A Revision of Panicum Sect. Dichotomomiflora (Poaceae - Panicoideae - Paniceae)

DARWINIANA, nueva serie 10(2): 325-364.
2022
Versión de registro, efectivamente publicada el 13 de septimebre de 2022
DOI: 10.14522/darwiniana.2022.102.1048
A REVISION OF PANICUM SECT. DICHOTOMIFLORA (POACEAE:

PANICOIDEAE: PANICEAE)
Fernando O. Zuloaga ID
Instituto de Botánica Darwinion (IBODA, ANCEFN-CONICET), Labardén 200, CC 22, B1642HYD, San Isidro, Buenos
Aires, Argentina; fzuloaga@darwin.edu.ar
Abstract. Zuloaga, F. O. 2022. A revision of Panicum sect. Dichotomiflora (Poaceae: Panicoideae: Paniceae).
Darwiniana, nueva serie 10(2): 325-364.
Panicum sect. Dichotomiflora comprises a complex of annual and perennial species, distributed
all over the world. They are characterized by its glabrous, ellipsoid to ovoid or lanceolate spikelets,
with the lower glume 1/6 to 1/4(-1/3) the length of the spikelet, nerveless to 3-nerved, and upper
glume and lower lemma subequal, 7-11(-13)-nerved. A total of 20 species are recognized in the
section: Panicum aquaticum, P. dichotomiflorum, P. gilvum, P. impeditum, P. lacustre, P. laevinode,
P. luzonense, P. madipirense, P. mlahiense, P. obseptum, P. paludosum, P. perangustatum, P. pilgerianum,
P. porphyrrhizos, P. schinzii, P. subalbidum, P. sublaeve, P. sumatrense, P. trichonode, and P. vaseyanum.
Also, lectotypes are designated for Panicum hygrophilum, P. ingens, P. lacustre, P. longiramum,
P. oryzetorum, P. madipirense, P. porphyrrhizos, P. praelongum, P. proliferum var. longijubatum, and
P. telmatodes. A geographic analysis of the species is presented, together with illustrations, and a key for
the species. Additional comments on morphology of sect. Dichotomiflora are also included.
Keywords. Panicum sect. Dichotomiflora; Poaceae; taxonomy.
Resumen. Zuloaga, F. O. 2022. Revisión de las especies de Panicum sect. Dichotomiflora (Poaceae: Panicoideae:
Paniceae). Darwiniana, nueva serie 10(2): 325-364.
Panicum sect. Dichotomiflora incluye especies anuales y perennes distribuidas en todo el Mundo.
Sus taxones se caracterizan por las espiguillas glabras, elipsoides a ovoides o lanceoladas, con la gluma
inferior 1/6 a 1/4(-1/3) del largo de la espiguilla, enervia a 3-nervia, y gluma superior y lemma inferior
subiguales, 7-11(-13) nervias. Se reconocen un total de 20 especies en la sección: Panicum aquaticum,
P. dichotomiflorum, P. gilvum, P. impeditum, P. lacustre, P. laevinode, P. luzonense, P. madipirense,
P. mlahiense, P. obseptum, P. paludosum, P. perangustatum, P. pilgerianum, P. porphyrrhizos, P. schinzii,
P. subalbidum, P. sublaeve, P. sumatrense, P. trichonode, y P. vaseyanum. Se designan lectotipos
para Panicum hygrophilum, P. ingens, P. lacustre, P. longiramum, P. oryzetorum, P. madipirense,
P. porphyrrhizos, P. praelongum, P. proliferum var. longijubatum, y P. telmatodes. Se presenta la
distribución geográfica de las especies; también se incluyen ilustraciones, una clave de diferenciación
de las especies, y observaciones de la morfología de la sección Dichotomiflora.
Palabras clave. Panicum sect. Dichotomiflora; Poaceae; taxonomía.
INTRODUCTION nearly 450 species distributed worldwide (Clayton

& Renvoize, 1986; Zuloaga, 1987), and inhabiting
Panicum L., as traditionally circumscribed, habitats from sea level to approximately 2500 m.
was one of the largest genera of the Poaceae, with The main character placing species in the genus
Original recibido el 23 de mayo de 2022, aceptado el 19 de julio de 2022

Editor Asociado: Carolina Inés Guerreiro 325
DARWINIANA, nueva serie 10(2): 325-364. 2022
was the spikelet structure, with a lower glume and Megathyrsus (Pilg.) B.K. Simon & S.W.L.
present, usually shorter than the upper glume and Jacobs (Zuloaga et al., 1998; Aliscioni et al., 2003;
lower lemma, the latter subequal, a lower flower Freckmann & Lelong, 2003; Simon & Jacobs,
present or absent, the upper anthecium indurate 2003; Barkworth, 2006). Also, several new genera,
and abaxially convex, and a caryopsis with a from species previously placed in Panicum, have
punctiform to oblong hilum. However, these been established (Bess et al., 2006; Zuloaga et al.,
characters are also present in other members of 2010, 2015; Scataglini & Zuloaga, 2013). Finally,
the Paniceae and Paspaleae. On the other hand, a Zuloaga et al. (2018) restricted the genus, within
wide array of differential characters were present subtribe Panicinae to its type subgenus (Soreng
in Panicum sensu lato, including photosynthetic et al., 2022), with approximately 163 species, and
pathway, with species C3, C4 and intermediate all remaining species classified as “incertae sedis”
C3/C4, anatomical features associated with the within tribes Panicinae and Paspaleae. Within
Kranz syndrome, and chromosome numbers. These Panicum s. str., Zuloaga et al. (2018) propose seven
characters were evaluated by Giussani et al. (2001), sections as follows: sect. Arthragrostis (Lazarides)
Aliscioni et al. (2003) and Morrone et al. (2012), Zuloaga, endemic of Australia and southeast Asia,
who concluded that Panicum was a polyphyletic sect. Yakirra (Lazarides & R.D. Webster) Zuloaga,
genus, and that many of its species were in need endemic of Australia, sect. Rudgeana (Hitchc.)
of transfer to other genera of grasses. As a result, Zuloaga, endemic of America, and four pantropical
subgenera were raised or reinstated at the generic sections: sect. Dichotomiflora (Hitchc.) Hitchc. &
level, such as Dichanthelium (Hitchc. & Chase) Chase ex Honda, sect. Hiantes Stapf, sect. Panicum,
Gould, Phanopyrum Raf., Steinchisma Raf., and sect. Repentia Stapf.
Sections of Panicum can be distinguished by the following key
1. Rachilla prominent between the glumes and lower lemma ...................................................................... 2

1. Rachilla not prominent betwen the glumes and lower lemma .................................................................. 3
2(1). Upper anthecium stipitate, with a pair of winged appendages ........................................... Sect. Yakirra
2. Upper anthecium stipitate or not, without a pair of winged appendages ....................... Sect. Arthragrostis
3(1). Plants with an heterogenous stipe in the base of the upper anthecium .......................... Sect. Rudgeana
3. Plants not stipitate, if a stipe if rarely present, it is homogenous in the base of the upper anthecium ........ 4
4(3). Plants annual or perennial, if perennial shortly rhizomatous, without prominent rhizomes ................. 5
4. Plants perennial, with prominent rhizomes .............................................................................................. 6
5(4). Lower glume 1/3 or less the length of the spikelet, truncate to obtuse or subacute. Plants aquatic or
palustrial, usually present in open and humid environments ........................................ Sect. Dichotomiflora
5. Lower glume 1/2 or more the length of the spikelet, acute. Plants of open and dry enviroments, growing
in campos, savannas or cerrados .............................................................................................. Sect. Panicum
6(4). Lower glume 1/4 to 1/3 the length of the spikelet, occasionally up to 1/2; spikelets no gaping at maturity
.................................................................................................................................................... Sect. Repentia
6. Lower glume 1/2 to 4/5 the length of the spikelet; spikelets gaping at maturity ................... Sect. Hiantes
The aim of this work is to complete a worldwide HISTORY AND MORPHOLOGY

revision of species of Panicum sect. Dichotomiflora,
including section characters, key to species, Panicum L., Section Dichotomiflora was
description, distribution, and illustrations of initially treated as a group by Hitchcock & Chase
16 species. (1910), who considered four species within it:
326
F. O. Zuloaga. Revision of Panicum sect. Dichotomiflora
P. vaseyanum Scribn., P. bartowense Scribn. Distribution and ecology. All the species
& Merr., P. dichotomiflorum Michx., and P. distribution, here discussed, are natural. Five
elephantipes Nees [= Louisiella elephantipes (Nees species of sect. Dichotomiflora are restricted to
ex Trin.) Zuloaga]. Later, Honda (1930) considered America: P. aquaticum Poir., with two varieties, P.
this group at the sectional level. Stapf (1920) treated dichotomiflorum, with two subspecies, P. lacustre
African species of this section, P. porphyrrhizos Hitchc. & Ekman, P. sublaeve Swallen, and P.
Steud. and P. schinzii Hack., within sect. Coloratae vaseyanum.Two species are endemic of Australia: P.
(= sect. Repentia), while Pilger (1931, 1940), laevinode Lindl. and P. obseptum Trin. Three species
considered in sect. Dichotomiflora the following grow in Asia: P. luzonense J. Presl, P. paludosum
species: P. dichotomiflorum, P. elephantipes Nees Roxb., and P. sumatrense Roth ex Roem. & Schult.;
(= Louisiella elephantipes), P. longijubatum Stapf while ten species are restricted to Africa, three of
(= P. subalbidum Kunth), and P. paludosum Roxb. them: P. gilvum Launert, P. impeditum Launert,
Zuloaga et al. (2018) concluded, in their molecular and P. schinzii Hack., growing in southern Africa;
analysis based on nine species of the group, that P. subalbidum in Africa and Madagascar, and the
Dichotomiflora is a monophyletic section and remaining species in central Africa: P. madipirense
tentatively place 17 species within it. Renvoize, P. mlahiense Renvoize, P. perangustatum
Renvoize, P. pilgerianum (Schweick.) Clayton, P.
Species of sect. Dichotomiflora are characterized porphyrrhizos, and P. trichonode Launert & Renvoize.
as being annual, or less frequently perennial species, All species are typically found in open and humid
with culms erect to geniculate ascending, decumbent habitats, occasionally on forest edges, are common
and rooting at the lower nodes; internodes are in margins of lakes, rivers and ponds.
hollow, with or without aerenchyma, and nodes
are conspicuously pilose to glabrous. Sheaths are
open, striate, pilose, with or without urticant hairs, CHROMOSOME NUMBERS
or glabrous. Ligules are membranouswith a ciliate
margin. Blades are linear to lanceolate, flat, less Panicum aquaticum, n = 36 (Davidse & Pohl,
frequently involute, pilose or glabrous, rounded to 1972); 2n = 36 (Davidse & Pohl, 1974; Mehra &
cordate at the base, the apex acuminate. Inflorescence Chaudhary, 1974); 2n = 72 (Pohl & Davidse, 1971).
is an open to contracted, terminal panicle, exserted Panicum dichotomiflorum, n = 18 (Davidse &
or partially included on the upper blades, with Pohl, 1974, under P. chloroticum; Dubcovsky &
branches alternate to opposite, spikelets paired or Zuloaga, 1991); 2n = 36 (Brown, 1948), 2n = 54
solitary and appressed or loose on the branches, (Church, 1929; Gould, 1958, 1968; Probatova
pedicels claviform, scabrous to smooth; axillary & Sokolovskaya, 1983; Hamoud et al., 1994).
panicles are present or absent. Spikelets are long Panicum luzonense, 2n = 18 (Veldkamp, 1996).
ellipsoid to ellipsoid, ovoid or lanceolate, glabrous, Panicum paludosum, 2n = 36, 54 (Veldkamp, 1996).
pale to greenish or tinged with purple, solitary or Panicum schinzii, 2n = 18, 40 (Veldkamp, 1996).
paired, the lower glume usually 1/6 to 1/4, less Panicum sublaeve, n = 18 (Dubcovsky & Zuloaga,
frequently up to 1/3 the length of the spikelet, 1991).
nerveless to 3(-7)-nerved, acute to truncate; upper Panicum sumatrense, 2n = 14, 36, 40, 54 (Veldkamp,
glume and lower lemma subequal, membranous, 1996).
7-11(-13)-nerved, acute; lower palea as long as the
upper lemma, hyaline, glabrous, to reduced or absent;
lower flower staminate, anthers 3, or absent; upper MATERIAL AND METHODS
anthecium as long as or shorter than the upper glume
and lower lemma, indurate, smooth, shiny, glabrous; Morphological and anatomical analyses were
upper lemma 5-7-nerved. The caryopsis is ellipsoid based on herbarium specimens from B, BA,
to ovoid or obovoid, with the hilum punctiform to BAA, BAF, BR, C, COL, CORD, F, G, GH,
oblong, rarely linear, embryo 1/3 to more than 1/2 ICN, K, LE, LIL, M, MA, MBM, MEXU, MO,
the length of the caryopsis. NY, P, SI, US, VEN, W, and WIS (Thiers, 2022).
327
Types were studied in person or with images glabrous; axillary inflorescences present or absent.
available online at the JStor Global Plants website Spikelets ellipsoid to long ovoid or lanceolate, pale
(http://plants.jstor.org) and/or at the websites of to greenish or tinged with purple, glabrous, solitary
aforementioned herbaria. or paired; lower glume 1/6 to 1/4(-1/3) the length of
the spikelet, nerveless to 1-3(-7)-nerved, apex acute
to truncate; upper glume and lower lemma subequal,
TAXONOMIC TREATMENT upper glume 7-11(-13)-nerved, lower lemma
7-11(-13)-nerved; lower palea elliptic, hyaline,
Panicum sect. Dichotomiflora (Hitchc.) Hitchc. & glabrous, as long as the lower lemma to reduced
Chase ex Honda, J. Fac. Sci. Univ. Tokyo, Sect. 3, or absent; lower flower staminate or absent; upper
Bot. 3(1): 244, 246. 1930. Panicum [unranked] anthecium ellipsoid to long ovoid, pale to dark
Dichotomiflora Hitchc., N. Amer. Fl. 3(2): 200, 202. brown at maturity, glabrous, smooth, indurated,
1915. Panicum group Dichotomiflora Hitchc. & as long as the upper glume and lower lemma or
Chase, Contr. U.S. Natl. Herb. 15: 28, 47. 1910, nom. shorter, occasionally early deciduous at maturity;
inval. Type species: Panicum dichotomiflorum Michx. upper lemma 5-7-nerved, upper palea 2-nerved,
Plants annual or perennial, the latter shortly inclosed by the lemma. Caryopsis ellipsoid to
rhizomatous, with culms erect to decumbent, ovoid or obovoid; hilum punctiform to oblong,
rooting and branching at the lower nodes, hollow, ocassionally linear, embryo 1/3 to little more than
with or without aerenchyma. Ligules membranous 1/2 the length of the caryopsis.
with a ciliate margin. Blades oblong-lanceolate to
lanceolate, linear-lanceolate or linear, flat, cordate to Section with 20 species distributed in all
rounded at base, pilose to glabrous. Inflorescence a continents; they grow in open and humid places,
terminal panicle, lax to contracted, multiflowered to being common elements of margins of rivers,
few flowered, fully exserted or partially included on swamps, lakes, and also present in forest edges. A
the upper blades; peduncle and main axis pilose to few species are fully aquatic.
Key to the species of Panicum sect. Dichotomiflora
1. Lower palea conspicuous, as long as the lower lemma; lower flower staminate, anthers 3 ..................... 2
1. Lower palea reduced; lower flower absent ................................................................................................ 7
2(1). Hilum linear, as long as the caryopsis; culms with aerenchyma ..................................... P. pilgerianum
2. Hilum punctiform to oblong, short; culms without aerenchyma ............................................................... 3
3(2). Upper glume and lower lemma 11-13-nerved ....................................................................... P. sublaeve
3. Upper glume and lower lemma 7-9(-11)-nerved ....................................................................................... 4
4(3). Lower glume 1/4 to 1/3 the length of the spikelet, 1-3-nerved ............................................................. 5
4. Lower glume 1/6 to 1/5 the length of the spikelet, nerveless to 1-nerved ................................................. 6
5(4). Plants annual; blades lanceolate, 0.8-1.2 cm wide, cordate. Africa................................. P. madipirense
5. Plants perennial; blades linear, 0.3-0.6 cm wide, rounded at base. America .......................... P. aquaticum
6(4). Nodes conspicuously pilose; blades linear-lanceolate, up to 0.5 cm wide; axillary inflorescences absent;
plants perennial .......................................................................................................................... P. trichonode
6. Nodes glabrous; blades lanceolate, up to 1.5 cm wide; axillary inflorescences present; plants annual ......
......................................................................................................................................................... P. schinzii
7(1). Inflorescence pauciflowered, partially included on the upper sheaths, 2-10 cm long .......................... 8
7. Inflorescence multiflowered, exserted, up to 50 cm long ........................................................................ 11
8(7). Upper glume 7-nerved .......................................................................................................................... 9
8. Upper glume 9-11(-13)-nerved ............................................................................................................... 10
9(8). Axillary inflorescences present; upper anthecium early deciduous at maturity; plants annual. America
................................................................................................................................................... P. vaseyanum
328
9. Axillary inflorescences absent; upper anthecium not early deciduous; plants perennial. Australia ...........
...................................................................................................................................................... P. obseptum
10(8). Lower glume nerveless to 1-nerved; spikelets long ellipsoid, 2.5-3.4 mm long ................... P. gilvum
10. Lower glume 3-5-nerved; spikelets lanceolate, 3.5-4.5 mm long ................................. P. perangustatum
11(7). Plants perennial ................................................................................................................................ 12
11. Plants annual ......................................................................................................................................... 14
12(11). Spikelets 2 mm long; upper glume and lower lemma 7-nerved. Cuba .............................. P. lacustre
12. Spikelets 2.5-4 mm long; upper glume and lower lemma 7-9-nerved. Africa ...................................... 13
13(12). Culms with aerenchyma, decumbent and rooting at the lower nodes; nodes sharply demarcated ....
.................................................................................................................................................. P. subalbidum
13. Culms without aerenchyma, geniculate to erect, not decumbent and rooting at the lower nodes; nodes not
sharply demarcated .............................................................................................................. P. porphyrrhizos
14(11). Upper glume 11-15-nerved .......................................................................................... P. sumatrense
14. Upper glume 7-9-nerved ....................................................................................................................... 15
15(14). Spikelets 2.2-2.5 mm long; lower glume up to 1/3 the length of the spikelet, 3-5-nerved ............. 16
15. Spikelets 2.5-4.2 mm long; lower glume 1/5 to 1/3 the length of the spikelet, nerveless to 3-nerved ......
...................................................................................................................................................................... 18
16(15). Sheaths covered with urticant hairs; blades with tubercullate hairs all over its surface; upper glume
and lower lemma with anastomosed nerves. Asia ....................................................................... P. luzonense
16. Sheaths and blades glabrous; upper glume and lower lemma without anastomosed nerves ................. 17
17(16). Blades lanceolate, 3-6(-10) × 0.3-0.4 cm; panicles up to 17 cm long. Africa ................. P. mlahiense
17. Blades oblong-lanceolate to lanceolate, 10-45 × 0.5-1.4 cm; panicles up to 30 cm long. America ..........
.......................................................................................................................................... P. dichotomiflorum
18(15). Plants with aerenchyma, aquatic; culms with aerenchyma ............................................ P. paludosum
18. Plants not without aerenchyma, growing in humid areas, not aquatic; culms without aerenchyma .........
..................................................................................................................................................................... 19
19(18). Panicle with the main axis pilose; peduncle with tubercullate hairs; axillary inflorescences absent.
Australia ....................................................................................................................................... P. laevinode
19. Panicle with the main axis glabrous; peduncle without tubercullate hairs; axillary inflorescences present
..................................................................................................................................................................... 20
20(19). Blades 10-45 × 0.5-1.5 cm; panicles up to 30 cm long; spikelets 2.4-3 mm. America ......................
........................................................................................................................................... P. dichotomiflorum
20. Blades 4-11 × 0.3-0.5 cm; panicles up to 14 cm long; spikelets (2.8-)3.1 mm. Africa ............ P. impeditum
1. Panicum aquaticum Poir. both surfaces glabrous, the lower margins ciliate o
Plants perennial, shortly rhizomatous, 80-200 cm not. Inflorescence a terminal exserted, lax panicle,
tall; culms decumbent and rooting at the lower 15-26(-30) × 8-15 cm; peduncle terete, glabrous;
nodes, then erect; internodes terete, glabrous, main axis terete, glabrous, pulvini brownish,
hollow; nodes brownish, glabrous. Sheaths striate, glabrous, first order branches alternate, the upper
8-15 cm long, usually longer than the internodes, ones whorled or alternate, axis of the branches and
densely papillose-pilose all over its surface or pedicels triquetrous, scabrous, spikelets paired
densely hispid toward the upper portion or glabrous, and congested on second order branches, pedicels
the margins membranous. Ligules membranous- 2-4 mm long. Spikelets long ovoid, 2.7-3.5 × 1
ciliate, 1-4 mm long; collar pale to brownish, pilose mm, greenish or tinged with purple, glabrous;
or glabrous. Blades linear, 9-25 × 0.3-0.6 cm, flat, lower glume 0.8-1.4 mm long, ca. 1/3 the length
ascendent, rounded at the base, the apex acuminate, of the spikelet, 1-3-nerved, acute to obtuse, upper
adaxial surface papillose-pilose, more densely so glume and lower lemma subequal, 9-nerved; lower
toward the base, abaxial surface sparsely pilose or palea elliptic, 2.4 × 0.8 mm, glabrous, hyaline;
329
lower flower staminate, stamens 3, anthers 1.2- P 00740938!, MO-321061!, NY 01767173!,

1.4 mm long; upper anthecium long ovoid, 1.9-2.2 × S04-321!, US 00139898!, fragment).
0.8 mm, whitish, smooth, shiny, indurated, glabrous.
Caryopsis ellipsoid, 1.3 × 0.6 mm, whitish; hilum Distribution and ecology. México and the
oval, embryo 1/3 the length of the caryopsis. Caribbean to Argentina; it grows in margins of lakes
and rivers, from sea level up to 1300 m a.s.l.
Observation. This species is morphologically
similar to P. dichotomiflorum, the latter an anual Representative specimens examined
species, with spikelets 2-3 mm long and lower ARGENTINA. Corrientes. 3 km E de Ituzaingó,
flower absent. ruta 12, 7-IV-1978, Ahumada 2143 (MO).
BELIZE. 9 mi W of Belize on the Western
Key to the varieties highway to Belmopan, 50 m, 21-III-1987, Davidse
1. Sheaths densely hispid; panicles less than 25 cm & Brant 32947 (MO, NY).
long ........................... P. aquaticum var. aquaticum BRAZIL. Bahia. Near Salvador, low moist
1. Sheaths densely papillose; panicles ca. 30 cm ground near tidal marsh, 23-XII-1924, Chase 8022
long ......................... P. aquaticum var. cartagoense (F, MO, NY, US)
COSTA RICA. Guanacaste. Road to Hacienda
1a. Panicum aquaticum Poir. var. aquaticum, Inocentes, 12 km from CIA, 330 m, 27-VII-1971,
Encycl., Suppl. 4: 281. 1816. TYPE. Puerto Rico, Pohl 12651 (F, MO)
without locality, 1797, M. Ledru s.n. (lectotype CUBA. Pinar del Río. 13 km south of Pinar del
P 00371641!, designated as “type” by Hitchcock & Río, 25-XI-1926, Hitchcock 23290 (US).
Chase, 1910: 48; isolectotypes FI, US 00148147). GUATEMALA. Jutiapa. Vicinity of Jutiapa,
850 m, 24-X-1940, Standley 76308 (MO).
Panicum chloroticum var. sylvestre Nees ex HONDURAS. El Paraíso. 32 km W of Danli
Trin. Gram. Pan.: 236. 1826. TYPE. Brazil. along highway 4, 700 m, 21-VII-1970, Pohl &
Bahia. Almada, L. Riedel s.n. (holotype LE- Davidse 12386 (F, MO).
TRIN-0633.06!; isotype US-974404!). MÉXICO. Quintana Roo. En brecha a Vallarta,
Panicum chloroticum Nees ex Trin. var. sylvestre a 5 kms al W de Puerto Morelos, 31-VIII-1980,
Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829, nom. Sousa 11207 (MEXU).
illeg. hom., non Trin. 1826. TYPE. Brazil. “Habitat NICARAGUA. Zelaya. Puerto Isabel, scrubby
in sylvis ad Almada, Ferradas et in via Felisbertia growth near seabeach, 3-I-1970, Seymour 2895
districtus Insulanorum prov. Bahiensis; (Martius et (GH, MO).
Maximil. Princ. Neovid.)”, C. F. P. von Martius s.n. PERU. Without locality, Dombey s.n. (P),
(lectotype M, designated, as “type”, by Hitchcock Gaudichaud s.n. (P).
& Chase, 1910: 49; isolectotype US 00148356). PUERTO RICO. Floresta del Río Piedras,
Panicum hygrophilum Salzm. ex Steud., Syn. Pl. vicinity of San Juan, NE of Río Piedras, Chase 6778
Glumac. 1: 71. 1853. TYPE. Brazil. Bahia: (RB, US).
without locality, 1827, P. Salzmann s.n. (lectotype SURINAM. Near Dohsen savanna, Krukoff 130
here designated, P 00740831!; isolectotypes (NY).
G 00099659!, G 00099660!, GH!, HAL0133162!, TRINIDAD-TOBAGO. Pitch Lake, 7-XII-1912,
K 000003808!, K 000003809!, MO-105084!, Hitchcock 10099 (G, MO, NY, US).
MPU 024441!, MPU 024442!, MPU 024443!, VENEZUELA. Apure. Without locality, Ramia
P 00740830!, P 00740831!, P 00740832!, US!, 7195 (VEN).
W 18890239825!, W 19160024808!).
Panicum proliferum var. strictum Griseb., Cat. 1b. Panicum aquaticum Poir. var. cartagoense
Pl. Cub.: 232. 1866. TYPE. Cuba. Without Davidse, Novon 2: 103. 1992. TYPE. Costa Rica.
locality, 1860-1864, C. Wright 3456 (holotype Cartago: 0.5 km E of Planta Radiográfica along
unknown; isotypes G 00099640!, G 00099641!, railroad, 2 km W of Paraíso, 1300 m, 6 Feb 1969,
GH 00135362!, K 000309158!, K 000309159!, plants 1-2 meters tall, rooted in a dense tangle of
330
herbage, R. W. Pohl & G. Davidse 11700 (holotype P. dichotomiflorum Michx. var. bartowense (Scribn.
ISC-v-0000567!; isotypes F!, MO-183078!). & Merr,) Fernald, Rhodora 38(455): 387. 1936.
TYPE. United States of America. Florida: Polk
Distribution and ecology. This variety is County, Bartow, 29-IX-1898, R. Combs 1220
restricted to edges of forests in Costa Rica. (lectotype US 00133099!, designated as “type”
by Hitchcock & Chase, 1910: 52; isolectotype
2. Panicum dichotomiflorum Michx. ISC-v-0000569!).
Plants anual, caespitose, 10-140 cm tall; culms
decumbent, rooting and densely branching at the Panicum proliferum Lam. var. pilosum Griseb.,
lower nodes to erect, simple or branching; internodes Cat. Pl. Cub.: 232. 1866. TYPE. Cuba, Ad
terete, 3-16 cm long, hollow, glabrous; nodes lagunas, C. Wright 1865 (lectotype GOET
brownish, compressed, glabrous. Sheaths striate, 006760!, designated by L. Catasús Guerra,
4-15 cm long, pale or tinged with purple, glabrous to 2004: 85; isolectotypes GOET 006761!, GOET
densely hirsute, the margins membranous. Ligules 006762!, US 00139897!, fragment ex GOET).
membranous-ciliate, up to 2 mm long; collar
brownish, glabrous. Blades oblong-lanceolate to Distribution and ecology. Subspecies restricted
lanceolate, 10-45 × 0.5-1.4 cm, flat, subcordate, the to southern United States and the Caribbean;
apex attenuate, glabrous, the margins scaberulous. it grows at margins of swamps and in flooded
Inflorescence a lax, diffuse, terminal panicle, habitats.
4-30 × 2-20 cm, first order branches opposite to
alternate, less frequently whorled, divergent and Observation. It differs from the type subspecies
distant, spikelets paired or solitary on third order by its densely hirsute sheaths and shorter spikelets,
branches; main axis scaberulous, pulvini glabrous; usually 2-2.2 mm long.
pedicels triquetrous 2-7 mm long, scabrous; axillary
panicles usually present, similar to the terminal Representative specimens examined
one. Spikelets long ovoid, (2-)2.4-3 × 0.9-1.1 mm, BAHAMAS. Grand Bahama. About Freeport
glabrous, acuminate, pale or tinged with purple; dump north of Airport, 25-I-1976, Correll &
lower glume 0.7-1.2 mm long, 1/4 to 1/3 the length Popenoe 46671 (MO).
of the spikelet, ovate, 1-3-nerved; upper glume and CAICOS ISLAND. Pine Cay, in clumps in
lower lemma subequal, acuminate, upper glume drying open pond of coppice on norhteast side,
9-nerved, lower lemma 7-9-nerved; lower palea Correll 43251 (NY).
elliptic, 2.1-2.5 × 0.5-0.7 mm, hyaline, glabrous CUBA. Habana. Barabanó, in coastal swamps,
to absent; lower flower absent; upper anthecium 10-XII-1920, Ekman 12602 (G, US).
ellipsoid, 2-2.5 × 0.7-1.1 mm, pale, dark brown at JAMAICA. St. Elizabeth. Near the town of
maturity, glabrous, smooth, indurated; upper lemma Black River, in a swamp along the Black River,
7-nerved. Caryopsis elipsoid, 1.5-1.6 × 0.9-1 mm, 22-X-1912, Hitchcock 9645 (MO, NY, US).
brownish; hilum oblong, embryo half the length of UNITED STATES OF AMERICA. Florida.
the caryopsis. Miami, Hitchcock s.n., Amer. Gr. Hb. 17 (US, W).
Key to the subspecies 2b. Panicum dichotomiflorum Michx. subsp.

1. Sheaths glabrous; spikelets 2.4-3 mm long dichotomiflorum, Fl. Bor.-Amer. 1: 48. 1803.
............ P. dichotomiflorum subsp. dichotomiflorum TYPE. United States of America. “Hab. in
1. Sheaths densely hirsute; spikelets 2.2-2.4 mm occidentalibus montium Alleghanis”, A. Michaux
long ........... P. dichotomiflorum subsp. bartowense s.n. (holotype P-MICHX!; isotype P 00740942!).
(Fig. 1).
2a. Panicum dichotomiflorum Michx. subsp.
bartowense (Scribn. & Merr.) Freckmann & Panicum miliaceum Walter, Fl. Carol.: 72. 1788, nom.
Lelong, Sida 20(1): 171. 2002. P. bartowense Scribn. illeg. hom., non L., 1753. TYPE. United States of
& Merr., Circ. Div. Agrostol. U.S.D.A. 35: 3. 1901. America. South Carolina (holotype unknown).
331
Fig. 1. Panicum dichotomiflorum. A, Habit. B, Spikelet, ventral view. C, Spikelet, dorsal view. D, Lower palea.
E, Upper anthecium, ventral view. F, Upper anthecium, dorsal view. (From Ahumada 610, CTES).
332
Panicum geniculatum Muhl., Cat. Pl. Amer. Sept.: 9. Hitchcock & Chase, 1910: 49; isolectotypes LINN-
1813. TYPE. United States of America. Without HS 118-84!, US 00139933!, fragment ex P).
locality, Muhlenberg Hb. Fol. 181 (holotype PH Panicum chloroticum Nees ex Trin. var. luxurians
00047147!; isotype US 00148518!, fragment, Döll, in C. Martius, Fl. Bras. 2 (2): 198. 1877.
probable isotype, BM 001042409!). TYPE. Holotype unknown.
Panicum multiflorum Poir., in Lamarck, Encycl. Panicum proliferum Lam. var. richardii Döll, in C.
Suppl. 4: 282. 1816. TYPE. United States of Martius, Fl. Bras. 2(2): 200. 1877. TYPE. Brazil.
America: “cette plante croit a la Caroline; elle m’a Pará: “prope urbem Para, in littore arenoso (L. C.
ete communiquee par M. Bosc.”, L. A. G. Bosc Richard), L. C. Richard s.n. (holotype unknown;
s.n. (lectotype P 00740941!, designated as “type” isotype W 19040013871!).
by Hitchcock & Chase, 1910: 48; isolectotypes Panicum amplectens Chapm., Bot. Gaz. 3(3): 20.
US 00139776!, US 00139777!, fragments). 1878. TYPE. United States of America. “South
Panicum brachiatum Bosc ex Spreng. Syst. Veg. Florida”, J. L. Blodgett s.n. (holotype unknown).
[Sprengel] 1: 321. 1825, nom. illeg. hom., non Panicum francavillanum E. Fourn., Mexic. Pl. 2:
Poiret, 1816. TYPE. Bermuda. “Ins. Bermud” 25. 1886. P. francavillanum E. Fourn. ex Hemsl.,
(holotype B-W-18843-01 0!; isotype US Biol. Cent.-Amer. Bot. 3: 489. 1885. nom. nud.
00148189!, fragment ex Willd. herb.). TYPE. México: Tabucaya, J. W. Schaffner 301
Panicum chloroticum Nees ex Trin., Gram. Panic.: (lectotype P 00740939!, designated as “type”
236. 1826. P. chloroticum var. agreste Nees ex Trin., by Hitchcock & Chase, 1910: 50; isolectotype
Gram. Panic.: 236. 1826. Panicum proliferum Lam. W 0021637!).
var. chloroticum (Nees ex Trin.) Hack., Repert Spec. Panicum proliferum Lam. var. geniculatum Alph.
Nov. Regni Veg. 6(21-26): 343. 1909. P. aquaticum Wood, Amer. Bot. Fl. 2: 392. 1871. Panicum
var. chloroticum (Nees) R.C. Foster, Rhodora 68: dichotomiflorum Michx. var. geniculatum (Alph.
320. 1966. TYPE. Brazil: “V. utriusque spp. Brasil. Wood) Fernald, Rhodora 38(455): 387, pl. 441,
(N. ab Esenb. Langsdorff)”, G. H. von Langsdorff fig. 2. 1936. TYPE. United States of America
s.n. (holotype LE-TRIN-0633.01!). (holotype unknown).
Panicum chloroticum Nees ex Trin. var. agreste Panicum dichotomiflorum Michx. var. imperiorum
Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829. TYPE. Fernald, Rhodora 44(526): 380. 1942. TYPE.
Brazil: “Habitat in graminosis cultis prope United States of America. Virginia: Greensville
Soteropolin et Oeiras provinciae Bahiensis et County, Fontaine Creek, southwest of Haley’s
Piauhiensis”, C. F. P. von Martius s.n. (lectotype Bridge, 14-X-1941, M. L. Fernald & Long
M, designated as “type” by Hitchcock & Chase, 13877 (holotype GH 00023012!; isotypes
1910: 49; isolectotype US 00148354!, fragment). GH 00023013!, NY 00413977!, PH 00018649!).
Panicum chloroticum Nees ex Trin. var. pingue Panicum dichotomiflorum Michx. var. purinatorum
Nees, Fl. Bras. Enum. Pl. 2(1): 164. 1829. Svenson, Rhodora 22(261): 154, fig. 1-5. 1920.
SYNTYPES. Brazil: “Habitat in cultis ad P. dichotomiflorum Michx. subsp. purinatorum
Soteropolin, provinciae Bahiensis; in campis (Svenson) Freckmann & Lelong, Sida 20(1): 171.
prope Almeirim et alibi ad flumen Amazonum, 2002. TYPE. United States of America. Maine:
provinciae Paraensis: statura maxima, culmo Barnstable, 13-IX-1919, M. L. Fernald 306 (holoty-
digitali”. In campis prope Almeirim et alibi ad pe GH 00024098!; isotypes BR 0000008408792!,
flumen Amazonum, C. F. P. von Martius s.n. NEBC 00028645!, NY 00413974!, P 00740940!,
(M!). Habitat in cultis ad Soteropolin. Provinciae PH 00018647!, PH 00018648!, US 00147806!).
Bahiensis”, C. F. P. von Martius s.n. (lectotype
M, designated as “type” by Hitchcock & Chase, Distribution and ecology. This subspecies
1910: 49; isolectotype US 00148355!, fragment). is widely distributed in America from Canada
Panicum retrofractum Delile ex Desv., Mém. Soc. and the United States to Argentina. It is usually
Agric. Angers 1: 200. 1831. TYPE. [United States found in open and humid habitats, being present
of America], “America borealis”, sin colector s.n. in margins of rivers and streams and in lower
(lectotype P 00661391!, designated as “type” by fields; it is also present as a weed.
333
The following names are nom. nud. which VENEZUELA. Anzoátegui. Pariaguán,
correspond to this subspecies: Laguna de los Patos, 17-VII-1946, Burkart 17289
(SI, VEN).
Panicum ammophilum Trin. ex Nees, Fl. Bras. Enum.
Pl. 2(1): 164. 1829. nom. nud. Panicum elliottii 3. Panicum gilvum Launert, Mitt. Bot.
Trin. ex Nees, Fl. Bras. Enum. Pl. 2(1): 170. 1829. Staatssamml. München 8: 153, tab. 1, fig. 9.
nom. nud, as a synonym of P. proliferum. 1970. TYPE. Namibia. Ikahandja Distr., Pad
Panicum proliferum Lam. var. xantochlorum Okahandja-Otjisazu, 12 km, 20-III-1912, M. K.
Hackel ex Bertoni, Anales Ci. Parag.: 150. 1918. Dinter 2544 (holotype Z 000080546!; isotypes
L 0044857!, M). (Fig. 2).
Representative specimens examined
ARGENTINA. Buenos Aires. Tigre: Paraná Panicum laevifolium Hack. var. contractum Pilg.,
Miní, Cabrera 1607 (F, LP, NY). Notizbl. Bot. Gart. Berlin-Dahlem 15: 448.
BAHAMAS. Near Nassau, 25-IV-1903, Curtiss 1940. Type. Namibia. Ikahandja Distr., Pad
177 (MO, US). Okahandja-Otjisasu, 12 km, 20-III-1912, M. K.
BERMUDA. Hamilton, Millspaugh 126 (US); Dinter 2544a (holotype B †; isotype Z).
roadside, Hungry Bay, Brown & Britton 1638 (NY).
BOLIVIA. Beni. Estancia El Porvenir, 50 km Plants annual, caespitose, 25-60 cm tall, culms
E of the Rio Mariqui on the road to Trinidad, decumbent, rooting at the lower nodes, and
17-XI-1985, Solomon 14771 (MO). geniculate, then ascending, branching at the lower
BRAZIL. Distrito Federal. West margin of and upper nodes; internodes terete, (3-) 5-10 cm
Lagoa Paranoa, Brasília, 11-III-1966, Irwin et al. long, hollow, glabrous; nodes compressed,
13875 (F, MO, NY, UB). brownish, glabrous. Sheaths (4-)5-13 cm long,
COLOMBIA. Meta. De Cumaral a San Nicolás, longer than the internodes, striate, glabrous; collar
10 km del desvío a San Nicolás, bosque secundario brown, glabrous. Ligules membranous-ciliate,
junto a arroyo y sabana, 480 m, 11-VI-1989, 0.9-2(-2.4) mm long. Blades linear to linear-
Zuloaga 3867 (COL, SI). lanceolate, 3-18(-22) × 0.3-0.7 cm, flat, acute,
CUBA. Pinar del Río. Remates, Ekman 11211 sparsely pilose toward the base, otherwise
(NY, US). glabrous, the margins pilose to scabrous.
ECUADOR. Archipielago de Colón. Isla Santa Inflorescence a terminal, contracted, partially
Cruz, 3-VIII-1974, van der Werff 1373 (MO). included in the upper blades panicle, 3-10 ×
FRENCH GUIANA. Pres de Loca-Loca, sur le 0.2-0.6(-5) cm, oblong to elliptic; main axis
Maroni, Schnell 11424 (P). triquetrous, glabrous, first order branches
GUADALUPE. Without locality, 1892, Duss appressed to ascendent, alternate, triquetrous,
3180 (MO). pulvini glabrous; spikelets paired or solitary on
HAITÍ. Massif da la Hatte, Miragoane, 15-XI-1926, pedicels 1-6 mm long, triquetrous, claviform.
Ekman 7233 (US). Spikelets long ellipsoid, 2.5-3.4 × 0.8-1 mm,
MÉXICO. Puebla. Laguna Epatlán, 11-XII-1942, glabrous, pale and tinged with purple; lower
Miranda 2496 (MEXU). glume. 0.8-1 mm long, 1/5 to 1/4 the length
PANAMÁ. Los Santos. 5 km SW of Chitre, of the spikelet, nerveless to 1-nerved, truncate
22-I-1966, Tyson et al. 3034 (MO). to obtuse; upper glume and lower lemma
PARAGUAY. Central. Cerca del lago Ipacaray, subequal, membranous, acute, upper glume
Hassler 12537 (BAF, G, GH, K, LIL, MO, NY). 9-11(-13)-nerved, lower lemma 9-nerved; lower
PERÚ. Loreto. Mazán, slight above Baradero de palea 1 × 0.5 mm, hyaline, glabrous; lower flower
Mazán, 11-X-1979, Rimachi 4657 (MO). absent; upper anthecium ellipsoid, 1.7-1.9 ×
SURINAM. 2 km above confluence with Lucie 0.7-0.8 mm, pale, indurated, glabrous, smooth
River, 3-VII-1963, Maguire et al. 54053 (F, MO, NY). and shiny, upper lemma 7-nerved. Caryopsis
URUGUAY. Artigas. Bella Unión, Rosengurtt broadly ellipsoid, 1.6-1.7 × 0.8-0.9 mm; hilum
B-7373 (US). oblong, embryo 1/2 the length of the caryopsis.
334
Fig. 2. Panicum gilvum. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, dorsal view. E, Spikelet,
ventral view. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Upper palea with caryopsis. I,
Caryopsis, embryo view. J, Caryopsis, hilum view. K, Base of the upper palea and anther (A-B, H-K from Smook 5807,
G; D-G from Dinter 2544, M).
335
Fig. 3. Panicum impeditum. A, Habit. B, Ligule region. C, Upper portion of a blade. D, Spikelet, ventral view.
E, Spikelet, dorsal view. F, Spikelet, dorsal view, without the upper glume. G, Lower palea. H, Upper anthecium,
ventral view. I, Upper palea, anthers, and lodicules. J, Gynoecium. (From Smook 6999, US).
336
Distribution and ecology. Originally described flower absent; upper anthecium ellipsoid, 2-2.1 ×
from Namibia, it also grows in Botswana, Kenya, 0.9-1 mm, smooth, shiny, glabrous and indurate;
and South Africa; introduced in Australia and upper lemma 5-nerved. Caryopsis ellipsoid, 1.3-1.4 ×
Europe; it is found in wet, sandy soils, frequently 0.9 mm; hilum punctiform, embryo 1/2 the length
in ephemeral rainwater pan. of the caryopsis.
Observation. Related to P. impeditum (see Distribution and ecology. This species grows
comments under this species). in wet places, around edges of dams and other
water surfaces. It is restricted to South Africa and
Representative specimens examined Namibia.
BOTSWANA. In the bed of a fossil river valley
at 20º43’S, 21º05’E, 23-III-1980, Smith 3300 (K); Observation. This species is related with P. gilvum
near the Tsau-Xaixai road, 19º55’S, 21º24’E, Launert, a species also from southern Africa, which
21-IV-1982, Smith 3846 (K). differs by having the panicles partially included on
KENYA. South Baringo, Gobat, 11-XI-1960, the upper leaves and shortly exserted, and spikelets
Knight 5086 (K). with the upper glume 9-11(-13)-nerved.
NAMIBIA. Okongo, II-1917, Dinter s.n. (K).
SOUTH AFRICA. 40 km S of Dewetsdorp Farm Representative specimens examined
Bloemspruit, 2-IV-1986, Smook 5807 (G). SOUTH AFRICA. Cape. 30.7 km NW of
Britstown at turn off to Jagskerm on main road to
4. Panicum im peditum Launert, Mitt. Bot. Prieska, 16-III-1988, Smook 6999 (US); 30.7 km
Staatssamml. München 8: 150. 1970. TYPE. SW of Britstown on road to Victoria West,
Namibia: Gideon Distr.: Mariental, 10-V-1955, 12-III-1998, Smook 6877 (P); 36.4 km N of Victoria
De Winter 3483 (holotype K 000255501!; West, Fram Rietpoort, 15-III-1988, Smook 6974 (P).
isotypes M 0103858!, PRE 0033208-0!). (Fig. 3). Northern Cape. Northwest of Kimberley,
Kopmansfonte in Agriculture Research Station Pan
Plants annual, caespitose, (15-)20-50 cm tall; area, 4-III-1998, Smook 10117 (C). Orange Free.
culms branching at the lower and middle nodes, 8 km from Luckhoff on road to Jacobsal, 30-III-1980,
erect; internodes 5-12 cm long, terete, hollow, Smook & Gibbs Russell 2453 (US).
striate, glabrous; nodes pale, glabrous. Sheaths
striate, 4-11 cm long, usually shorter than the 5. Panicum lacustre Hitchc. & Ekman, Man.
internodes, sparsely pilose toward the apex, Grasses W. Ind.: 253, fig. 205. 1936. TYPE.
otherwise glabrous; collar pale, glabrous. Ligules Cuba. Pinar del Río: Laguna de la Máquina,
membranous-ciliate, 0.6-1 mm long, greenish to 31-X-1923, E. L. Ekman 17878 (lectotype here
purple. Blades lanceolate, 4-11 × 0.3-0.5 cm, flat, designated US 00132924!; isolectotypes B 10
acute, glabrous. Inflorescence a terminal, oblong 0248983!, BAA 00000387!, BR 0000008408877!,
and exserted panicle, 7-14 × 3-5 cm; peduncle DAO 000465583!, G 00099619!, GH 00135365!,
terete, up to 12 cm long, glabrous, first order ISC-v-0000574!, LD 1010829!, LIL 000085!,
branches alternate, ascending, axis of the branches MICH 1127266!, MO 128363!, MVFA 0000436!,
triquetrous, glabrous, pulvini glabrous, spikelets NY 00071083!, NY 00803749!, P 00740958!,
appressed, paired or solitary on first and second R 000076339!, S13-12944!, SI 002808!,
order branches; pedicels 0.5-5 mm long, claviform, US 00132923!, W 1959000033!). (Fig. 4).
glabrous; axillary inflorescences present, similar
to ther terminal one. Spikelets long ellipsoid, Plants perennial, shortly rhizomatous, 100-
(2.8-)3.1 × 1-1.1 mm, greenish, glabrous; lower 150 cm tall; culms erect, with aerenchyma,
glume 0.7-0.8 mm long, 1/4 the length of the with adventitious roots at the lower nodes;
spikelet, nerveless to 3-nerved, acute; upper glume basal internodes short, terete, hollow, glabrous;
and lower lemma subequal, 9-nerved; lower palea nodes brownish, glabrous. Sheaths striate, with
2.3-2.4 × 0.9-1 mm, hyaline, glabrous, lower anastomosed veins, glabrous, the lower sheaths
337
Fig. 4. Panicum lacustre. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, lateral view. E, Spikelet,
ventral view. F, Spikelet, dorsal view. G, Upper anthecium, dorsal view. H, Upper anthecium, ventral view. I, Upper
lemma, lodicules and gynoecium. J, Upper lemma, lodicules and androecium. (From Ekman 17878, US).
338
shorter than the internodes, pale to purplish, Panicum withei J.M. Black, Trans. & Proc. Roy.
the upper ones with membranous margins, the Soc. South Australia 41: 632, tab. 39. 1917.
upper ones shortly pilose or glabrous. Ligules TYPE. Australia. Strzelecki Creek, south-west
membranous-ciliate, ca. 2 mm long; collar of Innamincka, Cooper Creek, at Cuttapin
brownish, glabrous. Blades linear-lanceolate, the corner, 27-IX-1916, S. A. White s.n. (lectotype
lower ones reduced, the upper ones ca. 20 × 0.2 cm, AD 99140233!; isolectotype K 000674341!).
the lower margins involute, then flattened, narrowed
at the base, the apex long subulate, adaxial surface Plants annual, caespitose, 60-80(-120) cm tall,
densely hispid toward the base, sparsely hispid at cataphylls pilose; culms erect, internodes cylindrical,
the apical portion, the abaxial surface scabrous, 4.5-12 cm long, hollow, glabrous or pilose; nodes
the margins scabrous. Inflorescence a terminal, brownish, pilose. Sheaths 3-10 cm long, striate,
exserted and multiflowered panicle 20-30 cm densely covered with tubercullate caducous hairs or
long; peduncle terete, scabrous; main axis wavy, glabrous; collar pale, glabrous. Ligules membranous-
scabrous, pulvini brownish, glabrous, first order ciliate, 1-1.2 mm long. Leaves lanceolate, 6-20 ×
branches alternate to opposite, axis of the branches 0.4-0.6 cm, flat, with tubercullate caducous hairs on
and pedicels triquetrous, scabrous, spikelets paired the lower portion and the basal margins, otherwise
and congested on short pedicels at the upper portion glabrous, acute or acuminate. Inflorescence a lax,
of the branches. Spikelets narrowly ellipsoid, 2 × diffuse, many flowered panicle, 10-50 × 8-25 cm;
0,8 mm, glabrous, greenish or tinged with purple; peduncle up to 20 cm long, with tubercullate caducous
lower glume 0.5 mm long, 1/4 the length of the hairs, main axis pilose, pulvini brownish, spikelets
spikelet, 3-nerved, ovate to truncate; upper glume paired and adpressed on the branches, pedicels 1-6 mm
and lower lemma subequal, acute, or the upper long, claviform, scaberulous; axillary inflorescences
glume shorter and obtuse, 7-nerved; lower palea absent. Spikelet long ovoid, 2.5-3.4 × 0.9-1 mm,
and lower flower absent; upper anthecium ellipsoid, glabrous, pale; lower glume 0.6-1.1 mm long, 1/4
1.8 × 0.7 mm, whitish, smooth and glabrous, shiny. to 1/3 the length of the spikelet, acute or truncate,
Caryopsis not seen. glabrous, 1-3-nerved; upper glume and lower lemma
subequal, acute, 7-9-nerved; lower palea 1-1.8 × 0.5-
Distribution and ecology. Species endemic of 0.6 mm, hyaline, glabrous; lower flower absent; upper
Cuba, where it is found in margins of lagoons anthecium ellipsoid, 1.8-2 × 0.8-1 mm, shorter than
and ponds. the upper glume and lower lemma, pale, olivaceous at
maturity, indurated, smooth and shiny, upper lemma
Observation. This species is characterized by 5-7-nerved, with simple papillae toward the apex.
being a perennial species, with the lower blades Caryopsis ellipsoid, 1.7 × 0.8 mm; hilum oblong,
reflexed and falling, and spikelets with the upper embryo less than half the length of the caryopsis.
glume and lower lemma 7-nerved, the lower palea
and lower flower absent. Distribution and ecology. Species endemic
of Australia, it grows in humid woodlands, and
Representative specimens examined semiarid and arid grasslands.
CUBA. Pinar del Río. Pinar del Río, in a small
Laguna at km 12 of the carretera to La Colona, Observation. Webster (1987) stressed that this
28-X-1923, Ekman 17822 (G); La Grifa, at a Laguna species is an important forage species, widely
north of town, 14-VI-1920, Ekman 11271 (G). distributed in arid and semiarid central Australia.
6. Panicum laevinode Lindl., Three Exped. Representative specimens examined

Austral. 1: 238. 1838. TYPE. Australia. Mitchell’s AUSTRALIA. Northern Territory. On nutwood
expedition s.n. specimen no. 3 (lectotype CGE! Downs Station Queensland, 2-V-1947, Blake 17591
[right-hand specimen), designated by S. Jacobs, (US); about 24 miles NE of Anthony Lagoon,
Kew Bull. 40(3): 662. 1985; isolectotype 15-V-1947, Blake 17794 (US). Queensland.
W 0023593!]. Frensham, near Kynuna, 13-V-1936, Blake 11491 (K);
339
64 miles south of Richmond, 20-VI-1954, Lazarides Plants annual, 30-90(-150) cm tall, culms
4462 (US); 26 miles NW of McKinlay, 20-III-1954, branching at the base and at the upper nodes, erect
Lazarides 4455 (US). or geniculate ascending; internodes 2-8 cm long,
terete, hollow, striate, pilose; nodes densely pilose,
7. Panicum luzonense J. Presl, Reliq. Haenk. 1(4-5): compressed. Sheaths striate, open, 1.5-7 cm long,
308. 1830. TYPE. Philippines. Luzon, without covered with urticant hairs, as long as or shorter
locality, T. Haenke s.n. (lectotype PR-612343!, than the internodes. Ligules membranous-ciliate,
designated by J. F. Veldkamp as “holotype”, 1-1.5 mm long, with long hairs beneath at the
Blumea 41: 195. 1996; isolectotypes LE 800.1!, base of the blade; collar pilose. Blades lanceolate,
fragment, W 0021811!, fragment). (Fig. 5). 4-11(-40 × 0.4-0.7(-1.6) cm, flat, subcordate, its
surface covered with tubercullate hairs, the margins
Panicum caesioglaucum Nees ex Steud., Syn. Pl. ciliate. Inflorescence a lax, terminal panicle,
Glumac. 1: 75. 1853. TYPE. Indonesia, Java 12-40 × 10-24 cm; main axis triquetrous, wavy,
(not located). hispid, covered with tubercullate hairs, pulvini
Panicum extensum Steud., Syn. Pl. Glumac. 1: hispid, first and second order branches divergent,
72. 1853, nom. illeg. hom., non Desvaux, alternate, scabrous and hispid, spikelets paired on
1831. TYPE. Philippines. Without locality, H. second order branches, pedicels 2-10 mm long,
Cuming 652 (lectotype P!, designated by J. F. scabrous, claviform; axillary panicles present,
Veldkamp, Blumea 41: 195. 1996; isolectotypes similar to the terminal one. Spikelets broadly
BM!, G 00374956!, K 000290312!, L 0044818!, ellipsoid, 2-2.5 × 1-1.1 mm, glabrous, greenish to
W 0021810!, W 18890086675!). pale; lower glume 0.7-0.8 mm, 1/3 the length of the
Panicum reticulatum Thwaites ex Trimen, J. Bot. spikelet, acute, 5-nerved; upper glume and lower
23: 271. 1885, nom. illeg. hom., non Torr., 1853. lemma subequal, membranous, 7-9-nerved, the
Panicum caesium Nees ex Hook. f., Fl. Brit. India nerves anatomosed toward the apex, acute; lower
7(21): 48. 1897[1896], nom. illeg. hom., non Nees, palea 2-2.1 × 0.6-0.7 mm, hyaline, glabrous; lower
1841. Panicum cruciabile Chase, J. Arnold Arbor. flower absent; upper anthecium ellipsoid, 1.8-1.9 ×
20: 309. 1939. TYPE. Sri Lanka: Hewesse, Pasdun 0.8-0.9 mm, whitish, indurate, glabrous; upper
Korle, 1868, G. H. K. Thwaites 3890 (lectotype PDA, lemma 5-nerved. Caryopsis broadly ellipsoid, 1.3-
designated by A. Chase, J. Arnold Arbor. 20: 309. 1.4 × 0.9-1 mm, whitish; hilum punctiform, embryo
1939; isolectotypes G 00176537!, G 00176538!, 1/3 the length of the caryopsis.
K 000245241!, K 000245242!, P 00740923!,
US-1715318!, US-1298826!, US-Z1445622!, Distribution and ecology. Asia tropical and
W 18890163875!). Syntype. Culloden Estate, near temperate, China and eastern Asia, India, Sri Lanka,
Kalutara, 1881, W. Ferguson s.n (not located). Indo-China, Malesia, Papuasia and Australia. It is
Panicum cambogiense Balansa, J. Bot. (Morot) 4(7): a frequent, and variable, species in open and humid
142. 1890. TYPE. Cambodia, without locality, A. places, also present in beaches, in roadsides and
Godefroy 62 (lectotype L 0044815!, designated by a weed in tea and rubber plantations (Veldkamp,
J. Veldkamp, Blumea 41: 195. 1996; isolectotypes 1996), between sea level and 1300 m a.s.l.
K 000290303!, fragment, P!). Syntype. Cambodia.
Without locality, 5-VI-1875, A. Godefroy 257 Observations. This species is characterized (Chase,
(L 0044814!). 1939) by the presence of short irritating hairs;
Panicum oryzetorum Balansa, J. Bot. Morot. 4: 141. also axillary inflorescences are conspicuous in all
1890. TYPE. Vietnam, Tonkin, Ouonbi, 1-XI-1885, specimens. When describing Panicum oryzetorum,
B. Balansa 472 (lectotype here designated G Balansa cited two syntypes from Vietnam, B.
00176526!; isolectotypes G 00374960!, L 0056274!, Balansa 472 and 1628. Of these, the specimen
L 0056275!, P 01859733!, US 00139821! ex G, US Balansa 472 from G is selected as the lectotype,
00139822 ex L). Syntype. Vietnam, Tonkin, Ouonbi, since is a complete specimen and fully agrees with
Nov 1885, B. Balansa 1628 (G 00374958!, L the protologue of the species. For comments on other
0056276!, P 01859731!, US 00139822! fragment ex L). typifications see Veldkamp (1996).
340
Fig. 5. Panicum luzonense. A, Habit. B, Ligule region. C, Spikelet, lateral view. D, Upper glume. E, Lower glume.
F, Lower palea. G, Upper anthecium, dorsal view. H, Upper anthecium, ventral view. I, Upper palea with caryopsis,
embryo view. J, Caryopsis, hilum view. (A-B from Balansa 1628, G; C-J from Balansa 472, G).
341
Fig. 6. Panicum madipirense. A, Habit. B, Ligule region. C, Spikelet, lateral view. D, Spikelet, dorsal view. E, Lower palea,
anthers and lodicules. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Upper palea with gynoecium
and lodicules. I, Lodicules and stamens caryopsis, embryo view. J, Caryopsis, hilum view. (From Barker s.n., US).
342
Representative specimens examined pale or tinged with purple, solitary, glabrous; lower
CAMBODIA. Chhnang, VII-1921, Petelot glume 0.9 mm long, less than 1/3 the length of the
251 (P). spikelet, 1(-3)-nerved, acute; upper glume and lower
MALAYSIA. Kampong Naka, Kedah, 9-IX-1933, lemma subequal, acute, upper glume 9-11-nerved,
Holttum s.n. (K); Kedah, 4-VIII-1919, lower lemma 9-nerved; lower palea elliptic, 2.9 ×
Vesterdal 4 (C); Kelantan, Sungal Lebir at 0.8 mm, hyaline, glabrous; lower flower staminate,
Bukit Batu Papan, 7-VII-1935, Henderson s.n. anthers 3; upper anthecium ellipsoid, 2.7 × 1 mm,
(P 01931258); Kampong Naka, Kedah, 9-IX-1933, shorter than the upper glume and lower lemma,
Henderson s.n. (P 01931259); Singapore, pale, glabrous, indurated, with papillae at the apex
Raffles College Ground, 14-XI-1932, Nor s.n. of lemma and palea. Caryopsis not seen.
(P 01931260).
PHILIPPINES. Rizal. Antipolo, X-1913, Ramos Distribution and ecology. Africa, where it is
s.n. (MO-755528); Taytay, V-1913, Merrill 1238 found in swamps and damp areas.
(G, MO, P); Luzon, X-1913, Ramos 2057 (G); Sulu,
Tawitawi, VII-1924, Ramos s.n. (P 01931252); Representative specimens examined.
Pampanga, Arayat, IX-1905, Merrill 1229 (G). MALAWI. South Nyasa, Damo, I-1929, Barker
THAILAND. Chantaburi, Soi Dao, 7-VI-1963, s.n. (P, US 2975890).
Larsen 9864 (C); Phitsanulok, Nakawn Thai TANZANIA. Near Ukenyule, Mbeya, I-1963,
Road, 24-VII-1966, Larsen et al. 850 (K); N Proctor 2370 (US).
Srithamarat, Kapiat, 28-VI-1956, Snan 715 (K);
Kuan Karong, 40 km northwest of Satun, 9. Panicum mlahiense Renvoize, Kew Bull.
Charoenphol et al. 3789 (C). 34(3): 554. 1979(1980). TYPE. Tanzania.
Ulanga District, Mlahi, Kilombero River, 8º17´S,
8. Panicum madi pirense Mez, Bot. Jahrb. Syst. 37º05´E, 5-V-1977, K. Vollesen 4563 (holotype
57: 189. 1921. TYPE. Tanzania. Mbeya District, C 10001151!; isotype K 000255579!). (Fig. 7).
Kiwere, 28-II-1909, E. Zimmerman s.n., Inst.
Amani 2582 (lectotype here designated EA Plants annual, 30-60 cm tall, culms erect,
000000481; isolectotypes K 000255581!, branching at the middle and upper nodes; internodes
US 00148269!). Syntype: Tanzania. Mbeya cylindrical, 3-15 cm long, striate, hollow, glabrous;
District, E. Zimmerman s.n., Inst. Amani 2583 (K nodes brownish, glabrous. Sheaths striate, 3-5 cm
000255581!, US 00148269!, another fragment). long, shorter than the internodes, glabrous. Ligules
(Fig. 6). membranous-ciliate, 1-1.5 mm long, the ciliate
portion short; collar glabrous. Blades lanceolate,
Plants annual, caespitose, 30-120 cm tall, culms 3-6(-10) × 0.3-0.4 cm, rounded at the base, flat
erect, branching at the base; internodes terete, or with involute margins, acuminate, glabrous.
4-15 cm long, glabrous, hollow; nodes brownish, Inflorescence a terminal, lax panicle, 5-17 × 5-12 cm;
compressed, glabrous. Sheaths 4-12 cm long, as peduncle 5-15 cm long, glabrous; main axis
long as or shorter than the internodes, 4-12 cm triquetrous, glabrous, pulvini glabrous, first order
long, densely hispid to glabrous; collar pale, branches alternate, divergent or reflexed, spikelets
glabrous. Ligules membranous-ciliate, 1-1.2 mm solitary or paired on the branches, pedicels 2-7 mm
long. Blades lanceolate, 12-26 × 0.8-1.2 cm, long, claviform, scaberulous; axillary panicles
cordate, acuminate, glabrous or hispid toward the present, similar to the terminal one. Spikelets long
base, the margins ciliate. Inflorescence terminal, ovoid, 2.2-2.5 × 0.8-1.2 mm, pale to purplish,
exserted; peduncle ca. 15-20 cm long, glabrous; glabrous; lower glume 0.8-0.9 mm long, ca. 1/3
panicle lax, open, multiflowered, 14-25 × 5-12 cm, the length of the spikelet, 3-5-nerved, acute to
pyramidal, first order branches alternate, truncate; upper glume and lower lemma subequal,
divergente, axis of the branches scaberulous; membranous, upper glume 9-nerved, lower lemma
pedicels claviform, 5-12 mm long, scaberulous. glumiform, 7-9-nerved; lower palea 1.2-1.3 x 0.4-
Spikelets long ellipsoid, 3.2-4 × 0.9-1 mm, 0.5 mm, hyaline, glabrous; lower flower absent;
343
Fig. 7. Panicum mlahiense. A, Habit. B, Ligule region. C, Upper portion of a blade. D, Spikelet, ventral view. E,
Spikelet, dorsal view. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Upper palea, lodicules,
anthers and gynoecium. I, Caryopsis, embryo view. J, Caryopsis, hilum view. (A-C from Phipps & Vesey-FitzGerald
3220, K; D-J from Webster A251, K).
344
upper anthecium ovoid, 1.7-2 × 0.8-1 mm, pale, Distribution and ecology. It is endemic to
glabrous, indurated, smooth and shiny, upper Australia, where it grows in humid areas at edge
lemma 7-nerved, with simple papillae toward the of lagoons.
apex. Caryopsis ovoid, 1.3-1.4 × 1-1.1 mm, whitish,
brown at maturity; hilum oblong, embryo less than Representative specimens examined
1/2 the length of the caryopsis. AUSTRALIA. New South Wales. Ca. 1 mile by
road NNE of Cattai, 7.5 miles NE of Windsor,
Distribution and ecology. Species restricted 21-I-1969, Coveny 774 (K). Queensland.
to Zambia and Tanzania, it grows (according to Stanthorpe, 11-III-1931, Hubbard 5732 (K).
Renvoize, 1989) in riverine thickets and laterite
pans, between 280-1800 m a.s.l. 11. Panicum paludosum Roxb., Fl. Ind. 1: 310.
1820. Panicum decompositum var. paludosum
Representative specimens examined (Roxb.) Trimen, Syst. Cat. Fl. Pl. Ceylon: 105.
ZAMBIA. Moporokoso, Mugombwe, 1885. Panicum repens var. paludosum (Roxb.)
15-IV-1961, Phipps & Vesey-FitzGerald 3220 (K); Kuntze, Revis. Gen. Pl. 3(3): 363. 1898. Panicum
Abercorn, 27-III-1959, Webster A251 (K); Uningi proliferum var. paludosum (Roxb.) Stapf, Fl.
Pans, Abercorn, 5-III-1965, Richards 17933 (K). Cap. 7. 407. 1899. Lousiella paludosa (Roxb.)
Landge, Phytotaxa 512(2): 109. 2021. TYPE.
10. Panicum obseptum Trin., Gram. Panic.: 149. [India]. “It grows generally in sweet water
1826. TYPE. Australia. “V. spp. nov. Holl.”, J. amongs the Circar mountains”, W. Roxburgh s.n.
Lindley s.n. (holotype LE 0857-01!). (Fig. 8). (lectotype K 00113133!, designated by I. M.
Turner et al., Gard. Bull. Singapore 71(1): 28.
Plants perennial, shortly rhizomatous, 20-50 cm 2019; possible isolectotypes BM 000795696!,
tall, stolons present or absent; culms decumbent at BM 000959566!, US 00139833!, fragment ex BM).
the lower nodes, then erect, internodes cylindrical, (Fig. 9).
2.5-8 cm long, hollow, glabrous; nodes brownish,
pilose. Sheaths 2-3 cm long, usually shorter than Panicum telmatodes Balansa, Bull. Soc. Bot.
the internodes, striate, with long whitish hairs France 19: 324. 1872. TYPE. New Caledonia.
at the base and toward the apex; collar pale or Marais des environs de Boural, 4-IV-1869,
tinged with purple. Ligules membranous-ciliate, B. Balansa 895 (lectotype here designated
ca. 0.5-1 mm long. Leaves linear-lanceolate, P 00625052!; isolectotypes P 00625050!,
2.5-11 × 0.2-0.3 cm, flat or with involute borders, P 00625051!).
reflexed, membranous, glabrous. Inflorescence a
shortly exserted or partially included panicle, 3-4 × Plants annual, (20-)30-150 cm tall, aquatics, with
1.5-2 cm, contracted; peduncle striate, glabrous, culms with aerenchyma, decumbent and rooting
main axis striate, glabrous, pulvini glabrous, at the lower nodes, floating, then erect; internodes
spikelets paired on first order branches, axis of 4-20 cm long, compressed, glabrous, hollow, with
the branches scaberulous; pedicels 1-5 mm long, aerenchyma; nodes conspicuous, brownish, glabrous.
claviform, scaberulous; axillary inflorescences Sheaths striate, shorter than the internodes, 5-15 cm
absent. Spikelets long ellipsoid, 2.7-3 × 0.9-1 mm, long, glabrous; collar brownish to black, glabrous.
glabrous, pale and tinged with purple; lower Ligules membranous ciliate, 3-4 mm long. Blades
glume 0.4-0.5 mm long, 1/4 the length of the lanceolate, (5-)12-40 × (0.5)-0.7-2 cm, flat, rounded
spikelet, acute to truncate, nerveless to 1-nerved; to subcordate at base, glabrous, the midnerve manifest.
upper glume and lower lemma subequal, acute, Inflorescence a terminal, lax and multiflowered
7-nerved, membranous; lower palea and lower panicle, 6-25 × 6-15 cm. exserted; peduncle terete,
flower absent; upper anthecium long ellipsoid, glabrous, main axis glabrous, lower branches whorled,
2.4-2.6 × 0.9-1 mm, pale, smooth, indurated then alternate to subopposite, pulvini glabrous,
and glabrous, brownish at maturity, not early spikelets paired and appressed on triquetrous, scabrous
deciduous. Caryopsis not seen. branches, pedicels 2-6 mm long, claviform, scabrous.
345
Fig. 8. Panicum obseptum. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, dorsal view. E, Spikelet,
ventral view. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Upper palea with lodicules,
anthers and gynoecium. I, Lodicules and anthers. (From Hubbard 5732, K).
346
Fig. 9. Panicum paludosum. A, Habit. B, Ligule region. C, Spikelet, ventral view. D, Spikelet, dorsal view. E, Upper
palea with lodicules and anthers. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Caryopsis,
hilum view. I, Caryopsis, embryo view. (From Balansa 4578, P).
347
Spikelets lanceolate, 3-4(-4.2) × 0.9-1.1 mm, CHINA. Kwangtung. Guangdong, Pakhoi,

glabrous, greenish; lower glume 0.6-1.3 mm long, 20-IX-1921, Hitchcock 19234 (US); Kowloon,
1/4 the length of the spikelet, the apex truncate to 22-V-1941, Taam 2059 (G).
acute, nerveless to 1-nerved; upper glume and lower INDIA. Belatal, 2-IV-1962, Bhallachoryya s.n.
lemma subequal, acuminate, 7-9-nerved; lower (MO-3738145); Tirap River Valley, 12.4 mile mark
palea and lower flower absent; upper anthecium on Ledo Road, 6-IX-1945, Juan s.n. (G 00375026).
long ellipsoid, 2.4-2.6 × 0.9-1 mm, shorter than JAPAN. Mie, Honshu, Kuwana-gun, 31-VII-1964,
the upper glume and lower lemma, pale, indurated, Shimizu 14370 (C).
smooth and glabrous; upper lemma 5-nerved. NEW CALEDONIE. Diahot, Vieillard 1470 (P).
Caryopsis not seen. PHILIPPINES. Luzon. Cagayan, III-1909, Ramos
7429 (P); Benguet, Baguio, III-1904, Elmer 5972 (G).
Distribution and ecology. This species grows SRI LANKA. Ruhuna National Park, Block 2; at
in India, Sri Lanka, China, Taiwan, Thailand, Walaskema, 31-VII-1969, Cooray s.n. (P 001883373).
Malesia, Vietnam, New Guinea, Japan, the TAIWAN. Tapieh, 29-IV-1903, Faurie 731 (P);
Philippines, Australia and New Caledonia. It is Tomita-cho, Taihoku-shi, 14-IX-1932, Tanaka &
found floating in shallow water, ditches, and rice Shimada 11191 (G).
fields, between sea level and 2000 m a.s.l. THAILAND. Chiengmai, Doi Sutep, Sorensen et
al. 4475 (C).
Observation. Recently, Landge & Shinde (2021) VANUATU. Lac au pied du volcan de Tanua,
transferred this species to the genus Louisiella 7-V-1970, Schmid 3197 (P); Eramanga, IX-1858,
Hubbard & Leonard, based on morphological Cuming 18 (G).
characters and without a molecular analysis of VIETNAM. Taai Wong Mo Shan, near Chuk-phai,
this species or its comparison with both species Ha-coi, Tonkin, V-1939, Tsang s.n. (C, P 01883469);
of Louisiella. It should be noted that the presence Son Tay, 10-VIII-1885, Balansa 1627 (P); Hanoi,
of culms with aerenchyma, and a linear hilum 28-V-1891, Balansa 4578 (P); Ouombi, 19-XI-1885,
(which does not look linear in the published Balansa 471 (P).
figure of the authors) is also present in species
of sect. Dichotomiflora. For this reason, in this 12. Panicum perangustatum Renvoize, Kew
contribution P. paludosum is considered with this Bull. 44(3): 545. 1989. TYPE. Zambia. Kasama,
section and Panicum, until there is additional Misamfu, 22-I-1961, E. A. Robinson 4293
evidence that supports this segregation. (holotype K!; isotype PRE 0592240-0!). (Fig. 10).
Shouliang & Renvoize (2006) cited P.
dichotomiflorum as apparently an introduced Plants annual, 20-30 cm tall, culms geniculate
species in China, including P. paludosum at base, then ascending, conspicuously branching
as a synonym. However, P. paludosum is at the lower and middle nodes; internodes 2-8 cm
distinguished from P. dichotomiflorum by long, hollow, striate, glabrous, greenish to purplish;
being a perennial, aquatic species, with culms nodes pale, glabrous. Sheaths 2-6 cm long, as equal
with aerenchyma, and spikelets 3-4.2 mm long or shorter than the internodes, striate, pilose on the
(vs. plants annual, non aquatic, with culm upper margins, otherwise glabrous, greenish or
herbaceous, and spikelets 2-3 mm long in purple. Ligules membranous-ciliate, 1 mm long;
P. dichotomiflorum). collar pale, glabrous to sparsely pilose. Blades linear,
Veldkamp (1996) indicated that it is a 4-10 × 0.2-0.5 cm, flat to involute, acute, the lower
“favourite fodder of elephants and cattle. In India margins pilose, otherwise glabrous. Inflorescence
flour is used for making cakes”. a terminal, contracted panicle, 4-5 × 0.5-0.7 cm,
shortly exserted and partially included in the upper
Representative specimens examined leaves; main axis triquetrous, glabrous, first order
AUSTRALIA. Queensland. Without locality, branches appressed, pulvini glabrous, spikelets
Dietrich 1548 (P); Bauple, Wide Bay, 10-VI-1945, paired and appressed on first order branches,
Clemens s.n. (SI-51923). pedicels 2-6 mm long, claviform, glabrous.
348
Fig. 10. Panicum perangustatum. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, lateral view. E,
Spikelet, ventral view. F, Spikelet, dorsal view. Upper anthecium, dorsal view. G, Lower lemma and upper anthecium,
dorsal view. H-L, Upper anthecium, dorsal view I, Upper anthecium, ventral view J, Upper palea, lodicules and
anthers. K, Upper palea, lodicules and gynoecium. M, Upper anthecium, ventral view. N, Caryopsis, hilum view. O,
Upper palea and caryopsis, embryo view. (From Robinson 4293, K).
349
Fig. 11. Panicum pilgerianum. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, dorsal view. E, Spikelet,
ventral view. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Lower palea with lodicules and
anthers. I, Upper palea with caryopsis. J, Caryopsis, embryo view. K, Caryopsis, hilum view. (From Seydel 2211, BR).
350
Spikelets lanceolate, (3.5-)3.7-4.5 × 0.9-1 mm, nerveless to 1-nerved; upper glume and lower
acuminate, pale, glabrous; lower glume 0.9- lemma subequal, 7-9-nerved, acuminate, glabrous;
1.2 mm long, nearly 1/4 the length of the spikelet, lower palea elliptic, 4.2-4.3 × 1.1-1.2 mm,
3-5-nerved, glabrous; upper glume and lower lemma hyaline, glabrous; lower flower staminate, anthers
subequal, upper glume 9-11-nerved, membranous, 3-3.2 mm long; upper anthecium long ovoid, 2.5-
lower lemma 7-9-nerved, membranous; lower 2.6 × 0.8-0.9 mm, shorter than the upper glume
palea 2.5-2.8 × 0.7 mm, hyaline, glabrous; lower and lower lemma, pale, glabrous, not indurated
flower absent; upper anthecium long ellipsoid, 2.2- at maturity; upper lemma 7-nerved. Caryopsis
2.3 × 0.7-0.8 mm, shorter than the upper glume oblong, 2.6-2.8 × 0.8 mm, whitish to olivaceous;
and lower lemma, not indurated, pale, glabrous. hilum linear, embryo more than 1/2 the length of
Caryopsis obovoid, 2 × 0.7 mm, whitish; hilum the caryopsis.
oblong, embryo little less than half the length of
the caryopsis. Distribution and ecology. It is restricted to South
Africa, Namibia, and Botswana. This species grows
Distribution and ecology. Species endemic of in water, in seasonally flooded areas.
Zambia, where it is found in wet places; only known
from the type specimen. Observation. Panicum pilgerianum is
morphologically characterized by being an aquatic
13. Panicum pilgerianum (Schweick.) Clayton, plant, with culms with aerenchyma, and spikelets
Kew Bull. 42(2): 402. 1987. Acroceras long ovoid, the caryopsis with a linear hilum.
pilgerianum Schweick., Notizbl. Bot. Gart.
Berlin-Dahlem 14(122): 199. 1938. Psilochloa Representative specimens examined
pilgeriana (Schweick.) Launert, Mitt. BOTSWANA. Samedupe Bridge, 5-II-1977,
Bot. Staatssamml. München 8: 156. 1970. Smith 1899 (K); Content Farm, 10-III-1978, Hansen
TYPE. Namibia. Damaraland, Ovikokorero, 3370 (C, K).
25-XI-1914, M. K. Dinter 3395 (holotype NAMIBIA. Okahandja, Pretorius 26 (K);
B 10 0167254!; isotypes K!, PRE 0031873-0!, 13-III-1940, Volk 2917 (K); Omuramba, 23-IV-1950,
US-766249!, fragment ex B). (Fig. 11). Schweickerdt 2199 (K); Waterberg, V-1949,
Liehenberg 4740 (K); Okatjongeama, 28-III-1960,
Plants annual, 40-60(-120) cm tall, branching Seydel 2211 (BR, K).
at the upper nodes; culms erect, geniculate and
rooting at the lower nodes; internodes with 14. Panicum porphyrrhizos Steud., Syn. Pl.
aerenchyma, 4-14(-20) cm long, hollow, glabrous; Glumac. 1: 72. 1853. TYPE. Ethiopia. “In
nodes brownish, glabrous. Sheaths 3-11(-17) cm planitie montana Schire”, 10-X-1840, G. H.
long, as long as or shorter than the internodes, W. Schimper 1808 (lectotype here designated
striate, glabrous. Ligules membranous-ciliate, P 01933656!; isolectotype W 0023588!). Other
1.2-2 mm long; collar glabrous. Leaves lanceolate, syntype. Ethiopia. Prope Gafta, 18-IX-1842,
4-30 × 0.3-1.1 cm, flat, rounded at the base, the G. H. W. Schimper 1230 (BM 000923133!,
apex acuminate, glabrous. Inflorescence a terminal, BR 0000008367433!, G 00022434!, G 00022435!,
contracted to lax panicle, 23-30 × 4-20 cm; GOET 006787!, L 0044835!, L 0044836!,
main axis triquetrous, wavy, glabrous, first order M 0103981!, STU 000128!, STU 000129!,
branches adpressed, axis of the branches striate, US 00902247!, US 00147972!, US 00147973!,
glabrous, spikelets solitary or paired on first and W 0023589!, W 18890236397!). (Fig. 12).
second order branches; pedicels 2-7 mm long,
claviform, striate, glabrous; axillary panicles Panicum praelongum Mez, Bot. Jahrb. Syst. 57:
present, similar to the terminal one. Spikelets long 186. 1921, nom. illeg. hom., non Steud., 1853.
ovoid, 4-5.3 × 1-1.1 mm, acuminate, glabrous, TYPE. Sudan, without locality, Roriani 8
greenish; lower glume 0.9-1.4 mm long, 1/5 to (lectotype here designated B 10 0168705!;
1/4 the length of the spikelet, hyaline, truncate, isolectotype B 10 0168704!).
351
Fig. 12. Panicum porphyrrhizos. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, ventral view. E, Spikelet,
dorsal view. F, Spikelet, lateral view. G, Upper anthecium, dorsal view. H, Upper anthecium, ventral view. I, Upper palea
with lodicules and anthers. J, Stamen. K, Upper palea with lodicules and caryopsis. (From Schweinfurth 1160, G).
352
Plants perennial, shortly rhizomatous, 60-120 cm Representative specimens examined.

tall, culms simple, erect or geniculate at the base, EGYPT. Cairo, Gay s.n. (US 04303922).
internodes 5-20 cm long, cylindrical, hollow, ETHIOPIA. Amozai, 19-IX-1862, Schimper s.n.
glabrous, without aerenchyma; nodes compressed, (US 04303921); about 200 km SW of Addis Adeba,
dark, glabrous, not sharply demarcated. Sheaths 23-VIII-1965, de Wilde et al. 7875 (C)
3-11 cm long, shorter than the internodes, striate, GHANA. Adisabel Estate, Cape Coast, 16-V-1965,
glabrous. Ligules membranous-ciliate, 1-1.3 mm Hall 3050 (US).
long, the ciliate portion short. Blades lanceolate, KENYA. 20 miles east of Nairobi, 16-VII-1951
10-27 × 0.5-0.9 cm, flat, acuminate, the lower Bogdan 3137 (K).
half and lower margins with tubercullate hairs, MALAWI. Nyasaland, ca. 2 miles north of
otherwise glabrous. Inflorescence a terminal, Kasungu, 14-I-1959, Robson 1172 (K, P).
exserted, lax and multiflowered panicle, 20-40 × SUDAN. Callabat, Matamma, 1865,
15-20 cm; peduncle 15-20 cm long, terete, Schweinfurth 1144 (P), 1160 (G).
glabrous; main axis striate, glabrous, the branches TANZANIA. Selous Game Reserve, Kingupira,
alternate to opposite, distant, divergent, glabrous, 27-II-1976, Vollesen 3291 (C).
pulvini glabrous, spikelets distant on second UGANDA. Kotido, Karamoja, 3-VI-1940,
and third order branches, pedicels 2-6 mm Thomas 3696 (K, MO).
long, claviform, scabrous. Spikelets long ovoid, ZAMBIA. Fort Jameson, I-1963, Verboom 501
2.9-3.5(-3.8) × 1.1-1.2 mm, glabrous, acuminate, (C, US).
greenish to purplish; lower glume 0.9-1 mm long, ZIMBAWE. Sengwa Research Station, 13-II-1977,
1/4 the length of the spikelet, acute, 1-3-nerved; Guy 2471 (K).
upper glume and lower lemma subequal,
membranous, upper glume 9-nerved, lower lemma 15. Panicum schinzii Hack. ex Schinz., Verh. Bot.
7-nerved; lower palea 2-2.4 × 0.5-0.8 mm, hyaline, Vereins Prov. Brandenburg 30: 142. 1888. TYPE.
glabrous; lower flower absent; upper anthecium Namibia, Olukonda, 1886, H. Schinz 641 (lectotype
long ovoid, 2-2.4 × 0.9-1 mm, glabrous, smooth, Z 000080584!, designated as “holotype” by
indurated, brownish at maturity; upper lemma Veldkamp, 1996: 204; isolectotypes K 000255500!,
5-nerved, palea and lemma with papillae toward US 00139969!, fragment ex W). (Fig. 13).
the apex. Caryopsis not seen.
Panicum laevifolium Hack., Bull. Herb. Boissier 3(8):
Distribution and ecology. Africa. This is a 378. 1895. TYPE. South Africa. Transvaal. Hogge
widely distributed African species, from Ethiopia Veld inter Porter et Trigardsfont, A. Rehmann
and Sudan in the north to Botswana, Zimbabwe 6614 (lectotype here designated K 000255496!;
and Mozambique in the south; it is found in wet possible isolectotype W 0177664!). Other syntypes:
habitats, near swamps and rivers, between 400 and South Africa. Transvaal. Pretoria, Kuduspoort, A.
2200 m a.s.l. Rehmann 4697 (W). Transvaal. Boshveld inter
Elandsriver et Klippan, A. Rehmann 5123 (W).
Observation. When describing P. porphyrrhizos, Donkershoek, A. Rehmann 6552 (K 000255498!, W).
Steudel mentioned two syntypes, both collected by Panicum laevifolium Hack. var. amboense Hack.,
Schimper in Ethiopia, #1230 and #1808. Of these, Vierteljahrsschr. Naturf. Ges. Zurich 56: 71. 1911.
the specimen Schimper 1808, P 01933656 of the TYPE. Namibia. Amboland: swischen Ondonga
Steudel herbarium, is here designated as lectotype und Uukuanyama, XI-1906, M. Rautanen 593 (K
of the species; it is a complete specimen which 0002555499!, US-80727!, fragment, possible type
agrees with the protologue of the species. W 19080009792! labeled as M. Rautanen 5).
There are two syntypes of P. praelongum Mez
in B; of these the specimen B 10 0168705 is here Plants annual, (30-)40-180 cm tall; culms
selected as lectotype of the species; it is a more erect to decumbent and geniculate at base, simple
complete specimen and has Mez script with the or branching, internodes terete, 3-30 cm long,
designation of a new species. hollow, glabrous; nodes brownish, glabrous.
353
Fig. 13. Panicum schinzii A, Habit. B, Ligule region. C, Spikelet, lateral view. D, Spikelet, ventral view. E, Lower
palea with lodicules and anthers. F, Upper anthecium, dorsal view. G, Upper anthecium, ventral view. H, Upper palea
with caryopsis, embryo view. I, Caryopsis, hilum view. (From Drummond 5330, P).
354
Sheaths striate, open, glabrous, 2-15 cm long; ZIMBABWE. Hunyani, 3-II-1932, Stent
collar pale, glabrous. Ligules membranous-ciliate, 5557 (K); Mensa Pan, ESE of Chirundu bridge,
1.5-2 mm long. Blades lanceolate, 10-25(-35) × 29-I-1958, Drummond 5320, 5330 (P); Melsetter,
1-1.5 cm, flat, subcordate, the lower margins Pasture Research Substation, 12-II-1950, Williams 61
ciliate, otherwise glabrous. Inflorescence a lax, (K, US), Williams 27247 (P); Salisbury, Salisbury
open and terminal panicle, (13-)20-27 × (-5)10- Experimental Station, 1-II-1961, Phipps 2846 (P); Mana
20 cm; main axis triquetrous, glabrous, pulvini Pools National Park, 21-III-1998, Poilecot 7771 (G).
glabrous, first order branching alternate, distant,
ascending, spikeletes dispersed on first and second 16. Panicum subalbidum Kunth in Révis. Gramin.
order branches; pedicels 2-7 mm long, scaberulous, 2: 397, tab. 112. 1831. TYPE. Senegal. Dagana,
claviform; axillary inflorescences present, similar Walo, IX-1925, F. M. R. Leprieur s.n. (holotype
to the terminal one. Spikelets ellipsoid, 2.5-2.8 × P 00442196!; possible isotype B 10 0168673!).
0.9-1 mm, glabrous, pale to greenish; lower (Fig. 14).
glume 0.6-0.7 mm long, 1/5 to 1/4 the length of
the spikelet, acute to truncate, 1-nerved, upper Panicum glabrescens Steud., Syn. Pl. Glumac.
glume and lower lemma subequal, 7-9-nerved, 1: 71. 1853. TYPE. Senegal. Without locality,
membranous, acute; lower palea elliptic, 2.5- F. M. R. Leprieur s.n. (holotype P 00731447!).
2.6 × 0.9-1 mm, hyaline, glabrous, lower Panicum kermesinum Mez, Bot. Jahrb. Syst.
flower staminate, anthers 3, 1.5 mm long; upper 57: 189. 1921. TYPE. Cameroon. Adamana,
anthecium long ellipsoid, 2-2.1 × 0.9-1 mm, pale, Ledermann 4115 (holotype B †; isotype? US
black at maturity, indurated, smooth and shiny, 00148215!, fragment ex B. Hitchcock mentioned
upper lemma 5-nerved. Caryopsis ellipsoid, 1.7- in this fragment that “this seems to be the type,
1.8 × 0.9-1 mm, whitish; hilum oblong, embryo Ledermann 4225 not in Berlin”, 1935).
less than half the length of the caryopsis. Panicum longijubatum (Stapf) Stapf, Fl. Trop. Afr. 9:
718. 1920. Panicum proliferum var. longijubatum
Distribution and ecology. It is widespread in Stapf, Fl. Cap. 7: 406. 1899. TYPE. South Africa.
South Africa and Namibia, reaching Botswana and Eastern Region: near Durban, Williamson 21
Zimbabwe; introduced in Malesia (Java), Australia (lectotype here designated K 000255491!;
and New Zealand. It grows in open, and seasonally isolectotype PRE 0038229-0!). Other syntypes.
flooded, fields, commonly as a weed, between 500 South Africa, Coast Region. Komgha division: by
and 2000 m a.s.l. the Kei River, near Komgha, 1-XII-1891, H. G.
Flanagan 953 (GRA 0000092-0!, K 000255495!).
Observation. The lectotype here designated of Eastern Region: Natal, near the Umzimkulu
P. laevifolium is Rehmann 6614, not 6646 as it was River, 1840, Drège s.n. (K 000255493!). Eastern
originally included in the type diagnosis. Region, common near Umpumulo, Buchanan 267
(K 000255494!, W 19160023954!).
Representative specimens examined Panicum ingens Peter, Repert. Spec. Nov. Regni
AUSTRALIA. New South Wales. Glen Innes, Veg. Beih. 40(1): 40, tab. 24, fig. 2. 1929. TYPE.
8-IV-1931, Hubbard 8210 (CANB, P). Tanzania. “Deutsch-Ostafrika: Ngulu: Gweko
BOTSWANA. Content Farm, 27-I-1978, Hansen westl. [to] Igalula im Wasser. ...”, 15-I-1926,
3341 (K). G. A. Peter 34926 (holotype B †; lectotype
NEW ZEALAND. Kekerengu Ecological here designated W 1960020754!; isolectotype
District, 1-I-2014, Jones s.n. (AK). G 00022441!).
SOUTH AFRICA. 28 miles east of Bethlehem, Panicum longiramum Peter, Repert. Spec. Nov.
16-III-1964, Oakes 493 (SI); Tweespruit, Regni Veg. Beih. 40(1): 44, tab. 29, fig. 1. 1929.
Thaba Pitswa, 18-II-1994, Du Preez 2702 (C); TYPE. Tanzania. “Deutsch-Ostafrika: Ngulu:
Nylstroom, Mosdene Farm, 4-II-1986, Smook Malongwe Nyahua ...”, 9-I-1926, G. A. Peter
2428 (C). Transvaal. Apiesrivier, 14-I-1894, 34539 (holotype B †; lectotype here designated
Schlechter 4166 (G, P). W 19600020878!).
355
Fig. 14. Panicum subalbidum. A, Habit. B, Ligule region. C, Inflorescence branch. D, Spikelet, ventral view. E,
Spikelet, dorsal view. F, Upper anthecium, ventral view. G, Upper palea with lodicules and gynoecium. (From
Friis & Vollesen 318, C).
356
Panicum bicuspidatum A. Camus, Bull. Soc. Bot. Representative specimens examined

France 99: 64. 1952. TYPE. Madagascar. Prov. BURUNDI. Muramvya, entrée Teza, 5-VI-1975,
Antsiranana: collines et plateaux calcaires de Reekmans 5239 (BR).
l’Ankarana, XII-1937/I-1938, H. Humbert CHAD. Baguirmi, Nigui, 10-VIII-1903,
18855 (holotype P 00450278!). Chevalier 9453 (P); Fort Lamy, 15-IX-1938, Eloueh
263 (P).
Plants perennial, shortly rhizomatous, 30-100 cm COTE D’IVOIRE. Route entre Sikensi et IV
tall, with adventitious roots and rooting at the lower Dancé, without collector 381A (P).
nodes, culms decumbent at the base, then erect, DEMOCRATIC REPUBLIC OF CONGO.
1-2 m tall; internodes 2-18 cm long, cylindrical, Kahuzi, Marais Musisi, 2-VI-1959, Leonard 4479
with aerenchyma, hollow, glabrous; nodes (BR); plaine de la Ruzizi, Luvungi, 27-II-1950,
compressed, dark, sharply demarcated, glabrous. Germain 6318 (BR); Yangambi, 11-XI-1947,
Sheaths 7-20 cm long, longer than the internodes, Leonard 1518 (C).
striate, glabrous; collar brown, glabrous. Ligules ETHIOPIA. Shoa, in the upper part of the
membranous-ciliate, 1-3 mm long. Blades Mugher river system, about 50 km N of Addis
lanceolate, 30-40 × 0.8-1.2 cm, flat, glabrous, the Adeba, 12-XI-1972, Friis et al. 1126 (C).
lower margins ciliate. Inflorescence a terminal, M adagascar . M a h a j a n g a . N e a r
multiflowered, lax and open panicle, 15-35 × 10-25 Antanimbary, environs of Maevatanana, 13-II-2013,
cm, partly or fully exserted, the branches naked at Vorontsova et al. 913 (K).
the base, spikelets paired and appressed on second MALI. De Sebé a Mopti, 1899, Chevalier 2199
order branches; peduncle glabrous, main axis and (K); Macina, de Randiagara a Mopti, 7-IX-1910,
pulvini glabrous; branches triquetrous, scabrous; Chevalier 24910 (P).
pedicels 2-5 mm long, triquetrous, scabrous and RUANDA. Rutsiro, 28-VII-1974, Lambinon
claviform. Spikelets lanceolate, 2.6-3 × 0.8-1 mm, 74/595 (BR).
glabrous, greenish to purple; lower glume 0.7-0.8 SENEGAL. Basse Casamance, Djifanghor,
mm long, 1/4 to 1/3 the length of the spikelet, acute 4-VIII-1978, Berghen 261a (BR).
to obtuse, 1-3-nerved; upper glume and lower SOUTHERN SUDAN. Between Gilo and Mt.
lemma subequal, 7-9-nerved; lower palea and Konoro, 18-XI-1980, Friis & Vollesen 318 (C).
lower flower absent; upper anthecium ellipsoid, TANZANIA. Arusha National Park, near Meru,
1.8-2.5 × 0.8-1.1 mm, shorter than the upper glume 8-VII-1970, Hansen 323 (C).
and lower lemma, indurate, brownish at maturity,
shiny. Caryopsis long ellipsoid, 2 × 1 mm, 17. Panicum sublaeve Swallen, Contr. U.S.
whitish; hilum oblong, embryo ca. 1/2 the length Natl. Herb. 29(9): 424. 1950. Panicum rigidum
of the caryopsis. Swallen, Ann. Missouri Bot. Gard. 30(2): 215.
1943, nom. illeg. hom., non Balfour, 1884.
Distribution and ecology. Common African TYPE. Panamá. Sabana de Juan Corso, near
grass, also growing in Madagascar, usually Chepo, 1-X-1911, H. Pittier 4526 (holotype
found growing in water, edges of rice paddies, US 00147999!). (Fig. 15).
0-2100 m a.s.l.
Plants anual, caespitose, 15-65 cm tall; culms
Observations. This species is recognisable by geniculate to erect, branching; internodes terete,
its aquatic habit, with culms with aerenchyma, hollow, glabrous; nodes brownish, glabrous.
decumbent and rooting at the lower nodes, and Sheaths longer than the internodes, striate,
dark nodes. glabrous. Ligules membranous-ciliate, ca. 1.6 mm
The specimen Williamson 21, from K, is long, the membranous portion 0.3 mm long; collar
designated as lectotype of P. proliferum var. brownish, glabrous. Blades linear-lanceolate,
longijubatum Stapf, among the syntypes originally (3-)13-24 × (0.2-)0.5-1.2 cm, rounded to subcordate
cited in this variety, since it is a full specimen and at the base, the apex attenuate, flat, the adaxial
agrees with the protologue. surface sparsely pilose, abaxial surface glabrous.
357
Fig. 15. Panicum sublaeve. A, Habit. B, Spikelet, ventral view. C, Spikelet, dorsal view. D, Lower palea. E, Upper
anthecium, dorsal view. F, Upper anthecium, ventral view. G, Apex of the upper anthecium. H, Caryopsis, embryo view.
I, Caryopsis, hilum view. (From Zuloaga 4354, SI).
358
Inflorescence a terminal, exserted and lax panicle, 18. Panicum sumatrense Roth ex Roem. &
15 × 10 cm; peduncle terete, 5-20 cm long, Schult., Syst. Veg. (ed. 15bis) 2: 434. 1817.
glabrous; main axis wavy, scaberulous to glabrous, TYPE. Indonesia. Sumatra, B. Heyne s.n.
pulvini glabrous, first order branches alternate, axis (lectotype B 10 0366235!, designated by J. F.
of the branches and pedicels triquetrous, scabrous, Veldkamp et al., Blumea 34: 82. 1989).
spikelets solitary and disperse on the branches;
axillary panicles present. Spikelets long ovoid, 3.5- Panicum attenuatum Willd, Enum. Pl.: 1033. 1809,
4 × 0.7-1.2 mm, glabrous, greenish; lower glume nom. illeg. hom., non Moench, 1802. Panicum
ovate, 1.2-1.5 mm long, acute, 5-7-nerved; upper proliferum var. attenuatum Schrad., Linnaea
glume and lower lemma subequal, 11-13-nerved, 12: 428. 1838. TYPE. Place or origin unknown,
acute, membranous; lower palea elliptic, 2.7-3.3 × J. T. Klein 195 (B-W-18740-09-0!; US 00148155!,
1 mm, hyaline, glabrous; lower flower staminate, fragment ex B).
anthers 3; upper anthecium long ovoid, 2.5-2.8 × Panicum psilopodium Trin., Gram. Panic.: 217.
0.9-1.2 mm, shorter than the upper glume and lower 1826. Panicum sumatrense subsp. psilopodium
lemma, whitish, smooth, indurated, shiny; upper (Trin.) de Wet, J. Agric., Tradit. Bot. Appl.
lemma 7-nerved, apiculate and with conspicuous 30: 159. 1983. TYPE. India. Without locality,
prickles toward the apex and papillae all over its J. Lindley s.n. (holotype LETRIN-0908.01!;
surface. Caryopsis long ellipsoid, 1.9 × 0.8 mm, isotype L 0044865!, fragment).
whitish; hilum oblong, embryo more than half the Panicum simplex Rottler ex Trin., Gram. Panic.:
length of the caryopsis. 216. 1826. TYPE. “V. sp. Ind. Or.”, Swartz s.n.
(LE).
Distribution and ecology. Species distributed Panicum crispum Llanos, Fragm. Fl. Filip.: 41.
from Mexico and Mesoamerica up to Venezuela; it 1851. TYPE. Philippines, Benguet, Luzon,
grows in open and humid habitats, from sea level 1-V-1914, E. D. Merrill 1764 (neotype L,
to 1100 m a.s.l. designated by J. F. Veldkamp, Blumea 41:
206. 1996; isoneotypes GH 00091557!,
Observation.This species is characterized by NY 00328834!, SING 0054929!).
having spikelets with the upper glume and lower Panicum psilopodium Trin. var. coloratum Hook.
lemma 11-13-nerved, upper anthecium shorter and f., Fl. Brit. India 7(21): 47. 1897. SYNTYPES.
apiculate, with conspicuous prickles at the apex. Burma. Shan Hills, Collett s.n. (K?). India?. Mt.
Abu, Duthie s.n. (K).
Representative specimens examined. Panicum psilopodium Trin. var. epaleatum Keng
COSTA RICA. Guanacaste. Nicoya Peninsula, ex S.L. Chen, T.D. Zhuang & X.L. Yang, Bull.
5 km SSW of Veinte Siete, 17-VIII-1968, Pohl & Bot. Res., Harbin 4(2): 124. 1984. TYPE.
Davidse 10941 (F). China. Guizhou: Du Shan, 7-IX-1930, Y. Tsiang
EL SALVADOR. La Libertad. Along CIA, km 38, 6912 (JSIB).
ca. 10 km W of La Libertad, open roadside near Panicum longiloreum M.M. Rahman, Kew
Pacific Ocean, 10 m, 17-VIII-1971, Pohl 12774 Bull. 44(3): 485. 1989. TYPE. Thailand.
(F, MO). Sena, Ayuthia, 23-IX-1930, A. F. G. Kerr
HONDURAS. Francisco Morazán. Vicinity of 19709 (holotype K 000290301!; isotypes
El Zamorano, 17-X-1951, Swallen 10839 (MO, US). BM 000959569!, BK 257944!, BRI-AQ269481!,
MEXICO. Chiapas. Small forested stream L 0044813!).
and adjacent savanna 2 km SE of Arriaga,
27-VIII-1974, Breedlove 36852 (MEXU, MO). Plants annual, 10-90 cm tall; culms erect to rooting
PANAMA. Canal Zone. Monagre Beach 5 mi and branching at the lower nodes, internodes 3-10 cm
SE of Chitre, 22-I-1966, Tyson et al. 3030 (MO). long, terete, hollow, glabrous; nodes brownish,
VENEZUELA. Guárico. Miranda, 10 km al N glabrous to sparsely pilose. Sheaths striate, with
de Camaguan, 6-VIII-1989, Zuloaga et al. 4354 sparse tubercullate, caducous hairs to glabrous,
(SI, VEN). 4-7 cm long, shorter or longer than the internodes.
359
Ligules membranous-ciliate, 0.6-1.7 mm long; PHILIPPINES. Luzon. Baguio, Benquet,

collar brownish, glabrous. Blades linear to VIII-1928, Clemens 18291 (C); Manila, VI-1911,
lanceolate, (5-)10-35 × 0.4-1.4 cm, flat, rounded Merrill 597 (G).
at base, acuminate, sparsely ciliate toward the SRI LANKA. Eastern Province. Amparai,
base to glabrous, the margins membranous, Lahugala-Kitulana Sanctuary at edge of the
smooth. Inflorescence a contracted to lax panicle, Lahugala Tank, 2-XII-1974, Davidse 8960 (CANB,
6-40 × 2-15 cm; main axis wavy, smooth to K, MO, US); near Padikema, Ruhuna National Park,
scaberulous, pulvini dark, glabrous; first order 6-XII-1969, Cooray s.n. (MO-4378481).
branches alternate, ascending to appressed, axis THAILAND. Ban Kao, 12-XI-1961, Larsen
of the branches wavy, scaberulous; spikelets paired 8159 (C).
or solitary, pedicels 2-9 mm long, scaberulous. VIETNAM. Chapa, VII-1927, Petelot 5063 (US).
Spikelets ellipsoid, (2.2-)3-3.5(-4) × 1.1-1 mm,
glabrous; lower glume 0.6-1.1 mm long, 1/4 to 19. Panicum trichonode Launert & Renvoize,
1/3 the length of the spikelet, 1-3-nerved, acute Prodr. Fl. Südwestafrika 34(160): 226. 1970.
to truncate at the apex; upper glume and lower TYPE. Zambia. 80 km W of Chingola,
lemma subequal, acute, upper glume 11-15-nerved, 18-XII-1963, E. A. Robinson 6124 (holotype
membranous, lower lemma 9-11-nerved; lower K 000282461!; isotype SRGH 0106321-0!).
palea almost as long as the lower lemma, hyaline, (Fig. 16).
glabrous; lower flower staminate; upper anthecium
ellipsoid, 2.3-2.6 × 0.9-1 mm, shorter than the Plant perennial, caespitose and shortly
upper glume and lower lemma, indurate, dark rhizomatous, 30-110 cm tall, shortly rhizomatous,
brown, shiny, glabrous. Caryopsis ellipsoid, 1.7- cataphyls pilose; culm erect, simple, internodes
1.9 × 0.9-1 mm; hilum oblong, embryo ca. 1/2 the 4-15 cm long, cylindrical, hollow, striate, glabrous;
length of the caryopsis. nodes densely pilose, with conspicuous white short
hairs. Sheaths striate, as long as or longer than the
Common names. “Blue panic”, “little millet”, internodes, 8-25 cm long, covered with tubercullate
“sama” (Veldkamp, 1996). caducous hairs; collar brown, pilose. Ligules
membranous-ciliate, 0.6-0.8 mm long. Blades linear-
Distribution and ecology. This species grows lanceolate, 12-25 × 0.3-0.5 cm, rounded at the base,
in Tropical Asia (India, Pakistan, China, Sri Lanka the apex acuminate, with caducous tubercullate
to Vietnam, and Malesia). It is common in open hairs all over its surface and margins, the lower ones
fields and forest edges and in rice fields and grassy ciliate. Inflorescence a terminal lax panicle, 3-20 ×
roadsides, between sea level and 1800 m a.s.l. 1.5-5 cm; peduncle terete, glabrous, up to 25 cm
long, first order branches appressed, the branches
Observations. Bor (1960) mentioned this is the triquetrous, smooth; pulvini glabrous, spikelets
valid name for P. psilopodium. However, the lower appressed on first and second order branches;
glume is more than 1/2 the length of the spikelet, pedicels 1-2.5 mm long, triquetrous, scaberulous;
therefore not pertaining to sect. Dichotomiflora. axillary inflorescences absent. Spikelets long
ellipsoid, 2.5-3 × 1-1.1 mm, glabrous, greenish or
Representative specimens examined. tinged with purple; lower glume 0.4-0.8 mm long,
CHINA. Yunnan. Lou-pou, Maire s.n. 1/5 to 1/4 the length of the spikelet, truncate to
(P 01907989); Hainan, Pak shik Ling and vicinity, acute, nerveless to 1-nerved; upper glume and lower
Ku Tung Village, 21-V-1933, Lei 685 (P); Yun-Nan, lemma subequal, 7-9-nerved, membranous; lower
18-II-1906, Ducloux 4193 (P). palea 2.7 × 0.7 mm, narrowly ellipsoid, glabrous,
INDIA. Assam, below Kohima, Naga Hills, hyaline; lower flower staminate, anthers 1.3 mm
2-VII-1950, Chand 3229 (US); Bombay, Borivali long; upper anthecium narrowly ellipsoid, 2.2-2.6 ×
National Park, 11-VIII-1952, Fernandez 101 (K). 1 mm, indurated, pale, smooth, shiny. Caryopsis
NEPAL. Near Tilhar, 19-V-1954, Stainton et al. ellipsoid, 1.7 × 0.9 mm; hilum oblong, embryo 1/3
2772 (G). the length of the caryopsis.
360
Fig. 16. Panicum trichonode. A, Habit. B, Ligule region. C, Detail of node. D, Spikelet, lateral view. E, Lower glume.
F, Upper glume. G, Lower palea with lodicules. H, Upper anthecium, dorsal view. I, Upper anthecium, ventral view.
(From Smith 1413, K).
361
Distribution and ecology. It is restricted to glabrous, first order branches alternate to opposite,
southern tropical Africa (Botswana, Zambia and ascendent, appressed, spikelets paired or solitary
Zimbabwe), and southern Africa (Namibia). It is on short second order branches; pedicels short,
found in open, damp grasslands on clay soils; 1000- claviform, scabrous; axillary inflorescences present.
2000 m a.s.l. Spikelets long ellipsoid, 2.2-3.2 × 1.1-1.4 mm,
acute, greenish, glabrous; lower glume 1/4 to 1/3
Representative specimens examined the length of the spikelet, ovate, hyaline, 1-nerved,
BOTSWANA. On the sides of Karongana river, obtuse to truncate; upper glume and lower lemma
19º45’S, 22º24’E, 18-VIII-1975, Smith 1413 (K); subequal, membranous, acute, 7-nerved; lower palea
Okovango River, 14-II-1979, Smith 2640 (K). and lower flower absent; upper anthecium long
NAMIBIA. Kunene Banks, III-1923, Barnard ovoid, 2-2.7 × 1-1.2 mm, smooth, shiny, glabrous,
26 (K). pale, dark brown at maturity, with verrucose papillae
ZAMBIA. Barotseland, Mongu, 20-I-1966, toward the apex of the palea, early deciduous at
Robinson 6816 (K); Kalomo, near Senkobo, Astle maturity. Caryopsis ellipsoid; hilum oblong, embryo
284b (K); two miles from Namwala, 10-XII-1962, 1/2 the length of the caryopsis.
van Rensburg 1051 (K); Dambo centre, I-1979,
Heery 25 (K). Distribution and ecology. Species endemic of
ZIMBABWE. Salisbury, 7-II-1919, Eyles 1524 (K); Mexico, where it grows in humid áreas between
without precise locality, IV-1920, Eyles 2205 (K); 1700-2700 m a.s.l.
Guruve, 3-I-1979, Nyariri 614 (K).
Observation. Panicum vaseyanum is well defined by
20. Panicum vaseyanum Scribn. ex Beal, having contracted panicles, axillary inflorescences
Grass. N. Amer. 2: 140. 1896. TYPE. Mexico. present, spikelets with the upper glume and lower
Chihuahua. Base of Sierra Madre, 30-IX-1887, lemma 7-nerved, lower palea and lower flower
C. G. Pringle 1415 (lectotype US 00148082!, absent, and upper anthecium dark brown and easily
designated as “type” by Hitchcock & Chase, deciduous at maturity.
1910: 47; isolectotypes COLO 00392639!,
COLO 00392621!, E 00373819!, F 0046899F!, Representative specimens examined
GH 00024127!, K 000309146!, MEXU MEXICO. Aguas Calientes. Near Aguas
00003724!, MICH 1108751!, MIN 1001936!, Calientes, 2-X-1910, Hitchcock s.n., AGH 15 (F,
MO-128362!, MSC 00922516!, NDG 06888!, MEXU, MO, NY, P) Chiapas. San Cristóbal de las
NY 00381781!, NY 00381782!, PH 00018737!, Casas, west edge of San Cristóbal de las Casas, plant
RSA 0000465!, US 00148081!, US 00037314!, in marsh, 2195 m, 29-X-1981, Breedlove & Davidse
UVMVT 027972!, W 19160022600!). 53971 (MO). Chihuahua. Pine-Oak región, Sierra
Madre Occidental, west of Casas Grandes, five
Plants annual, caespitose, 15-70 cm tall; culms miles south of Hernandez, 18-IX-1960, Reeder et al.
geniculate, decumbent and rooting at the lower 3526 (MEXU, US). Durango. Reserva La Michila,
nodes, then erect, branching at the upper nodes; 80 km al SE de Durango, 2500 m, 6-X-1980, Carrillo
internodes compressed, with aerenchyma, pale to 9 (MEXU). Guanajuato. About 6 km W of San
purplish, glabrous, hollow; nodes dark, glabrous. Felipe, on dry slopes approaching Cerro del Fraile,
Sheaths striate with anastomosed veins, glabrous, 24-X-1952, Sohns 399 (US). Jalisco. Guadalajara,
the margins membranous. Ligules membranous- road to Barranca Oblato, 6100 ft, 27-IX-1910,
ciliate, 0.5-2 mm long; collar brownish to purplish, Hitchcock 7315 (US). Mexico. Ciudad de Progreso,
glabrous. Blades lanceolate, 3-25 × 0.2-0.6 cm, flat 2700 m, 30-IX-1950, 2700 m, 30-IX-1950, Matuda
or with involute borders toward the base, rounded 19747 (MEXU, MO, US). Michoacán. 25 km E
at base, the apex attenuate, greenish to purplish, of Morelia, on highway 15, 2000 m, 20-X-1976,
glabrous, the margins scabrous. Inflorescence a Brunken & Perino 448 (MO). Puebla. Autopista de
contracted, shortly exserted panicle 2-9 ×1-1.5 cm; Puebla a Orizaba, al NE de Puebla, 1.4 km al E de
main axis wavy, scaberulous to glabrous, pulvini la Puente Amaluca, 2260 m, Vibrans 2453 (MEXU).
362
SPECIES EXCLUDED FROM SECT. Bor, N. L. 1960. Grasses of Burma, Ceylon, India and Pakistan.
DICHOTOMIFLORA Oxford: Pergamon Press.
Brown, W. V. 1948. A cytological study in the Gramineae.
Zuloaga et al. (2018) considered P. bechuanense American Journal of Botany 35: 382-395.
Brem. & Oberm. within sect. Dichotomiflora, while Chase, A. 1939. Papuan grasses collected by L. J. Brass, II.
in fact this is a species of sect. Repentia Stapf. Journal of the Arnold Arboretum 20: 304-316.
Veldkamp (1989) considered as synonym of P. Church, G. L. 1929. Meiotic phenomena in certain
sumatrense the following species: Gramineae. II. Paniceae and Andropogoneae. Botanical
Panicum albidulum Steud., Syn. Pl. Glumac. 1: 69. Gazette 88: 63-84.
1853. TYPE. Sudan. Cordofan, Abu Gerad, 22 Sep Clayton, W. D. & S. A. Renvoize. 1986. Genera graminum.
1839, C. G. T. Kotschy 42 (holotype P 00731470!; Grasses of the world. Kew Bulletin, Additional Series
isotypes BR 0000008767523!, L 0044866!, 13: 1-389.
M 0103913!, W 0023395!, W 18890236406!, Davidse, G. & R. W. Pohl. 1972. Chromosome numbers
W 18890236407!, W 19040010290!). and notes on some Central American grasses. Canadian
In fact, P. albidulum is a synonym of P. laetum Journal of Botany 50: 273-283.
Kunth, species of sect. Panicum. Davidse, G. & R. W. Pohl. 1974. Chromosome numbers,
Panicum cristatellum Keng, Sinensia 11: 412. meiotic behavior, and notes on tropical American
1940. TYPE. China. Jiangsu: Jiangyin, June 1922, grasses (Gramineae). Canadian Journal of Botany 52:
A. Allison 202 (holotype US 00148386!). 317-328.
This species, mentioned as a possible synonym of Dubcovsky, J. & F. O. Zuloaga. 1991. Números cromosómicos
P. sumatrense (Shouliang & Renvoize, 2006), clearly de especies sudamericanas de Panicum (Poaceae:
does not belong to sect. Dichotomiflora, since it has a Paniceae). Boletín de la Sociedad Argentina de Botánica
lower glume 3/4 the length of the spikelet. It is most 27: 201-206.
likely related to species of sect. Hiantes. Freckmann, R. W. & M. G. Lelong. 2003. Nomenclatural
changes and innovations in Panicum and Dichanthelium
(Poaceae: Paniceae). Sida 20: 161-174.
ACKNOWLEDGMENTS Giussani, L. M.; J. H. Cota-Sánchez, F. O. Zuloaga & E.
A. Kellogg. 2001. A molecular phylogeny of the grass
This work was supported by Agencia Nacional de subfamily Panicoideae (Poaceae) shows multiple origins of
Promoción Científica y Técnica, Argentina (ANPCyT) C4 photosynthesis. American Journal of Botany 88: 1993-
grant 2418 and CONICET grant 785. We thank 2012.
Francisco Rojas and Marcelo Moreno for the excellent Gould, F. W. 1958. Chromosome numbers in southwestern
illustrations, and anonymous reviewers for helpful grasses. American Journal of Botany 45: 757-768.
comments on the manuscript. Gould, F. W. 1968. Chromosome numbers of Texas grasses.
Canadian Journal of Botany 46: 1315-1325.
Hamoud, M. A., S. A. Haroun, R. D. Macleod & A. J. Richards.
BIBLIOGRAPHY 1994. Cytological relationships of selected species of
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A Revision of Panicum Sect. Dichotomomiflora (Poaceae - Panicoideae - Paniceae)

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A Revision of Panicum Sect. Dichotomomiflora (Poaceae - Panicoideae - Paniceae)

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DARWINIANA, nueva serie 10(2): 325-364.

A REVISION OF PANICUM SECT. DICHOTOMIFLORA (POACEAE:

Keywords. Panicum sect. Dichotomiflora; Poaceae; taxonomy.

Palabras clave. Panicum sect. Dichotomiflora; Poaceae; taxonomía.

INTRODUCTION nearly 450 species distributed worldwide (Clayton

Original recibido el 23 de mayo de 2022, aceptado el 19 de julio de 2022

Sections of Panicum can be distinguished by the following key

1. Rachilla prominent between the glumes and lower lemma ...................................................................... 2

The aim of this work is to complete a worldwide HISTORY AND MORPHOLOGY

Key to the species of Panicum sect. Dichotomiflora

lower flower staminate, stamens 3, anthers 1.2- P 00740938!, MO-321061!, NY 01767173!,

Key to the subspecies 2b. Panicum dichotomiflorum Michx. subsp.

6. Panicum laevinode Lindl., Three Exped. Representative specimens examined

Spikelets lanceolate, 3-4(-4.2) × 0.9-1.1 mm, CHINA. Kwangtung. Guangdong, Pakhoi,

Plants perennial, shortly rhizomatous, 60-120 cm Representative specimens examined.

Panicum bicuspidatum A. Camus, Bull. Soc. Bot. Representative specimens examined

Ligules membranous-ciliate, 0.6-1.7 mm long; PHILIPPINES. Luzon. Baguio, Benquet,

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