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

Aegilops Linnaeus 1753 (Goat Grass) References: Tucker (1996)=Z. 1 Spikelets cylindric; glumes with 4 awns; rachis disarticulating at maturity..........................................................Ae. cylindrica 1 Spikelets nearly ovate; glumes with 1 awn; rachis not disarticulating at maturity.................................................. Ae. neglecta * Aegilops cylindrica Host, Jointed Goat Grass. Mt, Pd (VA): disturbed areas; uncommon, introduced from s. Europe. [= C, F, G, HC, K, Z] * Aegilops neglecta Req. ex Bertoloni, Small Goat Grass. Cp (VA): disturbed areas; rare, introduced from s. Europe. Reported from Arlington County, VA. [= Z; Ae. ovata Linnaeus – C, G, HC, apparently misapplied; Ae. geniculata Roth – K, apparently misapplied] * Aegilops triuncialis Linnaeus. {MD} [= K] {not keyed at this time; synonymy incomplete}

Agropyron Gaertner 1770 (see Elymus, Elytrigia, Pascopyrum)

Agrostis Linnaeus 1753 (Bentgrass)

(also see Lachnagrostis and Polypogon) A genus of about 220 species, primarily temperate. References: Tucker (1996)=Z. 1 Palea 1/2-3/4 as long as the lemma, 0.6-1.2 mm long; plants introduced, often (though not always) in disturbed habitats;

plants flowering (collectively) June-October; [subgenus Agrostis]. 2 Ligule mostly 0.5-2 mm long, truncate; panicle branches naked towards the base, diffuse when in fruit, the spikelets

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

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

plant with rhizomes, without stolons.........................................................................................................A. gigantea 3 Leaves mostly 1-3 mm wide; inflorescence narrowly ovoid, the branches ascending at maturity, usually tan; plant

without rhizomes, with or without stolons. 4 Stolons well developed; leaves mostly < 5 cm long................................................A. stolonifera var. palustris 4 Stolons poorly developed or absent; leaves mostly > 7 cm long ......................... A. stolonifera var. stolonifera

1 Palea < 2/5 as long as the lemma, 0-0.5 mm long; plants native, typically in more or less natural habitats; plants flowering (collectively) March-November; [subgenus Vilfa]. 5 Lemma usually awned (sometimes unawned), the awn inserted near the apex, 4-10 mm long, straight, very delicate and

flexuous; annual, flowering April-June.......................................................................................................... A. elliottiana 5 Lemma awned or not, the awn (when present) inserted either near the middle of the lemma or near the apex, 0-6 mm

long, straight or bent, neither delicate nor flexuous; perennial, flowering (collectively) March-November. 6 Lemma with a 3-6 mm long, geniculate awn inserted near the middle; [of high elevation rock outcrops] ................. .................................................................................................................................................................A. mertensii 6 Lemma awnless or with a 0-3 mm long, straight awn inserted near the tip; [of various habitats].

7 Spikelets 1.2-2 mm long; anthers 0.3-0.6 mm long; plants flowering March-July........................... A. hyemalis 7 Spikelets (1.8-) 2.2-3.5 (-3.7) mm long; anthers (collectively) 0.3-1.2 mm long; plants flowering June-

November. 8 Leaves mostly involute, 1-2 (-3) mm wide; panicle branches mostly forking well beyond the middle....... .....................................................................................................................................................A. scabra 8 Leaves flat, 2-6 mm wide; panicle branches mostly forking at or below the middle.

9 Lemma 1.8-3 mm long, minutely but copiously scabrous (at 20× or more); anthers 0.7-1.2 mm long; spikelets (2.3-) 2.7-3.5 (-3.7) mm long, usually clustered near the tips of the branchlets; panicle branches scabrous; culms to 15 dm tall; [of wet savannas and other wet habitats of the Coastal Plain] ................................................................................................................... A. altissima

9 Lemma 1.4-2 mm long, glabrous; anthers 0.3-0.6 mm long; spikelets (1.8-) 2.2-2.7 (-3.2) mm long, usually not clustered near the tips of the branchlets; panicle branches glabrous to scabrous; culms to 10 dm tall; [of various habitats, nearly throughout our area]..........................................A. perennans

POACEAE 838

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

ponds, edges of swamp forests; rare (NC Watch List, VA Watch List). October-November. MA (?) and NJ south to se. LA, primarily on the Coastal Plain. [= F, HC, Z; < A. perennans – RAB, FNA GW, K; = A. perennans var. elata (Pursh) A. Hitchcock – C, G, S] * Agrostis capillaris Linnaeus, Rhode Island Bentgrass, Colonial Bentgrass, Browntop. Mt (NC, SC, VA), Pd, Cp (VA): meadows, roadsides, disturbed areas; uncommon, introduced from Europe (and possibly n. North America). June-August. [= C, FNA, K, Z; = A. tenuis Sibthorp – RAB, G, HC, S, W; > A. tenuis var. tenuis – F]

Agrostis elliottiana J.A. Schultes, Elliott's Bentgrass, Southern Bentgrass. Pd, Cp (GA, NC, SC, VA), Mt (GA, NC, SC): dry soils of barrens, fields, and rock outcrops; uncommon (VA Watch List). April-June. MD west to s. OH, and e. KS, south to panhandle FL and c. TX. [= RAB, C, F, FNA, G, HC, K, S, W, Z] * Agrostis gigantea Roth, Redtop, Black Bentgrass. June-October. Mt, Pd (GA), {provinces} (SC, VA). [= C, F, FNA, K, W, Z; < A. stolonifera – RAB, GW; = A. stolonifera Linnaeus var. major (Gaudin) Farwell – G; = A. alba – HC, misapplied; >< A. alba – S, misapplied]

Agrostis hyemalis (Walter) Britton, Sterns, & Poggenburg, Ticklegrass, Small Bentgrass. Mt, Pd, Cp (GA, NC, SC, VA: roadsides, other disturbed habitats; common. March-July. [= F, FNA, K, Z; < A. hyemalis – RAB (also see A. scabra); = A. hyemalis var. hyemalis – C, G; = A. hiemalis – GW, HC, orthographic variant; < A. hiemalis – S, W, orthographic variant (also see A. scabra var. scabra]

Agrostis mertensii Trinius, Arctic Bentgrass. Mt (NC, VA): in thin soil of high elevation rocky summits; rare (NC Rare). July-August. Circumboreal, in North America south to ME (Mt. Katahdin), NH (White Mountains), VT, NY (Adirondack Mountains), WV (Spruce Knob), TN (Roan Mountain, Mt. Leconte), NC (Roan Mountain, Big Yellow Mountain, Black Mountains), Québec, British Columbia, CO, UT (?), and AK. [= C, FNA, K, W, Z; > A. borealis Hartman – RAB, HC, S; > A. borealis Hartman var. americana (Scribner) Fernald – F, G]

Agrostis perennans (Walter) Tuckerman, Upland Bent, Autumn Bentgrass. Mt, Pd, Cp (GA, NC, SC, VA): . August-October. [= HC, Z; < A. perennans – RAB, FNA, GW, K, W (also see A. altissima); = A. perennans var. perennans – C, G, S; > A. perennans var. perennans – F; > A. perennans var. aestivalis Vasey – F]

Agrostis scabra Willdenow, Fly-away Grass, Rough Bentgrass. (GA, NC, SC, VA). (VA Watch List). June-November. [= FNA, GW, K, Z; < A. hyemalis – RAB, W; = A. hyemalis (Walter) Britton, Sterns, & Poggenburg var. scabra (Willdenow) Blomquist – C; > A. scabra var. scabra – F; = A. hyemalis (Walter) Britton, Sterns, & Poggenburg var. tenuis (Tuckerman) Gleason – G; = A. scabra var. scabra – HC] * Agrostis stolonifera Linnaeus var. palustris (Hudson) Farwell, Creeping Bentgrass. June-October. [= C; < A. stolonifera – RAB, FNA GW, W (also see A. gigantea); = A. alba Linnaeus var. palustris (Hudson) Persoon – F, misapplied; = A. stolonifera var. compacta Hartman – G; = A. palustris Hudson – HC, Z; < A. stolonifera – K; < A. alba – S, misapplied] * Agrostis stolonifera Linnaeus var. stolonifera. June-October. [= C, G; < A. stolonifera – RAB, FNA, GW, W (also see A. gigantea); = A. alba Linnaeus var. alba – F, misapplied; = A. stolonifera – HC, Z; < A. stolonifera – K; >< A. alba – S, misapplied] * Agrostis canina Linnaeus, Brown Bentgrass, Velvet Bentgrass, ranges south to DE, se. PA (Rhoads & Klein 1993), WV, and TN (Kartesz 1999). [= C, FNA, K] {not keyed at this time; synonymy incomplete}

Aira Linnaeus (Hair Grass) References: Tucker (1996)=Z. 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 Maiden-cane. 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

Andropogon Linnaeus 1753 (Broomsedge, Bluestem)

(also see Bothriochloa and Schizachyrium) A genus of about 100-110 species, mainly tropical. The difference between this treatment and that in RAB may cause some users to react with skepticism, dismay, or alarm, but I am confident that it represents a much truer description of the genus. Campbell's work (1983, et seq.) has greatly clarified the taxonomy of Andropogon in e. North America. Great confusion and disagreement were previously the rule in dealing with the A. virginicus-A. glomeratus complex. Campbell's careful morphologic work has provided workable technical characters which distinguish the taxa he recognizes. I have generally followed Campbell (1983, et seq.) in his circumscriptions of taxa. I disagree, however, with his strongly morphologic species concept and the basis for his decisions regarding the rank of the taxa (species, variety, and "variant," an informal, English name for a subvarietal entity). Campbell bases the rank recognition of taxa on their "morphological distance" from one another, as determined by the sum of non-matching characters out of 33 characters analyzed. In general, he regards species as separated by a morphologic distance of 9 or more, varieties by 6 or more, and variants by 3 or more. Such an approach fails to take into account additional evidence of the ecological preferences, geographic distributions, reproductive isolation, evolutionary pathways, and population biology of the taxa. Evidence presented by Campbell (1983) is useful in determining a more meaningful assignment of taxonomic rank. For instance, he states that "there are ample opportunities for gene flow between taxa because they frequently grow together and flower at the same time of day and (mostly) at the same time of year. I have observed two taxa growing within one to three meters of one another over our hundred times. In only five of these opportunities for hybridization were there plants whose intermediate morphology suggested that they were hybrids. In the rare instances where hybridization does take place, there are few mature hybrid individuals. I have found only twelve putative hybrid individuals in the five localities where hybridization is suspected. The parents outnumber these hybrids by between five and one hundred or more to one." In discussing A. virginicus var. glaucus (here treated as A. capillipes) he states "the drylands variant ... produces generally shorter raceme sheaths, racemes, and spikelets; its flowers are more frequently chasmogamous, and unlike the wetlands variant, it has no hairs below the raceme sheath. In addition, it grows in better-drained soil and has a narrower geographic range ... I have seen these taxa growing within one to three meters of one another at three localities in northwestern Florida. At only one of these was there difficulty in classifying any individual: a single plant on a slope between a bog inhabited by the wetlands variant and a roadside lined with the drylands variant..." Some additional examples would be Campbell's discussion of several closely related taxa in the A. glomeratus complex. "Andropogon glomeratus var. glomeratus and the robust variant of var. pumilus have inflorescences so similar in shape that most previous workers have united them and have overlooked the differences between them. The robust variant is taller, usually with rather smooth sheaths and with shorter, more ciliate, and darker ligules, narrower raceme sheaths, and lower glume keels that are scabrous below the middle. Although both taxa grow in wet sites, the robust variant is weedier, shows a greater tolerance for drier conditions and various soil types, and has a wider geographic range." His discussion of A. glomeratus var. glaucopsis and var. hirsutior is also worth repeating. "Often the two grow together in populations of thousands of individuals. Because they grow together so frequently and are morphologically so alike, the possibility that they are not distinct taxa but merely genetic segregates of one another has been carefully considered. Based on observations of several hundred seedlings grown from seeds from both taxa (growing together in nature), there is no evidence for [mere] genetic segregation. The glaucousness/greenness and pubescence/glabrousness of the stem sheaths are discernible in the seedlings within a few weeks of germination." Taxa differing in numerous morphologic characters, with different (though overlapping) geographic ranges, with different ecological preferences (often rather narrowly segregated by hydrology), and (when they do occur in proximity to one another) showing little or no sign of introgression or hybridization are probably better treated as biological species. Thus, I have treated a number of Campbell's varieties as species. Several of his "variants" also seem to warrant taxonomic recognition, at varietal or specific rank; in fact, he subsequently elevated several (Campbell 1986). References: Campbell (1983)=Z; Campbell in FNA (2003a). Key adapted in part from Z. Identification notes: A thorough understanding of the architecture of the inflorescences of Andropogon is necessary in order to identify them successfully. The parts will be described, beginning from the apex of a branch of the inflorescence. Spikelets occur in pairs, the sessile spikelet (usually just referred to as the spikelet) and the pedicelled spikelet, which is usually vestigial or absent (except in A. gerardii) and sterile (except in A. gerardii, where it is staminate). The first or lower glume of the sessile spikelet has two keels, and the presence and location of antrorse prickle hairs (scabrousness) is an important character in the A. glomeratus complex. The length of the sessile spikelet is an important character; it should be measured exclusive of the awn, borne at the apex of the lemma. Awn length is also a useful taxonomic character. The pedicelled spikelet is borne on the pedicel, which is attached at the base of the sessile spikelet and typically angles away from it at about a 45 degree angle. The rachis internode extends from the base of one sessile spikelet to the next sessile spikelet above, breaking apart (upon dehiscence) just below the next spikelet and remaining attached to the sessile spikelet below. The dispersal unit consists of a sessile spikelet sitting in the V shape formed by (on one side) the pedicel and pedicelled spikelet and (on the other side) the rachis

POACEAE 841

internode. Both the pedicel and the rachis internode are usually pubescent with long hairs, and the color of those hairs and their distribution are useful characters. While the dispersal units are still attached to one another, the rachis internodes form a continuous and more-or-less straight rachis. The dispersal units attached together in an unbranched sequence are termed a raceme, whose length is a useful character. Two or more racemes are attached digitately at the summit of the peduncle (in Schizachyrium only a single raceme is found). The number of racemes attached is an important character. A raceme sheath subtends the peduncle, often more or less surrounding the peduncle and the racemes. The length of the peduncle (distance between the points of attachment of the raceme sheath and the racemes) is an important character. The length and width (at its widest point) of the raceme sheath are very useful characters, used throughout the key. The racemes, peduncle and subtending raceme sheath make up an inflorescence unit. The overall inflorescence is more-or-less complexly branched; its overall size and shape are very useful in recognizing the various taxa, but variation in such a subjective (and environmentally plastic) character has added to the taxonomic confusion in Andropogon. The use of inflorescence shape in the key has been minimized, but is often mentioned in the discussion of each species. The number of inflorescence units per plant varies from species to species, in some species rarely exceeding 10, in others ranging upwards to 500 or 600. The absence or presence of hairs immediately below the raceme sheath is useful in some groups. There are several important characters of the foliage. A. capillipes and A. glaucopsis have culm sheaths and leaf blades that are strongly glaucous; this is usually very obvious, but can be tested for by running the finger along the surface of the leaf (a white coating of wax will come off on the finger). The key often calls for the ligule length; measure the longest portion of the undivided portion of the ligule. The ligule often has an erose or ciliate upper margin; measure the length of the ciliations. The length of leaf blade is measured from the ligule to the leaf apex; do not include the leaf sheath, which is often long and (especially late in the year) only loosely sheathing the culm or even divergent it. Whether the culm is antrorsely scabrous or smooth is better determined by touch than by sight. Choose several mid-culm sheaths, run one's finger downwards and upwards along the sheath surface (near the collar is best). If the sheath is antrorsely scabrous one will feel a somewhat greater resistance to moving the finger downwards than upwards. 1 Pedicellate spikelet staminate, as large as the sessile, fertile spikelet; sessile spikelets > 7 mm long; [section Andropogon] .... ...................................................................................................................................................................................A. gerardii 1 Pedicellate spikelet sterile, vestigial or absent; sessile spikelets < 7 mm long; [section Leptopogon].

2 Leaves strongly glaucous (often nearly white with a powdery wax that can be rubbed off on the fingers), glabrous. 3 Ligules (0.9-) 1.5 (-2.0) mm long, with ciliations 0-0.2 mm long; leaf blades usually (33-) avg. 40 (-75) cm long;

pubescence beneath raceme sheaths moderate to dense; raceme sheaths (2.0-) 2.4-3.6 (-4.4) cm long, (1.3-) 2.0-2.5 (-3.0) mm wide.......................................................................................................................................A. glaucopsis

3 Ligules (0.2-) 0.4 (-0.5) mm long, with ciliations 0.3-1.2 mm long; leaf blades (12-) avg. 19 (-38) cm long; pubescence beneath raceme sheaths absent to dense; raceme sheaths (2.1-) 2.9-4.3 (-6.0) cm long, (2.7-) 3.1-3.8 (-5.5) mm wide. 4 Summit of branchlet below attachment of raceme sheath pubescent with hairs 2-4 mm long; raceme sheaths

(2.4-) 3.2-4.8 (-6.0) cm long; spikelets (3.0-) 3.5-3.9 (-4.4) mm long; racemes (1.5-) 2.0-3.0 (-4.0) cm long; leaves 2.5-6.5 mm wide, averaging 5 mm; upper floret lemma awn 0.9-2.1 mm long, averaging 1.4 mm .........

.............................................................................................................A. capillipes var. 1 ["wetland variant"] 4 Summit of branchlet below attachment of raceme sheath glabrous; raceme sheaths (2.1-) 2.6-3.8 (-4.9) cm

long; spikelets (2.6-) 3.2-3.5 (-3.9) mm long; racemes (1.4-) 1.7-2.4 (-3.2) cm long; leaves 2-5 mm wide, averaging 3.5 mm; upper floret lemma awn 0.6-1.5 mm long, averaging 1.1 mm..............................................

............................................................................................................. A. capillipes var. 2 ["dryland variant"} 2 Leaves green (to somewhat glaucous, but never powdery white), pubescent or glabrous.

5 Upper culm sheaths distinctly broadened and strongly overlapping, often largely hiding the raceme sheaths before senescence (but in some forms with the raceme sheaths strongly exserted); culms mostly < 1 m tall (to 1.4 m tall) .

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

> 1 m tall (except A. perangustatus, A. tracyi, and small forms of A. virginicus). 6 Many or all peduncles longer than the subtending raceme sheaths at maturity, racemes then fully exserted

above the apex of the raceme sheath. 7 Inflorescence branches arching outwards in pronounced curves; racemes (1.2-) 1.5-2.1 (-2.6) cm long;

awn (0.2-) avg. 0.7 (-1.1) cm long; spikelets (4.1-) 4.4-4.6 (-5.0) mm long...................A. brachystachyus 7 Inflorescence branches erect; racemes (2.2-) 2.6-6 cm long; awn 0.5-2.0 cm long; spikelets (4.3-) 4.9-6.5

(-7.5) mm long. 8 Lower glumes more or less folded; stamen 1; racemes (2.2-) 2.6-4.3 (-5.3) cm long; awn (0.5-) avg.

0.8 (-1.6) cm long; spikelets (4.3-) 4.9-5.4 (-6.1) mm long ................................................A. arctatus 8 Lower glumes flat; stamens 3; racemes 3-6 cm long; awn 1-2 cm long; spikelets (4.5-) 5-6.5 (-7.5)

mm long ....................................................................................................A. ternarius var. ternarius 6 Peduncles all shorter than the subtending raceme sheaths at maturity, at least the bases of the racemes not

exserted above the apex of the raceme sheath.

POACEAE 842

9 Inflorescence units with (2-) 4-7 (-13) racemes; raceme sheaths (4.1-) 5.3-8.0 (-10-1) mm wide; hairs of the rachis internode and pedicel yellow-tawny when dry ............................................................A. mohrii

9 Inflorescence units with 2-5 (-7) racemes; raceme sheaths (1.5-) 2.0-4.8 (-6.3) mm wide; hairs of the rachis internode and pedicel gray to whitish when dry. 10 Postflowering peduncles < 10 mm long.

11 Culm sheaths antrorsely scabrous (often hirsute as well); leaf blades usually > 35 cm long. 12 Ligules (0.6-) 0.8 (-1.3) mm long (usually < 1 mm long), with ciliations 0.2-0.9 mm long;

raceme sheaths (1.5-) 2.0-2.5 (-3.0) mm wide (usually < 2.5 mm wide); keels of first glume often scabrous below the middle.................................................... A. tenuispatheus

12 Ligules (0.7-) 1.2 (-2.2) mm long (usually > 1 mm long), with ciliations 0.0-0.3 mm long; raceme sheaths (2.0-) 2.4-3.4 (-4.7) mm wide (usually > 2.5 mm wide); keels of first glume scabrous only above the middle, smooth below. 13 Inflorescences oblong to obpyramidal; spikelets (3.8-) 4.1-4.4 (-5.0) mm long;

anthers usually not marcescent within spikelet; mature peduncles (4 -) 11-35 (-60) mm long (usually some of them > 10 mm long) ......... A. glomeratus var. glomeratus

13 Inflorescences (linear to) oblong; spikelets (3.4-) 3.6-3.8 (-4.6) mm log; anthers usually marcescent within spikelets; peduncles (2-) 3-5 (-8) mm long........................

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

var. pumilus). 14 Leaves glabrous.

15 Ligules (0.8-) 1.1 (-1.5) mm long, with ciliations 0-0.1 mm long; basal leaves often filiform, < 1.5 mm wide, strongly erect ............................................A. perangustatus

15 Ligules (0.2-) 0.5 (-0.8) mm long, with ciliations 0.2-1.3 mm long; basal leaves usually > 2 mm wide, soon arching. 16 Culm internodes green (or glaucous just below the node only); raceme sheaths

(2.2-) 2.5-3.8) (-4.5) cm long, (1.7-) 2.4-3.1) (-4.0) mm wide; peduncles (1-) 4-9 (-30) mm long; racemes 2 (-3) per inflorescence unit .......................................

................................................................................. A. virginicus var. decipiens 16 Culm internodes glaucous; raceme sheaths (2.8-) 3.3-4.7 (-6.7) cm long, (3.0-)

3.2-3.8 (-5.2) mm wide; peduncles (2-) 3-4 (-6) mm long; racemes 2-4 (-7) per inflorescence unit, at least some inflorescence units (especially at culm and branch apices) with 3 or more racemes................................................................

.................................................. A. virginicus var. virginicus ['smooth variant'] 14 Leaves pubescent, at least on the margin near the collar.

17 Keels of first glume often scabrous below the middle; leaves usually > 44 cm long... .......................................................................................................... A. tenuispatheus 17 Keels of first glume scabrous only above middle; leaves usually < 31 cm long.

18 Pubescence of young culm sheaths appressed; spikelets usually > 4 mm long; hairs on rachis internode and sterile pedicel dense, long; callus hairs 1.5-5 mm long ................................................................................................A. longiberbis

18 Pubescence of young culm sheaths spreading; spikelets mostly < 4 mm long; hairs on rachis internode and sterile pedicel rather sparse and short; callus hairs 1-2.5 mm long. 19 Raceme sheaths (2.2-) 2.5-3.8 (-4.5) cm long, (1.7-) 2.4-3.1 (-4.0) mm

wide; racemes 2 (3) per inflorescence unit; spikelets (3.0-) 3.3-3.6 (-4.0) mm long ........................................................... A. virginicus var. decipiens

19 Raceme sheaths (2.3-) 3.4-5.2 (-6.7) cm long, (2.7-) 3.3-4.0 (-5.5) mm wide; racemes 2-5 (-7) per inflorescence unit; spikelets (2.9-) 3.7-3.9 (-4.7) mm long................................................... A. virginicus var. virginicus

10 Postflowering peduncles > 15 mm long. 20 Culm sheaths antrorsely scabrous (often hirsute as well).

21 Ligules (1.0-) 1.2 (-2.0) mm long, with ciliations 0-0.3 mm long; keels of first glume scabrous only above middle................................................ A. glomeratus var. glomeratus

21 Ligules (0.6-) 0.8 (-1.3) mm long, with ciliations 0.2-0.9 mm long; keels of first glume often scabrous below middle .................................................................... A. tenuispatheus

20 Culm sheaths not scabrous (often hirsute). 22 Culms < 1.2 m tall; leaf blades < 30 cm long and < 3 mm wide; inflorescence units rarely

> 20/culm. 23 Raceme sheaths (2.2-) 2.5-3.8 (-4.5) cm long; spikelets (3.0-) 3.3-3.6 (-4.0) mm

long; leaf blades (2.5-) 3.6 (-5.5) mm wide ..................... A. virginicus var. decipiens 23 Raceme sheaths (2.6-) 4.1-6.6 (-8.5) cm long; spikelets (3.0-) 3.4-5.1 (-5.5) mm

long; leaf blades (0.8-) 1.8 (-3.0) mm wide. 24 Ligules (0.8-) 1.1 (-1.5) mm long, with ciliations 0-0.1 mm long........................

POACEAE 843

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

22 Culms usually > 1.2 m tall; leaf blades often > 30 cm long and > 3 mm wide; inflorescence units usually > 20/culm. 25 Inflorescence branches arching outwards in pronounced curves; awn mostly < 1 cm

long; spikelets (4.1-) 4.4-4.6 (-5.0) mm long; anther > 1.7 mm long........................... ........................................................................................................A. brachystachyus 25 Inflorescence branches erect; awn mostly >1 cm long; spikelets (3.0-) 3.3-3.8 (-4.5)

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

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

26 Raceme sheaths (1.7-) 2.4-3.1 (-4.0) mm wide; keels of first glume scabrous only above middle; culms < 1.7 m tall; leaves < 35 cm long and 5.5 mm wide...

................................................................................. A. virginicus var. decipiens * Andropogon arctatus Chapman, Florida Bluestem. Cp (NC): moist disturbed ground; rare, apparently introduced from farther south (NC Watch List). This curious record (the specimen at GH, collected by Randolph and Randolph in 1922 in Pamlico County, NC, annotated as A. arctatus by Campbell) is likely a waif. The species is native to pinelands from n. FL west to w. panhandle of FL and adjacent s. AL, south to s. FL. [= FNA, HC, K, S, Z] Andropogon brachystachyus Chapman, Shortspike Bluestem. Cp (GA, SC): moist to wet pinelands, natural pond margins, bogs, disturbed roadsides; rare (GA Special Concern). Se. SC (McMillan et al. 2002) south to FL, south to s. FL, west to e. FL Panhandle. A. brachystachyus is considered by some to range north to NC. [= FNA, K, Z; = A. brachystachys – GW, HC, S, orthographic variant] Andropogon capillipes Nash var. 1, Wetland White Bluestem. Cp (GA, NC, SC): wet savannas, ditches adjacent to savannas, depressional wetlands; common. September-October. S. NJ south to s. FL and west to e. TX; also in the Bahamas (Sorrie & LeBlond 1997). Campbell (1983) informally describes two "variants" of this species (which he treats at the varietal level, as A. virginicus var. glaucus). A. capillipes is clearly a species distinct from A. virginicus; moreover, the substantial morphological and ecological differences between Campbell's two "variants" (which he describes as nearly always sharply distinct, even when growing in close proximity) warrant recognition as good species, or at least as varieties. [< A. virginicus – RAB; < A. virginicus var. glaucus Hackel – F, FNA; < A. capillipes – GW, HC, K, S; = A. virginicus var. glaucus "wetlands variant" – Z; = A. virginicus var. dealbatus Mohr ex Hackel] Andropogon capillipes Nash var. 2, Dryland White Bluestem. Cp (GA, NC, SC): dry to mesic pine flatwoods, sandhills, adjacent roadbanks; uncommon. September-October. Se. NC south to s. FL and west to s. AL. See A. capillipes var. 1 for discussion of these two taxa. The type of A. capillipes (collected by A.H. Curtiss in FL) is of this taxon; Nash (1900) states that it occurs "in dry soil, North Carolina to Florida" and emphasizes that it is "abundantly distinct from A. virginicus, to which it is related." [< A. virginicus – RAB; < A. virginicus var. glaucus Hackel – F, FNA; < A. capillipes – GW, HC, K, S; = A. virginicus var. glaucus "drylands variant" – Z] Andropogon elliottii Chapman. Cp, Pd, Mt (GA, NC, SC, VA): dry to moist forests, woodlands, fields, and disturbed areas; common (uncommon in Mountains). September-October. Widespread in se. United States, from s. NJ west to s. IN, s. IL, s. MO, south to s. FL and TX. Campbell (1983) argued that the name A. elliottii should be replaced by A. gyrans; Ward (2004c) argues for retention of the traditional A. elliottii. [= HC; > A. elliottii – RAB, S; > A. campyloracheus Nash – RAB, S; = A. gyrans Ashe – C, W; = A. gyrans var. gyrans – FNA, K, Z; > A. elliottii var. elliottii – F, G; > A. elliottii var. gracilior Hackel – F, G; > A. elliottii var. projectus Fernald & Griscom – G] Andropogon floridanus Scribner, Florida Bluestem. Cp (GA): longleaf pine sandhills; rare. September-October. S. GA west to FL Panhandle, south to s. FL. Reported for e. and s. GA (FNA, Jones & Coile 1988). [= FNA, HC, K, S] {not keyed at this time} Andropogon gerardii Vitman, Big Bluestem, Turkeyfoot. Cp, Pd, Mt (GA, NC, SC, VA): in a wide variety of habitats, usually rather dry, such as sandhills, glades, cliffs, and rock outcrops, in the Piedmont in woodlands, former prairie-like sites, woodlands, open forests, and river-scour grasslands, in the Mountains in glades, riverside scour areas, and rarely in grassy balds, ascending to at least 1600 m over mafic rocks (on Old Field Bald, Watauga and Ashe counties, NC); common. July-October. Québec west to Saskatchewan, south to FL and AZ. Some favor treating A. hallii Hackel as a subspecies of A. gerardii (Wipff 1996c). I do not agree, but if that course is followed, then our eastern taxon should be known as A. gerardii ssp. gerardii. [= RAB, C, FNA, G, GW, HC, K, W; > A. gerardii var. gerardii – F; = A. provincialis Lamarck – S] Andropogon glaucopsis Elliott, Chalky Bluestem. Cp (GA, NC, SC, VA): wet savannas, pine flatwoods, ditches, wet disturbed sites; uncommon (VA Watch List). September-October. Se. VA south to c. peninsular FL and west to e. TX. The extent of the western Gulf Coastal Plain distribution (to the West Gulf Coastal Plain of w. LA and e. TX) is based on specimens (at BRIT) and sight records (B. Sorrie, pers. comm.). Although sometimes included in the past in either A. glomeratus or A. virginicus, this species is distinctive and easily recognized in the field (even from a car at 60 m.p.h.) by the combination of blue color, height of well over 1 m (taller than the other glaucous bluestems), and semi-bushy inflorescence. [= GW, K; < A. virginicus – RAB; = A. virginicus var. glaucopsis (Elliott) A.S. Hitchcock – F, HC; = A. glomeratus var. glaucopsis (Elliott) A.S. Hitchcock – FNA, Z; < A. glomeratus – S]

POACEAE 844

Andropogon glomeratus (Walter) Britton, Sterns, & Poggenburg var. glomeratus. Cp, Pd, Mt (GA, NC, SC, VA): swamps, wet savannas, pine flatwoods, wet disturbed sites; common. September-October. S. MA south to c. peninsular FL and west to s. MS, primarily on the Coastal Plain, but scattered inland to w. PA, WV, c. KY, c. TN and AR. [= FNA, K, Z; < A. virginicus – RAB; = A. virginicus var. abbreviatus (Hackel) Fernald & Griscom – C, F, G, GW; < A. glomeratus – HC, S, W] Andropogon glomeratus (Walter) Britton, Sterns, & Poggenburg var. hirsutior (Hackel) C. Mohr. Cp (GA, NC, SC, VA): wet savannas, pine flatwoods, adjacent ditches, other wet disturbed sites; common. September-October. E. MD south to c. peninsular FL west to se. LA. This taxon should be recognized at the specific level, but the appropriate combination has not been made. [= FNA, K, Z; < A. virginicus – RAB; ? A. virginicus var. glaucopsis (Elliott) A.S. Hitchcock – G, misapplied; = A. virginicus var. hirsutior (Hackel) A.S. Hitchcock; < A. glomeratus – HC, S] Andropogon longiberbis Hackel, Longbeard Bluestem. Cp (GA, NC, SC): dry sandy soils of sandhills and dunes; rare (GA Special Concern, NC Watch List). September-October. Se. NC south to s. and w. FL, and in the Bahamas. [= FNA, HC, K, S, Z] Andropogon mohrii (Hackel) Hackel ex Vasey, Tawny Bluestem, Bog Bluestem. Cp (GA, NC, SC, VA): wet savannas, sphagnous bogs; rare (GA Special Concern, NC Rare, VA Rare). September-October. Se. VA south to n. FL, west to LA. [= RAB, C, F, G, GW, HC, K, S; = A. liebmannii Hackel var. pungensis (Ashe) C.S. Campbell – FNA, Z] Andropogon perangustatus Nash, Narrow-leaved Bluestem. Cp (GA, NC, SC, VA): clay-based Carolina bays and boggy wetlands; rare (NC Watch List, VA Watch List). August-October. E. VA south to c. peninsular FL, east to e. TX. Growth form, general appearance, and habitat (dense bluish tussocks with very narrow leaves and long ligules, growing in wet areas such as clay-based Carolina bays) make A. perangustatus readily recognizable. [= HC, S; = A. gyrans Ashe var. stenophyllus (Hackel) C.S. Campbell – FNA, K, Z; = A. elliottii Chapman var. stenophyllus (Hackel) D.B. Ward] Andropogon tenuispatheus (Nash) Nash. Cp (GA, NC, SC, VA), Pd (GA, NC, SC): maritime wet grasslands, brackish marsh edges, moist disturbed sites; common (VA Watch List). September-October. Se. VA and c. OK south to s. FL and w. TX, also south into Central America and the Caribbean. [< A. virginicus – RAB; = A. glomeratus (Walter) Britton, Sterns, & Poggenburg var. pumilus Vasey ex Dewey – FNA, K, Z ("robust variant"); < A. glomeratus – HC, S] Andropogon ternarius Michaux var. ternarius, Splitbeard Bluestem. Cp, Pd, Mt (GA, NC, SC, VA): dry to moist soils; common (uncommon in Mountains). September-October. Var. ternarius ranges from DE west to KY and s. MO, south to FL and TX. Var. cabanisii (Hackel) Fernald & Griscom is endemic in s. and c. peninsular FL. [= FNA, K, Z; < A. ternarius – RAB, C, G, W; > A. ternarius var. ternarius – F; > A. ternarius var. glaucescens (Scribner) Fernald & Griscom – F; = A. ternarius – HC, S] Andropogon tracyi Nash, Tracy's Bluestem. Cp, Pd (GA, NC, SC): dry sandy or clayey soils of sandhills, disturbed sites; rare (NC Watch List). September-October. E. NC south to s. FL and west to MS. [= FNA, HC, K, S, Z] Andropogon virginicus Linnaeus var. decipiens C.S. Campbell, Deceptive Bluestem. Cp (GA, NC, SC, VA): savannas, flatwoods, maritime wet grasslands, disturbed pinelands; uncommon (VA Watch List). September-October. Se. VA south to s. FL and west to w. FL; also in the Bahamas (Sorrie & LeBlond (1997). [= FNA, K, Z (1986); < A. virginicus – RAB, S; < A. virginicus var. virginicus – F, G, HC; = A. virginicus var. virginicus – Z (1983 – "deceptive variant")] Andropogon virginicus Linnaeus var. virginicus, Old-field Broomstraw, Broomsedge, "Sedge Grass", "Sage Grass". Cp, Pd, Mt (GA, NC, SC, VA): old fields, roadbanks, disturbed sites; common. September-October. Widespread, from MA west to MI and e. KA, south to FL and e. TX, and in the Caribbean and Central America. Campbell (1983) recognized 3 "variants" within A. virginicus var. virginicus; the "deceptive variant" he later (1986) described formally as var. decipiens (see above). The "old-field variant" is the common "variant" in our area, occurring abundantly throughout the state. It has green stem internodes and the leaves usually pubescent, at least on the margins near the collar. The "smooth variant" is known only from the Coastal Plain and is apparently rare in our area, known from NC and SC (Berkeley and Marion counties; P. McMillan, pers. comm.). It has glaucous stem internodes and glabrous leaves. It is unclear whether the "smooth variant" warrants taxonomic recognition. [= FNA, K, Z ("oldfield variant" and "smooth variant"); < A. virginicus – RAB, S, W; < A. virginicus var. virginicus – C; < A. virginicus var. virginicus – G, HC (also see var. decipiens); >< A. virginicus var. virginicus – F; > A. virginicus var. tetrastachyus (Elliott) Hackel – F]

Anthenantia Palisot de Beauvois (Silkyscale) A genus of 3 species, of se. North America (or 4-5 species of se. North America and tropical America, if Leptocoryphium is included in Anthenantia). Clayton & Renvoize (1986) state that "Anthenantia is the etymologically correct version of three alternative spellings given by Beauvois." References: Wipff in FNA (2003a); Crins (1991)=Z; Kral (2004)=Y; Clayton & Renvoize (1986). 1 Leaves mostly 3-5 mm wide, ascending to erect, not squarrose (lacking a sharp bend outward at the summit of the sheath),

medium green, with a very short taper to a blunt or rounded apex, the lower sheaths crowded and keeled (therefore distichous), generally suffused with purple; spikelets usually purple (fading tan); leaf margins scaberulous .................A. rufa

1 Leaves mostly 5-10 mm wide, spreading, usually squarrose (with a sharp bend outward at the summit of the sheath), yellowish green, with a long taper to a sharp apex, the lower sheaths not crowded, keeled, or distichous, green; spikelets usually green (fading yellow); leaf margins papillose-ciliate towards the base ........................................................... A. villosa

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

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

POACEAE 845

Se. NC south to n. FL and west to w. LA. A. rufa inhabits much wetter habitats than the similar A. villosa, and is more typical of the outer Coastal Plain. Plants without culms are reminiscent of the Liliaceae. [= FNA, Y; = Anthaenantia rufa – RAB, GW, HC, K, S, Z, orthographic variant]

Anthenantia villosa (Michaux) Palisot de Beauvois, Green Silkyscale. Cp (GA, NC, SC): sandhills, especially in submesic swales; uncommon (rare in the outer Coastal Plain). September-October. Se. NC south to s. FL and west to e. TX. A. villosa is found in drier habitats than A. rufa, most typically in upland swales in the sandhills. Kral (2004) has segregated a new species, A. texana Kral, of the w. Gulf Coastal plain, previously confused with A. villosa. [= Y; < Anthaenantia villosa – RAB, HC, K, S, Z, orthographic variant; < Anthenantia villosa – FNA]

Anthoxanthum Linnaeus (Vernal Grass) References: Tucker (1996)=Z. 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 appressed-

pubescent................................................................................................................................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

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varietal status for A. stricta and A. beyrichiana. [= K, Y; < A. stricta – RAB, FNA, GW, HC, S, V, Z; = A. stricta Michaux var. beyrichiana (Trinius & Ruprecht) D.B. Ward – X] Aristida condensata Chapman, Big Three-awn. Cp (GA, NC, SC): dry sandy soils of sandhills; rare (NC Watch List, SC Rare). August-October. Sc. NC south to s. FL, west to s. MS (Sorrie & Leonard 1999). [= RAB, FNA, HC, K, S, Z] Aristida curtissii (A. Gray ex S. Watson & Coulter) Nash, Curtiss's Three-awn. Cp, Pd (GA, NC, SC, VA): roadsides, disturbed areas, bare eroding soil; uncommon. August-October. ME west to WY, south to FL, AR, OK, and CO, perhaps largely or entirely adventive in our area. See Z for a discussion of the rationale for reducing A. curtissii to a variety of A. dichotoma. C reduces it to a variety of the more western A. basiramea Engelmann ex Vasey. For now, and for simplicity, I prefer to retain the two as species. [= RAB, G, HC, S; = A. basiramea Engelmann ex Vasey var. curtissii (A. Gray ex S. Watson & Coulter) Shinners – C; = A. dichotoma Michaux var. curtissii A. Gray – F, FNA, K, W, Z] Aristida dichotoma Michaux, Fork-tip Three-awn. Cp, Pd, Mt (GA, NC, SC, VA): roadsides, fields, disturbed areas, bare eroding soil; common. August-October. ME west to WI, south to FL and TX. See A. curtissii for comments. [= RAB, C, G, HC, S; = A. dichotoma var. dichotoma – F, FNA, K, W, Z] Aristida lanosa Muhlenberg ex Elliott, Woollysheath Three-awn. Cp, Pd (GA, NC, SC, VA): dry sandy soils of sandhills and fields; common, rare in Piedmont (VA Watch List). August-October. NJ south to FL, west to TX, north in the interior to MO and OK. [= RAB, C, FNA, G, HC, K, S, Z; > A. lanosa var. lanosa – F; A. lanosa var. macera Fernald & Griscom – F] Aristida longespica Poiret var. geniculata (Rafinesque) Fernald, Eastern Slim-spike Three-awn. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): disturbed areas; common? August-October. The distribution and habitats of the 2 varieties in our area are poorly known, pending further field and herbarium investigation. [= C, F, FNA, HC, K, Z; < A. longespica – RAB, W; > A. intermedia Scribner & Ball – F, G, S; > A. longespica – G] Aristida longespica Poiret var. longespica, Eastern Slim-spike Three-awn. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): disturbed areas; uncommon? August-October. The distribution and habitats of the 2 varieties in our area are poorly known, pending further field and herbarium investigation. [= C, F, FNA, HC, K, Z; < A. longespica – RAB, G, W; = A. longespica – S] Aristida mohrii Nash, Mohr's Three-awn. Cp (GA, SC): sandhills; rare. August-October. Panhandle FL and sw. GA west to s. AL; apparently disjunct in SC (Chesterfield and Richland counties). [= FNA, HC, K, S, Z] Aristida oligantha Michaux, Prairie Three-awn. Pd, Cp, Mt (GA, NC, SC, VA): roadsides, fields, disturbed areas; common. August-October. VT west to SD, south to FL and TX, scattered elsewhere as a weed. [= RAB, C, F, FNA, G, HC, K, W, S, Z] Aristida palustris (Chapman) Vasey, Longleaf Three-awn. Cp (GA, NC, SC): wet pine savannas, limesink depressions; uncommon. August-October. Se. NC south to FL, west to TX; apparently disjunct on the Cumberland Plateau of KY. [= C, FNA, K, S, Z; = A. affinis (Schultes) Kunth – RAB, F, G, GW, HC, misapplied] Aristida purpurascens Poiret, Arrowfeather. Cp, Pd, Mt (GA, NC, SC, VA): dry habitats, especially in dry sandy soils; common. August-October. MA west to WI and KS, south to FL and TX. In the Sandhills occurring in two forms, one green, the other strongly glaucous-blue. [= RAB, C, G, HC, S, W; > A. purpurascens var. purpurascens – F; > A. purpurascens var. minor Vasey – F; = A. purpurascens var. purpurascens – FNA, K, Z] * Aristida purpurea Nuttall var. longiseta (Steudel) Vasey, Red Three-awn. Cp (SC): disturbed areas; rare, adventive from further west. August-October. Also reported from NC, but the collection is from a Soil Conservation Service test nursery, and there is no evidence of naturalization. [= C, FNA, K, Z; > A. longiseta var. robusta Merrill – F; = A. longiseta Steudel – G, HC] Aristida simpliciflora Chapman, Southern Three-awn, Chapman's Three-awn. Cp (GA, NC): wet pine savannas; rare (GA Special Concern, NC Rare). Sw. GA west through the FL Panhandle and c. AL to s. MS (Sorrie & Leonard 1999), and south into central Peninsular Florida; also in se. NC, where apparently disjunct (it should be searched for in SC). A. simpliciflora was believed to be a Gulf Coastal Plain endemic until found by R. LeBlond in 1999 in wet savannas in se. NC (Green Swamp savannas, Brunswick County; Old Dock Savanna, Columbus County; and The Neck Savanna, Pender County). It is reported for sw. GA (Jones & Coile 1988, Kartesz 1999). Harper also reports it for c. GA. [= FNA, HC, K, S, Z] Aristida spiciformis Elliott, Bottlebrush Three-awn, Spike Three-awn. Cp (GA, NC?, SC): wet pine savannas and seepage areas; rare (NC Watch List). August-October. E. SC (McMillan et al. 2002) south to FL, west to MS. Allred (1986) also reports this species from NC, but the documentation is unknown to me. [= RAB, FNA, GW, HC, K, S, Z] Aristida stricta Michaux, Carolina Wiregrass, Pineland Three-awn. Cp (NC, SC), Pd (NC): Coastal Plain pinelands of nearly all sorts, ranging from the driest white-sand sandhills to seasonally saturated pine savannas dominated by a mixture of longleaf pine and pond pine, largely or entirely replaced in the wettest savannas by Sporobolus teretifolius, Sporobolus pinetorum, Muhlenbergia expansa, Ctenium aromaticum, and Calamovilfa brevipilis; also in Piedmont areas adjacent to the Coastal Plain and formerly supporting fire-maintained longleaf pine woodlands; common, rare in Piedmont. September-November. Ne. NC (south of Albemarle Sound and the Roanoke River), south to ne. SC (Lee and Kershaw counties). A. stricta was the keystone species of much of the upland Coastal Plain of the Carolinas. Its flammable foliage facilitated the spread of lightning-set fires that maintained the biologically rich pine savanna, sandhill, and pine flatwood ecosystems once widespread in our area. Though still locally common in parts of the Sandhill region and in portions of Brunswick, Pender, Onslow, and Carteret counties, NC, A. stricta is much rarer than formerly. The conversion of vast acreages of former pinelands to agriculture, pine tree farms, and developed areas has taken its toll over the years. In the twentieth century, suppression of fire has also led to the destruction of A. stricta. More recently, pine-straw raking is leading to the serious decline of A. stricta in its few remaining strongholds on public lands. A. stricta has little tolerance for ground disturbance. See Peet (1993) for discussion of the taxonomy and ecology of this species. Ward (2001) proposes varietal status for A. stricta and A. beyrichiana. [= K, Y; < A. stricta – RAB, FNA, GW, HC, S, V, Z (also see A. beyrichiana); = A. stricta var. stricta – X]

POACEAE 848

Aristida tenuispica A.S. Hitchcock, Southern Arrowfeather. Cp (GA, NC, SC): sandy habitats in the Coastal Plain; uncommon? August-October. NC south to FL and west to MS. [= HC, S; = A. purpurascens Poiret var. tenuispica (A.S. Hitchcock) Allred – FNA, K, Z] Aristida tuberculosa Nuttall, Seabeach Needlegrass. Cp (GA, NC, SC, VA): sandhills, coastal dunes (in VA), other dry, sandy habitats such as sandy roadsides; common, rare in VA (VA Watch List). August-October. Se. NH south to NJ and disjunct in e. VA in the outer Coastal Plain; from sc. NC south to Panhandle FL and west to s. MS (Sorrie & Leonard 1999), mostly in the inner Coastal Plain; and also near the Great Lakes in sw. MI, n. IN, n. IL, s. WI, se. MN, and e. IA. The curious trimodal distribution is unexplained. [= RAB, C, F, FNA, G, HC, K, S, Z] Aristida virgata Trinius. Cp (GA, NC, SC, VA), Pd (GA), Mt (NC): moist to wet savannas, mountain bogs (Henderson Co., NC), other moist habitats; common. August-October. S. NJ south to FL, west to TX, primarily on the Coastal Plain. [= RAB, C, F, G, GW, HC, S; = A. purpurascens Poiret var. virgata (Trinius) Allred – FNA, K, Z] * Aristida adscensionis Linnaeus, Sixweeks Three-awn, has been reported as an introduction (from western United States) in SC (FNA, Kartesz 1999). {investigate} [= F, FNA, G, K] {not keyed at this time; synonymy incomplete} Aristida basiramea Engelmann ex Vasey, Forktip Three-awn. Occurs in VA, SC, etc. (FNA). [= F, FNA, G, K; = A. basiramea var. basiramea – C] {not keyed at this time; synonymy incomplete} Aristida gyrans Chapman, Corkscrew Three-awn. Cp (GA): dry pinelands; rare (GA Special Concern). E. GA and w. Panhandle FL, south to s. FL. In Bryan, Long, and Montgomery counties in e. GA (Sorrie 1998b), and in wc. GA (J. Allison, pers. comm.). [= FNA, K, S] {not keyed at this time; synonymy incomplete} Aristida ramosissima Engelmann ex A. Gray. East to Panhandle FL, c. TN, and e. KY (FNA) and might occur in our area. It is similar to A. oligantha and will key to it; it differs in having the central awn of the lemma 9-30 mm long (vs. 30-70 mm long), and the awn of the second glume 3-7 mm long (vs. 7.5-17 mm long). [= C, F, FNA, G, K, S] {not keyed at this time; synonymy incomplete} Allred (1986) reports the collection of several additional non-native species from our area, including A. divaricata Willdenow from sw. United States (from a Soil Conservation Service test nursery in Chapel Hill, NC) and uncertainly identified material of an Australian species (from a wool-combing mill at Jamestown, SC). There is no evidence that either are naturalized.

Arrhenatherum Palisot de Beauvois (False Oatgrass) References: Tucker (1996)=Z. 1 Base of culm consisting of a series of adjacent (moniliform) corms ca. 1 cm in diameter...................A. elatius var. bulbosum 1 Base of culm not swollen or cormose ....................................................................................................... A. elatius var. elatius * Arrhenatherum elatius (Linnaeus) J. & K. Presl var. bulbosum (Willdenow) Spenner, Tuber Oatgrass, Onion Couch. (VA): habitat in our area not known; abundance not known, introduced from Europe. This variety was apparently cultivated for the edible tubers in Bronze Age Europe (Tucker 1996). Cited for VA in HC. [= C, F, G, HC, K, Z; < A. elatius – GW, W; = A. elatius var. tuberosum Thiel. – S] * Arrhenatherum elatius (Linnaeus) J. & K. Presl var. elatius, Tall Oatgrass. Pd (GA, NC, VA), Mt, Cp (NC, VA): meadows, fields, roadsides; common, introduced from Europe. May-June. [= C, F, G, HC, K, S, Z; < A. elatius – RAB, GW, W]

Arthraxon Palisot de Beauvois (Basket Grass) References: van Welzen (1981)=Y; Thieret in FNA (2003a); Kiger (1971)=Z. Identification notes: Sometimes confused (especially before flowering) with Microstegium, but Arthraxon has distinctly cordate-clasping leaves, which Microstegium lacks. Also vegetatively similar to Oplismenus. * Arthraxon hispidus (Thunberg) Makino var. hispidus, Basket Grass. Cp, Pd, Mt (GA, NC, SC, VA): moist ditches, bottomlands, disturbed areas; common, native of se. Asia. September-October. Like Microstegium, Arthraxon appears to be steadily increasing its abundance in our area. [= FNA, Y; < A. hispidus – C, GW, K, Z; > A. hispidus var. cryptatherus (Hackel) Honda – RAB, F, G, HC, W]

Arundinaria Michaux (Cane) Both species of Arundinaria were much reduced by the foraging of free-range livestock in the eighteenth and early nineteenth centuries and by fire suppression in the late nineteenth century and throughout the twentieth century. "Canebrakes," large areas dominated by cane, were described in many historical accounts and apparently occupied large parts of the landscape of the

POACEAE 849

Coastal Plain, also occurring in the Piedmont and low Mountains. References: Tucker (1988)=Y; McClure (1973)=Z; McClure (1963); Judziewicz et al. (2000)=X; Triplett, Weakley, & Clark (2006)=Q. The key adapted from Y and Z. 1 Primary branches with 0-1 compressed basal internodes; culm internodes usually sulcate; culm leaves deciduous; culms to

10 m tall; rhizomes lacking air canals.......................................................................................................................A. gigantea 1 Primary branches with 2-5 compressed basal internodes; culm internodes usually terete; culm leaves persistent to tardily

deciduous; culms to 4 m tall; rhizomes with or without longitudinal air canals (visible in cross-section as a cylinder of hollow canals 1 mm or less from the outer surface). 2 Foliage blades chartaceous, deciduous, abaxial surfaces pilose or glabrous, weakly tessellate; primary branches usually

< 35 cm long, basal nodes not developing secondary branches; top knot blades 12-22.5 cm long; rhizomes with or without air canals ......................................................................................................................................A. appalachiana

2 Foliage blades coriaceous, persistent, abaxial surfaces densely pubescent or glabrous, strongly tessellate; primary branches usually > 50 cm long, basal nodes developing secondary branches; top knot blades 20-30 cm long; rhizomes with air canals ......................................................................................................................................................... A. tecta

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

slopes; common. The short plants (often only knee-high, though sometimes head-high) on mountain slopes south of Asheville are autumn-deciduous, whereas both "rivercane" appears to be evergreen. [= Q; < A. gigantea (Walter) Walter – RAB, GW; < A. gigantea ssp. tecta (Walter) McClure – K, X, Z; < A. tecta – HC, S, Y; = A. tecta var. decidua Beadle in L.H. Bailey; apparently and implicitly included in the concept of either A. tecta or A. gigantea by earlier authors]

Arundinaria gigantea (Walter) Walter, Giant Cane, River Cane. Mt, Pd, Cp (GA, NC, SC, VA): swamps, floodplains; common. April-July. Widespread in se. North America, ranging from s. DE s. OH, south to FL and e. TX. There has been much disagreement over the recognition of one, two, or several taxa of cane in the Southeastern United States. This species reaches heights of 6-7 (-10) m and is supposed to flower only once every 40-50 years. A. macrosperma Michaux is controversial, sometimes considered to be a synonym of A. gigantea or to represent hybridization or introgression between A. gigantea and A. tecta. [= F, HC, Q, S, Y; < A. gigantea – RAB, C, GW (also see A. tecta); = A. gigantea ssp. gigantea – K, Z; > A. gigantea ssp. gigantea – X; > A. gigantea (Walter) Walter ssp. macrosperma (Michaux) McClure – X; > A. macrosperma Michaux]

Arundinaria tecta Walter, Switch Cane, Small Cane. Cp, Pd (GA, NC, SC, VA): savannas, pocosins, canebrakes, generally (but not solely) in wetlands; common. April-July. Primarily a Southeastern Coastal Plain endemic: e. VA to FL and s. AL. A. tecta is a smaller plant than A. gigantea (normally 1-2 m tall, but reaching heights of up to 4 m where fire-suppressed), and flowers more frequently, supposedly every 3-4 years (Tucker 1988), probably actually in response to fire. [= Q; < A. gigantea – RAB, C, GW; < A. tecta – F, HC, S, Y; < A. gigantea ssp. tecta (Walter) McClure – K, X, Z]

Arundo Linnaeus (Giant Reed) A genus of 3 species, widespread in the tropics, subtropics and warm-temperate areas. References: Allred in FNA (2003a). * Arundo donax Linnaeus, Giant Reed. Cp, Pd (GA, NC, SC, VA), Mt (NC, VA): disturbed areas; uncommon, introduced from the Old World. September-October. Var. versicolor, with leaves transversely striped white and green, is better considered only as a form or cultivar. [= RAB, F, FNA, K, S; > A. donax var. donax – HC; > A. donax var. versicolor (P. Miller) Stokes – HC]

Avena Linnaeus 1753 (Oats) References: Tucker (1996)=Z. 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].

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2 Panicle branches deciduous; disarticulation occurring at the base of the branch (the branch therefore falling whole); spikelets 2-3 per branch, appressed to the branch; [native species of limestone habitats, also with introduced populations]; [subgenus Bouteloua]..............................................................................B. curtipendula var. curtipendula

2 Panicle branches persistent; disarticulation occurring above the glumes (the individual florets therefore falling); spikelets >6 per branch, pectinately disposed; [rare introductions]; [subgenus Chondrosum]. 3 Panicle branches terminating in a spikelet ..................................................................................................B. gracilis 3 Panicle branches extending beyond the base of the terminal spikelets.................................... B. hirsuta var. hirsuta

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

and bluffs over limestone or serpentine, limestone glades; uncommon. July-September. S. CT west to MT, south to VA, e. TN, nw. GA, AL, panhandle FL, TX, AZ, and CA; also in Central and South America. The older literature refers to B. curtipendula as introduced in SC, but the single specimen documenting its occurrence there appears to be from experimental plantings at Clemson University; there is apparently no evidence of its establishment. B. curtipendula occurs on serpentine in the Piedmont of Georgia (Allison, pers. comm.). Var. caespitosa Gould & Kapadia is cespitose rather than rhizomatous and occurs in sw. United States. [= C, FNA, K, Y; < B. curtipendula – RAB, F, G, HC, S, W] * Bouteloua dactyloides (Nuttall) J.T. Columbus, Buffalo Grass. Mt (VA), Pd, Cp (GA): disturbed areas; rare, introduced from w. North America. [= Z; = Buchloe dactyloides (Nuttall) Engelmann – C, F, FNA, G, HC, K] * Bouteloua gracilis (Willdenow ex Kunth) Lagasca y Segura ex Griffiths, Blue Grama. Cp (SC): disturbed areas; rare, introduced from western North America. Reported for SC (Gould 1979). [= F, FNA, K, Y; > Bouteloua gracilis var. gracilis – HC] * Bouteloua hirsuta Lagasca y Segura var. hirsuta, Hairy Grama. Cp (GA, SC): disturbed areas; rare, introduced from western North America. Reported for SC and GA (Kartesz 1999). [= K, Y; < Bouteloua hirsuta – F, HC; = Bouteloua hirsuta ssp. hirsuta – FNA]

Brachiaria (see Urochloa)

Brachyelytrum Palisot de Beauvois (Shorthusk) The only other species of the genus is B. japonicum Hackel, of s. Japan, Korea, and ec. China (Saarela et al. 2003, Tucker 1988). References: Saarela et al. (2003)=Z; Tucker (1988)=Y; Stephenson (1971); Voss (1972); Campbell, Garwood, & Specht (1986). Key based in part on Saarela et al. (2003). 1 Lemmas hirsutulous or minutely scabrous, the longest hairs (0.06-) 0.08-0.14 (-0.2) mm long (not evident at 10×); lemma

(0.7-) 0.8-1.2 (-1.4) mm wide; widest leaf blade (8-) 10-14 (-16) mm wide; second glume (0.6-) avg. 1.2 (-3.0) mm long; [plants of the Mountains] ........................................................................................................................................B. aristosum

1 Lemmas hirsute with hairs (0.2-) 0.4-0.8 (0.9) mm long (easily seen at 10×); lemma (0.8-) 1.1-1.5 (-1.8) mm wide; widest leaf blade (9-) 11-17 (-20) mm wide; second glume (0.2-) avg. 2.2 (7.0) mm long; [plants widely distributed in our area] ......

...................................................................................................................................................................................B. erectum Brachyelytrum aristosum (Michaux) Trelease in Branner & Coville, Northern Shorthusk. Mt (GA, NC, VA): moist forests, mostly at moderate to high elevations, such as northern hardwoods and spruce-fir; rare (GA Special Concern, NC Watch List). July-August. Fairly widespread in ne. North America, south in the mountains to sw. NC and n. GA. In MI, B. septentrionale flowers about 10 days before co-occurring B. erectum, with strongly synchronized anthesis of each species occurring on a single day (Stephenson 1971)). Reputed intermediates and hybrids between the two taxa are apparently based on the use of ambiguous characters. [= Z; = Brachyelytrum septentrionale (Babel) G. Tucker – K, Y; < B. erectum – RAB, G, HC, S, W; = B. erectum var. septentrionale Babel – F; = B. erectum var. glabratum (Vasey ex Millspaugh) Koyama & Kawano – C] Brachyelytrum erectum (Schreber ex Sprengel) Palisot de Beauvois, Common Shorthusk. Mt, Pd (GA, NC, SC, VA), Cp (NC, SC, VA): mesic forests, in the Mountains at lower elevations than B. septentrionale); common (uncommon in Piedmont, rare in Coastal Plain). June-August. MA, NY, OH, MI, and s. WI south to FL and TX. [= K, Y, Z; < B. erectum – RAB, G, HC, S, W (also see B. septentrionale); = B. erectum var. erectum – C, F]

Brachypodium Palisot de Beauvois * Brachypodium sylvaticum (Hudson) Palisot de Beauvois ssp. sylvaticum, Slender False Brome. Pd (VA): roadsides and yards; rare, introduced from Europe. [= FNA; < B. sylvaticum – HC, K]

Briza Linnaeus (Quaking Grass) References: Tucker (1996)=Z.

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]

Catapodium Link 1827 References: Soreng et al. (2003)=Z. * 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): savanna-pocosin 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]

Chloris Swartz (Finger-grass, Chloris) (also see Eustachys)

Key based partly on C. 1 Inflorescence verticillate, typically the panicle branches in 2-5 verticels; perennial; fertile lemma inconspicuously appressed-

pilose; spikelets not imbricate............................................................................................................................. Ch. verticillata 1 Inflorescence digitate, the panicle branches in a single verticel at the apex of the culm; annual; lemma conspicuously long-

ciliate; 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, pocosin-

savanna ecotones, seepage bogs, sandhill-pocosin ecotones, sandhill seeps; common (VA Rare). June-August (or later in response to late summer fires). Southeastern Coastal Plain endemic: se. VA south to FL and west to LA and e. TX (Singhurst, Keith, & Holmes 2005). The entire plant is aromatic and numbs the mouth, tongue, and lips when chewed, hence the specific epithet and common names. Like many species of the longleaf pine ecosystem, toothache grass generally flowers only following fire (MacRoberts & MacRoberts 1992). Sterile clumps can be recognized by the rather broad, bicolored leaves (bluish on the upper surface, bright green on the lower surface). [= RAB, C, F, FNA, G, GW, HC, K; = Campulosus aromaticus (Walter) Trinius – S]

Ctenium floridanum (A.S. Hitchcock) A.S. Hitchcock, Florida Toothache Grass. Cp (GA): dry pinelands, sandhills, upper ecotones of pineland pools; rare (GA Special Concern). June-September. Southeastern Coastal Plain endemic: se. GA to ne. FL. Like Ct. aromaticum, generally flowering only following fire. [= FNA, GW, HC, K; = Campulosus floridanus A.S. Hitchcock – S]

Cynodon L.C. Richard (Bermuda Grass) A genus of ca. 9 species, native to the tropical Old World. References: Barkworth in FNA (2003a). * Cynodon dactylon (Linnaeus) Persoon var. dactylon, Bermuda Grass, Scutch Grass. Cp, Pd, Mt (GA, NC, SC, VA): lawns, gardens, roadsides, pastures, fields, disturbed areas; common, introduced from Eurasia. May-October. [= FNA; < C. dactylon – RAB, C, F, G, HC, K, W; < Capriola dactylon (Linnaeus) Kuntze – S]

Cynosurus Linnaeus 1753 (Dog's-tail, Dogtail Grass) References: Tucker (1996)=Z. 1 Panicle linear-oblong, 1-10 (-14) cm long, 0.4-1 cm wide; leaves 1-3 (-4) mm wide; perennial; fertile lemma 3-4 mm long,

plus a 0-1 mm long mucro; [section Cynosurus] ......................................................................................................C. cristatus 1 Panicle ovoid, 1-4 (-8) cm long, 0.7-2 cm wide; leaves (2-) 3-10 mm wide; annual; fertile lemma 4.5-7 mm long, plus a 6-16

mm long awn; [section Falona] ..............................................................................................................................C. echinatus * Cynosurus cristatus Linnaeus, Crested Dog's-tail. Mt (NC), {VA}: lawns, roadsides; rare, introduced from Eurasia. June-July. [= RAB, C, F, G, HC, K, Z] * Cynosurus echinatus Linnaeus, Rough Dog's-tail. Cp (NC, SC, VA), Pd (GA, NC, SC, VA), Mt (GA): lawns, roadsides; rare, introduced from Eurasia. May-June. [= RAB, C, F, HC, K, Z]

Dactylis Linnaeus 1753 (Orchard Grass) References: Tucker (1996)=Z. * 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 long-

pilose. 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 rosette-forming, 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, plano-convex 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 papillose-

based 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 bluish-glaucous..............................................................................................................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 white-

beige 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 spreading-pilose 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]

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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

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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 well-drained 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 –

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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}

Dinebra Jacquin (Viper Grass) References: Barkworth in FNA (2003a). * 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]

Eleusine Gaertner (Yard Grass) References: Hilu in FNA (2003a). Key based on FNA. 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, <1.5 mm wide; lower glume 0.7-1.7 mm long...................

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 greenish-gray 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 0.6-1.3 mm wide; panicle branches usually whorled at the lowest 2 nodes .................

....................................................................................................................... E. pilosa 30 First glume 0.5-1.5 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 wool-combing 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 wool-combing 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 <2.5 mm long............................................................................................E. petraea

2 Keel of the fertile lemma glabrous; spikes 7-20; [aliens, in disturbed situations]. 4 Spikelets >2.4 mm long; sterile floret oblanceolate, acute ...............................................................E. distichophylla 4 Spikelets <2.1 mm long; sterile floret widely cuneate, truncate....................................................................E. retusa

* Eustachys distichophylla (Lagasca y Segura) Nees, Weeping Finger-grass. Cp (GA): disturbed areas; rare, introduced from South America. In GA and FL (Kartesz 1999). [= FNA, K, Z; = Chloris distichophylla Lagasca y Segura – HC]

Eustachys floridana Chapman, Florida Finger-grass. Cp (GA): sandhills, pine flatwoods; rare (GA Special Concern). E. GA south to c. peninsular FL, west to w. Panhandle FL and s. AL. [= FNA, K, Z; = Chloris floridana (Chapman) Wood – HC, S] {synonymy}

Eustachys glauca Chapman, Saltmarsh Finger-grass. Cp (GA, NC, SC): marshes and marsh edges; rare. June-October. Se. NC south to FL and west to s. AL. [= FNA, K, Z; = Chloris glauca (Chapman) Wood – RAB, GW, HC, S]

Eustachys petraea (Swartz) Desvaux, Dune Finger-grass. Cp (GA, NC, SC): dune slacks and sand flats, sometimes in disturbed areas; common (uncommon north of GA). (May-) June-October. NC (Dare County) south to FL and west to TX. [= FNA, K, Z; = Chloris petraea Swartz – RAB, GW, HC, S] * Eustachys retusa (Lagasca y Segura) Kunth, Argentine Finger-grass. Cp (GA, SC): sandy fields; rare, introduced from Argentina. June. [= FNA, K, Z; ? Chloris argentina (Hackel) Lillo & Parodi – RAB, G, HC] * Eustachys caribaea (Sprengel) Herter, Chickenfoot Grass. Cp (GA): disturbed areas; rare, native of South America. [= FNA, K; = Chloris capensis – HC, misapplied] {add to synonymy; not keyed at this time}

Eustachys neglecta (Nash) Nash. Native. Pinelands, sandy fields. N. and peninsular FL, se. AL, and e. TX. [= FNA, K; = Chloris neglecta Nash – HC, S] {synonymy incomplete; not keyed at this time}

Festuca Linnaeus (Fescue) (also see Vulpia)

A genus of qbout 350 species, nearly cosmopolitan in temperate regions. References: Darbyshire (1993)=X; Aiken & Darbyshire (1990)=Y; Tucker (1996)=Z; Soreng & Terrell (1998). Key based in part on C and Y. 1 Leaves 0.2-3 mm wide, often involute; [subgenus Festuca].

2 Plant loosely tufted, often rhizomatous; basal sheaths disintegrating into fibers; spikelets 6-13 mm long ............F. rubra 2 Plant tufted, lacking rhizomes; basal sheaths persistent, remaining firm and entire; spikelets 3-9 mm long.

3 Lemmas 2.3-4.0 (-4.4) mm long, awnless, or with a minute projection to 0.4 mm long; anther 1.5-2.2 (-2.5) mm long; spikelets 3.0-6.0 (-6.5) mm long .....................................................................................................F. filiformis

3 Lemmas 3.8-5.5 mm long, with an awn 0.5-2.5 mm long; anther (2.3-) 2.5-3.0 mm long; spikelets 5.5-9.0 mm long .................................................................................................................................................... F. trachyphylla

1 Leaves 3-12 mm wide, flat. 4 Larger lemmas 5.5-10 mm long; leaf blades auriculate at the base; anthers 2-4 mm long.................... [see Schedonorus] 4 Larger lemmas 3.3-5.2 mm long; leaf blades not auriculate at the base; anthers 0.8-1.5 mm long; [subgenus Subulatae,

section Obtusae]. 6 Principal lowermost panicle branches with 8-20 spikelets clustered at the end; spikelets broadly ovate, 4-6 mm

wide......................................................................................................................................................... F. paradoxa 6 Principal lowermost panicle branches with 2-7 spikelets scattered along the outer half; spikelets narrowly ovate, 2-

4 mm wide .......................................................................................................................................F. subverticillata * Festuca filiformis Pourret, Hair Fescue, Fineleaf Sheep Fescue. Mt, Pd (NC, VA): lawns, disturbed areas; rare, introduced from Eurasia. May-June. [= C, K, Y, Z; ? F. capillata Lamarck – RAB, F, HC; ? F. ovina Linnaeus var. capillata (Lamarck) Alefeld – G; ? F. tenuifolia Sibthorp – W]

Festuca paradoxa Desvaux, Cluster Fescue. Cp, Pd, Mt (GA, NC, SC, VA): bottomlands, uplands over mafic rock; common. May-July. PA west to WI and IA, south to SC, c. GA, and e. TX. [= RAB, C, F, G, GW, HC, K, W, Z; ? F. shortii Kunth ex Wood – S, misapplied]

Festuca rubra Linnaeus, Red Fescue. Mt (NC, SC, VA), Pd, Cp (GA, NC, VA): roadsides, fields, disturbed areas, pastures, grassy balds; common. May-July. In our area, this species is considered to be partly native and partly introduced. This species is circumboreal, extending south in North America to GA and MO. Many varieties or subspecies have been described in the F. rubra complex; it is uncertain how they might apply in our area. [= RAB, C, G, HC, S, Y, Z; > F. rubra var. rubra – F; > F. rubra var. commutata Gaudin – F; ? F. rubra ssp. rubra – K]

Festuca subverticillata (Persoon) Alexeev, Nodding Fescue. Mt, Pd, Cp (GA, NC, SC, VA): moist to wet forests, woodlands, and disturbed areas; common. May-June. ME, Québec, and Manitoba south to FL and e. TX. [= C, K, Y, Z; ? F. obtusa Biehler – RAB, F, G, GW, HC, S, W] * Festuca trachyphylla (Hackel) Krajina, Hard Fescue. Pd (GA, NC, VA), Mt (NC, VA), Cp (NC, SC, VA): meadows, pastures, disturbed areas; uncommon, introduced from Eurasia. May-June. The nomenclatural debate about the application of the name F. trachyphylla is summarized in Darbyshire & Pavlick (1997). [= C, K, Y, Z; < F. ovina – RAB, S, W, in the broad

POACEAE 887

sense (misapplied as to our material); < F. ovina var. ovina – F, G, HC; < F. ovina var. duriuscula (Linnaeus) W.D.J. Kock – F, G, HC, misapplied as to our material] * Festuca thurberi Vasey. Introduced in SC. [= K] {not keyed; investigate}

Festuca versuta Beal, Texas Fescue. Native, east to TN. [= K] {not keyed; investigate}

Glyceria R. Brown (Mannagrass) (also see Torreyochloa)

A genus of about 40 species, nearly cosmopolitan. References: Tucker (1996)=Z. 1 Spikelets (10-) 15-40 mm long, linear, subterete, 5-15× as long as wide; [section Glyceria].

2 Lemma (6-) 7-8.5 (-10) mm long, acute to acuminate; palea longer than the lemma, extending 1.5-3 mm beyond the lemma apex.....................................................................................................................................................G. acutiflora

2 Lemma 2.5-5.3 mm long, obtuse to notched; palea about as long as the lemma (ranging from shorter than the lemma and included, to projecting up to 1 mm beyond the lemma apex). 3 Anthers 0.5-0.8 mm long; lemma mostly 2.5-3.5 mm long................................................................... G. arkansana 3 Anthers 1.0-2.0 mm long; lemma mostly 3.6-5.5 mm long.............................................................G. septentrionalis

1 Spikelets 2.5-8 mm long, ovate to oblong, 1.5-3× as long as wide; [section Hydropoa]. 4 Inflorescence compact (at maturity), the branches stiffly ascending to appressed, the tips never nodding; ligule < 1 mm

long. 5 Inflorescence branches elongate, appressed; lower internodes of the inflorescence 2-8 cm long; spikelets with 3-4

flowers, 3.5-4 mm long; lemma 1.9-2.8 mm long; leaves 2-5 mm wide; [of the Mountains, rarely elsewhere] ......... ................................................................................................................................................................ G. melicaria 5 Inflorescence branches short, stiffly ascending; lower internodes of the inflorescence 0.8-2.0 (-2.5) cm long;

spikelets with 4-7 flowers, 4-8 mm long; lemma 3.0-3.7 mm long; leaves 3-10 mm wide; [of the Coastal Plain, rarely disjunct inland to the Mountains of VA]............................................................................................G. obtusa

4 Inflorescence lax and diffuse (at maturity), the branches spreading to somewhat ascending, the tips often nodding or drooping; ligule 1-6 mm long. 6 Veins of the lemma visible, but not raised; lemma 2.3-4.0 mm long; ligule 2-6 mm long.

7 Lemma 2.9-4.0 mm long, projecting conspicuously beyond the palea; spikelets 4-8 mm long, with 5-10 flowers...........................................................................................................................................G. canadensis

7 Lemma 2.3-2.9 mm long, more-or-less equal to the palea; spikelets 3-5 mm long, with (2-) 3-5 (-6) flowers ... ................................................................................................................................................................ G. laxa

6 Veins of the lemma prominently raised; lemma 1.4-3.0 mm long; ligule 1-4 mm long. 8 Lemma 1.4-2.1 mm long, usually green; first glume 0.5-1.0 mm long; second glume 0.8-1.3 mm long; culms

mostly 5-12 dm tall; leaves 2-5 (-8) mm wide ...................................................................G. striata var. striata 8 Lemma 2.5-3.0 mm long, purple or green; first glume 1.2-1.9 mm long; second glume 1.5-2.4 mm long;

culms mostly 10-20 dm tall; leaves 5-12 mm wide. 9 Glumes obtuse, subequal (the first 1.2-1.9 mm long, the second 1.5-2.4 mm long); spikelets 4-6.5 mm

long, (4-) 5-9-flowered; lemmas purplish, contrasting with the pale glumes; ligule 2-4 (-5) mm long; [of various montane wetlands of VA and possibly NC]...............................................G. grandis var. grandis

9 Glumes acute, unequal (the first 1.4-1.6 mm long, the second 1.8-2.1 mm long); spikelets mostly 3-4 mm long, 2-4-flowered; lemmas dark green (or purplish?); ligule 1.5-3 mm long; [of high elevation seepages in the Great Smoky Mountains of NC and TN]........................................................ G. nubigena

Glyceria acutiflora Torrey. Mt (GA, VA): shallow water and wet mucky soils in mountain ponds, wet pastures; rare (GA

Special Concern). June-July. ME west to MI, south to DE, VA, nw. GA (Jones & Coile 1988), e. TN, and MO; also in e. Asia. [= C, F, G, GW, HC, K, W, Z; = Panicularia acutiflora (Torrey) Kuntze – S]

Glyceria arkansana Fernald, Arkansas Mannagrass. Cp (VA): swamps; rare. May-June. IL south to LA and AR; disjunct in se. VA and w. NY. The appropriate treatment of this taxon needs further investigation. [= F, HC, K, Z; < G. septentrionalis – C, G; = G. septentrionalis A.S. Hitchcock var. arkansana (Fernald) Steyermark & Kučera]

Glyceria canadensis (Michaux) Trinius, Rattlesnake Mannagrass. Mt (NC, VA), Cp (VA): bogs, seepages, and wet meadows; rare. June-July. Newfoundland west to MN, south to NJ, VA, nw. NC, and IL. [= C, F, G, K, Z; = G. canadensis var. canadensis – HC, W]

Glyceria grandis S. Watson var. grandis, American Mannagrass. Mt (NC?, VA), Cp (VA): wet, mucky soils of open wetlands; rare. Nova Scotia west to AK, south to VA, IA, NM, and OR. Attributed to w. NC by Tucker (1996). [= K; < G. grandis – C, F, G, GW, HC, Z; = Panicularia grandis (S. Watson) Nash – S; ? G. maxima (Hartman) Holmberg ssp. grandis (S. Watson) Hultén]

Glyceria laxa (Scribner) Scribner, Lax Mannagrass. Mt (NC, VA): bogs; rare. June-July. Prince Edward Island south to NC, mostly Appalachian. Though often described as a hybrid of G. canadensis and either G. striata var. striata and/or G. grandis var. grandis, G. laxa ranges south of the distribution of both G. canadensis and G. grandis var. grandis. It is best

POACEAE 888

considered as a species, perhaps of hybrid origin. [= F, G, K; = G. canadensis (Michaux) Trinius var. laxa (Scribner) A.S. Hitchcock – RAB, HC; = G. ×laxa – C; < G. canadensis – GW]

Glyceria melicaria (Michaux) F.T. Hubbard, Northeastern Mannagrass. Mt (GA, NC, VA), Pd (NC): mountain swamp forests and seepages; uncommon (GA Special Concern). June-August. Nova Scotia west to Québec, south to n. GA (Jones & Coile 1988) and KY. [= RAB, C, F, G, GW, HC, K, W, Z; = Panicularia melicaria (Michaux) A.S. Hitchcock – S]

Glyceria nubigena W.A. Anderson, Smoky Mountain Mannagrass. Mt (NC): moderate to high elevation seepages in the Great Smoky Mountains, sometimes in areas appearing dry (such as heath balds), nearly endemic to Great Smoky Mountains National Park; rare (US Species of Concern, NC Rare). June-July. Endemic to the Great Smoky Mountains of w. NC and e. TN. G. nubigena has nearly the same range as Rugelia nudicaulis, but is more restricted to seepage. The distinctions and relationship between this taxon and G. grandis need further investigation. [= RAB, HC, K, W, Z]

Glyceria obtusa (Muhlenberg) Trinius, Coastal Mannagrass. Cp (NC, SC, VA), Mt (VA): blackwater swamp forests, wet meadows, freshwater marshes; uncommon. June-September. Nova Scotia south to SC, on or near the Coastal Plain. [= RAB, C, F, G, GW, HC, K, W, Z; = Panicularia obtusa (Muhlenberg) Kuntze – S]

Glyceria septentrionalis A.S. Hitchcock, Floating Mannagrass, Eastern Mannagrass. Cp, Pd (GA, NC, SC, VA), Mt (NC, VA): shallow water, wet mucky soils, floodplain sloughs, cypress ponds; uncommon (GA Special Concern). May-June. MA west to MN, south to SC, ne. GA, and TX. [= RAB, F, GW, HC, K, W, Z; < G. septentrionalis – C, G (also see G. arkansana); = Panicularia septentrionalis (A.S. Hitchcock) Bicknell – S]

Glyceria striata (Lamarck) A.S. Hitchcock var. striata, Fowl Mannagrass. Mt, Pd, Cp (GA, NC, SC, VA): wet meadows, seepages, bogs, marshes, swamp forests; common. April-June. Newfoundland west to British Columbia, south to FL and CA. Var. stricta (Scribner) Fernald is more northern. [= C, F, G, HC, Z; < G. striata – RAB, GW, K, W; = Panicularia striata (Lamarck) A.S. Hitchcock – S; = G. striata ssp. striata]

Gymnopogon Palisot de Beauvois (Beard Grass, Skeleton Grass) A genus of about 15 species, in temperate and tropical areas of the Americas. References: Smith (1971)=Z. Identification notes: When sterile, Gymnopogon is sometimes confused with Dichanthelium. Gymnopogon differs in having the sheaths conspicuously overlapping (vs. not overlapping in Dichanthelium) and leaves that are definitely cordate-clasping and of stiff texture (only a few Dichanthelium have this combination). 1 Awn of the lemma 4.5-12 mm long; inflorescence branches with spikelets distributed from the tip nearly to the base; leaves

5-15 mm wide; [of the Coastal Plain, Piedmont, and Mountains]...........................................................................G. ambiguus 1 Awn of the lemma 0.8-1.6 (-3.5) mm long; inflorescence branches with spikelets distributed from the tip nearly to the base

(G. chapmanianus) or to roughly the midpoint, the basal portion naked (or some branches rarely with a few spikelets) (G. brevifolius); leaves 2-8 mm wide; [of the Coastal Plain and lower Piedmont]. 2 Spikelets 1-flowered; first glume 2.3-3.7 mm long....................................................................................... G. brevifolius 2 Spikelets 2-4-flowered; first glume 3.8-5 mm long ............................................................................... G. chapmanianus

Gymnopogon ambiguus (Michaux) Britton, Sterns, & Poggenburg, Eastern Beard Grass. Cp, Pd (GA, NC, SC, VA), Mt

(NC, SC, VA): prairies, glades, barrens, dry pinelands and woodlands, dry fields; common (rare in Mountains). August-October. S. NJ west to KY, OH, and MO, south to FL and TX. [= RAB, C, F, G, HC, K, S, W, Z]

Gymnopogon brevifolius Trinius, Pineland Beard Grass. Cp, Pd (GA, NC, SC, VA), Mt (GA): pine savannas, sandhills, dry woodlands, prairies, calcareous glades; common (rare in lower Piedmont) (GA Special Concern, VA Watch List). August-October. S. NJ south to FL, west to LA and AR; disjunct in the Highland Rim of KY and TN. [= RAB, C, F, G, HC, K, S, Z]

Gymnopogon chapmanianus A.S.Hitchcock, Chapman's Beard Grass. Cp (GA): sandhills and other xeric, sandy habitats; rare (GA Special Concern). Se. GA south to FL. [= HC, K, S; > G. chapmanianus – Z; G. floridanus Swallen – Z]

Hackelochloa Kuntze (Pitscale Grass * Hackelochloa granularis (Linnaeus) Kuntze, Pitscale Grass. Cp (GA): disturbed areas; rare, introduced from the Old World. Reported for sw. GA and other Gulf Coast states (Thieret in FNA 2003a, Jones & Coile 1988, Kartesz 1999). [= FNA, HC, K; = Rytilix granularis (Linnaeus) Skeels – S; = Mnesithea granularis (Linnaeus) Koning & Sosef]

Hainardia W. Greuter (Thintail) References: Tucker (1996)=Z. * Hainardia cylindrica (Willdenow) W. Greuter, Thintail. Cp (SC): waste areas around wool-combing mills; rare, introduced from the Old World. April-June. [= K, Z; = Lepturus cylindricus (Willdenow) Trinius – RAB; = Monerma cylindrica (Willdenow) Cosson & Durieu – HC]

POACEAE 889

Heteropogon Persoon (Tanglehead)

A genus of about 10 species, pantropical and extending into subtropical and warm temperate areas. References: Barkworth in FNA (2003a). * Heteropogon melanocarpus (Elliott) Elliott ex Bentham, Sweet Tanglehead. Cp (GA, NC, SC): sandy roadsides, disturbed areas; rare, probably naturalized from further south (or even from the Old World). September-October. The species is widespread in the Old World and New World tropics, north in North America to se. NC. [= RAB, FNA, HC, K, S]

Hierochloe R. Brown (Holy Grass, Sweet Grass, Vanilla Grass)

A genus of about 30 species, temperate and boreal in the northern and southern hemispheres. Tucker (1996) and Soreng et al. (2003) propose the inclusion of Hierochloe into a more broadly circumscribed Anthoxanthum. References: Tucker (1996)=Z; Soreng et al. (2003)=Y.

Hierochloe odorata (Linnaeus) Palisot de Beauvois, Holy Grass, Sweet Grass, Vanilla Grass. Mt (NC, VA): fens, wet calcareous medaows, high elevation pastures and openings; rare (NC Rare). April-May. A circumboreal species and subspecies, widespread in n. Eurasia and n. North America, ranging south in North America to NJ, MD, PA, OH, IN, IL, IA, SD, CO, UT, NM, and CA, with several disjunct occurrences in North Carolina, in Long Hope Valley, Ashe County, the Nantahala River Bogs, Macon County, and Pond Mountain, Ashe County. The report by S ("recorded by Chapman from Statesville, N.C.") can be discounted; the record reflects a collection made in the mountains by Mordecai E. Hyams, a botanist based in Statesville. Belden et al. (2004) document the first occurrence in Virginia. The sweet, vanilla-like odor of this grass is responsible for various folk uses – by Native Americans for making fragrant baskets, in Scandinavia strewn on church floors on festival days. Kartesz (1999) maps the NC occurrence of Hierochloe as H. hirta ssp. arctica; the reasons for this are unknown. {investigate} [= C, F, G, HC; ? H. odorata ssp. odorata – K; ? H. hirta (Schrank) ssp Borbás ssp. arctica (J. Presl) G. Weimarck – K; Torresia odorata (Linnaeus) A.S. Hitchcock – S; = Anthoxanthum nitens (Weber) Y. Schouten & Veldkamp – Z; ? Anthoxanthum nitens (Weber) Y. Schouten & Veldkamp spp. nitens – Y; > H. odorata var. fragrans (Willdenow) Richter (the North American plants)]

Holcus Linnaeus (Velvet Grass, Soft Grass) References: Tucker (1996)=Z. 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 well-established 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]

Koeleria Persoon (Junegrass, Koeleria) A genus of about 60 species, north and south temperate.

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 wool-combing 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)

Limnodea L.H. Dewey 1894 References: Brandenburg & Thieret (2000)=Z. * 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]

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* 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

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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]

Miscanthus Andersson (Eulalia) References: Barkworth in FNA (2003a). * 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].

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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,

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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}

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* 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 cordate-clasping). 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 well-developed........................................................................................................................... [see Piptatherum racemosum]

Oryzopsis asperifolia Michaux, Rough-leaved Ricegrass, Whiteseed Mountain-ricegrass. Mt (VA): high elevation pine-

oak/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 C3 photosynthesis. Eriochloa and Urochloa (including Brachiaria) have C4 photosynthesis, with PEP-ck decarboxylation. Panicum and Setaria (Paspalidium) have C4 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" C3 and C4 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 C3, with the outer parenchymatous sheath lacking specialized chloroplasts", etc. The argument that Phanopyrum also has C3 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 oblong-lanceolate, acuminate spikelets are (2.0-) 2.6-3.6 mm long, culms to 2 m long, and leaves 4-20 mm wide. Plants with geniculate bases, enlarged lower nodes and sheaths, and panicles with included peduncles and divergent branches have been recognized as var. geniculatum (A. Wood) Fernald. Plants with spikelets similar to those of var. puritanorum, but with culm and leaf features of var. dichotomiflorum, have been recognized as var. imperiorum Fernald, and are known only from se. VA. Recognition of any infraspecific taxa in this morphologically complex species is risky business. [= HC, K; < P. dichotomiflorum – RAB, C, GW, S, Z; > P. dichotomiflorum var. dichotomiflorum – F, G, W; > P. dichotomiflorum var. geniculatum – F, G, W; > P. dichotomiflorum var. imperiorum – F; Panicum s.s.] Panicum flexile (Gattinger) Scribner, Wiry Panic Grass. Pd (GA, NC, SC, VA), Mt (GA, NC,VA): glades and openings over mafic rocks, damp sandy meadows, open woods; rare (NC Rare). July-October. NY, sw. Québec, S. Ontario, and ND south to FL and TX. First reported for SC by Nelson & Kelly (1997). [= RAB, C, F, FNA, G, HC, K, S, W, Y, Z; Panicum s.s.] Panicum gattingeri Nash, Gattinger's Panic Grass. Mt, Pd (NC, VA), {GA}: damp or dry, usually calcareous sandy soils of fields, roadsides, shores, and cultivated ground; frequent in Mountains, uncommon in Piedmont (VA Watch List). August-October. NY, sw. Québec, and MN south to NC, TN, GA, AL, and AR. [= RAB, F, HC, K, S; < P. capillare – C, Y; = P. philadelphicum Bernhardi ex Trinius ssp. gattingeri (Nash) Freckmann & Lelong – FNA; = P. capillare Linnaeus var. campestre Gattinger – G, W; Panicum s.s.] Panicum hemitomon J.A. Schultes, Maidencane. Cp (GA, NC, SC, VA), Mt (VA): lake, pond, and river shores, swamp borders, marshes, ditches, often in shallow water; common (VA Rare). June-July. Coastal Plain from s. NJ south to FL, west to TX; also TN; South America. Often forming dense colonies in the low margin and shallow waters of limesink ponds. [= RAB, C, F, FNA, G, GW, HC, K, S, W, Z; not Panicum] Panicum lithophilum Swallen, Flatrock Panic Grass. Pd (GA, NC, SC), Mt (NC): soil islands on granitic flatrocks and domes; rare (NC Rare). August-October. Restricted to granite outcrops in NC, SC, and ec. GA. There is some question about the distinctness of this taxon from P. philadelphicum; Zuloaga & Morrone (1996) did not consider it separable from P. philadelphicum. [= RAB, HC, K; = P. philadelphicum Bernhardi ex Trinius ssp. lithophilum (Swallen) Freckmann & Lelong – FNA; < P. capillare Linnaeus var. sylvaticum Torrey – W; < P. philadelphicum – Y; Panicum s.s.] Panicum longifolium Torrey var. combsii (Scribner & Ball) Fernald, Combs Panic Grass. Cp (GA, NC, SC, VA): pond shores, depression meadows, cypress savannas, marshes, low woods; uncommon (VA Watch List). July-October. Scattered on the outer Coastal Plain from se. MA, NJ, se. VA, se. NC, e. SC, e. GA, and FL, west to se. LA. First glumes of var. combsii typically are longer than 1.5 mm long, while those of var. longifolium are shorter than 1.5 mm long. [= RAB, F, G; = P. rigidulum Bosc ex Nees ssp. combsii (Scribner & Ball) Freckmann & Lelong – FNA; = P. rigidulum Bosc ex Nees var. combsii (Scribner & Ball) Lelong – K, Z; < P. longifolium – C; = P. combsii Scribner & Ball – HC, S; not Panicum] Panicum longifolium Torrey var. longifolium, Long-leaved Panic Grass. Cp, Pd (GA, NC, SC, VA), Mt (NC, SC, VA): wet sandy or peaty soils of bogs, savannas, pond shores, depression meadows; common (uncommon in Piedmont, rare in Mountains). July-October. Nova Scotia, NH, MA, PA, and IN south to FL, west to TX. See note under var. combsii. [= RAB, G; = P. rigidulum Bosc ex Nees ssp. pubescens (Vasey) Freckmann & Lelong – FNA; = P. rigidulum Bosc ex Nees var. pubescens (Vasey) Lelong – K, W, Z; < P. longifolium – C, GW; = P. longifolium – HC, S; > P. longifolium var. longifolium – F; > P. longifolium var. pubescens (Vasey) Fernald – F; not Panicum] * Panicum miliaceum Linnaeus ssp. miliaceum, Broomcorn Millet, Proso Millet, Hog Millet. Cp (NC), Mt (VA): planted in wildlife food plots, sometimes persistent or self-sowing; rare, introduced, native of Eurasia. July-October. [= C, FNA, K; < P. miliaceum – F, G, HC, S, Y; Panicum s.s.] Panicum philadelphicum Bernhardi ex Trinius, Woodland Panic Grass. Pd, Mt (GA, NC, SC, VA), Cp (VA): glades, barrens, desiccated pondshores, riversides, or other rocky or dry sandy soil of open woods and roadsides; frequent (rare in SC). Nova Scotia west to WI, south to GA and e. TX. Plants formerly known as P. tuckermanii Fernald, ranging from se. Canada south to n. VA and OH, are distinguished by included or short-exerted peduncles less than one-third as long as the panicles. [= RAB, C, G, K, S; > P. philadelphicum – F, HC; > P. tuckermanii Fernald – F, HC; = P. philadelphicum Bernhardi ex Trinius ssp. philadelphicum – FNA; < P. capillare Linnaeus var. sylvaticum Torrey – W; < P. philadelphicum – Y (also see P. lithophilum); Panicum s.s.] *? Panicum repens Linnaeus, Torpedo Grass. Cp (GA, NC, SC): disturbed coastal sands, in area where ship's ballast was deposited; rare, apparently introduced. First reported for NC by Leonard (1971b). [= FNA, GW, HC, K, S; Panicum s.s.] Panicum rigidulum Bosc ex Nees var. condensum (Nash) Mohlenbrock, Dense Panic Grass. Cp (GA, NC, SC, VA): marshes, meadows, low woods, ditches, stream and pond shores, freshwater tidal shores; occasional. September-October. Coastal Plain south from se. MA to FL, west to se. TX and AR; West Indies. Usually readily identified by its tall stature and compact inflorescence, somewhat resembling a large P. hemitomon, with which it occasionally occurs. [= P. agrostoides Sprengel var. condensum (Nash) Fernald – RAB, F; < P. rigidulum – C, GW; < P. rigidulum Bosc ex Nees ssp. rigidulum – FNA; < P. agrostoides – G; = P. condensum Nash - HC, S; < P. rigidulum var. rigidulum – K, Z; not Panicum] Panicum rigidulum Bosc ex Nees var. elongatum (Pursh) Lelong, Tall Flat Panic Grass. Pd, Cp, Mt (GA, NC, SC, VA): marshes, low woods, ditches, swamps, shores, meadows; occasional (common in Piedmont). August-October. CT and NY west to IN, south to GA, LA, and ne. TX. [= K, W, Z; = P. stipitatum Nash – RAB, F, HC, S; = P. rigidulum Bosc ex Nees ssp. elongatum (Pursh) Freckmann & Lelong – FNA; < P. rigidulum – C, GW; = P. agrostoides Sprengel var. elongatum (Pursh) Scribner – G; not Panicum] Panicum rigidulum Bosc ex Nees var. rigidulum, Redtop Panic Grass. Cp, Pd (GA, NC, SC, VA): wet sandy or peaty soils low woods, meadows, marshes, shores, swamps, ditches; frequent. July-October. ME and MI south to FL and TX; also in CA and British Columbia; Central America. [= W; = P. agrostoides Sprengel var. agrostoides – RAB, G; < P. rigidulum Bosc

POACEAE 902

ex Nees ssp. rigidulum – FNA; < P. rigidulum var. rigidulum – K, Z; < P. rigidulum – C, GW; > P. agrostoides var. agrostoides – F, HC; > P. agrostoides var. ramosius (C. Mohr) Fernald – F, HC; = P. agrostoides – S; not Panicum] Panicum tenerum Beyrich ex Trinius, Southeastern Panic Grass. Cp (GA, NC, SC): limesink ponds, depression meadows, cypress savannas, wet pinelands, bogs; rare (GA Special Concern, NC Rare). June-September. Coastal Plain from se. NC to FL, west to e. TX; West Indies. The rhizomes produce lines of closely spaced culms. Though 0.5-1 m tall, the culms are narrow and inconspicuous. [= RAB, FNA, GW, HC, K, S, Z; not Panicum] Panicum verrucosum Muhlenberg, Warty Panic Grass. Cp, Pd, Mt (GA, NC, SC, VA): wet pinelands, marshes, shores, ditches; common (occasional in Piedmont, rare in Mountains). August-October. MA and PA west to MI and IN, south to FL and se. TX. Spikelets deep green, the warty surface unique among Panicum in our region. [= RAB, C, F, FNA, G, GW, HC, K, S, W, Z; not Panicum] Panicum virgatum Linnaeus var. cubense Grisebach, Blunt Panic Grass. Cp (GA, NC, SC, VA?): wet to dry sandy pinelands; occasional (frequent in the Sandhills). June-October. Coastal Plain from MA to FL, west to MS; also in MI; West Indies. [= F, HC, S; < P. virgatum – RAB, C, FNA, G, GW, W, Z; < P. virgatum var. virgatum – K] Panicum virgatum Linnaeus var. virgatum, Switchgrass. Cp, Pd, Mt (GA, NC, SC, VA): dry or wet sandy soils of pinelands, fresh and brackish marshes, shores; common (occasional in Mountains). June-October. Sw. Québec and ND south to FL and TX, west to NV; Bermuda; Central and South America. [= F, HC, S; < P. virgatum – RAB, C, FNA, G, GW, W, Z; < P. virgatum var. virgatum – K; not Panicum] * Panicum antidotale Retzius. Cp (SC): Native of India. Reported for NC and SC (FNA, Kartesz 1999). {investigate} [= FNA, HC, K; not Panicum] {not keyed at this time} * Panicum bergii Arechav. Cp (GA): disturbed areas; rare, native of South America. Reported for sc. GA (HC) and AL (Kartesz 1999). {investigate} [= FNA, HC, K; Panicum s.s.] {not keyed at this time} * Panicum bisulcatum Thunberg. Cp (GA, SC): disturbed areas; rare, native of Asia. Reported introduction in SC, GA, and PA (Kartesz 1999), and as a ballast plant for se. PA (Philadelphia) (Rhoads & Klein 1993, as P. acroanthum Steudel). [= FNA, K; ? P. acroanthum Thunberg] {not keyed at this time} Panicum brachyanthum Steudel. Cp (GA): {habitat unknown}; rare. Sw. GA west to c. TX. [= FNA, HC, K] Panicum dichotomiflorum Michaux var. puritanorum Svenson, Puritan Panic Grass. Alleged to be in VA northward (FNA). [= F, G, HC, K; < P. dichotomiflorum – C; = P. dichotomiflorum Michaux ssp. puritanorum (Svenson) Freckmann & Lelong – FNA] * Panicum miliaceum Linnaeus ssp. ruderale (Kitag.) Tzvelev, Panic Millet. [= FNA, K; = P. miliaceum ssp. spontaneum (Kit.) Tzvelev – C; < P. miliaceum – F, G, HC] {not keyed at this time} Panicum virgatum Linnaeus var. spissum Linder ranges south to PA, MD, and DE (Kartesz 1999). [= F, HC, K; < P. virgatum – C, FNA, G; not Panicum] {not keyed at this time}

Parapholis C.E. Hubbard (Sickle Grass) References: Tucker (1996)=Z. * 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's-tail." [< 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 under-represented 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 well-developed..................................................................................................................................................... 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}

Polypogon Desfontaines References: Tucker (1996)=Z; Barkworth in FNA (in prep.). 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]

Puccinellia Parlatore (Alkali Grass, Goosegrass) A genus of about 80 species, north temperate.

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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}

Reimarochloa A.S. Hitchcock A genus of about 4 species, of the New World tropics. Reimarochloa oligostachya (Munro ex Bentham) A.S. Hitchcock. AL, FL; Cuba. [= HC, K]

Rhynchelytrum Nees (see Melinis)

Ripidium Trinius (Ravenna-grass) References: Hodkinson et al. (2002). * 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

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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 ]

Schizachne Hackel (False Melic) A monotypic genus, circumboreal in Asia and North America.

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

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Schizachyrium littorale (Nash) Bicknell, Seaside Little Bluestem. Cp (GA, NC, VA): coastal dunes and maritime dry grasslands, often with Uniola paniculata, Panicum amarum, and other dune plants; common. August-October. E. MA south to NC (or SC?), and inland on the shores of the Great Lakes. In NC, Sch. littorale is present and abundant on dunes of barrier islands from Shackleford Banks, Carteret County south to Brunswick County, near the SC border, and entirely absent from the Outer Banks (from Cape Lookout, Carteret County, north through Hyde County to Dare County). [= FNA, GW, K; < Andropogon scoparium Michaux – RAB; = Sch. scoparium var. littorale (Nash) Gould – C, Z; = Andropogon scoparius Michaux var. littoralis (Nash) A.S. Hitchcock – F, G; = Andropogon littoralis Nash – HC, S; < Sch. scoparium (Michaux) Nash ssp. littorale (Nash) Gandhi & Smeins – Y] Schizachyrium sanguineum (Retzius) Alston var. hirtiflorum (Nees) Hatch, Hairy Crimson Bluestem. Cp (GA): {habitat}; rare. Sw. GA and FL west to AZ and south through Central America to South America. [= FNA, K; = Andropogon hirtiflorus (Nees) Kunth – HC, S; ? Sch. sanguineum var. brevipedicellatum (Beal) Hatch] Schizachyrium scoparium (Michaux) Nash var. scoparium, Common Little Bluestem. Cp, Pd, Mt (GA, NC, SC, VA): in a wide range of moist to dry habitats; common. (June-) August-October. New Brunswick west to Alberta, south to FL and Mexico. One of the most ubiquitous plants in the modern landscape of our area, occurring throughout in the majority of habitats. This species is extremely variable, some of the variability correlated with habitat and geography; the recognition of infraspecific taxa is warranted. [= C, FNA, Z; < Andropogon scoparius Michaux – RAB (also see Sch. littorale); = Sch. scoparium – GW; > Andropogon scoparius var. scoparius – F, G, HC; > Andropogon praematurus Fernald – F, G; > Andropogon scoparius var. polycladus Scribner & Ball – F; > Andropogon scoparius var. frequens F.T. Hubbard – F; = Sch. scoparium ssp. scoparium – K, Y; < Andropogon scoparius – S, W] Schizachyrium scoparium (Michaux) Nash var. stoloniferum (Nash) J. Wipff, Creeping Little Bluestem. Cp (GA, SC): fall-line sandhills in the inner Coastal Plain, perhaps in other dry habitats, the habitat and range in our area requiring further study; uncommon? August-October. SC and GA south to FL and west to MS. See Wipff (1996a) for additional discussion. [= FNA, K, Z; = Sch. stoloniferum Nash – GW; = Andropogon stolonifer (Nash) A.S. Hitchcock – HC, S; < Sch. scoparium ssp. littorale (Nash) Gandhi & Smeins – Y] Schizachyrium tenerum Nees, Slender Bluestem. Cp (GA): longleaf pine savannas; uncommon. GA west to e. TX. [= FNA, K; = Andropogon tener (Nees) Kunth – HC, S] Schizachyrium maritimum (Chapman) Nash. AL, FL west to LA. [= K; = Andropogon maritimus Chapman] {not keyed at this time; add to synonymy} Schizachyrium niveum (Swallen) Gould, Pinescrub Bluestem, is reported for Lowndes Co. in sc. GA (Kral 1973), but the report has been discounted by later authors (Wipff in FNA 2003a). Not keyed. [= FNA, K; = Andropogon niveus Swallen – HC, S] Schizachyrium scoparium (Michaux) Nash var. divergens (Hackel) Gould, Pinehill Bluestem. East to c. TN, AL, KY. [= FNA, K; = Andropogon scoparius Michaux var. divergens Hackel; = Andropogon divergens – HC; < Andropogon scoparius – S]

Sclerochloa Palisot de Beauvois References: Tucker (1996)=Z; Brandenburg, Estes, & Thieret (1991). * Sclerochloa dura (Linnaeus) Palisot de Beauvois. Mt (VA): disturbed areas; rare, introduced from Mediterranean Europe. A monotypic genus, native to s. Europe. [= C, HC, K, Z]

Secale Linnaeus 1753 (Rye) References: Tucker (1996)=Z. * 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 drooping-secund; 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 cross-

section, 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 short-pedicelled 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)

Thinopyrum (Prat) Á. Löve References: Tucker (1996)=Z; Barkworth (1997)=Y. * 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, clay-based 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]

Triticum Linnaeus 1753 (Wheat) References: Tucker (1996)=Z; Zohary & Hopf (1994). * 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 triangular-

lanceolate, 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, pond-shores, 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 reddish-brown, 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, reddish-

brown, 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]

Danae Medikus (Alexandrian Laurel, Danaë) A monotypic genus, a shrub, of sw. Asia.

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 upwardly-scabrous 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]

TRILLIACEAE Lindley 1846 (Trillium Family)

References: Farmer & Schilling (2002).

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 slow-growing 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 greenish-white 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 linear-

lanceolate (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 cross-section); 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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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