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LEPTOCHLOA (POACEAE: CHLORIDOIDEAE) IN COLOMBIA PDF

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LEPTOCHWA (POACEAE: CHLORIDOIDEAE) COLOMBIA IN Snow Neil Paul M. Peterson Herbarium Pacificum Department ofBotany Museum Bishop Museum National ofNatural History 1525 Bernice Street Smithsonian Institution Honolulu, HI 968 U.S.A. Washington, DC 2001 3-70 1 7, 12, U.S.A. [email protected] [email protected] Diego Giraldo-Canas Leptochloa P. Beauv., sensu lato (including Diplachne P. Beauv.) is a genus of approximately 32 annual or perennial species (Snow 1997). The native range of the genus pantropical into warmer temperate is regions, with several species being weedy and widely distributed. Although Parodi McNeill (1927), (1979), Phillips (1982), Nicora and Rugolo de Agrasar (1987), and Nicora (1995) recognized Diplachne as a separate genus, we McVaugh agree with Clayton and Renvoize Nowack Snow (1983), (1986), (1994), Renvoize (1998), and number 1998a) (1997, that the of florets, the size of the spikelets, shape of the caryopsis, arrangement of on spikelets the panicle branches secund or not secund), and other morphological characters (i.e., inter- grade strongly and are unable to support generic separation (Snow 1997). Moreover, based on numerical a analysis of morphological characters, Phillips (1982) found that species placed in Diplachne and Leptochloa overlapped one another in a principal coordinates scatter plot. Although it is likely (next paragraph) that some may elements of Leptochloa need to be segregated in the future with further analyses, in our opinion it is unlikely that such a generic segregate would correspond closely to Diplachne as recognized by the authors cited above. Thus time seems more at this it appropriate to recognize Leptochloa in broad sense (Peterson its Snow et 1997; 1997). al. The monophyly of Leptochloa sensu lato was not supported when tested cladistically with morphological and anatomical data, depending on the number of outgroups included (Snow 1997). Moreover, monophyly its has not been tested in breadth using molecular data, and the relationship of Leptochloa with other genera is from (Columbus far certain et al. 2007; Peterson et al. 1997, 2007). For example, in a restriction fragment cpDNA New analysis of using a few representatives of World genera (Duvall et 1994), Leptochloa dubia al. [L. (Kunth) Nees] was member common a of a basal clade, sharing a ancestor with Scleropogon Phil, Dasyochloa Willd. ex Rydb., Munroa Torr., and Erioneuron Nash. In a strict consensus tree with bootstrap analysis, species common of Leptochloa did not form a particularly strong clade with the other genera, and instead shared a ancestor in a clade that included the Muhlenbergiinae, Munroinae, Scleropogon, Sporobolus R. Br., Eleusine & & Roem. Tripogon Roem. Schult., and Eustachys Desv. (Duvall et al. 1994). In a Gaertn., Tridens Schult., phylogenetic study of Chloridoideae based on matK sequences, Leptochloa (L. dubia) formed a clade with & & Other Coelachyrum Hochst. Nees, and Astrebla Muell. emerged as sister to these (Hilu Alice 2001). F. Cynodon genera in the C clade of Hilu and Alice (2001) included Brachyachne (Benth.,) Stapf, Chloris Sw., x Dinebra Enteropogon Nees, Eustachys Desv., Lepturus R. Br., Lintonia Stapf, Microchloa R. Rich., Jacq., Eleusine, & and Fourn. Benth. In another phylogenetic study Oxychloris Lazarides, Tetrapogon Desf., Trichloris E. Br., on combined trnL-F and ITS sequences, the placement of species of Lep- Chloridoideae based analysis of of tochloa dubia, L.jusca (L.) Kunth, and L. panicea (Retz.) Ohwi] in separate clades suggests a polyphyletic [L. formed origin (Columbus et 2007). In a phylogenetic analysis of rpsl6 sequences, Leptochloa dubia a clade al. waxy whereas based on analysis of sequences, Leptochloa with Eleusine coracana (L.) Gaertn.], in a tree [E. and formed a clade with Eleusine coracana and Dactyloctenium aegyptium (L.) Willd. D. radulans (R. Br.) P. & Beauv. (Ingram Doyle 2007). In summary, the monophyly of Leptochloa has not been corroborated with (Columbus with other genera remain obscure molecular and phylogenetic relationships of Leptochloa data, genus et al. 2007, Peterson et al. 2007). For these reasons Peterson et al. (2007) treated the as incertae sedis within the tribe Cynodonteae. New which Leptochloa represented by 17 species in the World (13 of these native), six of occur in is Our taxonomic treatment contains keys Colombia (Penaloza-Jimenez 2002; Peterson et 2001, 2007). et al. al. and synonymies. Only determining the species, descriptions, distributions, specimens studied, illustrations, for Snow synonymy Peterson synonyms used frequently in Colombian literature are given. For further see (1997), et COAH, and Soreng This study based on the examination of herbarium specimens from (2001), et (2008). is al. al. CAUP, COL, HUA, MO, and US, including the type specimens of most species studied. The present study aims Program and to contribute to the knowledge of the Colombian grasses by collaborating with the Biodiversity Colombian Flora Inventories. Leptochloa Beauv, Ess. AgrostOgr. 71. 1812. Type: Leptochloa virgata (L.) P. Beauv. P. and and rhizomes but sometimes geniculate root- Plants annual or perennial; caespitose, lacking stolons cm ing the lower nodes in mesic habitats. Flowering culms 30-150 tall, ascending to erect, glabrous to at than internodes with sometimes pilose, arising from fibrous roots. Leaves cauline; sheaths longer or shorter smooth margins; ligules a membrane or a ciliate membrane; blades narrowly lanceolate to linear, flat, often primary branches exserted lower inrolled on drying, not pungent. Inflorescence a panicle of spicate (or main main primary branches occasionally inserted proximally) along an elongated axis; axis scabrous; main branches spaced or sometimes subdigitate, ascending to spreading from axis, terminating in a spike- glabrous or scabrous. Cleistogamous spikelets absent or present in axils of lower sheaths (in L. dubia or let; lemma above glumes, and palea falling as a unit; callus rarely in L.jusca subsp./asricularis). Disarticulation glabrous sparsely sericeous. Spikelets solitary, laterally compressed or terete, subsessile to shortly (<2 to mm) glumes than unequal, smooth, pedicellate, distant but often imbricate, often secund; shorter spikelets, glumes upper glumes shorter than lower lemma, 1-3-veined, glabrous or slightly scabrous; lower 1-veined; unawned; pronounced between glabrous; florets 2-13 per spikelet, sterile florets often pres- rachilla florets, lemma 3-veined above (and uppermost often reduced to small, awnless rudiment); ent fertile florets floret and sometimes membranous, smooth, glabrous or hairy along veins (rarely 4-5-veined in L. dubia at base), between apex emarginate or lobed, awnless, mucronate, or awned; paleas glabrous to sericeous, veins, entire, unawned, membranous, margins not enfolding fruit, smooth; lodicules truncate; stamens 2 or 3; anthers (sometimes weakly caryopses dorsiven- yellow or reddish-purple; stigmas 2. Fruit with adnate pericarp so); compressed compressed (Snow 1998a). Base chromosome number of x = 10. or laterally trally was Comments.— and Diplachne were established by Palisot de Beauvois (1812). Leptochloa Leptochloa Kunth based on three species, while Diplachne was based on D. fascicularis (Lam.) P. Beauv. [= L.jusca (L.) Snow et Leptochloa Colombia al., in 863 subsp. fascicularis (Lam.) N. Snow]. Leptochloa was later lectotypified with L. virgata (L.) Beauv. by Nash P. which was Nominum (1913), ineffective according Index Genericorum example to Art. 10.5 In addition, 6. both genera were placed in the same "cohors," which suggested that Palisot de Beauvois regarded them as more or less closely related. In his "Tabula Methodica" he mentioned that the lemmas of Diplachne were shortly awned, whereas those of Leptochloa were said to be awnless. However, this feature varies considerably and lemma the of Leptochloa can be entire, emarginate, lobed, awned, mucronate, or unawned. The micromor- lemma phological characters of the vary little in Leptochloa (Snow 1996), and with few exceptions, cork cells are present, silica cells are absent, chloridoid bicellular microhairs are present, prickles are present, and macrohairs are present. However, the occurrence of papillae on long cells and short cells is more variable, may suggesting their presence or absence be phylogenetically informative within the genus (Snow 1996). — Ecology and geographic distribution. The native range of Leptochloa nearly worldwide in warm-temperate is to tropical regions. Its ecological distribution mostly includes semi-open vegetation such as grasslands. The most grow species frequently in periodically disturbed or seasonally saturated areas such as riparian cor- ephemeral impoundments, ridors, pools, or although artificial L. dubia generally prefers well drained soils. All widespread annual species are considered weedy in agricultural situations (Snow 1997) and some have & become established on continents far beyond their native range (Snow Simon 1999; Snow 2004). KEY TO THE OF LEPTOCHLOA COLOMBIA SPECIES IN Sheaths sparsely to moderately pilose, th< 1 . Sheaths glabrous to sparsely the ha hairy, 1 . membranous Ligules and neither 2. cili membranous Ligules . on florets slightly diverging from anthe rachilla at s; Plants annual; 2.2-3.8 ligules long. Culms more round 5. or less in cross section; panicle branches rr mm mm long; anthers 0.6-0.8 long -C). ChlorisdubiaKumhJ mm Perennials, mostly erect or infrequently decumbent or sprawling. Culms (5-)30-110 cm 1.0-4.5 tall, wide, unbranched or only as tillers from very base, round or flattened below; nodes glabrous, internodes cm 3-11 long, herbaceous, solid (or occasionally hollow by virtue of aging). Leaf sheaths longer or shorter than internodes, sparsely pilose, especially below, and occasionally pilose (sometimes densely near the so) collar, the hairs occasionally from papillose bases, the margins glabrous or somewhat pilose; collars green mm or tan; ligules mostly 1.0-1.5 long, membranous, truncate, blades (2-)8-35 cm ciliate apically; long, mm 2-8 wide, mostly somewhat linear or lanceolate, scabrous above at base or with a sparse covering of pilose hairs, glabrous to minutely scabrous below, but drying involute, midvein mostly prominent flat or sometimes two not. Panicles of types, the apical ones generally exserted at maturity and the lateral ones completely hidden in lower leaf sheaths; apical panicles 10-45 cm long, (2-)3-25 cm wide; the branches cm (2-)5-15, (1.5-)3-19 long, alternate or infrequently subdigitate, ascending to reflexed, often somewhat mm lax, minutely scabrous, the branch axils pilose or merely scabrous. Spikelets 4.0-12.0 long, imbricate of the Botanical Research Institute of Texas 2(2) Snow et Leptochloa Colombia al., in mostly to distant, sessile; florets 4-13; callus glabrous or with a few short hairs; 1 mm membranous, long, narrowly triangular or ovate, scabrous along midvi mm acute; upper glumes 3.3-6.0 membranous, long, ovate to lanceolate, scabrous on midvein (and some- mm times with or 2 additional veins and on lemmas 1 at least basally) papillate edges, acute; 3.5-5.0 long, membranous, ovate to obovate or widely obovate, light brown, bronze, light to very dark green, the lateral veins usually prominent but sometimes not so, sericeous at least along lower portions of lateral veins and sometimes on midvein and between apex veins, the usually broadly awn- bifid, acute, obtuse, or truncate, less or mucronate; paleas subequal to lemma, lanceolate, membranous above but somewhat cartilaginous near base, distinctly ciliate on edges, sometimes sericeous between veins, apex acute; stamens anthers 3; mm mm mm (0.3-)1.0-1.6 long, yellow. Caryopses 1.5-2.3 long, 0.9-1.0 wide, narrowly elliptic, elliptic or very widely obovate in hilar profile, depressed obovate in transverse section, with a broad, shallow groove on smooth, brown; hilar side, pericarp weakly adnate endosperm. the to USA Distribution and habitat.—Native in the from Arizona to Oklahoma and Texas, southern Florida, common in Mexico, sporadically in the Caribbean and in Mesoamerica through more common Bolivia, again from Bolivia and Chile through Paraguay, Uruguay, and southern and much Brazil of Argentina; in a variety and of soil vegetation types, but most frequently on well-drained slopes, 100-2600 m, but occasionally to m 3150 in Peruvian and Bolivian Andes. Syn. PI. Glumac. 1:247. 1855. Type: PALESTINE: F. Hasselquist s.n. (lectotype: LINN 92.21!, designated by Phillips, Fl. Trop. East Gram. Afr. 2:281. 1974). Annuals, ascending sometimes and Culms cm to erect, geniculate rooting lower nodes. 130 (when at to tall mm 1-8 erect); wide, branched or unbranched; nodes glabrous; internodes (0.5-)3-26 cm long, hollow, herbaceous sometimes or firm. Leaf sheaths longer or shorter than the internodes, glabrous on and sides mm margins; mostly brown 5-8 collars light or green; ligules long, hyaline, apically attenuate but often mm becoming lacerated due to mechanical damage; blades (3-)5-50 cm 2-6 long, wide, linear, gla- flat brous to somewhat scabrous above and below. Panicles 15-150 cm mostly 2-30 cm long, wide, partially included at base to completely exserted; the branches (1.5-)3-20 cm long, (3-)5-35 per panicle, alternate, sometimes reflexed or steeply but mostly somewhat erect ascending, minutely scabrous, the branch stiff, mm axils glabrous. Spikelets 5-12(-14) more long, short pedicellate, or less secund, sometimes distant near base of branches but overlapping near branch tips; florets (3-)6-12; callus glabrous or hairy; lower glumes mm membranous, 1.0-3.5(-4.9) long, lanceolate to ovate, usually scabrous on the midvein, broadly acute mm to acute, awnless or infrequently shortly awned; upper glumes 1.8-5.5 long, elliptic, usually ovate or widely sometimes ovate, obovate, scabrous on midvein, obtuse (rarely) or acute at apex, rarely short-awned; mm lemmas 2.3-6.0 membranous, long, elliptic, ovate, or lanceolate, the lateral veins distinct and often more slightly excurrent, or less sericeous on lateral veins and the midvein; apex truncate, obtuse, to acute mm or acuminate and sometimes awn awnless, mucronate, awned, bifid, or the to 3.5 long; paleas subequal or slightly exceeding lemma, more or less sericeous on veins; apex acute or obtuse; stamens 2 or mostly 1, mm mm mm anthers 0.2-2.7 Caryopses 3; long. 1.0-2.4 long, 0.7-1.2 wide, elliptic, ovate, or obovate in hilar profile, transversely elliptic to depressed obovate in transverse section, hilar groove lacking, smooth sometimes or slightly rugose, brown; pericarp weakly adnate the endosperm. to common Distribution.—Widespread and abundant warm-temperate and to in between tropical areas, New approximately 49°N and 40°S in the World and 40°N and 42°S in Old World; mostly below 2000 m. — Comments. Populations of the Leptochloa fusca complex often appear morphologically distinct in lo- . Journal of the Botanical Research Institute of Texas 2(2) 866 and names been forms throughout the Americas, Africa, and dozens of have applied to regional cal areas, Since morphological characters intergrade when considered globally most local variants do Australia. all not merit taxonomic recognition (Snow 1997). However, herbarium work and multivariate statistical studies and support (Snow, unpubl.) based on eleven population samples (n = 30) from America, Africa, Australia, recognition of four subspecies. The subspecies, which are usually easy to distinguish, are: L.jusca subsp. the New polymorphic paleotropical taxon adventive in a few areas in the World (Nicora 1995); L.jusca fusca, a known much and Simon subsp. muelkri (Benth.) N. Snow, from of the interior of Australia (Sharp 2002); and fusca subsp. uninervia, native to the neotropics but adventive elsewhere; L. fusca 3sp. fascicularis, L. New native to the temperate and tropical regions of the World. COLOMBIA KEY TO SUBSPECIES OF LEPTOCHLOA FUSCA IN panicle branches often inserted upper sheath; uppermost leaf blade lengtl usu- in ng length of panicle; leaf sheaths sometimes mottled with anthocyanin pigments; len Lowermost panicle branches generally exserted from sheath; uppermost leaf blade length generally not ex . ceeding terminal panicle; leaf sheaths only rarely mottled with anthocyanin pigments; lemmas smoky white o times dark green or plumbeous, but usually n< mm Culms (3.5-)15-130 cm l-5(-8) wide, often branching, round; nodes Annuals, ascending to erect. tall, 3-18 cm Leaf sheaths mostly longer than the internodes, herbaceous, hollow. glabrous; internodes long, mm cm 3-45 on and margins; green or brown; ligules 5-7 long, hyaline; blades glabrous sides collar light mm when becoming long, 2.5-7 wide, linear, usually sparsely scabrous above and below, flat but involute cm cm included midvein prominent. Panicles (1.5-)10-72 long, (l-)4-22 wide, generally partially dry, cm ascending, below; the branches 3-35, (0.5-)3-22 long, alternate, erect, 5-12 mi or slightly flexuous, minutely scabrous, the axils glabrous. Spikelets mm lower g" cate, pedicels less than 1 long; florets 6-12; callus glabrous; lanceolate or somewhat asymmetric, scabrous on midvein, acute to aris mm membranous, scabro upper glumes 2.5-5.0 long, elliptic to ovate, mm lemmas membranous, short-awned; 2.5-5 long, lanceolate, light to white, the lateral veins pronounced and extending to edges often as sn veins and often midvein, glabrous between veins; apex acute to lateral : lemma, awns subequal sericeou< 3.5 long; paleas generally to elliptic, to mm mm stamens anthers 0.2-0.5 long, yellow. Caryopses 1-2 long, 0. 3; groove smooth, brown; peric transversely in transverse section, hilar lacking, in hilar profile, elliptic weakly adnate the endosperm. to Washing and habitat—Massachusetts (USA) and southern Ontario (Canada) west to Distribution 40° open mesic brackish marshes along the east (USA) and south about latitude S in Argentina; areas, to from 1850 US seaboard, agricultural lands, ruderal sites along roads, and saline flats. Elevation sea level to Specimens examined. COLOMBIA. La Guajira: Costa del Caribe, llanura literal arenosa y arida entre Mayapo y El Pajaro, 29 Jan 1! Dugand 6667 (COL, US). A. Novon D-F). 2b. Leptochloa fusca subsp. uninervia Presl) N. Snow, 8:79. 1998b. (Fig. 2, Megasta (J. Annuals or occasionally weakly perennial in regions lacking erect or ascending. Culms (15-)25-110 frosts, mm cm 2-5 cm tall, wide, often branching, round; nodes glabrous; internodes 2-11 long, herbaceous, hollow. Leaf sheaths longer or shorter than the internodes, often flattened below, glabrous on sides and margins; mm mm 5-8 cm collars mostly tan or green; ligules long, hyaline; blades (2-)5-37 long, 2.0-5.5 wide, linear, usually densely scabrous above, sparsely to densely scabrous below, flat but becoming involute when dry, 868 cm cm midvein prominent. Panicles 10-57 long, (0.5-)3-18 wide, usually exserted; the branches (3-)10-60, cm (0.3-)2-ll long, the branches mostly alternate, ascending to erect, stiff or slightly flexuous, minutely mm scabrous, the axils glabrous. Spikelets 5-10 long, rarely distant to normally imbricate (sometimes tightly mm so), pedicels mostly less than 1 long; florets (3-)6-10; callus sparsely sericeous; lower glumes 1.0-2.6 mm long, membranous, narrowly triangular, lanceolate, or ovate, glabrous or scabrous on midvein, acute to mm aristate or mucronate; upper glumes 1.8-2.8 long, membranous, obovate to widely ovate, glabrous or mm lemmas membranous, scabrous on midvein, obtuse, acute, or rarely mucronate; 2.0-3.6 long, ovate or elliptic, light brown to very dark green or somewhat plumbeous, lateral veins more or less prominent and generally extending to edges, sparsely sericeous below on lateral veins and often midvein, glabrous between veins; apex broadly acute, more commonly obtuse to truncate, sometimes bifid or mucronate; paleas subequal or slightly longer than lemma, membranous, elliptic, sericeous along veins; apex obtuse; stamens 3; anthers mm mm mm 0.4-1.0 long, yellow. Caryopses 1.0-1.5 long, 0.7-0.8 wide, elliptic, ovate, or obovate in hilar smooth brown; transversely in transverse section, hilar groove lacking, or slightly rugose, profile, elliptic pericarp weakly adnate endosperm. to — New Distribution and habitat. In the World mostly south of latitude 37°N, south to Argentina, occa- sionally adventive in Old World (Snow and Simon 1999); open mesic areas, agricultural lands, saline flats, mangrove swamps. from 1200 m. Elevation sea level to is Specimens examined. COLOMBIA. Atlantico: Entre Palmar de Varela y Ponedera, bosques xerofiticos, 10 m, 27 Ago 1960, A. Dugand 5304 (COL, US). Magdalena: Cienaga, alrededores de Aguacoca, 20 m, 21 Feb 1959, R. Romero-Castaneda 7268 (COL, US). Novon 3. Leptochloa panicea subsp. brachiata (Steud.) N. Snow, 8:79. 1998b. (Fig. 3, A-E). Leptochloabmchiata GUADALOUPE US Steud., Syn. Glumac, 209. 1854. Type: [French Republic]: Duchassaing s.n. (holotype: P!; fragm.!). Pi. SOUTH Pers., Syn Pi. 1: 87. 1805. Leptochloa filiformis (Pers.) P Beauv., Ess. Agrostogr.: 71, 163, 166. 1812. Type: lis mm cm Annuals, ascending Culms (10-)30-150 mostly than 5 wide, frequently branched; to erect. tall, less cm nodes glabrous; internodes mostly 5-30 long, hollow, herbaceous or sometimes firm. Leaf sheaths lon- ger or shorter than the internodes, sparsely to moderately pilose (hairs with papillose bases); collars often mm mm cm magenta; ligules 1.0-3.2 long, membranous but erose apically; blades 6-25 long, 2-21 wide, cm cm glabrous sparsely pilose on both surfaces. Panicles 8-30 long, usually 5-35 wide, linear, flat, to cm occasionally partially included at base; the branches (l-)3-19 long, (3-)10-100 per panicle, alternate, ascending (mostly) to somewhat reflexed apically (but otherwise firm), minutely scabrous, the branch axils mm glabrous. Spikelets 2-4 long, short pedicellate, more or less secund, sometimes distant near base of mm branches but overlapping near branch tips; florets 2-5(-6); callus glabrous; lower glumes 1.6-4 long, mm membranous, lanceolate, usually scabrous on the midvein, acute, awnless; upper glumes 1.6-3.6 long, mm lemmas membranous, more on lanceolate, awnless; 1.3-1.7 long, lanceolate, or less sericeous lateral veins and the midvein; apex acute, awnless; paleas subequal to lemma; apex acute or obtuse; stamens 3; mm mm mm more rounded anthers 0.2-0.3 long. Caryopses 0.9-1.2 long, 0.5-0.6 wide, or less in hilar profile, widely depressed obovate to obdeltoid in transverse section, hilar groove narrow and shallow, smooth or sometimes slightly rugose, brown to reddish; pericarp adnate to the endosperm. — Distribution and habitat. Southern third of United States through Mesoamerica and the West Indies to southern Brazil and Argentina; disturbed, mesic, and agricultural sites, to ca. 2800 m. — much warm Comments. Leptochloa panicea a polymorphic species occurring throughout of the is temperate and tropical regions, and comprised of three subspecies (Snow 1997, 1998a). Snow and Davidse is New (1993) reported that the name then in use for the widespread World taxon, L. filiformis (Lam.) P. Beauv, homonym was an should be replaced by mucronata (Michx.) Kunth. Although not illegitimate later that L. New the do Nowack (1994, 1995) recently placed World specimens as a subspecies of the oth- first to so, erwise Old World species panicea, and made the new combination L. panicea subsp. mucronata. Included L. in Nowack's concept of L. panicea subsp. mucronata were two taxa formerly recognized as L. filiformis vars. Snow and attenuata, which (1997, 1998b) treated (respectively) as L. panicea subsp. brachiata (Steud.) filiformis Snow et Leptochloa Colombia al., in A. Habit. B. Inflorescence. C. Portion of the leaf sheath with a covering of papillose-based e;. I 870 N. Snow and panicea subsp. mucronata (Michx.) N. Snow. The much more widespread taxon, referred to L. & new name (Snow was earlier as L.filiformis or L. mucronata Davidse, 1993), given the L. panicea subsp. (Snow brachiata 1998b). Drawing on and experience exa field eluded that the phenetic variation sufficiently c is L. panicea subsp. brachiata is present in Colombia. — Vernacular name. Pasto paja-mona (Valle del Cauca). COLOMBIA. km Specimens examined. Antioquia: Mun. Necocli, 42 carret mm cm Annuals, often geniculate below but generally Culms (7-)45-110 1-7 wide, branch- erect. tall, cm ing (sometimes profusely), round; nodes glabrous; internodes (l-)4-10 long, herbaceous, hollow. somewhat Leaf sheaths generally longer than the corresponding internodes, flatted below, glabrous, the margins sometimes very sparsely ciliate along the lower half; collars sometimes magenta; ligules 2.2-3.8 mm mm long, membranous, truncate, somewhat erose apically; blades 4-20 cm long, mostly 4-8 wide, and linear or the shorter ones lanceolate, glabrous to minutely scabrous above below. Panicles (l-)20-35 cm cm cm long, (3-)4-12 wide, generally exserted; the branches (20-)40-90, 2.5-7 long, alternate, mostly mm 4-5 ascending, to weakly flexuous, minutely scabrous, the axils glabrous. Spikelets long, generally stiff mm somewhat imbricate, pedicles 1-2 long; florets (4-)5 or 6(-7); callus glabrous; lower glumes 0.9-1.9 mm long, membranous, narrowly lanceolate to lanceolate or sometimes falcate, minutely scabrous on the mm midvein, acute; upper glumes 1.8-2.3 long, membranous, ovate, minutely scabrous on the midvein, mm acute to obtuse; lemmas 2.4-3.0 long, membranous, narrowly elliptic to ovate, tan, maroon, or green, at base of lateral veins and/or midvein; apex acute to broadly acute, awnless or mucronate; paleas subequal to ca. 2/3 as long as the lemma, membranous, ovate to elliptic, glabrous throughout; apex obtuse; stamens mm mm mm maroon anthers 0.6-0.8 long, or yellow. Caryopses 1.1-1.4 long, 0.7 wide, elliptic in hilar 3;

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