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}
References: Tucker (1996)=Z.
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]
References: Tucker (1996)=Z.
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.
References: Tucker (1996)=Z.
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]
References: Tucker (1996)=Z.
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]
References: Tucker (1996)=Z.
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]
References: Tucker (1996)=Z.
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]
References: Tucker (1996)=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
2 Spikelets 0.8-3.2 mm long.
3 Spikelets 2.1-3.2 mm long.
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
3 Spikelets 0.8-2.0 mm long.
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
1 Spikelets 2.4-4.5 mm long.
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
1 Spikelets 0.9-2.3 mm long.
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
8 Spikelets glabrous, 1.1-1.8 mm long; blade surfaces pubescent or glabrous.
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
17 Spikelets glabrous, 1.1-1.8 mm long; blade surfaces pubescent or glabrous.
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
15 Spikelets glabrous, 1.1-1.8 mm long; blade surfaces pubescent or glabrous.
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]
References: Tucker (1996)=Z.
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)]
References: Tucker (1996)=Z.
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.
[= K] {not keyed}
* 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}
References: Tucker (1996)=Z.
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.
29 Spikelets 2.5-4.1 mm long.
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
29 Spikelets 1.3-2.5 mm long.
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]
{synonymy incomplete}
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]
{synonymy incomplete}
* 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;
synonymy incomplete}
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]
References: Tucker (1996)=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.
[= K] {not keyed}
* 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.

POTAMOGETONACEAE 931
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)

POTAMOGETONACEAE 932
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;
synonymy incomplete}
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).

POTAMOGETONACEAE 933
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

TRILLIACEAE 942
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

TRILLIACEAE 943
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}

TRILLIACEAE 944
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}

TRILLIACEAE 945
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

XYRIDACEAE 948
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]

XYRIDACEAE 949
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]
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BIBLIOGRAPHY 1005
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BIBLIOGRAPHY 1006
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

BIBLIOGRAPHY 1007
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

BIBLIOGRAPHY 1008
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

BIBLIOGRAPHY 1009
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

BIBLIOGRAPHY 1010
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

BIBLIOGRAPHY 1011
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

BIBLIOGRAPHY 1012
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

BIBLIOGRAPHY 1013
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

BIBLIOGRAPHY 1014
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

BIBLIOGRAPHY 1015
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