................. 358 

Astranthium.............................. 127 

Astrolepis .................................... 44 

Athyrium ..................................... 52 

Atriplex ..................................... 287 

Aucuba...................................... 410 

Aureolaria................................. 528 

Avena ........................................ 849 

Axonopus .................................. 849 

Azolla .......................................... 20 

AZOLLACEAE........................... 20 

Baccharis .................................. 127 

Bacopa ...................................... 543 

Balduina ................................... 128 

BALSAMINACEAE ................. 224 

Bambusa ................................... 850


Baptisia......................................360 

Barbarea....................................239 

Bartonia ....................................412 

Bassia ........................................288 

BATACEAE ..............................224 

Batis...........................................224 

Begonia .....................................225 

BEGONIACEAE ......................225 

Bejaria.......................................325 

Belamcanda...............................794 

Bellis..........................................128 

BERBERIDACEAE..................225 

Berberis .....................................225 

Berchemia .................................592 

Berlandiera ...............................128 

Berteroa.....................................239 

Beta ...........................................289 

Betula ........................................229 

BETULACEAE.........................227 

BIBLIOGRAPHY....................950 

Bidens........................................128 

Bigelowia...................................130 

Bignonia....................................231 

BIGNONIACEAE.....................231 

BLECHNACEAE........................21 

Blechnum ....................................21 

Blephilia....................................447 

Boechera ...................................239 

Boehmeria.................................671 

Boerhavia ..................................507 

Bolboschoenus ..........................716 

Boltonia.....................................130 

BORAGINACEAE....................232 

Borago.......................................233 

Borreria.....................................623 

Borrichia ...................................131 

Bothriochloa .............................850 

Botrychium..................................37 

Botrypus ......................................38 

Bouchetia ..................................656 

Bouteloua ..................................850 

Boykinia ....................................649 

Brachyelytrum...........................851 

Brachypodium...........................851 

Brasenia ....................................258 

Brassica.....................................241 

BRASSICACEAE .....................237 

Braya .........................................241 

Brickellia...................................131 

Brintonia ...................................132 

Briza ..........................................851 

Brodiaea ....................................696 

BROMELIACEAE....................707 

Bromus ......................................852 

Broussonetia .............................499 

Brunnichia ................................563 

Buchloe .....................................854 

Buchnera...................................529 

Buckleya....................................638 

Buddleja ....................................654 

BUDDLEJACEAE....................257 

Buglossoides..............................233 

Bulbostylis .................................716 

Bunias........................................242 

Bupleurum...................................87 

Burmannia ................................708 

BURMANNIACEAE.................708 

Butia ..........................................706 

BUXACEAE..............................257 

Buxus.........................................257 

Cabomba....................................258 

CABOMBACEAE .....................257 

Cacalia.......................................132 

CACTACEAE............................258 

Cakile.........................................242 

Calamagrostis............................854 

Calamintha................................447 

Calamovilfa ...............................855 

Calepina.....................................243 

Calibrachoa ...............................656 

Calla ..........................................702 

Callicarpa ..................................447 

Callirhoe....................................488 

Callisia.......................................710 

CALLITRICHACEAE ..............260 

Callitriche..................................543 

Callitropsis...................................57 

Calluna ......................................326 

CALOCHORTACEAE..............709 

Calopogon .................................821 

Calotis........................................132 

Caltha ........................................580 

CALYCANTHACEAE ..............260 

Calycanthus...............................260 

CALYCERACEAE ....................260 

Calycocarpum............................497 

Calydorea...................................794 

Calylophus.................................514 

Calyptocarpus............................132 

Calystegia ..................................300 

Camassia ...................................687 

Camelina ...................................243 

Camellia.....................................667 

Campanula ................................261 

CAMPANULACEAE ................260 

Campanulastrum.......................262 

Campsis .....................................231 

Canna ........................................709 

CANNABACEAE......................264 

Cannabis....................................265 

CANNACEAE ...........................709 

Capnoides ..................................408 

CAPPARACEAE.......................266 

CAPRIFOLIACEAE.................266 

Capsella .....................................243 

Capsicum ...................................656 

Cardamine .................................243 

Cardaria.....................................246 

Cardiospermum.........................643 

Carduus .....................................132 

Carex .........................................717 

Carphephorus............................133 

Carpinus ....................................230 

Carthamus .................................134 

Carum ......................................... 87 

Carya......................................... 441 

CARYOPHYLLACEAE............ 269 

Cassia........................................ 363 

Castanea ................................... 397 

Castilleja ................................... 530 

Casuarina ................................. 284 

CASUARINACEAE ................. 284 

Catalpa...................................... 232 

Catapodium............................... 855 

Catharanthus............................ 105 

Caulophyllum ........................... 226 

Cayaponia................................. 311 

Cayratia..................................... 683 

Ceanothus ................................. 592 

Cedrus......................................... 61 

CELASTRACEAE.................... 284 

Celastrus ................................... 284 

Celosia......................................... 80 

CELTIDACEAE....................... 286 

Celtis ......................................... 265 

Cenchrus................................... 855 

Centaurea ................................. 134 

Centaurium............................... 412 

Centella....................................... 87 

Centrosema ............................... 363 

Centunculus.............................. 503 

Cephalanthus............................ 623 

CEPHALOTAXACEAE ............. 57 

Cephalotaxus .............................. 57 

Cerastium.................................. 271 

Ceratiola ................................... 326 

CERATOPHYLLACEAE......... 286 

Ceratophyllum .......................... 286 

Cercis ........................................ 363 

Cestrum..................................... 656 

Chaenomeles............................. 598 

Chaenorrhinum ........................ 544 

Chaerophyllum ........................... 88 

Chaetopappa ............................. 135 

Chaiturus .................................. 447 

Chamaecrista............................ 364 

Chamaecyparis ........................... 57 

Chamaedaphne......................... 326 

Chamaelirium........................... 814 

Chamaemelum.......................... 135 

Chamaesyce .............................. 345 

Chamerion ................................ 515 

Chaptalia................................... 135 

Chasmanthium ......................... 856 

Cheilanthes ................................. 44 

Chelidonium ............................. 534 

Chelone..................................... 544 

CHENOPODIACEAE.............. 287 

Chenopodium............................ 289 

Chevreulia................................. 136 

Chimaphila ............................... 326 

Chionanthus ............................. 511 

Chloris ...................................... 857 

Chondrilla................................. 136 

Chondrophora .......................... 136 

Chorispora ................................ 246


Chromolaena.............................136 

Chrysanthemum........................136 

CHRYSOBALANACEAE.........293 

Chrysogonum............................136 

Chrysoma ..................................137 

Chrysopogon .............................857 

Chrysopsis .................................137 

Chrysosplenium ........................649 

Cichorium .................................138 

Ciclospermum .............................88 

Cicuta ..........................................88 

Cimicifuga.................................580 

Cinna.........................................857 

Cinnamomum ...........................472 

Circaea ......................................515 

Cirsium......................................139 

Cissus ........................................684 

CISTACEAE .............................294 

Citrullus ....................................311 

Citrus.........................................632 

Cladanthus ................................140 

Cladium.....................................754 

Cladrastis ..................................365 

Claytonia ...................................570 

Cleistes ......................................822 

Clematis.....................................580 

CLEOMACEAE........................298 

Cleome.......................................298 

Clerodendrum ...........................447 

Clethra.......................................299 

CLETHRACEAE......................299 

Cliftonia ....................................313 

Clinopodium..............................448 

Clintonia....................................807 

Clitoria ......................................365 

CLUSIACEAE ..........................299 

Cnicus........................................141 

Cnidoscolus...............................347 

Cocculus....................................497 

Cocos .........................................706 

Coeloglossum ............................823 

Coelorachis ...............................858 

Coincya......................................246 

Coix ...........................................858 

COLCHICACEAE ....................709 

Colchicum .................................709 

Collinsia ....................................545 

Collinsonia ................................449 

Colocasia...................................702 

Comandra..................................638 

Commelina ................................711 

COMMELINACEAE................710 

COMPOSITAE .........................118 

Comptonia.................................501 

Conioselinum ..............................89 

Conium........................................89 

Conoclinium..............................141 

Conopholis ................................530 

Conradina .................................450 

Conringia ..................................246 

Consolida ..................................583 

Convallaria................................933 

CONVALLARIACEAE.............714 

CONVOLVULACEAE..............299 

Convolvulus...............................301 

Conyza .......................................141 

Coptis.........................................583 

Corallorhiza...............................823 

Coreopsis ...................................141 

Coriandrum .................................89 

CORNACEAE ...........................306 

Cornus .......................................306 

Coronilla....................................365 

Coronopus .................................246 

Corrigiola ..................................272 

Cortaderia..................................858 

Corydalis....................................409 

Corylus.......................................230 

Cosmos.......................................145 

Cota............................................145 

Cotinus.........................................81 

Cotula ........................................145 

Crassula.....................................308 

CRASSULACEAE.....................308 

Crataegus...................................598 

Crepis.........................................146 

Crinum.......................................697 

Crocanthemum..........................294 

Crocosmia..................................794 

Crocus........................................795 

Croomia.....................................938 

Croptilon....................................146 

Crossopetalum...........................284 

Crotalaria ..................................365 

Croton........................................347 

Crotonopsis................................348 

Cruciata.....................................623 

CRUCIFERAE..........................237 

Cryptogramma.............................45 

Cryptotaenia ................................89 

Ctenium .....................................858 

Cucumis.....................................311 

Cucurbita...................................312 

CUCURBITACEAE..................311 

Cudrania....................................499 

Cullen ........................................366 

Cunila ........................................450 

Cunninghamia ............................58 

Cuphea.......................................483 

CUPRESSACEAE.......................57 

Cuscuta......................................301 

CUSCUTACEAE.......................313 

Cuthbertia..................................712 

Cyclachaena ..............................146 

Cycloloma..................................291 

Cydonia......................................605 

Cymbalaria ................................545 

CYMODOCEACEAE................714 

Cymophyllus..............................754 

Cynanchum ...............................106 

Cynodon.....................................859 

Cynoglossum .............................233 

Cynosciadium..............................89 

Cynosurus..................................859 

CYPERACEAE......................... 715 

Cyperus ..................................... 755 

Cypripedium.............................. 824 

Cyrilla ....................................... 313 

CYRILLACEAE ....................... 313 

Cyrtomium .................................. 23 

Cystopteris................................... 52 

Cytisus....................................... 366 

Dactylis ..................................... 859 

Dactyloctenium......................... 859 

Dalea......................................... 367 

Dalibarda .................................. 605 

Danae........................................ 933 

Danthonia ................................. 860 

Dasiphora ................................. 605 

Dasistoma ................................. 530 

Datura....................................... 656 

Daubentonia ............................. 368 

Daucus ........................................ 90 

Decodon .................................... 483 

Decumaria ................................ 425 

Delphinium ............................... 583 

Dendrolycopodium ..................... 31 

Dennstaedtia ............................... 22 

DENNSTAEDTIACEAE............ 22 

Deparia ....................................... 53 

Deschampsia............................. 860 

Descurainia............................... 246 

Desmanthus .............................. 368 

Desmazeria ............................... 861 

Desmodium ............................... 368 

Deutzia ...................................... 426 

Diamorpha................................ 308 

Dianthus ................................... 273 

DIAPENSIACEAE................... 314 

Diarrhena ................................. 861 

Dicentra .................................... 409 

Dicerandra................................ 450 

Dichanthelium.......................... 861 

Dichelostemma ......................... 938 

Dichondra ................................. 302 

Dicliptera .................................... 68 

Didiplis...................................... 484 

Diervilla .................................... 315 

DIERVILLACEAE................... 315 

Digitalis..................................... 545 

Digitaria.................................... 876 

Dinebra ..................................... 877 

Dioclea ...................................... 372 

Diodia........................................ 623 

Dionaea..................................... 317 

DIONAEACEAE ...................... 316 

Dioscorea .................................. 786 

DIOSCOREACEAE ................. 786 

Diospyros .................................. 318 

Diphasiastrum ............................ 31 

Diphylleia.................................. 226 

Diplazium.................................... 54 

Diplotaxis.................................. 247 

DIPSACACEAE ....................... 316 

Dipsacus.................................... 316 

Dirca ......................................... 668


Distichlis....................................878 

Ditrysinia...................................348 

Dittrichia ...................................147 

Dodecatheon .............................573 

Doellingeria...............................147 

Draba.........................................247 

Dracocephalum.........................451 

Dracopis ....................................148 

Drosera......................................317 

DROSERACEAE......................317 

Drymaria ...................................273 

Drymocallis ...............................606 

DRYOPTERIDACEAE...............23 

Dryopteris....................................23 

Duchesnea.................................606 

Dulichium .................................761 

Dyschoriste..................................68 

Dysphania .................................291 

Dyssodia ....................................148 

EBENACEAE ...........................318 

Echinacea..................................148 

Echinochloa ..............................878 

Echinocystis ..............................312 

Echinodorus..............................689 

Echinops....................................149 

Echium......................................234 

Eclipta .......................................149 

Edgeworthia ..............................668 

Egeria........................................792 

Eichhornia ................................927 

ELAEAGNACEAE ...................319 

Elaeagnus..................................319 

ELATINACEAE .......................319 

Elatine .......................................319 

Eleocharis .................................761 

Elephantopus ............................149 

Eleusine.....................................879 

Eleutherococcus........................113 

Elionurus ..................................879 

Elliottia......................................327 

Ellisia ........................................429 

Elodea........................................792 

Elsholtzia...................................451 

Elymus.......................................879 

Elytraria ......................................69 

Elytrigia.....................................881 

Emilia........................................150 

Endodeca...................................116 

Enemion....................................584 

Enteropogon..............................881 

Epidendrum...............................825 

Epifagus ....................................530 

Epigaea......................................327 

Epilobium..................................516 

Epipactis....................................825 

EQUISETACEAE.......................26 

Equisetum ...................................26 

Eragrostis ..................................882 

Eranthis.....................................584 

Erechtites ..................................150 

Eremochloa...............................885 

Erianthus ..................................885 

Erica ..........................................327 

ERICACEAE.............................320 

Erigenia.......................................90 

Erigeron.....................................150 

ERIOCAULACEAE..................787 

Eriocaulon.................................787 

Eriochloa ...................................885 

Eriogonum.................................563 

Eriophorum ...............................766 

Erodium.....................................417 

Eruca .........................................248 

Erucastrum................................248 

Eryngium.....................................90 

Erysimum ..................................248 

Erythrina ...................................372 

Erythronium..............................808 

Eschscholzia..............................534 

Eubotrys.....................................327 

Eulophia ....................................825 

Euonymus..................................285 

Eupatoriadelphus .. See Eutrochium 

Eupatorium................................152 

Euphorbia..................................348 

EUPHORBIACEAE .................344 

Eurybia ......................................157 

Eustachys...................................885 

Eustoma.....................................412 

Euthamia...................................159 

Eutrochium................................160 

Evax...........................................161 

Evolvulus...................................303 

Exochorda .................................606 

FABACEAE ..............................353 

Facelis .......................................161 

FAGACEAE..............................397 

Fagopyrum ................................564 

Fagus.........................................398 

Falcaria .......................................92 

Fallopia .....................................564 

Fatoua .......................................500 

Festuca ......................................886 

Ficaria .......................................585 

Ficus ..........................................500 

Filago.........................................161 

Filipendula ................................606 

Fimbristylis................................766 

Firmiana....................................489 

Flaveria .....................................161 

Fleischmannia...........................161 

Floerkea.....................................478 

Foeniculum .................................92 

Forestiera ..................................511 

Forsythia ...................................511 

Fothergilla.................................423 

Fragaria.....................................606 

Frangula....................................593 

Franklinia .................................667 

Frasera ......................................413 

Fraxinus ....................................512 

Froelichia ....................................80 

Fuirena......................................768 

Fumaria.....................................410 

FUMARIACEAE...................... 408 

Funastrum ................................ 106 

Gaillardia.................................. 162 

Galactia..................................... 372 

Galanthus ................................. 697 

Galax......................................... 314 

Galearis..................................... 825 

Galeopsis................................... 451 

Galinsoga.................................. 162 

Galium ...................................... 624 

Gamochaeta.............................. 162 

GARRYACEAE ........................ 410 

Gaultheria................................. 327 

Gaura ........................................ 517 

Gaylussacia............................... 328 

GELSEMIACEAE.................... 411 

Gelsemium ................................ 411 

Genista ...................................... 373 

Gentiana ................................... 413 

GENTIANACEAE.................... 411 

Gentianella ............................... 414 

Gentianopsis ............................. 415 

GERANIACEAE ...................... 417 

Geranium.................................. 418 

Geum......................................... 607 

Gillenia ..................................... 609 

Ginkgo......................................... 60 

GINKGOACEAE........................ 60 

Gladiolus................................... 795 

Glandularia............................... 675 

Glaucium .................................. 535 

Glaux ........................................ 503 

Glebionis................................... 164 

Glechoma.................................. 451 

Gleditsia .................................... 373 

Glottidium................................. 374 

Glyceria..................................... 887 

Glycine ...................................... 374 

Glycyrrhiza ............................... 374 

Gnaphalium.............................. 164 

Gomphrena ................................. 81 

Gonolobus................................. 106 

Goodyera................................... 826 

Gordonia................................... 667 

Gossypium................................. 489 

GRAMINEAE........................... 836 

Grammitis.................................... 41 

Gratiola..................................... 545 

Grindelia................................... 164 

GROSSULARIACEAE............. 419 

Guilleminea ................................ 81 

Guizotia..................................... 164 

Gutierrezia ................................ 165 

Gymnocarpium ........................... 54 

Gymnocladus ............................ 374 

Gymnopogon............................. 888 

Gymnostyles.............................. 165 

Gypsophila ................................ 273 

Habenaria ................................. 826 

Hackelia.................................... 234 

Hackelochloa............................ 888 

HAEMODORACEAE .............. 789


Hainardia ..................................888 

Halesia ......................................664 

Halodule....................................714 

HALORAGACEAE...................421 

HAMAMELIDACEAE.............423 

Hamamelis ................................424 

Haplopappus .............................165 

Harperocallis.............................939 

Hartwrightia..............................165 

Hasteola ....................................165 

Hedeoma ...................................452 

Hedera.......................................113 

Hedyotis.....................................627 

Helenium...................................165 

Helianthus.................................167 

Heliomeris.................................172 

Heliopsis....................................172 

HELIOTROPACEAE...............424 

Heliotropium.............................424 

Helleborus.................................585 

Helminthotheca.........................172 

Helonias ....................................814 

HEMEROCALLIDACEAE......789 

Hemerocallis .............................789 

Hemicarpha...............................769 

Hepatica ....................................585 

Heracleum...................................92 

Herbertia ...................................795 

Herniaria...................................273 

Hesperis.....................................249 

Heteranthera .............................928 

Heteropogon..............................889 

Heterotheca...............................173 

Heuchera...................................649 

Hexalectris ................................826 

Hexastylis ..................................116 

Hibiscus.....................................489 

Hieracium .................................173 

Hierochloe.................................889 

HIPPOCASTANACEAE ..........425 

Holcus .......................................889 

Holosteum .................................273 

Honckenya ................................274 

Hordeum ...................................889 

HOSTACEAE ...........................790 

Hottonia ....................................574 

Houstonia..................................627 

Hovenia .....................................593 

Hudsonia...................................295 

Humulus....................................265 

Huperzia......................................32 

HYACINTHACEAE .................790 

Hyacinthoides ...........................790 

Hyacinthus ................................791 

Hybanthus.................................677 

Hydrangea.................................426 

HYDRANGEACEAE................425 

HYDRASTIDACEAE ...............427 

Hydrastis ...................................428 

Hydrilla .....................................792 

HYDROCHARITACEAE .........791 

Hydrocotyle .........................92, 114 

Hydrolea ............................428, 429 

HYDROLEACEAE ...................428 

HYDROPHYLLACEAE............428 

Hydrophyllum............................429 

Hygrophila...................................69 

Hylotelephium ...........................309 

Hymenocallis.............................697 

Hymenopappus..........................175 

HYMENOPHYLLACEAE ..........27 

Hymenophyllum ..........................27 

Hymenoxys ................................175 

HYPERICACEAE.....................432 

Hypericum .................................432 

Hypochaeris...............................175 

Hypopitys...................................329 

HYPOXIDACEAE ....................793 

Hypoxis......................................793 

Hyptis.........................................452 

Hyssopus....................................452 

Hystrix .......................................890 

Iberis..........................................249 

Ilex.............................................109 

Iliamna ......................................490 

ILLICIACEAE ..........................440 

Illicium ......................................441 

Impatiens ...................................224 

Imperata ....................................890 

Indigofera..................................374 

Inula ..........................................176 

Iodanthus...................................249 

Ionactis ......................................176 

Ipheion.......................................696 

Ipomoea.....................................303 

Ipomopsis...................................555 

Iresine..........................................81 

IRIDACEAE..............................794 

Iris..............................................795 

Isanthus .....................................453 

Isatis ..........................................249 

ISOETACEAE ............................28 

Isoetes ..........................................28 

Isolepis.......................................769 

Isopyrum....................................585 

Isotrema.....................................118 

Isotria.........................................827 

Itea.............................................441 

ITEACEAE................................441 

Iva..............................................176 

Ixeris..........................................177 

Jacquemontia ............................305 

Jamesianthus.............................177 

Jasione.......................................262 

Jasminum ..................................513 

Jeffersonia.................................226 

JUGLANDACEAE....................441 

Juglans ......................................445 

JUNCACEAE............................799 

JUNCAGINACEAE..................806 

Juncus .......................................799 

Juniperus.....................................58 

Justicia.........................................69 

Kallstroemia ..............................686 

Kalmia....................................... 330 

Kerria ........................................ 609 

Kickxia ...................................... 546 

Knautia ..................................... 317 

Kochia....................................... 292 

Koeleria..................................... 890 

Koelreuteria .............................. 643 

Kolkwitzia ................................. 481 

Kosteletzkya .............................. 490 

Krameria................................... 445 

KRAMERIACEAE ................... 445 

Krigia ........................................ 177 

Kuhnia ...................................... 178 

Kummerowia............................. 375 

Kyllinga..................................... 769 

Lablab ....................................... 375 

Lachnagrostis ........................... 890 

Lachnanthes ............................. 789 

Lachnocaulon........................... 788 

Lactuca ..................................... 178 

Lagenaria.................................. 312 

Lagerstroemia........................... 484 

Lagurus..................................... 890 

LAMIACEAE ........................... 445 

Lamiastrum............................... 453 

Lamium..................................... 453 

Lamprocapnos .......................... 410 

Landoltia................................... 702 

Lantana..................................... 675 

Laportea.................................... 672 

Lappula..................................... 234 

Lapsana..................................... 179 

LARDIZABALACEAE............. 472 

Larix............................................ 61 

Lathyrus.................................... 375 

LAURACEAE........................... 472 

Laurus....................................... 472 

Leavenworthia .......................... 249 

Lechea....................................... 296 

Leersia....................................... 891 

LEGUMINOSAE...................... 353 

Leiophyllum.............................. 331 

Leitneria.................................... 655 

LEITNERIACEAE................... 474 

Lemna ....................................... 702 

LEMNACEAE .......................... 806 

LENTIBULARIACEAE........... 474 

Leonotis..................................... 453 

Leontodon................................. 179 

Leonurus................................... 453 

Lepidium................................... 250 

Leptochloa ................................ 891 

Leptoloma ................................. 892 

Leptopus.................................... 540 

Lepuropetalon........................... 536 

Lespedeza.................................. 376 

Lesquerella ............................... 251 

Leucaena................................... 379 

Leucanthemum......................... 180 

Leucojum .................................. 698 

Leucospora ............................... 547 

Leucothoe ................................. 331


Liatris ........................................180 

Licania ......................................293 

Ligularia....................................183 

Ligusticum...................................92 

Ligustrum..................................513 

Lilaeopsis ....................................92 

LILIACEAE..............................806 

Lilium........................................809 

LIMNANTHACEAE.................478 

Limnobium................................792 

Limnodea ..................................892 

Limnophila................................547 

Limonium..................................555 

Limosella...................................547 

LINACEAE...............................478 

Linaria.......................................547 

Lindera......................................473 

Lindernia...................................479 

LINDERNIACEAE ..................479 

Linnaea .....................................481 

LINNAEACEAE.......................480 

Linum........................................478 

Liparis .......................................827 

Lipocarpha ................................770 

Liquidambar................................76 

Liriodendron .............................485 

Liriope .......................................934 

Listera........................................827 

Lithospermum ...........................234 

Litsea .........................................473 

Lobelia.......................................262 

Lobularia...................................251 

LOGANIACEAE.......................481 

Lolium .......................................892 

Lonicera ....................................266 

Lophiola ....................................819 

Lotus..........................................379 

Ludwigia....................................517 

Luffa..........................................312 

Lunaria......................................251 

Lupinus .....................................380 

Luziola.......................................893 

Luzula........................................805 

Lychnis......................................274 

Lycium.......................................657 

Lycopersicon .............................657 

LYCOPODIACEAE....................30 

Lycopodiella ................................33 

Lycopodium.................................34 

Lycopus .....................................454 

Lycoris.......................................698 

Lygodesmia ...............................183 

LYGODIACEAE.........................35 

Lygodium ....................................35 

Lyonia........................................331 

Lysimachia ................................503 

LYTHRACEAE.........................482 

Lythrum.....................................484 

Macbridea .................................455 

Macfadyena...............................232 

Macleaya ...................................535 

Maclura.....................................500 

Macranthera..............................531 

Macroptilium.............................380 

Macrothelypteris..........................50 

Madia.........................................183 

Magnolia ...................................486 

MAGNOLIACEAE ...................485 

Mahonia ....................................227 

Maianthemum ...........................934 

Malaxis ......................................828 

Malus.........................................609 

Malva.........................................491 

MALVACEAE...........................487 

Malvastrum................................491 

Malvaviscus ...............................492 

Manfreda ...................................688 

Manihot .....................................351 

MARANTACEAE .....................812 

Marrubium ................................455 

Marshallia .................................183 

Marsilea.......................................36 

MARSILEACEAE.......................35 

MARTYNIACEAE ....................494 

Matelea ......................................107 

Matricaria..................................185 

Matteuccia ...................................36 

Matthiola ...................................252 

Mayaca ......................................813 

MAYACACEAE ........................813 

Mazus ........................................539 

Mecardonia................................547 

Medeola .....................................811 

Medicago ...................................380 

Meehania...................................455 

Megalodonta..............................185 

Melampyrum .............................531 

Melanthera ................................185 

MELANTHIACEAE .................813 

Melanthium ...............................814 

MELASTOMATACEAE...........495 

Melia..........................................497 

MELIACEAE ............................497 

Melica ........................................893 

Melilotus....................................381 

Melinis.......................................893 

Melissa.......................................455 

Melochia....................................492 

Melothria ...................................312 

MENISPERMACEAE ..............497 

Menispermum............................498 

Mentha.......................................455 

MENYANTHACEAE................498 

Menyanthes ...............................498 

Menziesia...................................332 

Mercurialis ................................351 

Merremia ...................................305 

Mertensia...................................235 

Micranthemum..........................480 

Micromeria................................456 

Micropolypodium ........................41 

Microstegium.............................893 

Microthlaspi ..............................252 

Mikania .....................................186 

Milium....................................... 894 

Mimosa ..................................... 381 

Mimulus.................................... 539 

Minuartia.................................. 274 

Mirabilis.................................... 507 

Miscanthus ............................... 894 

Misopates .................................. 547 

Mitchella................................... 629 

Mitella....................................... 651 

Mitracarpus .............................. 629 

Mitreola..................................... 481 

Modiola..................................... 492 

Moehringia ............................... 275 

Moenchia .................................. 275 

Moldavica ................................. 456 

MOLLUGINACEAE ................ 499 

Mollugo..................................... 499 

Monarda ................................... 457 

Monotropa ................................ 333 

Monotropsis .............................. 333 

Montia....................................... 570 

MORACEAE ............................ 499 

Morella...................................... 501 

Morus........................................ 500 

Mosla ........................................ 459 

Mucuna..................................... 382 

Muhlenbergia ........................... 894 

Murdannia................................ 712 

Muscari..................................... 791 

Myosotis .................................... 235 

Myosoton................................... 275 

Myosurus .................................. 585 

Myrica....................................... 502 

MYRICACEAE......................... 501 

Myriophyllum ........................... 421 

MYRSINACEAE ...................... 502 

NAJADACEAE ........................ 817 

Najas ......................................... 817 

Nama......................................... 430 

Nandina .................................... 227 

Napaea ...................................... 492 

Narcissus................................... 698 

NARTHECIACEAE ................. 818 

Narthecium ............................... 819 

Nassella..................................... 897 

Nasturtium................................ 252 

Neeragrostis.............................. 897 

Nelumbo.................................... 506 

NELUMBONACEAE............... 506 

Nemastylis................................. 797 

Nemopanthus............................ 112 

Nemophila................................. 430 

Neobeckia ................................. 252 

Nepeta ....................................... 459 

Neptunia ................................... 382 

Nerium ...................................... 108 

Nestronia................................... 638 

Neviusia .................................... 610 

Nicandra ................................... 657 

Nicotiana................................... 657 

Nierembergia ............................ 658 

Nigella....................................... 585


Nolina........................................934 

NOLINACEAE .........................819 

Nothoscordum...........................696 

Nuphar ......................................507 

Nuttallanthus ............................548 

NYCTAGINACEAE..................506 

Nymphaea .................................509 

NYMPHAEACEAE ..................507 

Nymphoides...............................498 

Nyssa .........................................509 

NYSSACEAE ............................509 

Obolaria ....................................415 

Ocimum.....................................459 

Oclemena ..................................186 

Oenothera..................................522 

Oldenlandia...............................630 

OLEACEAE..............................510 

Oligoneuron ..............................186 

ONAGRACEAE........................514 

Onoclea .......................................37 

ONOCLEACEAE........................36 

Onopordum ...............................187 

Onosmodium .............................236 

OPHIOGLOSSACEAE...............37 

Ophioglossum .............................38 

Oplismenus................................897 

Opuntia......................................258 

Orbexilum .................................382 

ORCHIDACEAE ......................819 

Origanum ..................................459 

Ornithogalum............................791 

OROBANCHACEAE................525 

Orobanche.................................531 

Orontium...................................703 

Orthilia......................................333 

Orychophragmus ......................253 

Oryza .........................................897 

Oryzopsis ...................................897 

Osmanthus ................................514 

Osmorhiza ...................................93 

Osmunda .....................................40 

OSMUNDACEAE.......................40 

Ostrya ........................................231 

OXALIDACEAE.......................533 

Oxalis ........................................533 

Oxycaryum ................................770 

Oxydendrum..............................333 

Oxypolis.......................................93 

Pachysandra..............................257 

Packera......................................187 

Paederia ....................................630 

Paeonia......................................534 

PAEONIACEAE.......................534 

Palafoxia ...................................189 

Palhinhaea ..................................34 

PALMAE...................................705 

Panax ........................................115 

Panicum ....................................898 

Panphalea .................................189 

Papaver......................................535 

PAPAVERACEAE....................534 

Parapholis .................................902 

Parietaria...................................672 

Parkinsonia ...............................383 

Parnassia ...................................536 

PARNASSIACEAE ...................536 

Paronychia ................................276 

Parthenium................................189 

Parthenocissus ..........................684 

Pascopyrum ...............................902 

Paspalidium...............................902 

Paspalum ...................................902 

Passiflora...................................537 

PASSIFLORACEAE.................537 

Pastinaca .....................................94 

Paulownia..................................538 

PAULOWNIACEAE .................538 

Pavonia......................................492 

Paxistima...................................286 

Paysonia ....................................253 

Pectis..........................................190 

Pedicularis.................................532 

Pediomelum...............................384 

Pellaea .........................................45 

Peltandra ...................................703 

Pennisetum................................907 

Penstemon .................................548 

PENTHORACEAE ...................538 

Penthorum.................................538 

Pentodon....................................630 

Peperomia..................................541 

Perideridia...................................94 

Perilla ........................................459 

Peripleura..................................190 

Periploca....................................108 

Persea ........................................474 

Persicaria...................................565 

Petalostemon .............................384 

Petasites.....................................191 

Petrorhagia................................277 

Petroselinum................................94 

Petunia.......................................658 

Peucedanum................................94 

Phacelia.....................................430 

Phalaris .....................................907 

Phanopyrum..............................908 

Phaseolus...................................384 

Phegopteris..................................50 

Phemeranthus ...........................570 

Philadelphus..............................427 

Phlebodium..................................41 

Phleum.......................................908 

Phlox..........................................556 

Phoebanthus..............................191 

Phoradendron............................682 

Photinia .....................................610 

Phragmites.................................908 

Phryma ......................................539 

PHRYMACEAE ........................539 

Phyla..........................................675 

PHYLLANTHACEAE ..............540 

Phyllanthus................................540 

Phyllostachys.............................909 

Physalis......................................658 

Physaria .................................... 253 

Physocarpus.............................. 610 

Physostegia ............................... 459 

Phytolacca................................. 541 

PHYTOLACCACEAE.............. 541 

Picea............................................ 61 

Picris ......................................... 191 

Pieris ......................................... 333 

Pilea .......................................... 672 

Piloblephis ................................ 461 

Pilularia ...................................... 36 

Pimpinella................................... 94 

PINACEAE................................. 60 

Pinckneya ................................. 630 

Pinellia...................................... 704 

Pinguicula................................. 474 

Pinus ........................................... 62 

PIPERACEAE.......................... 541 

Piptatherum .............................. 910 

Piptochaetium........................... 910 

Piriqueta ................................... 669 

Pistia ......................................... 704 

Pisum ........................................ 385 

PITTOSPORACEAE................ 541 

Pittosporum............................... 542 

Pityopsis .................................... 191 

Plagiobothrys............................ 237 

Planera...................................... 670 

PLANTAGINACEAE............... 542 

Plantago.................................... 549 

PLATANACEAE ...................... 554 

Platanthera ............................... 828 

Platanus .................................... 554 

Platycladus.................................. 59 

Platycodon ................................ 264 

Platythelys................................. 831 

Plectocephalus .......................... 192 

Pleea.......................................... 939 

Pleioblastus............................... 910 

Pleopeltis..................................... 41 

Pluchea ..................................... 193 

PLUMBAGINACEAE.............. 554 

Poa ............................................ 910 

POACEAE ................................ 836 

Podophyllum............................. 227 

PODOSTEMACEAE................ 555 

Podostemum.............................. 555 

Pogonia..................................... 832 

Polanisia ................................... 298 

POLEMONIACEAE ................ 555 

Polemonium.............................. 559 

Polycarpon................................ 278 

Polygala .................................... 559 

POLYGALACEAE ................... 559 

POLYGONACEAE................... 562 

Polygonatum............................. 935 

Polygonella ............................... 566 

Polygonum................................ 567 

Polymnia................................... 194 

POLYPODIACEAE.................... 41 

Polypodium ................................. 42 

Polypogon ................................. 912


Polypremum ..............................666 

Polystichum.................................25 

Polytaenia....................................94 

Poncirus ....................................632 

Pontederia .................................928 

PONTEDERIACEAE...............927 

Ponthieva ..................................832 

Populus......................................633 

Porteranthus .............................611 

Portulaca...................................571 

PORTULACACEAE.................569 

Potamogeton .............................929 

POTAMOGETONACEAE........929 

Potentilla ...................................611 

Prenanthes ................................194 

PRIMULACEAE ......................573 

Proboscidea...............................494 

Prosartes....................................811 

Proserpinaca .............................423 

Prunella.....................................461 

Prunus.......................................612 

Pseuderanthemum ......................70 

Pseudognaphalium ...................195 

Pseudolycopodiella......................35 

Pseudosasa ................................912 

Pseudotaenidia............................94 

PSILOTACEAE..........................42 

Psilotum ......................................42 

Psoralea.....................................385 

Psoralidium...............................385 

Psychotria..................................631 

Ptelea.........................................632 

PTERIDACEAE .........................43 

Pteridium.....................................22 

Pteris............................................46 

Pterocaulon...............................196 

Pteroglossaspis..........................832 

Ptilimnium...................................95 

Puccinellia.................................912 

Pueraria ....................................385 

Punica .......................................484 

PUNICACEAE .........................574 

Pycnanthemum .........................461 

Pyracantha ................................615 

Pyrola ........................................334 

Pyrrhopappus............................196 

Pyrularia ...................................638 

Pyrus..........................................615 

Pyxidanthera.............................314 

Quercus .....................................399 

RANUNCULACEAE ................574 

Ranunculus...............................585 

Raphanus ..................................254 

Rapistrum..................................254 

Ratibida .....................................196 

Reimarochloa............................913 

Reseda .......................................591 

RESEDACEAE.........................591 

Reynoutria.................................568 

RHAMNACEAE .......................592 

Rhamnus ...................................593 

Rhapidophyllum........................706 

Rheum .......................................568 

Rhexia........................................495 

Rhizophora ................................595 

RHIZOPHORACEAE...............594 

Rhodiola ....................................309 

Rhododendron...........................334 

Rhodotypos ................................615 

Rhus.............................................82 

Rhynchelytrum ..........................913 

Rhynchosia................................386 

Rhynchospora............................770 

Ribes ..........................................419 

Richardia ...................................631 

Ricinus.......................................352 

Ripidium ....................................913 

Robinia ......................................386 

Rorippa ......................................254 

Rosa ...........................................616 

ROSACEAE ..............................595 

Rosmarinus................................464 

Rostraria....................................913 

Rotala.........................................485 

Rottboellia .................................913 

RUBIACEAE ............................622 

Rubus.........................................617 

Rudbeckia..................................197 

Ruellia .........................................70 

Rugelia.......................................199 

Rumex........................................569 

Ruppia .......................................932 

RUPPIACEAE ..........................932 

RUSCACEAE............................933 

Ruta ...........................................633 

RUTACEAE ..............................632 

Sabal ..........................................706 

Sabatia.......................................415 

Saccharum.................................913 

Sacciolepis .................................914 

Sageretia....................................594 

Sagina........................................278 

Sagittaria ...................................690 

SALICACEAE...........................633 

Salicornia ..................................292 

Salix...........................................635 

Salpichroa .................................661 

Salsola .......................................292 

Salvia .........................................464 

Salvinia........................................47 

SALVINIACEAE ........................47 

Sambucus ....................................72 

SAMOLACEAE ........................637 

Samolus .....................................637 

Sanguinaria...............................535 

Sanguisorba...............................619 

Sanicula.......................................96 

SANTALACEAE.......................637 

Santolina ...................................199 

SAPINDACEAE........................638 

Sapindus ....................................643 

Sapium.......................................352 

Saponaria ..................................278 

SAPOTACEAE..........................643 

Sarcocornia............................... 293 

Sarracenia................................. 645 

SARRACENIACEAE ............... 645 

Sassafras ................................... 474 

SAURURACEAE...................... 648 

Saururus ................................... 648 

Saxifraga................................... 651 

SAXIFRAGACEAE.................. 648 

Scandix ....................................... 97 

Sceptridium................................. 39 

Schedonorus ............................. 915 

Scheuchzeria............................. 935 

SCHEUCHZERIACEAE ......... 935 

Schisandra ................................ 653 

SCHISANDRACEAE............... 653 

Schizachne................................ 915 

Schizachyrium .......................... 915 

Schizaea ...................................... 48 

SCHIZAEACEAE ...................... 47 

Schoenocaulon ......................... 814 

Schoenolirion.................... 688, 791 

Schoenoplectus ......................... 779 

Schwalbea ................................. 532 

Scirpus ...................................... 781 

Scleranthus ............................... 279 

Scleria ....................................... 783 

Sclerochloa ............................... 916 

Sclerolepis................................. 200 

Scolymus................................... 200 

Scoparia .................................... 551 

Scrophularia ............................. 654 

SCROPHULARIACEAE.......... 653 

Scutellaria................................. 465 

Sebastiania................................ 352 

Secale ........................................ 916 

Securigera................................. 388 

Sedum ....................................... 309 

Selaginella .................................. 48 

SELAGINELLACEAE............... 48 

Senecio...................................... 200 

Senna ........................................ 388 

Serenoa ..................................... 707 

Sericocarpus ............................. 200 

Sesbania.................................... 389 

Sesuvium..................................... 75 

Setaria....................................... 916 

Seutera ...................................... 108 

Seymeria ................................... 532 

Sherardia .................................. 631 

Shortia ...................................... 315 

Sibara........................................ 255 

Sibbaldiopsis ............................. 619 

Sicyos ........................................ 313 

Sida ........................................... 492 

Sideritis ..................................... 468 

Sideroxylon ............................... 643 

Silene ........................................ 279 

Silphium.................................... 201 

Silybum ..................................... 203 

SIMAROUBACEAE................. 655 

Sinapis ...................................... 255 

Sisymbrium ............................... 255


Sisyrinchium .............................797 

Sium ............................................97 

Smallanthus ..............................203 

SMILACACEAE.......................936 

Smilax........................................936 

SOLANACEAE.........................655 

Solanum ....................................661 

Solidago.....................................204 

Soliva.........................................211 

Sonchus.....................................211 

Sophora .....................................389 

Sophronanthe............................552 

Sorbaria.....................................620 

Sorbus........................................620 

Sorghastrum..............................917 

Sorghum....................................918 

SPARGANIACEAE ..................938 

Sparganium...............................945 

Spartina.....................................918 

Spergula ....................................281 

Spergularia................................281 

Spermacoce ...............................631 

Spermolepis.................................97 

Sphaeralcea...............................493 

Sphagneticola............................212 

Sphenoclea ................................663 

SPHENOCLEACEAE ..............663 

Sphenopholis.............................919 

Spigelia......................................482 

Spinacia.....................................293 

Spinulum.....................................35 

Spiraea ......................................620 

Spiranthes .................................832 

Spirodela ...................................704 

Sporobolus.................................920 

Stachys ......................................468 

Staphylea...................................663 

STAPHYLEACEAE..................663 

Steinchisma...............................922 

Stellaria .....................................282 

STEMONACEAE .....................938 

Stenandrium................................71 

Stenanthium..............................815 

Stenotaphrum............................923 

Stephanandra............................622 

STERCULIACEAE ..................663 

Sternbergia................................699 

Stewartia....................................668 

Stillingia....................................352 

Stipa...........................................923 

Stipulicida .................................283 

Stokesia .....................................212 

Streptopus..................................811 

Striga .........................................532 

Strophostyles .............................389 

Stuartina....................................212 

Stuckenia...................................932 

Stylisma .....................................305 

Stylodon.....................................676 

Stylophorum..............................536 

Stylosanthes...............................390 

Styphnolobium ..........................390 

STYRACACEAE .......................664 

Styrax.........................................664 

Suaeda .......................................293 

Sullivantia .................................653 

Symphoricarpos.........................268 

Symphyotrichum .......................212 

Symphytum................................237 

SYMPLOCACEAE....................665 

Symplocarpus ............................704 

Symplocos..................................665 

Synandra ...................................470 

Syngonanthus............................789 

Syringa.......................................514 

Syringodium ..............................714 

Taenidia.......................................97 

Tagetes.......................................218 

Talinum .....................................573 

TAMARICACEAE ....................665 

Tamarix .....................................665 

Tanacetum.................................218 

Taraxacum ................................219 

TAXACEAE ................................65 

Taxodium.....................................59 

Taxus ...........................................66 

Teesdalia....................................256 

Tephrosia...................................390 

TETRACHONDRACEAE ........666 

Tetragonia ...................................76 

Tetragonotheca..........................219 

Tetraneuris ................................219 

Teucrium ...................................470 

Thalia.........................................812 

Thalictrum.................................589 

Thaspium.....................................98 

THEACEAE..............................667 

THELYPTERIDACEAE.............49 

Thelypteris ...................................50 

THEMIDACEAE ......................938 

Thermopsis ................................391 

Thinopyrum...............................923 

Thlaspi.......................................256 

Thuja ...........................................60 

THYMELAEACEAE ................668 

Thymophylla..............................219 

Thymus ......................................471 

Tiarella ......................................653 

Tilia............................................493 

TILIACEAE ..............................669 

Tillandsia...................................707 

Tipularia....................................835 

Tofieldia.....................................939 

TOFIELDIACEAE ...................939 

Tomanthera ...............................533 

Torenia ......................................540 

Torilis...........................................99 

Torreya ........................................66 

Torreyochloa .............................923 

Toxicodendron ............................83 

Trachelospermum .....................108 

Tradescantia..............................713 

Tragia ........................................352 

Tragopogon ...............................219 

Tragus....................................... 923 

Trapa......................................... 485 

Trautvetteria ............................. 591 

Trepocarpus................................ 99 

Triadenum ................................ 440 

Triadica..................................... 353 

Triantha .................................... 939 

Trianthema ................................. 76 

Tribulus..................................... 686 

Trichomanes ............................... 28 

Trichophorum........................... 785 

Trichostema .............................. 471 

Tricyrtis..................................... 812 

Tridens ...................................... 924 

Trientalis................................... 506 

Trifolium................................... 391 

Triglochin ................................. 806 

TRILLIACEAE ........................ 940 

Trillium..................................... 940 

Triodanis................................... 264 

Triosteum.................................. 269 

Triphora.................................... 835 

Triplasis .................................... 924 

Tripleurospermum.................... 220 

Tripsacum ................................. 924 

Trisetum.................................... 925 

Tristagma.................................. 696 

Triticum .................................... 925 

Triumfetta................................. 494 

TROPAEOLACEAE................. 669 

Tropaeolum............................... 669 

Tsuga .......................................... 65 

Tulipa........................................ 812 

TURNERACEAE ..................... 669 

Turritis ...................................... 256 

Tussilago................................... 220 

Typha ........................................ 945 

TYPHACEAE ........................... 945 

Ulex........................................... 394 

ULMACEAE............................. 669 

Ulmus........................................ 670 

UMBELLIFERAE ..................... 85 

Uniola ....................................... 925 

Urena ........................................ 494 

Urochloa ................................... 925 

Uropappus................................. 220 

Urtica ........................................ 673 

URTICACEAE ......................... 671 

Utricularia ................................ 475 

Uvularia .................................... 709 

UVULARIACEAE.................... 946 

Vaccaria.................................... 283 

Vaccinium................................. 338 

Vachellia Wight & Arnott 1834 394 

Valeriana .................................. 674 

VALERIANACEAE ................. 673 

Valerianella .............................. 674 

Vallisneria................................. 793 

Veratrum................................... 816 

Verbascum ................................ 654 

Verbena..................................... 676 

VERBENACEAE ..................... 674


Verbesina ..................................220 

Vernicia.....................................353 

Vernonia....................................221 

Veronica....................................552 

Veronicastrum...........................554 

Viburnum ....................................72 

Vicia ..........................................394 

Vigna .........................................396 

Viguiera.....................................223 

Vinca .........................................108 

Viola ..........................................677 

VIOLACEAE ............................677 

VISCACEAE.............................682 

VITACEAE ...............................682 

Vitex ..........................................471 

Vitis ...........................................684 

Vittadinia...................................223 

Vittaria ........................................46 

Vulpia........................................926 

Wahlenbergia............................264 

Waldsteinia................................622 

Warea.........................................256 

Websteria ...................................786 

Weigela ......................................316 

Wisteria......................................396 

Wolffia.......................................705 

Wolffiella...................................705 

Woodsia .......................................54 

WOODSIACEAE ........................51 

Woodwardia.................................21 

Xanthium...................................223 

Xanthorhiza...............................591 

Xanthosoma...............................705 

Xerophyllum..............................816 

XYRIDACEAE ..........................946 

Xyris...........................................946 

Yeatesia........................................71 

Youngia .....................................223 

Yucca .........................................688 

Zamia...........................................66 

ZAMIACEAE ............................. 66 

Zannichellia.............................. 949 

ZANNICHELLIACEAE .......... 949 

Zanthoxylum............................. 633 

Zea ............................................ 926 

Zenobia ..................................... 343 

Zephyranthes ............................ 699 

Zeuxine ..................................... 836 

Zigadenus ................................. 817 

Zinnia........................................ 224 

Zizania ...................................... 927 

Zizaniopsis ................................ 927 

Zizia ............................................ 99 

Ziziphus..................................... 594 

Zornia ....................................... 397 

Zostera ...................................... 950 

ZOSTERACEAE ...................... 950 

Zosterella .................................. 929 

Zoysia........................................ 927 

ZYGOPHYLLACEAE .............. 686

Part 7 - UNC Herbarium

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