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A phylogenetic evaluation of Panicum sects. Agrostoidea, Megista, Prionitia and

Tenera (Panicoideae, Poaceae): Two new genera, Stephostachys and Sorengia

Article  in  Taxon · October 2010

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TAXON 2 September 2010: 12 pp. Zuloaga & al. • Two new genera segregated from Panicum

A phylogenetic evaluation of Panicum sects. Agrostoidea, Megista,

Prionitia and Tenera (Panicoideae, Poaceae): Two new genera,
Stephostachys and Sorengia
Fernando O. Zuloaga, M. Amalia Scataglini & Osvaldo Morrone

Instituto de Botánica Darwinion, Labardén 200, Casilla de Correo 22, B1642HYD, San Isidro, Buenos Aires, Argentina
Author for correspondence: Fernando O. Zuloaga, fzuloaga@darwin.edu.ar

Abstract The taxonomic features of species of Panicum, previously classified in Panicum subg. Agrostoidea and Phanopyrum
including sects. Megista, Tenera, Prionitia and Agrostoidea, are here reviewed and compared with subg. Panicum and other
taxa of Paniceae. All taxa, previously considered in these subgenera and sections, are now included in the new genera Ste-
phostachys and Sorengia, which are based on morphological and molecular characters. Stephostachys includes one species,
S. mertensii (Roth) Zuloaga & Morrone, ranging from Mexico to Argentina, while Sorengia comprises seven species, S. anceps
(Michx.) Zuloaga & Morrone, S. caricoides (Nees ex Trin.) Zuloaga & Morrone, Sorengia longifolia (Torr.) Zuloaga & Morrone
S. petersonii (Hitchc. & Ekman) Zuloaga & Morrone, S. prionitis (Nees) Zuloaga & Morrone, S. stenodes (Griseb.) Zuloaga &
Morrone, and S. tenera (Beyr. ex Trin.) Zuloaga & Morrone, distributed from the United States to Bolivia and Brazil. Affini-
ties of the new genera with other genera of Paniceae are also discussed. Lectotypes are designated for Panicum caricoides var.
glabriusculum Döll, P. caricoides var. pilosum Döll, P. prionitis Nees, P. prionitis var. varium Kuntze, and P. stenodes Griseb.

Keywords Paniceae; Panicum; phylogeny; Poaceae; Sorengia; Stephostachys; taxonomy

Supplementary Material Appendices 2 and 3 are available in the free Electronic Supplement to the online version of this article
(http://www.ingentaconnect.com/content/iapt/tax).

INTRODUCTION
Panicum L., the most important genus of Paniceae, is a
large and taxonomically difficult group, distributed worldwide
mainly in tropical and subtropical regions (Webster, 1988).
Recent phylogenetic studies have demonstrated that the genus
is polyphyletic. In recent years, several new genera have been
recognized within Panicum s.l. in order to recognize mono-
phyletic taxa (Gómez-Martínez & Culham, 2000; Zuloaga
& al., 2000; Duvall & al., 2001, 2003; Giussani & al., 2001;
Aliscioni & al., 2003; Morrone & al., 2008a). Some of these
are completely new taxa; others were originally recognized as
subgenera. The largest of these is Panicum s.str. which corre-
sponds to Panicum subg. Panicum and includes approximately
100 species (Zuloaga & al., 2006, 2007; Morrone & al., 2007,
2008b; Sede & al., 2008, 2009b).
Among the “incertae sedis” species remaining in Pani-
cum, three sections of P. subg. Agrostoidea (Hitchc.) Zuloaga
(P. sect. Tenera (Hitchc. & Chase) Pilg., sect. Agrostoidea
(Hitchc.) C.C. Hsu, and sect. Prionitia Zuloaga), and P. sect.
Megista Pilg., were grouped together in the “ambiguous clade”
by Aliscioni & al. (2003). This clade also included dissimi-
lar genera, such as Altoparadisium Filg., Davidse, Zuloaga &
Morrone, Apochloa Zuloaga & Morrone, Arthropogon Nees,
Canastra Morrone, Zuloaga, Davidse & Filg., Cyphonanthus
Zuloaga & Morrone, Homolepis Chase, Mesosetum Steud.,
Phanopyrum (Raf.) Nash, Oncorachis Morrone & Zuloaga,
Oplismenopsis Parodi, and Tatianyx Zuloaga & Soderstr. While
most of these genera have been recently reviewed (Zuloaga
& Soderstrom, 1985; Filgueiras & al., 2001; Morrone & al.,
2001, 2007; Zuloaga & al., 2006; Sede & al., 2008, 2009a), the
classification of sections of Panicum in this clade remain to be
addressed. The purpose of this work is to establish phyloge-
netic relationships of the species in these sections, to discuss
their affinities with other genera of Paniceae and to classify
them within the tribe. In order to accomplish this, we sampled
additional species and analyzed them in conjunction with the
data published by Aliscioni & al. (2003).
MATERIALS AND METHODS
Morphological analyses. — Morphological studies were
based on herbarium specimens from ANSM, B, BAA, BAF,
BM, CEN, COL, CTES, F, G, GOET, IAN, IBE, IBGE, K, LE,
LIL, LPB, M, MEXU, MO, NY, P, PORT, R, RB, SI, SP, UB,
UMO, US, VEN, and W. All the specimens examined are listed
in Appendices 1 and 2.
Taxon sampling for molecular analysis. — A total of
138 taxa were used in the phylogenetic analysis. Of these, 14 se-
quences are new, including species of Panicum sects. Agrostoi-
dea (P. anceps), Megista (P. mertensii), Prionitia (P. prionitis)
and Tenera (P. caricoides, P. stenodes, P. tenerum). Several of
these specimens were collected in the field and dried in silica
gel. The herbarium vouchers used by Aliscioni & al. (2003) that
showed conflictive positions, such as P. stenodes, P. caricoides
and P. mertensii, were, whenever possible, re-extracted, re-
amplified and re-sequenced. Two sequences of P. caricoides

1
Zuloaga & al. • Two new genera segregated from Panicum
AY188458 and P. stenodes AY188487 obtained by Aliscioni &
al. (2003) and not confirmed in this study by a new sequence,
were excluded from the analysis. Details of the materials used
and voucher information are listed in Appendix 1.
DNA isolation, amplification and sequencing. — Total
genomic DNA was extracted using modified CTAB protocols
from Doyle & Doyle (1987) for silica samples and DNeasy
Plant Mini Kit (Qiagen) for herbarium material. NdhF ampli-
fication and sequencing were carried out using the following
pairs of primers specified by Olmstead & Sweere (1994) and
Clark & al. (1995): 5F-972R, 972F-1666R and 1666F-2110R.
For herbarium material, smaller fragments were amplified:
5F-536R and 536F-972R. PCR reactions were performed in a
25 μl final volume with 50–100 ng of template DNA, 0.2 μM
of each primer, 25 μM of DNTPs, 5 mM MgCl2, 1× buffer
and 1.5 units of Taq polymerase provided by Invitrogen. The
reaction conditions were: a first period of denaturation at 94°C
for 5 min, followed by 35 cycles of denaturation at 94°C for
30 s, annealing at 48°C for 1 min, and extension at 72°C for
1 min 30 s. Final extension at 72°C for 6 min terminated the
reactions. A negative control with no template was included
for each series of amplifications to eliminate the possibility
of contamination. PCR products were run out on a 1% TBE
agarose gel stained with ethidium bromide.
Automated sequencing was performed by Macrogen, Inc.
GenBank accession numbers are listed in Appendix 1. Se-
quence edition and assemblage were performed with BioEdit
v.5.0.9 (Hall, 1999). Alignments were manually performed
since no indels with ambiguous length were found.
The 14 sequences obtained, excluding two sequences of
Panicum sect. Tenera from Aliscioni that could not be cor-
roborated, were added to the 123-taxa Panicoid grass matrix of
Aliscioni & al. (2003). Also, Renvoizea sacciolepoidea, R. trinii
(Sede & al., 2008) and Cyphonanthus discrepans (Morrone &
al., 2007) were included in the matrix, giving a total of 138 taxa.
The aligned matrix was submitted to TreeBASE (http://
www.treebase.org) under the submission name M. Amalia
Scataglini and the reviewer’s P.I.N. code 14616.
Data analysis. — A parsimony analysis was performed
using TNT (Goloboff & al., 2008) under equal weights. Char-
acters were considered unordered, and uninformative char-
acters were excluded from the analysis. The search strategy
consisted of heuristic searches performed using 1000 series
of random addition sequences followed by TBR branch rear-
rangements and retaining 10 trees per series. The trees found
were saved in memory and, additionally, TBR swapped, re-
taining a maximum of 20,000 total trees. Branches with am-
biguous length of 0 or 1 under alternative optimizations were
collapsed, according to collapsing rule 1. A strict consensus
tree (Nelson, 1979; Rohlf, 1982) was generated from the most
parsimonious trees.
Gaps were considered as missing data, but optimization of
informative indels was performed in all the most parsimonious
trees using the command “Common Synapomorphies” of the
TNT, by which the common optimization to every individual
most parsimonious tree is represented in the consensus dia-
gram to determine whether they were synapomorphies and
2
TAXON 2 September 2010: 12 pp.
homoplasies in the clade of interest. Branch support was cal-
culated using Jackknife frequency values (Farris & al., 1996)
as implemented in TNT, with a character removal probability
of 36%, performing 10,000 replicates and a heuristic search
strategy of 10 series of addition sequences swapped with TBR,
saving two trees per series.
RESULTS
The ndhF complete data matrix consisted of 138 taxa with
2061 characters. There were 439 informative positions. The
total percentage of missing data was 1.88. Cladistic analysis
resulted in more than 20,000 most parsimonious trees, 1542
steps long with consistency index = 0.41 and retention index =
0.79. The strict consensus tree is shown in Fig. 1.
Monophyly of Paniceae was neither corroborated nor dis-
proved, species of the tribes Andropogoneae and Arundinelleae
being included in a polytomy with the x = 9 and x = 10 Paniceae
clades in a very well supported (99%) but unresolved group.
The x = 10 clade had 100% support and includes three sub-
clades with 90%, 89% and 67% support, respectively. Species
of Panicum sects. Megista, Prionitia, Agrostoidea and Tenera
appeared in the subclade that had 67% support. Several distinct
clades were identified within this subclade, but the relationships
at its base were unresolved. The specimens of Panicum mer-
tensii, the only species of P. sect. Megista, formed one of the
distinct clades with 100% support. The other taxa that were the
focus of this study, P. sects. Agrostoidea, Prionitia and Tenera,
were in a clade that had 62% support and included represen-
tatives of Phanopyrum, Canastra, and Apochloa. The other
lineages at the unresolved base of the 67% subclade involved
genera that were not the primary focus of this study (Homolepis,
Oplismenopsis, Altoparadisum-Arthropogon, Onchorachis-
Cyphonanthus, Mesosetum-Tatianyx). Within the 62% clade,
species of P. sects. Agrostoidea, Prionitia and Tenera formed a
monophyletic clade, with 99% support. Panicum sect. Prionitia
was monophyletic, with 94% support, and sister to the remain-
ing species. Panicum caricoides and P. stenodes, previously
classified in P. sect. Tenera, formed a strongly supported clade,
with a value of 98%. The previous clade is the sister group
of another clade, including P. tenerum and species formerly
placed in P. sect. Agrostoidea. Panicum tenerum, the type of
P. sect. Tenera, appeared in a monophyletic group, with 98% of
support, with P. anceps and P. longifolium, species of P. sect.
Agrostoidea.
Within the 62% clade, Panicum caricoides and P. stenodes
have a synapomorphic indel, consisting of a 15 bp long gap
(positions 1522 to 1536 of the alignment). In P. tenerum and
P. anceps this gap is 6 bp long (1522 to 1527).
DISCUSSION
As established in previous treatments (Zuloaga & al.,
2006, 2007; Morrone & al., 2007, 2008b; Sede & al., 2008,
2009b), Panicum has to be restricted to its type subgenus, i.e.,
TAXON 2 September 2010: 12 pp. Zuloaga & al. • Two new genera segregated from Panicum

Thysanolaena maxima
Danthoniopsis dinteri
Zeugites pittieri
Chasmanthium laxum subsp. sessiliflorum

Arundinelleae + Andropogoneae
100
100

x=9
87 Paniceae
99

Anthaenantia lanata
Panicum hylaeicum
90 83 Panicum pilosum
Hymenachne donacifolia
75 78 Hymenachne grumosa
88 58 Hymenachne pernambucense
98 Plagiantha tenella
Otachyrium versicolor
84
Steinchisma decipiens
97 62 Steinchisma laxa
93 Steinchisma hians
99 86 Steinchisma spathellosa
Ichnanthus pallens
Renvoizea sacciolepoidea
70 Renvoizea trinii
Streptostachys asperifolia
99 Axonopus anceps
100 Ophiochloa hydrolithica
x=10 89
Echinolaena inflexa
Ocellochloa stolonifera
Paniceae 80
54 Ocellochloa chapadense
100 99 Ocellochloa piauiense
Panicum validum
89 Anthaenantiopsis rojasiana
95 Panicum tuerckheimii
100 Hopia obtusa
55 Paspalum vaginatum
90 Paspalum conjugatum
72 Paspalum glaziovii
56 Paspalum arundinellum
Paspalum remotum
Homolepis glutinosa
Oplismenopsis najada
Altoparadisium chapadense
100 Arthropogon villosus
Cyphonanthus discrepans
100 Oncorachis ramosa
Mesosetum chaseae
Ambiguous clade 67
99 Tatianyx arnacites
Panicum mertensii
Panicummertensii Z33397
100 Panicum mertensii Z37795 Section Megista
Panicum mertensii M s/n
Phanopyrum gymnocarpon
Canastra lanceolata
Apochloa euprepes
62 99 Apochloa subtiramulosa
Panicum petersonii
80 94
Panicum prionitis Section Prionitia
Panicum prionitis M6195
Panicum caricoides B55335
Panicum caricoides M4969
Panicum stenodes G1177
99
98 Panicum stenodes F9554
Section Tenera pp.
Panicum stenodes F9540
Panicum anceps B40
99 Panicum longifolium
Panicum tenerum
Panicum tenerum M5908 Section Tenera pp.
98 Panicum tenerum P12298 +
Panicum tenerum McK772
Panicum anceps Section Agrostoidea
77 Panicum anceps K61

Fig. . Consensus tree obtained in the phylogenetic analysis. Jackknife support values are shown below branches. Collection numbers are indi-
cated for the species sequenced in the present work. The black circle indicates the optimization of the informative indel for the clade of interest.
Optimization of chromosome numbers is shown white bars for x = 9 and black ones for x = 10.

3
Zuloaga & al. • Two new genera segregated from Panicum
include only species with the following characteristics: an-
nual or perennial cespitose plants with membranous-ciliate or
ciliate ligules, open and lax inflorescences, ellipsoid to long-
ellipsoid spikelets, with the upper anthecium indurated, without
multicellular microhairs, and with simple or compound papillae
at the top of the palea; also, all species of Panicum s.str. differ
in their C4 carbon fixation type, i.e., in having the NAD-me
subtype (Zuloaga, 1987).
Panicum sect. Megista, which includes only P. mertensii,
has several unique features that distinguish it from Panicum
s.str., and other genera of the “ambiguous clade”: in P. mer-
tensii, all branches of the inflorescence are whorled, spikelets
are obovoid, globose and the upper anthecium has stomata
and papillae on its surface and is greenish at the apex. The
inflorescence type is a special feature of P. mertensii, with
branches arranged in 4–8 nearly equidistant pseudoverticils,
each pseudoverticil containing 10–60 branches. Aliscioni &
al. (2003) postulated a relationship between P. mertensii and
P. caricoides and P. stenodes, a relationship that was not sup-
ported by this analysis nor by their morphological character-
istics (see description below).
Previous research (Zuloaga & al., 2000; Giussani & al.,
2001; Aliscioni & al., 2003) postulated that Panicum subg.
Agrostoidea is polyphyletic, and that its species are unrelated to
other species of Panicum; consequently, the sections included
in this subgenus should be segregated into independent genera
or assigned to other genera of Paniceae. Subsequently, species
of P. sects. Bulbosa, Discrepantia, and Obtusa were treated
as new genera of Paniceae (Bess & al., 2005; Morrone & al.,
2007; Zuloaga & al., 2007), since their morphology did not
match existing genera.
The species of Panicum sects. Tenera, Agrostoidea and
Prionitia form a strongly supported clade. These species also
shared several morphological characteristics: cespitose, short
to long-rhizomatous perennial plants, with simple culms and
mostly basal leaves, glabrous sheaths, blades and spikelets, the
latter with the lower glumes that are1/3 to 3/4 of the spikelet,
and (1–)3–5-nerved, upper glumes and lower lemmas that are
5–9-nerved, lower palea present, and upper anthecium smooth,
shiny, and indurate. All species are Kranz, of the NADP-me
subtype, and the basic chromosome number is mostly x = 10,
except in species of Agrostoidea, P. anceps and P. longifo-
lium, whose chromosome number is x = 9. As a result of the
segregation of this new genus Sorengia and those described
earlier (Cyphonanthus, Hopia, Zuloagaea), of the 13 species
originally classified in P. subg. Agrostoidea (Zuloaga, 1987),
only P. tuerckheimii Hack. and P. validum Mez remain as
“incertae sedis” species of Panicum s.l. within the ambigu-
ous clade. Panicum tuerckheimii, a species of the monotypic
P. sect. Tuerckheimiana (Hitchc.) Zuloaga, is an endemic spe-
cies of southern Mexico, Guatemala, and Belize, with several
characteristic features, such as a scale-like, nerveless lower
glume, lower palea reduced or absent, lower flower absent,
and upper anthecium cartilaginous with the lemma margins
closed, not exposing the tip of the palea. Panicum validum, the
only species of P. sect. Valida Zuloaga & Morrone, grows in
eastern Argentina, southeastern Brazil and western Uruguay,
4
TAXON 2 September 2010: 12 pp.
in margins of rivers and streams. It is a species with glabrous
spikelets, with the lower glume 3/4 the length of the spikelet,
(1–)3-nerved, and upper anthecium with multicellular micro-
hairs at the apex (Zuloaga & al., 1989). Based on molecular
characters, P. validum and P. tuerckheimii are related to the
genus Anthaenantiopsis Mez ex Pilg. The latter genus dif-
fers by its contracted inflorescences, with spikelets pilose,
the upper anthecium gaping at maturity, its apex with long
macrohairs.
The two species of Panicum sect. Prionitia, P. petersonii
and P. prionitis, formed a subclade within the clade which
contained P. sects. Tenera and Agrostoidea (Fig. 1). They have
a disjunct distribution. Panicum prionitis is a common species
of inundated habitats from Brazil to Argentina; P. petersonii
is an endemic Cuban species; the latter was collected last, in
western Cuba, more than a hundred years ago. Two authors
(OM and FOZ) recently looked for the species in Cuba, includ-
ing looking in the type locality, but were not able to find it,
probably because there has been severe modification of the en-
vironment. Both P. prionitis and P. petersonii are robust plants,
with solid, compressed culms, sheaths and blades keeled, ter-
minal inflorescences pyramidal, lax and open, with glabrous
spikelets on long pedicels, lower palea and lower flower pres-
ent, and upper anthecium glabrous, occasionally with prickle
hairs and bicellular microhairs at the apex, smooth and shiny.
These characteristics relate species of P. sect. Prionitia with
those of P. sect. Agrostoidea. The latter differ only by being
more delicate plants, with lower flower absent, presence of
conspicuous prickle hairs on the apex of the upper lemma,
and a basic chromosome number of x = 9. Also, P. prionitis
and P. petersonii are unique within this group because of the
presence of two bundle sheaths, an external mestome and one
internal parenchymatous sheath. They are the only NADP-me
species of the tribe Paniceae exhibiting this feature (Brown,
1977; Zuloaga & al., 1989: fig. 1F).
Affinities of P. tenerum with both species of P. sect. Agros-
toidea were first reported by Aliscioni & al. (2003). Panicum
tenerum is in a strongly supported clade, together with both
species previously included in Agrostoidea, P. anceps and
P. longifolium. They are all rhizomatous plants, with spikelets
with lower flower absent, and upper anthecium with conspicu-
ous prickle hairs at the apex of the lemma. Panicum tenerum
differs by having a contracted, few-flowered panicle, and a
basic chromosome number of x = 10 (Davidse & Pohl, 1972). Its
range extends from southeastern United States to Mesoamerica
and the Caribbean.
Finally, Panicum caricoides and P. stenodes, previously
included in P. sect. Tenera, share, besides the molecular char-
acters, similar contracted inflorescences, with panicles few-
flowered, spikelets glabrous, and upper anthecium smooth,
glabrous. Both species are present from Central America and
the Caribbean to Bolivia and Brazil.
Considering both the molecular and morphological char-
acteristics of the species that were the focus of this paper, we
are proposing that the species previously included in P. sects.
Agrostoidea, Prionitia, and Tenera, be placed in one new ge-
nus, Sorengia, and that P. mertensii, the only species of P. sect.
TAXON 2 September 2010: 12 pp.
Megista, be treated as the only member of another new genus,
Stephostachys. The key below will aid in identifying the genera
that we recognize. It is followed by formal presentation of the
names by the new generic treatment.
Key to differentiate Stephostachys and Sorengia
from Panicum and related genera of the
“ambiguous clade”
1. Inflorescence a single raceme with spikelets unilaterally
arranged . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Mesosetum
1. Inflorescence paniculate, with several branches and spike-
lets not unilaterally arranged . . . . . . . . . . . . . . . . . . . . . 2
2. Spikelets with a hairy callus. . . . . . . . . . . . . . . . . . . . . . 3
2. Spikelets without a hairy callus . . . . . . . . . . . . . . . . . . . 4
3. Spikelets laterally compressed, upper palea well devel-
oped; lower glume 1/2–3/4 the length of the spikelet,
3-nerved; intercostal areas of the upper glume flat; lower
flower staminate; distinctive Kranz cells absent . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arthropogon
3. Spikelets dorsally compressed; upper palea absent or re-
duced, lower glume absent o reduced, 1-nerved; intercostal
areas of the upper glume prominently sulcate; lower flower
absent; distinctive Kranz cells present. . Altoparadisium
4. Branches of the inflorescence in several whorls . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stephostachys
4. Branches of the inflorescences alternate to opposite, not
whorled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Upper anthecium stipitate, less than half the length of the
glumes and lower lemma . . . . . . . . . . . . . . Phanopyrum
5. Upper anthecium non-stipitate, as long as the glumes and
lower lemma or slightly shorter . . . . . . . . . . . . . . . . . . . 6
6. Caryopsis with a linear hilum . . . . . . . . . . . . . . . . . . . . 7
6. Caryopsis with a punctiform, rarely oblong, hilum . . . 10
7. Spikelets densely villous; pedicel falcate at the apex . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tatianyx
7. Spikelets glabrous; pedicel truncate at the apex . . . . . . 8
8. Lower and upper glumes subulate or awned; floating
plants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oplismenopsis
8. Lower and upper glumes muticous; terrestrial plants . . 9
9. Lower and upper glumes equal in length; rachilla not thick-
ened between the lower glume and the upper anthecium
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homolepis
9. Lower glume shorter than the upper glume; rachilla thick-
ened between the lower glume and upper anthecium . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oncorachis
10. Lower and upper glume awned; upper anthecium mem-
branous; fusoid cells present . . . . . . . . . . . . . . Canastra
10. Lower an upper glume muticous; upper anthecium indu-
rate; fusoid cells absent  . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Spikelet gibbous; lower glume usually absent; upper anthe-
cium gaping at maturity, with whitish curled macrohairs
at tip and base . . . . . . . . . . . . . . . . . . . . . . Cyphonanthus
11. Spikelet not gibbous; lower glume present, 1/2–3/4 the
length of the spikelet; upper anthecium not gaping at ma-
turity, glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Leaves flat or inrolled, pungent; ligules usually absent,
Zuloaga & al. • Two new genera segregated from Panicum
membranous-ciliate when present; vascular bundles with-
out specialized chloroplasts, with 5–9 mesophyll cells be-
tween contiguous vascular bundles . . . . . . . . . Apochloa
12. Leaves flat or filiform, not pungent; ligules present, mem-
branous to ciliate; vascular bundles with specialized chlo-
roplasts, with 2–3 mesophyll cells between contiguous
vascular bundles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Upper anthecium with simple or compound papillae at the
apex of lemma and palea; parenchymatous sheath with
specialized chloroplasts . . . . . . . . . . . . . . . . . . .Panicum
13. Upper anthecium smooth, shiny, without papillae and usu-
ally with prickle hairs towards the apex of the lemma; mes-
tome sheath with specialized chloroplasts . . . .  Sorengia
TAXONOMIC TREATMENT
Stephostachys Zuloaga & Morrone, gen. nov. ≡ Panicum sect.
Megista Pilg. in Notizbl. Bot. Gard. Berlin-Dahlem 11:
243. 1931 – Type: Panicum mertensii Roth in Syst. Veg.
2: 458. 1817 (≡ Stephostachys mertensii (Roth) Zuloaga
& Morrone).
Gramina perennia, rhizomatosa, rhizomata brevia. Ligulae
membranaceae. Laminae lineari-lanceolatae vel lanceolatae.
Paniculae nodis pseudoverticillatis, spiculae obovoideae, gla-
brae. Gluma infera 1/2–3/4 quam spiculam longiora, 3–5-ner-
via. Gluma supera 7–9 nervia, lemma inferum aequans, lemma
inferum 5–9 nervium. Flos inferus praesens. Anthoecium su-
perum ellipsoideum, induratum, stomatibus et papillis simpli-
cibus apicem ornatum.
Plants perennial, rhizomatous, rhizomes short. Culms
erect, 1–3 m tall, some culms decumbent and rooting at the
lower nodes; internodes 7–20 cm long, terete, solid toward
the basal portion, glabrous, pale; nodes purplish, glabrous.
Sheaths 7–15 cm long, shorter or longer than the internodes,
papillose-pilose all over the surface, the margins membranous;
auricles present in mature plants. Ligules 1.5–4.0 mm long,
membranous, laciniate; collar glabrous. Blades linear-lanceo-
late to lanceolate, (10–)22–40 cm long, (1.0–)1.5–3.5 cm wide,
flat, the base subcordate to rounded, glabrous, lower margins
ciliate, otherwise scabrous. Inflorescence a terminal, exerted
panicle, peduncles up to 13 cm long, glabrous; panicles lax,
20–60 cm long, 9–30 cm wide; main axis terete, striate, scab-
erulous to glabrous, pulvini pubescent or glabrous; branches
arranged in 4–8 distant pseudoverticils, each pseudoverticil
with 10–60 first order ascendant branches, axis of the branches
and pedicels scaberulous, pedicels 0.7–0.8 mm long. Spikelets
obovoid, 3.2–4.0 mm long, 1.2–2.0 mm wide, glabrous, green-
ish or tinged with purple, shiny, glumes and lower lemma with
evident nerves; lower glume 1.0–1.8 mm long, ovate, acute
to obtuse, 3–5-nerved, midnerve scabrous at the apex; upper
glume as long as the spikelet, acute, 7–9-nerved; lower lemma
broadly ovate, acute, 5–7(–9)-nerved; lower palea elliptic,
2.8–3.6 mm long, 1.7–1.9 mm wide, the margins denticulate;
lower flower staminate; upper anthecium ellipsoid, 2.7–3.3 mm
long, 1.3–1.7 mm wide, smooth and shiny, whitish, acuminate
or apiculate, with stomata and papillae at the apex; caryopsis
5
Zuloaga & al. • Two new genera segregated from Panicum TAXON 2 September 2010: 12 pp.

ovoid, 1.8 mm long, 1.3 mm wide; hilum punctiform, embryo
half the length of the caryopsis.
Etymology. – The new name refers to the whorled branches
of the inflorescence.
1. Stephostachys mertensii (Roth) Zuloaga & Morrone, comb.
nov. ≡ Panicum mertensii Roth in Roem. & Schult., Syst.
Veg. 2: 458. 1817 – Type: Guyana. Essequibo, 1809,
C.H. Mertens s.n. (BAA-1889!, fragment ex B, BM!, US-
29003015!, fragment ex BM). Figure 2.
= Panicum altissimum G. Mey., Prim. Fl. Esseq.: 63. 1818, nom.
illeg. (non DC. ex Hornem., 1813) ≡ Panicum megiston
Schult., Mant. 2: 248. 1824 ≡ Panicum elatius Kunth, Re-
vis. Gramin. 1: 38. 1829, nom. superfl. et nom. illeg. non
L. f., 1781 – Type: Guyana. Hof van Holland, Rodschied
s.n. (GOET, US-2903017!, fragment ex LE).

Fig. . Stephostachys mertensii

(Roth) Zuloaga & Morrone. A,
Apical portion of the florifer-
ous culms; B, spikelet, lateral
view; C, spikelet, lower glume
view; D, spikelet, upper glume
view; E, lower palea; F, upper
anthecium. A–F based on Rojas
9234 (SI).

6
 TAXON 2 September 2010: 12 pp.
= Panicum proximum Steud., Syn. Pl. Glumac. 1: 64. 1853 – Type:
French Guiana, without date and locality, G.F.W. Meyer 22
(holotype: P!; isotype: US-2903016!, fragment ex P).
= Panicum megiston f. pauciflora Hack. in Trab. Mus. Farma-
col. 21: 30. 1909 – Type: Orillas del Rio [fluminis Pilco-
mayo] Argentina or Paraguay, T. Rojas 129 (US-80797!,
fragment ex P).
Distribution and habitat. – Widely distributed from Mex-
ico and Cuba to Paraguay, Bolivia, and northern Argentina. It
grows in or on the edges of forest, usually in inundated areas.
Chromosome number. – 2n = 40 (Davidse & Pohl, 1971),
n = 20 (Davidse & Pohl, 1978).
Common name. – “Capim-lixa” (Smith & al., 1982).
added
Notes. – Panicum equisetum Nees ex Döll in Martius, Fl.
Bras. 2 (2): 206. 1877 and Panicum latifolium var. altissimum
Rupr. in Fournier, Mexic. Pl. 2: 32, 1886, were listed as manu-
script names in the synonymy of Panicum mertensii, and so
are not validly published names.
Sorengia Zuloaga & Morrone, gen. nov. ≡ Panicum [unranked]
Agrostoidea Hitchc. & Chase in Contr. U.S. Natl. Herb.
15: 29, 99. 1910 (‘Agrostoidia’) nom. illeg (non Nash) ≡
Panicum sect. Agrostoidea C.C. Hsu in J. Fac. Sci. Univ.
Tokyo, Sect. 3, Bot. 9: 116. 1965, nom. illeg. non Nash (Art.
53.4). ≡ Panicum subg. Agrostoidea Zuloaga in Soderstrom
& al., Grass. Syst. Evol.: 292. 1987 (‘Agrostoides’), nom.
illeg. non Nash (Art. 53.4) – Type: Panicum agrostoides
Hitchc. & Chase.
= Panicum [unranked] Tenera Hitchc. & Chase in Contr.
U.S. Natl. Herb. 15: 29, 97. 1910 [in Contr. U.S. Natl. Herb.
17(6): 462, 490.1915] ≡ Panicum sect. Tenera (Hitchc. &
Chase) Pilg. in Engler, Nat. Pflanzenfam. ed. 2, 14e: 17.
1940 – Type: Panicum tenerum Beyr. ex Trin. [≡ Sorengia
tenera (Beyr. ex Trin.) Zuloaga & Morrone].
= Panicum sect. Prionitia Zuloaga in Soderstrom & al., Grass.
Syst. Evol.: 294. 1987 – Type: Panicum prionitis Nees [≡
Sorengia prionitis (Nees) Zuloaga & Morrone].
Gramina perennia, rhizomatosa. Ligulae membranaceae,
breviter ciliata. Laminae filiformes, lineare vel lineare-lan-
ceolata. Spiculae ellipsoideae vel obovoideae, glabrae. Gluma
infera 1/2–2/3 quam spiculam longiora, 1–3(–5)-nervia. Gluma
supera 5–9-nervia, lemma inferum aequans, lemma inferum
5–9 nervia. Flos inferus praesens or absens. Anthoecium su-
perum ovoideum vel ellipsoideum, induratum.
Plants perennial, cespitose, short to long rhizomatous, the
rhizomes short or long. Culms simple or branching at the mid-
dle and upper nodes, internodes terete or compressed, hollow
to solid, pilose or glabrous; nodes glabrous or pilose. Sheaths
striate, keeled, usually shorter than the internodes, villous to
glabrous. Ligules membranous, laciniate or shortly pilose at
the apex. Blades filiform, linear to linear-lanceolate, flat or
with the margins involute, glabrous or pilose. Inflorescence
exerted, panicles lax to contracted, few or multiflowered; main
axis wavy, scabrous, glabrous, branches alternate or opposite,
spreading or ascending; axillary panicles present or absent.
Spikelets narrowly ellipsoid to obovoid, glabrous, greenish
or tinged with purple; lower glume 1–3(–5)-nerved, the apex
Zuloaga & al. • Two new genera segregated from Panicum
acute, 1/2 to 2/3 the length of the spikelet; upper glume and
lower lemma subequal, usually gaping at the apices, 5–9-nerved,
the nerves scabrous; lower palea reduced or nearly as long as
the lemma, membranous to hyaline, glabrous, 2-nerved; lower
flower staminate or absent. Upper anthecium ovoid to ellipsoid,
slightly shorter than the upper glume and lower lemma, smooth,
and shiny, indurate, apex of the lemma with or without two con-
spicuous prickle hairs. Caryopsis ellipsoid, hilum punctiform,
embryo less than 1/2 the length of the caryopsis.
Distribution. – An American genus with seven species
growing from southern United States and the Caribbean to
added
South America.
Etymology. – It is a pleasure to us to name this new genus
in honor of Robert J. Soreng, a renowned agrostologist and
friend, who also helped us a lot with useful comments on this
and other contributions. The proposed change in name avoids
confusion with Agrostideae or Agrostidoideae that might arise
from adopting the sectional epithet Agrostoidea as the name
of the genus. There has been some confusion in the literature
surrounding the publication of this infrageneric name. Hitch-
cock & Chase (in Contr. U.S. Natl. Herb. 15: 29, 99. 1910)
coined the epithet Agrostoidea (as “Agrostoidia”), as part of
their “further division [of “True Panicum”] into minor groups,
the names of which are the plurals of the characteristic species
of each group.” They went on to say (p. 17) that “these names
are not intended to be formal and should have no nomencla-
tural standing.” However, the names were clearly “accepted” by
Hitchcock & Chase (cf. Art. 34.1) and under current rules (Art.
35.3) would appear to be validly published but unranked names,
capable to serving as a basionym or replaced synonym. The
name was, however, unquestionably validly published in 1915
when Hitchcock (publishing alone) did not make any mention of
formal or other use of the name, but again did not assign a rank
to it and did not mention any authorship. In publishing P. sect.
Agrostoidea in 1969, Hsu was the first to assign a definite rank
to this subdivision of Panicum. See McNeill & al. (2010) for
further details regarding this nomenclatural problem.
1. Sorengia anceps (Michx.) Zuloaga & Morrone, comb. nov. ≡
Panicum anceps Michx., Fl. Bor.-Amer. 1: 48. 1803 – Type:
United States: in herbosis humidis Carolina, Virginiae,
Georgiae, A. Michaux s.n. (P-MICHX!, US-80457!, frag-
ment and photo ex P-MICHX).
1a. Sorengia anceps subsp. anceps
= Panicum rostratum Muhl. ex Willd., Enum. Pl.: 1032. 1809
– Type: “America borealis”, Muhlenberg s.n. (B-W, PH,
US-80959!, fragment and photo ex PH).
= Agrostis nutans Poir., Encycl. Suppl. 1: 255. 1810 ≡ Vilfa
nutans (Poir.) P. Beauv., Ess. Agrostogr.: 16, 148, 181. 1812
≡ Panicum nutans (Poir.) Desv. in Mem. Soc. Agric. An-
gers 1: 197. 1831 – Type: United States. “Cette plante a été
recuille dans la Caroline,” Bosc s.n. (not seen).
= Panicum anceps var. angustum Vasey in Dept. Agric. Bot.
Div. Bull. 8: 37. 1889 – Type: United States. Texas, without
locality, 1887, G.C. Nealley s.n. (holotype: US-80528!).
= Panicum anceps var. densiflorum Vasey in Dept. Agric. Bot.
7
 Zuloaga & al. • Two new genera segregated from Panicum
Div. Bull. 8: 37. 1889 – Type: United States. Texas: Har-
rison Co., Marshall, Nov 1884, J.F. Riggs 91 (holotype:
US-80531!).
Distribution and habitat. – This subspecies grows exclu-
sively in eastern and southeastern United States, where it is
common in sandy areas, pine savannahs, and forests.
Chromosome number. – 2n = 18, 36 (Burton, 1942; Brown,
1948, 1951; Gould, 1968).
Common name. – “Beaked panicgrass” (Freckmann &
Lelong, 2003).
1b. Sorengia anceps subsp. rhizomata (Hitchc. & Chase) Zu-
loaga & Morrrone, comb. nov. ≡ Panicum rhizomatum
Hitchc. & Chase in Contr. U.S. Natl. Herb. 15: 109, f. 104.
1910 ≡ Panicum anceps var. rhizomatum (Hitchc. & Chase)
Fernald in Rhodora 36(423): 73. 1934 ≡ Panicum anceps
subsp. rhizomatum (Hitchc. & Chase) Freckmann & Le-
long in Sida 20: 171. 2002 – Type: United States. South
Carolina: Orangeburg Co.: Orangeburg, 18 Aug 1905,
A.S. Hitchcock 450 (Amer. Gr. Nat. Herb. 55) (holotype:
US-592752!; isotype: NY!).
Distribution and habitat. – Restricted to the United States,
where it is found in the states of Alabama, Arkansas, Florida,
Georgia, Louisiana, Mississippi, North Carolina, South Caro-
lina, Texas and Virginia.
Chromosome number. – 2n = 18 (Brown, 1948, sub Pani-
cum rhizomatum).
2. Sorengia caricoides (Nees ex Trin.) Zuloaga & Morrone,
comb. nov. ≡ Panicum caricoides Nees ex Trin., Gram. Pan.:
149. 1826 – Type: Brazil. Pará: Almeirim, K.F.P. von Mar-
tius s.n. (LE!, M !, US-80557!, fragment ex LE). Figure 3.
= Panicum caricoides var. pilosum Döll in Martius, Fl. Bras.
2(2): 239. 1877 – Lectotype (designated here): French
Guiana, without locality, Leprieur s.n. (P!). Syntypes:
without locality, L.C. Richard s.n. (P!); Jelski s.n. (P).
= Panicum stenodoides F.T. Hubb. in Proc. Amer. Acad. Arts
49: 497. 1913 – Type: Belize: low pine ridge, Ycacos La-
goon, 5 Mar 1907, M.E. Peck 681 (GH, K!).
Distribution and habitat. – This species grows from Mex-
ico and the Caribbean to Brazil, common in humid savannahs
on sandy soils.
Chromosome number. – 2n = 20 (Davidse, 1994).
Sorengia caricoides is related to S. stenodes, the latter
comprises more robust plants, inflorescences with 10–25 spike-
lets and spikelets without a manifest internode between the
lower and upper glume. The specimen Leprieur s.n. has been
chosen as the lectotype of P. caricoides var. pilosum because
this specimen agrees with the protologue of the variety.
Notes. – Panicum junciforme Steud., Syn. Pl. Glumac. 1:
added
82. 1854, was listed as a manuscript name in the synonymy of
Panicum caricoides, and so is not a validly published name.
3. Sorengia longifolia (Torr.) Zuloaga & Morrone, comb. nov.
≡ Panicum longifolium Torr., Fl. N. Middle United States
1: 149. 1824 – Type: United States. New Jersey, Oct 1817,
J. Goldy s.n. (NY!, US-80744!, fragment ex NY).
8
TAXON 2 September 2010: 12 pp.
3a. Sorengia longifolia subsp. abscissa (Swallen) Zuloaga &
Morrone, comb. nov. ≡ Panicum abscissum Swallen in
J. Wash. Acad. Sci. 30: 215, f. 4. 1940 ≡ Panicum rigidulum
subsp. abscissum (Swallen) Freckmann & Lelong in Sida
20: 172. 2002 – Type: United States. Florida: Highlands
Co., Sebring, in dry sand at tourist camp near lake, 3 Oct
1925, P. Weatherwax s.n. (US-1259952!).
Distribution and habitat. – Endemic to central Florida,
where it frequently grows in moist or marshy environments
in sandy soils.
Common name. – “Redtop Panicum” (Freckmann & Le-
long, 2003).
3b. Sorengia longifolia subsp. combsii (Scribner & C.R. Ball)
Zuloaga & Morrone, comb. nov. ≡ Panicum combsii
Scribner & C.R. Ball in Bull. Div. Agrostol., U.S.D.A.
24: 42, f. 16. 1901 ≡ Panicum longifolium var. comb-
sii (Scribner & Ball) Fernald in Rhodora 36: 69. 1934 ≡
Panicum rigidulum var. combsii (Scribner & C.R. Ball)
Lelong in Brittonia 36: 263. 1984 ≡ Panicum rigidulum
subsp. combsii (Scribner & C.R. Ball) Freckmann & Le-
long in Sida 20: 172. 2002 – Type: United States. Florida:
Washington Co., Chipley, 20 Aug 1898, R. Combs 583
(US-80589!).
Distribution and habitat. – Endemic to southeastern
United States, restricted to the states of Alabama, Florida, and
Georgia. It grows on dry slopes, sand or margins of ponds and
wet woods.
3c. Sorengia longifolia Bosc ex Nees subsp. elongata
(Scribn.) Zuloaga & Morrone, comb. nov. ≡ Panicum
agrostoides var. elongatum Scribn. in Bull. Agric. Exp.
Sta. Univ. Tennessee 7: 42, t. 9, f. 34. 1894 ≡ Panicum
elongatum Pursh, Fl. Amer. Sept. 1: 69. 1814, nom. illeg.
(non Salisb. 1796) ≡ Panicum stipitatum Nash, Bull. Div.
Agrostol., U.S.D.A. 17(ed. 2): 56. 1901 ≡ Panicum rigidu-
lum var. elongatum (Scribn.) Lelong in Brittonia 36: 263.
1984 ≡ Panicum rigidulum subsp. elongatum (Scribn.)
Freckmann & Lelong in Sida 20: 172–173. 2002 – Type:
United States. Delaware: without locality, F.T. Pursh s.n.
(lectotype: K, designated by Hitchcock & Chase in Contr.
U.S. Natl. Herb. 15: 104. 1910; isolectotype: US-80630!,
fragment ex K).
Distribution and habitat. – Found in mountain regions of
eastern United States.
3d. Sorengia longifolia subsp. longifolia
= Panicum anceps var. pubescens Vasey in Dept. Agric. Bot.
Div. Bull. 8: 37. 1889 ≡ Panicum longifolium var. pubes-
cens (Vasey) Fernald in Rhodora 36: 69. 1934 ≡ Panicum
rigidulum var. pubescens (Vasey) Lelong in Brittonia 36:
263. 1984 ≡ Panicum rigidulum subsp. pubescens (Vasey)
Freckmann & Lelong in Sida 20: 173. 2002 – Type: United
States. Alabama: Mobile County, Mobile, C.T. Mohr s.n.
(lectotype: US-80530!, designated by Hitchcock & Chase
in Contr. U.S. Natl. Herb. 15: 105. 1910).
= Panicum pseudanceps Nash in Bull. Torrey Bot. Club 25:
TAXON 2 September 2010: 12 pp.
85. 1898 – Type: United States. Florida. Without locality,
1889, J.H. Simpson s.n. (NY!, US-80936!, fragment and
photo ex NY).
Distribution and habitat. – Grows primarily in moist sandy
soils, pine savannahs, and bogs of coastal areas of southeastern
United States.
3e. Sorengia longifolia subsp. rigidula (Bosc ex Nees) Zuloaga
& Morrone, comb. nov. ≡ Panicum rigidulum Bosc ex
Nees in Martius, Fl. Bras. Enum. Pl. 2: 163. 1829 – Type:
United States. Without locality, Bosc s.n. (B-W!).
Zuloaga & al. • Two new genera segregated from Panicum
= Panicum agrostoides Muhl., Descr. Gram.: 119. 1817, nom.
illeg., non Spreng. (1815) – Type: not designated; original
material: “Penns. Georgia, Carolina, Cherokee”, Muhlen-
berg s.n. (not seen).
= Panicum condensum Nash in Small, Fl. S.E.U.S. 93: 1327.
1903 ≡ Panicum agrostoides var. condensum (Nash)
Fernald in Rhodora 36: 74. 1934 – Type: United States.
Florida: Ditches and swampy places in pine barrens, near
Jacksonville, 16 Oct 1895, A.H. Curtiss 5576 (NY!, US-
742212!, US-80599!, US-742213!).
= Panicum agrostoides Hitchc. & Chase in Contr. U.S. Natl.
Fig. . Sorengia caricoides
(Nees ex Trin.) Zuloaga & Mor-
rone. A, Habit; B, portion of the
leaf in the ligular area; C, apical
portion of the floriferous culms;
D, spikelet, lower glume view;
E, spikelet, upper glume view;
F, upper anthecium, upper lem-
ma view; G, upper anthecium,
upper palea view; H, caryopsis,
embryo side; I, caryopsis, hilum
side. A–I based on Díaz &
Pereira 8981 (SI).
9
 Zuloaga & al. • Two new genera segregated from Panicum
Herb. 15: 100. 1910, nom. illeg., non Spreng. (1815), nom.
superfl. P. rigidulum Bosc ex Nees incl. – Type: “e Penn-
sylvannia,” Muhlenberg s.n. (B-W).
Distribution and habitat. – Southern Canada, the United
States, Mexico, Mesoamerica (Guatemala), and the Caribbean.
It is found in swamps, wet pine savannahs, wet woodlands and
margins of rivers, ponds and lakes.
Chromosome number. – 2n = 18 (Brown, 1948, sub. Pani-
cum agrostoides).
For a detailed analysis of the nomenclatural problems as-
updated by us
sociated with the name P. agrostoides see McNeill & al. (Taxon
……. 2010).
4. Sorengia petersonii (Hitchc. & Ekman) Zuloaga & Mor-
rone, comb. nov. ≡ Panicum petersonii Hitchc. & Ekman
in Hitchcock, Man. Grasses W. Ind. 243: 263, f. 220. 1936
– Type: Cuba. Pinar del Río: Mendoza, collected in tem-
porarily flooded places, 16 Nov 1923, E.L. Ekman 18085
(LIL!, MO!, NY!, US- 1295015!, 1296166!, 1502196!).
Distribution and habitat.– Endemic to Cuba, where it has
been sporadically collected in flooded areas of Pinar del Río.
added
Notes. – Sorengia petersonii is closely related to S. prio-
nitis, a South American species, the latter differing by having
larger spikelets.
5. Sorengia prionitis (Nees) Zuloaga & Morrone, comb. nov.
≡ Panicum prionitis Nees in Martius, Fl. Bras. Enum. Pl.:
162. 1829 – Lectotype (designated here): Brazil. Minas
Gerais: “Adamantum pr. Tejuco V. do principi.,” K.F.P. von
Martius s.n. (M!; isolectotypes: B!, BAA!, fragment ex B,
US-80930!, fragment ex M). Syntype: Uruguay. Montevi-
deo: Montevideo, F. Sellow s.n. (BAA!, MO!, US!).
= Coleataenia gynerioides Griseb., Symb. Fl. Arg. (in Abh.
Königl. Ges. Wiss. Göttingen 24): 308. 1879 ≡ Panicum gy-
nerioides (Griseb.) Pilg. in Notizbl. Bot. Gart. Berlin 104:
243. 1931 ≡ Panicum prionitis subsp. gynerioides (Griseb.)
Roseng. & al., Gram. Urug.: 341. 1970 – Type: Argentina.
Entre Ríos: Concepción del Uruguay, G.P. Lorentz s.n.
(BAA!, BAF!, GOET).
= Panicum prionitis var. pallidum Kuntze, Revis. Gen. Pl. 3(3):
363. 1898 – Type: Paraguay. Río Tebicuary, Sep 1893,
O. Kuntze s.n. (NY!).
= Panicum prionitis var. varium Kuntze, Revis. Gen. Pl. 3(3):
363. 1898 – Lectotype (designated here): Paraguay. Con-
cepción: Concepción, Sep 1892, O. Kuntze s.n. (NY!).
Syntype: Paraguay: “SudParaguay, IX-92,” O. Kuntze
s.n. (NY!).
Distribution and habitat. – From southern Brazil to Para-
guay, Uruguay and Argentina. It is frequent in lowlands and
margins of rivers and streams. Flowering from October to
January.
Common names. – “Paja brava,” “paja de techar” (Burkart,
1969); “capim-Santa-Fe” (Smith & al., 1982).
Chromosome Number. – 2n = 20 (Parodi, 1946; Núñez,
1952); 2n = 20, 40 (Bouton & al., 1981).
The specimen Martius s.n., from M, has been chosen as
the lectotype of P. prionitis because this specimen agrees
10
TAXON 2 September 2010: 12 pp.
completely with the protologue of the species. Similarly, the
specimen Kuntze s.n., from Concepción, Paraguay, has been
designated as the lectotype of Panicum prionitis var. varium
because it agrees with the protologue of the variety.
6. Sorengia stenodes (Griseb.) Zuloaga & Morrone, comb.
nov. ≡ Panicum stenodes Griseb., Fl. Brit. W. Ind.: 547.
1864 – Lectotype (designated here): Cuba: without locality,
C. Wright 792 (GOET!; isolectotype, US-81132! fragment
ex GOET and photo). Syntype: Jamaica: in savannahs,
Manchester, Purdie s.n. (K!, US! fragment ex K).
= Panicum caricoides Nees ex Trin. var. glabriusculum Döll
in Martius, Fl. Bras. 2(2): 239. 1877 – Lectotype (desig-
nated here): Brazil. Amazonas: Manaos, R. Spruce 39 (M!;
isolectotypes: G!, K!, P!, US-2903360!, fragment ex M).
Syntype: Brazil. Minas Gerais, J.E.B. Warming s.n. (C,
not seen).
Distribution and habitat.– From Mesoamerica and the
Caribbean to Bolivia and Brazil; it grows in inundated savan-
nahs.
The specimen Wright 792, from GOET, has been chosen
as the lectotype of P. stenodes because it agrees with the pro-
tologue of the species. Also, the specimen Spruce 39, from
M, has been designated as lectotype of P. caricoides var. gla-
briusculum since it agrees with the protologue of the variety.
7. Sorengia tenera (Beyr. ex Trin.) Zuloaga & Morrone, comb.
nov. ≡ Panicum tenerum Beyr. ex Trin. in Mem. Acad.
Imp. Sci. Saint-Petersbourg, Ser. 6, Sci. Math., Seconde Pt.
Sci. Nat. 3, 1(2–3): 341. 1834 – Type: United States. Geor-
gia: Georgia, in paludosis nemorum, 1834, H.C. Beyrich
62 (LE-TRIN-0978.01!, US-81935! fragment and photo
ex LE).
= Panicum anceps var. strictum Chapm., Fl. South. U.S.: 573.
1860 – Type: United States. Florida, without locality,
A.W. Chapman s.n. (K!, NY, US-80456!, fragment and
photo ex NY).
Distribution and habitat. – From southern United States
to Mexico, Mesoamerica and the Caribbean. It grows in open
and humid habitats.
Chromosome number. – n = 20 (Davidse & Pohl, 1972).
Common name. – “Blue-joint panicgrass” (Freckmann &
Lelong, 2003).
ACKNOWLEDGEMENTS
This study was supported in part by ANPCyT (Agencia Nacional
de Promoción Científica y Técnica, Argentina), grants 13374, 32664
and 1286 by CONICET, grant 5453 and by the National Geographic So-
ciety, grant 7792–05. We are grateful to Raúl Pozner for his assistance
with the Latin descriptions, and to Vladimiro Dudás for preparing
the illustrations. The authors sincerely thank Mary Endress, Robert
J. Soreng and two anonymous reviewers for comments that greatly
strengthened the manuscript. Finally, we are particular grateful to
John McNeill for very important and useful nomenclatural comments
which also strengthened the manuscript.
TAXON 2 September 2010: 12 pp.
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Appendix . List of taxa used in the ndhF analysis and GenBank accession numbers. Specimens sequenced for this paper are indicated with an asterisk.
Tribe Andropogoneae. Andropogon gerardii Vitman, AF117391 (Spangler & al. 1999). Bothriochloa bladhii (Retz.) S.T. Blake, AF117395 (Spangler & al. 1999).
Chionachne koenigii (Spreng.) Thwaites, AF117397 (Spangler & al. 1999). Cymbopogon flexuosus (Nees ex Steud.) Will. Watson, AF117404 (Spangler & al. 1999).
Phacelurus digitatus (Sibth. & Sm.) Griseb., AF117418 (Spangler & al. 1999). Sorghum bicolor (L.) Moench, U21981 (Clark & al. 1995). Zea mays L., acc. no.?
(Clark & al. 1995). Tribe Arundinelleae. Arundinella hirta (Thunb.) Tanaka, AF117393 (Spangler & al. 1999). Tribe Centotheceae. Chasmanthium laxum (L.)
H.O. Yates subsp. sessiliflorum (Poir.) L.G. Clark, U27296 (Clark & al. 1995). Zeugites pittieri Hack., U21987 (Clark & al. 1995). “Insertae sedis” tribe. Dan-
thoniopsis dinteri (Pilg.) C.E. Hubb., AYO29695 (Giussani & al. 2001). Tribe Paniceae, excluding Panicum. Acroceras zizanioides (Kunth) Dandy, AY029618
(Giussani & al. 2001). Altoparadisium chapadense Filg. & al., AY029619 (Giussani & al. 2001). Anthaenantia lanata (Kunth) Benth., AY029640 (Giussani & al.
2001). Anthaenantiopsis rojasiana Parodi, AY029620 (Giussani & al. 2001). Apochloa euprepes (Renvoize) Zuloaga & Morrone, AY029657 (Giussani & al. 2001).
A. subtiramulosa (Renvoize & Zuloaga) Zuloaga & Morrone, AY188490 (Aliscioni & al. 2003). Arthropogon villosus Nees, AY029622 (Giussani & al. 2001).
Axonopus anceps (Mez) Hitchc., AY029623 (Giussani & al. 2001). Canastra lanceolata (Filg.) Morrone & al., AY029621 (Giussani & al. 2001). Cenchrus ciliaris
L., AY029625 (Giussani & al. 2001). Chaetium bromoides (J. Presl) Benth. ex Hemsl., AY029626 (Giussani & al. 2001). Cyphonanthus discrepans (Döll) Morrone
& Zuloaga, DQ646392 (Morrone & al. 2007). Dichanthelium acuminatum (Sw.) Gould & Clark, AY188485 (Aliscioni & al. 2003). D. clandestinum (L.) Gould,
AY188461 (Aliscioni & al. 2003). D. cumbucana (Renvoize) Zuloaga, AY188464 (Aliscioni & al. 2003). D. koolauense (H. St. John & Hosaka) C.A. Clark & Gould,
AY029627 (Giussani & al. 2001). D. sabulorum (Lam.) Gould & C.A. Clark, AY029654 (Giussani & al. 2001). Digitaria setigera Roth ex Roem. & Schult.,
AY029629 (Giussani & al. 2001). Echinochloa colona (L.) Link, AY029631 (Giussani & al. 2001). Echinolaena inflexa (Poir.) Chase, AY029633 (Giussani & al.
2001). Eriochloa punctata (L.) Desv., AY029634 (Giussani & al. 2001). Homolepis glutinosa (Sw.) Zuloaga & Soderstr., AY029637 (Giussani & al. 2001). Hopia
obtusa (Kunth) Zuloaga & Morrone, AY029659 (Giussani & al. 2001). Hymenachne donacifolia (Raddi) Chase, AY029635 (Giussani & al. 2001). H. grumosa
(Nees) Zuloaga, AY188468 (Aliscioni & al. 2003). H. pernambucense (Spreng.) Zuloaga, AY188478 (Aliscioni & al. 2003). Ichnanthus pallens (Sw.) Munro ex
Benth., AY029638 (Giussani & al. 2001). Lasiacis sorghoidea (Desv.) Hitchc. & Chase, AY029639 (Giussani & al. 2001). Megathyrsus maximus (Jacq.) B.K. Simon
& S.W.L. Jacobs, AY029649 (Giussani & al. 2001). Melinis repens (Willd.) Zizka, AY029675 (Giussani & al. 2001). Mesosetum chaseae Luces, AY029641 (Gius-
sani & al. 2001). Moorochloa eruciformis (Sm.) Veldkamp, AY188452 (Aliscioni & al. 2003). Ocellochloa chapadense (Swallen) Zuloaga & Morrone, AY188486
(Aliscioni & al. 2003). O. piauiense (Swallen) Zuloaga & Morrone, AY029656 (Giussani & al. 2001). O. stolonifera (Poir.) Zuloaga & Morrone, AY188488 (Alis-
cioni & al. 2003). Oncorachis ramosa (Zuloaga & Soderstr.) Morrone & Zuloaga, AY029686 (Giussani & al. 2001). Ophiochloa hydrolithica Filg. & al., AY029642
(Giussani & al. 2001). Oplismenopsis najada (Hack. & Arechav.) Parodi, AY188453 (Aliscioni & al. 2003). Oplismenus hirtellus (L.) P. Beauv., AY029644 (Gius-
sani & al. 2001). Otachyrium versicolor (Döll) Henrard, AY029643 (Giussani & al. 2001). Parodiophyllochloa cordovensis (E. Fourn.) Zuloaga & Morrone,
AY188463 (Aliscioni & al. 2003). P. missiona (Ekman) Zuloaga & Morrone, AY188473 (Aliscioni & al. 2003). P. ovulifera (Trin.) Zuloaga & Morrone, AY029653
(Giussani & al. 2001). P. penicillata (Nees ex Trin.) Zuloaga & Morrone, AY188477 (Aliscioni & al. 2003). Paspalidium geminatum (Forssk.) Stapf, AY029662
(Giussani & al. 2001). Paspalum arundinellum Mez, AY029663 (Giussani & al. 2001). P. conjugatum Bergius, AY029669 (Giussani & al. 2001). P. glaziovii (A.G.
Burm.) S. Denham, AY029689 (Giussani & al. 2001). P. remotum J. Remy, AY029668 (Giussani & al. 2001). P. vaginatum Sw., AY029665 (Giussani & al. 2001).
Pennisetum alopecuroides (L.) Spreng., AY029672 (Giussani & al. 2001). P. setaceum (Forssk.) Chiov. AY029673 (Giussani & al. 2001). Phanopyrum gymnocar-
pon (Elliott) Nash, AY188469 (Aliscioni & al. 2003). Plagiantha tenella Renvoize, AY029674 (Giussani & al. 2001). Pseudechinolaena polystachya (Kunth) Stapf,
AY029676 (Giussani & al. 2001). Renvoizea sacciolepoidea (Renvoize & Zuloaga) Zuloaga & Morrone, EU107787 (Sede & al. 2008). R. trinii (Kunth) Zuloaga &
Morrone, EU107781 (Sede & al. 2008). Rupichloa acuminata (Renvoize) Salariato & Morrone AY029629 (Giussani & al. 2001). Sacciolepis indica (L.) Chase,
AY029677 (Giussani & al. 2001). Setaria lachnea (Nees) Kunth, AY029683 (Giussani & al. 2001). S. macrostachya Kunth, AY029678 (Giussani & al. 2001). S.
palmifolia (J. König) Stapf, AY029680 (Giussani & al. 2001). S. viridis (L.) Beauv., U21976 (Clark & al. 1995). Steinchisma decipiens (Nees ex Trin.) W.V. Br.,
AY188499 (Aliscioni & al. 2003). S. hians (Elliot) Nash, AY029685 (Giussani & al. 2001). S. laxa (Sw.) Zuloaga, AY029655 (Giussani & al. 2001). S. spathellosa
(Döll) Renvoize, AY188500 (Aliscioni & al. 2003). Stenotaphrum secundatum (Walter) Kuntze, AY029684 (Giussani & al. 2001). Streptostachys asperifolia Desv.,
AY029687(Giussani & al. 2001). Tatianyx arnacites (Trin.) Zuloaga & Soderstr., AY029688 (Giussani & al. 2001). U. plantaginea (Link) R.D. Webster, AY029693
(Giussani & al. 2001). Zuloagaea bulbosa (Kunth) Bess, AY029648 (Giussani & al. 2001). Panicum sect. Dichotomiflora (Hitchc.) Honda. Panicum aquaticum
Poir., AY029658 (Giussani & al. 2001). P. dichotomiflorum Michx., AY188466 (Aliscioni & al. 2003). P. elephantipes Nees ex Trin., AY029647 (Giussani & al.
2001). P. gouinii E. Fourn., AY188467 (Aliscioni & al. 2003). P. pedersenii Zuloaga, AY029646 (Giussani & al. 2001). P. repens L., AY029651(Giussani & al.
2001). Panicum sect. Panicum. P. bergii Arechav., AY188457 (Aliscioni & al. 2003). P. fauriei Hitchc., AY029650 (Giussani & al. 2001). P. miliaceum L., AY188472
(Aliscioni & al. 2003). P. nephelophilum Gaudich., AY029645 (Giussani & al. 2001). P. stramineum Hitchc. & Chase, AY188489 (Aliscioni & al. 2003). Panicum
sect. Rudgeana (Hitchc.) Zuloaga. P. cervicatum Chase, AY188459 (Aliscioni & al. 2003). P. rudgei Roem. & Schult., AY029661 (Giussani & al. 2001). Panicum
sect. Urvilleana (Hitchc.) Pilg. P. chloroleucum Griseb., AY188460 (Aliscioni & al. 2003). P. racemosum (P. Beauv.) Spreng., AY188481 (Aliscioni & al. 2003).
Panicum sect. Virgata (Hitchc. & Chase) Pilg. P. tricholaenoides Steud., AY188493 (Aliscioni & al. 2003). P. virgatum L., U21986 (Clark & al. 1995). Ungrouped
species of Panicum. P. mistasypus Zuloaga & Morrone, AY188474 (Aliscioni & al. 2003). P. olyroides Kunth, AY188475 (Aliscioni & al. 2003). Panicum “in-
certae sedis”: P. antidotale Retz., AY188456 (Aliscioni & al. 2003). P. hylaeicum Mez, AY188470 (Aliscioni & al. 2003). P. pilosum Sw., AY188480 (Aliscioni &
al. 2003). sect. Agrostoidea (Hitchc.) C.C. Hsu. P. anceps Michx., AY188455 (Aliscioni & al. 2003). *P. anceps Baker 40, GU253324. *P. anceps Kearney Jr. 61,
GU253325. P. rigidulum Bosc ex Nees, AY188482 (Aliscioni & al. 2003). Sect. Clavelligera Stapf. P. adenophorum K. Schum., AY188454 (Aliscioni & al. 2003).
P. claytonii Renvoize, AY188462 (Aliscioni & al. 2003). Sect. Megista Pilg. P. mertensii Roth, AY188471 (Aliscioni & al. 2003). *P. mertensii Zardini 33397,
GU253320. * P. mertensii Zardini 37795, GU253321. *P. mertensii Morrone s/n, GU253322. Sect. Monticola Stapf. P. millegrana Poir., AY029660 (Giussani &
al. 2001). P. sellowii Nees, AY188484 (Aliscioni & al. 2003). Sect. Parvifolia (Hitchc. & Chase) Pilg. P. cyanescens Nees ex Trin., AY188465 (Aliscioni & al.
2003). P. parvifolium Lam., AY188476 (Aliscioni & al. 2003). P. schwackeanum Mez, AY188483 (Aliscioni & al. 2003). P. trichanthum Nees, AY188492 (Aliscioni
& al. 2003). P. wettsteinii Hack., AY188497 (Aliscioni & al. 2003). Sect. Prionitia Zuloaga. P. petersonii Hitchc. & Ekman, AY188479 (Aliscioni & al. 2003). P.
prionitis Nees, AY029652 (Giussani & al. 2001). *P. prionitis Morrone 6195, GU253323. Sect. Tenera (Hitchc. & Chase) Pilg. *P. caricoides Nees ex Trin. Mor-
rone 4969, GU253330. *P. caricoides Breedlove 55335, GU253329. *P. stenodes Griseb. Gutiérrez 1177, GU253331. *P. stenodes Zuloaga 9540, GU253333. *P.
stenodes Zuloaga 9554, GU253332. P. tenerum Beyr. ex Trin., AY188491 (Aliscioni & al. 2003). *P. tenerum Pohl 12298, GU253327. *P. tenerum McKenzie 772,
GU253328. *P. tenerum Morrone 5908, GU253326. Sect. Tuerckheimiana (Hitchc.) Zuloaga. P. tuerckheimii Hack., AY188494 (Aliscioni & al. 2003). Sect.
Valida Zuloaga & Morrone. P. validum Mez, AY188495 (Aliscioni & al. 2003). Sect. Verrucosa (Hitchc.) C.C. Hsu. P. verrucosum Muhl., AY188496 (Aliscioni
& al. 2003). Tribe Thysanolaenae. Thysanolaena maxima (Roxb.) Kuntze, U21984 (Clark & al. 1995).

12
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