Phylogeny of The Paniceae (Poaceae, Panicoideae) Integrating Plastid DNA Sequences and Morphology Into A New Classification

Cladistics
Cladistics (2011) 1–24

10.1111/j.1096-0031.2011.00384.x
Phylogeny of the Paniceae (Poaceae: Panicoideae): integrating plastid

DNA sequences and morphology into a new classification
Osvaldo Morronea, Lone Aagesena,*, Maria A. Scataglinia, Diego L. Salariatoa,
Silvia S. Denhama, Maria A. Chemisquya, Silvana M. Sedea, Liliana M. Giussania,
Elizabeth A. Kelloggb and Fernando O. Zuloagaa
a
Instituto de Botánica Darwinion, Labardén 200, San Isidro, B1642HYD, Buenos Aires, Argentina; bUniversity of Missouri-St Louis,
One University Boulevard, St Louis, MO 63121, USA
Accepted 12 October 2011
Abstract
Included in the PACMAD clade of the family Poaceae (Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae, Aristidoideae,
Danthonioideae), the tribe Paniceae s.l. is one of the largest tribes of the subfamily Panicoideae, with more than 2000 species. This tribe
comprises a huge morphological, cytological and physiological diversity represented by different inflorescence types, several basic
chromosome numbers, and at least four major photosynthetic pathways. The tribe Paniceae has been the subject of molecular studies
that have confirmed its paraphyly: two major clades were recognized based on their basic chromosome numbers (x = 9, x = 10). The
x = 10 Paniceae clade is sister to the Andropogoneae–Arundinelleae s.s. clade (x = 10), while the combined x = 10 clade is sister to
the x = 9 clade that contains the remaining genera of Paniceae. As a result of a recent realignment within the tribe in terms of the
phylogenetic position of minor and major Paniceae genera, a reanalysis of the whole sampling is performed and new underrepresented
taxa are discussed. A total of 155 genera, currently considered within subfamily Panicoideae, are represented here by almost all genera
of Paniceae s.l., representatives of Andropogoneae and Arundinelleae s.s., and the endemic and small tribe Steyermarkochloeae; we
also included specimens of subfamily Micrairoideae, tribes Isachneae and Eriachneae. The sampling includes as outgroups 18 genera of
the PACMAD clade (excluding Panicoideae) and four genera from the BEP clade (Bambusoideae, Ehrhartoideae, Pooideae), rooting
with Bromus inermis. A matrix with 265 taxa based on the combined evidence from ndhF plastid sequences (2074 bp) and 57
morphological characters was subjected to parsimony analyses. Jackknife resampling was used to calculate group support. Most clades
are characterized by morphological, cytological, anatomical, and ⁄ or physiological characters. Major tribal changes are based on the
basic chromosome number; the pantropical x = 9 clade is here recognized as Paniceae s.s., while the American x = 10 Paniceae s.l. is
restricted to the reinstated tribe Paspaleae. The optimization of the photosynthetic pathway for the Paspaleae–Andropogoneae–
Arundinelleae s.s. clade, including the monotypic Reynaudia, shows a plesiomorphic C4 state while the ancestral state for Paniceae s.s. is
ambiguous. If Reynaudia were not included or placed elsewhere, the ancestral photosynthetic pathway for both the Paspaleae–
Andropogoneae–Arundinelleae s.s. clade and the Paniceae s.s. would be unambiguously C3. In order to explore character evolution
further, the morphological characters were mapped onto one of the most parsimonious trees. A relationship between photosynthetic
pathways and inflorescence morphology is suggested here for the first time. Based on the optimization of morphological characters and
additional data, we propose names for almost all inner clades at the rank of subtribe with a few groups as incertae sedis. With this
extensive sampling, we resolved the phylogenetic relationships and the assignation of synapomorphies, and improved the support in
subtribe sorting; consequently a robust circumscription of the tribe Paniceae s.l. is proposed.
Paniceae is one of the largest tribes of subfamily

Panicoideae (Poaceae), in which more than 2000
species are included. The Paniceae are mainly distributed
*Corresponding author:
E-mail address: laagesen@darwin.edu.ar in tropical and subtropical regions of both hemispheres,
Ó The Willi Hennig Society 2011

2 O. Morrone et al. / Cladistics (2011) 1–24
with only a few representatives in temperate and cold GPWG (2001) considered 3270 species grouped in 206
areas. genera and six tribes in the subfamily: Andropogoneae,
This tribe includes enormous morphological, cytolog- Arundinelleae, Hubbardieae, Isachneae, Paniceae and
ical and physiological diversity, represented by different Steyermarkochloeae (GPWG, 2001). Soreng and Davis
developmental patterns of spikelets, inflorescence types, (1998) suggested that tribes Centotheceae and Thysa-
several basic chromosome numbers, leaf anatomy, and nolaeneae should be included in subfamily Panicoideae.
at least four major photosynthetic pathways [Clayton Duvall et al. (2007), using two plastid markers (rbcL
and Renvoize, 1986; Watson and Dallwitz, 1992; and ndhF), proposed that Isachne R. Br. and Eriachne R.
GPWG (Grass Phylogeny Working Group), 2001]. Br. should be excluded from Panicoideae as both genera
Cereal and forage grasses [Panicum miliaceum L., are closely related to Micraira F. Muell. The three
Cenchrus americanus (L.) Morrone, Setaria italica (L.) genera were placed in subfamily Micrairoideae, which
P. Beauv., Paspalum dilatatum Poir.] and turf and was later emended by Sánchez-Ken et al. (2007) with the
ornamental species [Axonopus compressus (Sw.) P. addition of other genera of tribes Isachneae and
Beauv., Cenchrus purpureus (Schumach.) Morrone, Eriachneae. Recently, Sánchez-Ken and Clark (2010)
Stenotaphrum secundatum (Walter) Kuntze], as well as proposed an expanded classification for the Panicoideae
species used for biofuel (Panicum virgatum L.), make the with 12 tribes: Andropogoneae, Arundinelleae, Cento-
tribe Paniceae important in the economy of the human theceae, Chasmanthieae (including Bromuniola Stapf
community. Improvement of crops, developmental and C.E. Hubb. and Chasmanthium Link), Gynerieae,
studies, genome sequencing, and evolutionary studies Cyperochloeae (proposed to include Spartochloa C.E.
have been the subject of major research projects Hubb. and Cyperochloa Lazarides and L. Watson.),
involving panicoid species in recent years (e.g. McLaren, Hubbardieae, Paniceae, Steyermarkochloeae, Thysano-
2005; McLaughlin and Kszos, 2005; Reinheimer et al., laeneae, Tristachyideae (a new tribe with genera previ-
2005; Buell, 2009; Hua et al., 2009). ously included in the Arundinelleae) and Zeugiteae.
DNA sequencing has transformed traditional taxon- Tribes Paniceae and Andropogoneae have the great-
omy and has led to a robust new systematic classifica- est number of species and genera within the subfamily
tion for the family Poaceae (Barker et al., 1995; Clark Panicoideae. The tribe Paniceae has been the subject of
et al., 1995; Duvall and Morton, 1996; Soreng and molecular studies in the past decade that have confirmed
Davis, 1998; Hilu et al., 1999; GPWG, 2001). The most its paraphyly (Gómez-Martı́nez and Culham, 2000;
recent phylogeny of the entire grass family (GPWG, Duvall et al., 2001; Giussani et al., 2001; Mathews
2001), based on morphology and seven molecular data et al., 2002). The former Paniceae has been divided into
sets, divided Poaceae in two major clades: BEP (Bam- two major clades that differ in basic chromosome
busoideae, Ehrhartoideae, Pooideae) and PACMAD number (x = 9, x = 10); the x = 10 Paniceae clade is
(Panicoideae, Arundinoideae, Chloridoideae, Micrairoi- sister to the Andropogoneae–Arundinelleae clade (also
deae, Aristidoideae, Danthonioideae); the latter acronym x = 10) while the combined x = 10 clade is sister to the
as accepted by Duvall et al. (2007) and Sánchez-Ken x = 9 clade (Gómez-Martı́nez and Culham, 2000;
et al. (2007). These results opened up a new range of Giussani et al., 2001; Aliscioni et al., 2003). Increased
evolutionary hypotheses among subfamilies, tribes and taxonomic sampling in subsequent phylogenetic studies
genera of grasses, and represent a challenge to reassess within Paniceae allowed the recircumscription of minor
the classification within each of the clades. In this lineages. Panicum L. has proven to be a polyphyletic
regard, molecular phylogenetic studies re-evaluated the genus and was consequently restricted to C4 species with
traditional classification of major groups: subfamily an NAD–malic enzyme (NAD–ME) physiological and
Bambusoideae by Kelchner and Clark (1997), Zhang anatomical subtype (Aliscioni et al., 2003). As a result,
and Clark (2000), and Triplett and Clark (2010); several species of Panicum were segregated and new
subfamily Pooideae by Catalán et al. (1997), Davis genera were established: Apochloa Zuloaga and Mor-
and Soreng (2007), Döring et al. (2007), Soreng et al. rone, Canastra Morrone, Zuloaga, Davidse and Filg.,
(2007), and Schneider et al. (2009); subfamily Chlori- Coleataenia Griseb., Cyphonanthus Zuloaga and Mor-
doideae by Peterson et al. (2010, 2011); subfamily rone, Hopia Zuloaga and Morrone, Megathyrsus (Pilg.)
Panicoideae by Giussani et al. (2001), Aliscioni et al. B.K. Simon and S.W.L. Jacobs, Ocellochloa Zuloaga
(2003), and Sánchez-Ken et al. (2007). and Morrone, Parodiophyllochloa Zuloaga and Mor-
Subfamily Panicoideae has been consistently highly rone, Renvoizea Zuloaga and Morrone, Stephostachys
supported as monophyletic (GPWG, 2001; Vicentini Zuloaga and Morrone, and Zuloagaea Bess (Morrone
et al., 2008; Sánchez-Ken and Clark, 2010), and several et al., 2001, 2007, 2008; Simon and Jacobs, 2003; Bess
morphological synapomorphies characterize the group: et al., 2006; Zuloaga et al., 2007a; Sede et al., 2008,
the presence of a two-flowered spikelet, with the lower 2009a; Soreng, 2010).
floret staminate or neuter and the upper one perfect, and Because of the extensive recent realignment of genera
with simple starch grains (Kellogg and Campbell, 1987). within the Paniceae (Giussani et al., 2001; Aliscioni
O. Morrone et al. / Cladistics (2011) 1–24 3
et al., 2003; Donadı́o et al., 2009; Ibrahim et al., 2009; involucre of bristles below the spikelets, with four
Kellogg et al., 2009; Salariato et al., 2009, 2010; Sede genera: Cenchrus L., Pennisetum Rich., Plagiosetum
et al., 2009b; Chemisquy et al., 2010), a review of the Benth., and Paratheria Griseb.; (iii) ‘‘Chamaeraphis’’
whole tribe is needed, including a comprehensive sample with a fruiting glume less hardened, including seven
representing its morphological, anatomical and cytoge- small genera: Echinolaena Desv., Chamaeraphis R. Br.,
netic diversity. Stenotaphrum Trin., Spartina Schreb. (currently in
The aim of this work is to establish a robust Chloridoideae), Phyllorachis Trimen (now in Ehrhartoi-
circumscription of the tribe Paniceae and the major deae), Thuarea Pers., and Xerochloa R. Br.; and (iv) a
clades within it. In order to accomplish this objective, all heterogeneous group of seven genera, of which only
genera of the tribe were sampled. Relationships among Spinifex L. is still considered within the tribe. The other
genera are discussed based on the ndhF plastid gene, in six genera placed in Paniceae by Bentham (1881) were:
addition to cytological and morphological evidence. To Anomochloa Brongn. (now Anomochlooideae), Lepta-
resolve the monophyly of Paniceae we included, other spis R. Br. (Pharoideae), Lygeum Loefl. ex L. (Pooi-
than the well represented genera of Paniceae, Andro- deae), Olyra L. (Bambusoideae), Pharus P. Browne
pogoneae and Arundinelleae, ndhF sequences of the (Pharoideae), and Streptochaeta Schrad. ex Nees
endemic and small tribe Steyermarkochloeae of the (Anomochlooideae). The remaining tribes of the Panic-
subfamily Panicoideae, as well as representatives of aceae proposed by Bentham represent an artificial
subfamily Micrairoideae tribes Isachneae and Eriach- grouping with some representatives of the panicoid
neae, among others. Based on a combined morpholog- tribes Paniceae and Andropogoneae, but also from
ical and molecular analysis, a new classification for the other subfamilies such as Chloridoideae and Bambusoi-
tribe Paniceae is presented here. deae. Tribe Zoysieae included some panicoid genera
such as Anthephora Schreb., Neurachne R. Br., and
Taxonomic history of the tribe Paniceae Trachys Pers., and also several Chloridoideae genera;
tribe Maydeae mostly included andropogonoid genera,
In the middle of the 17th century, Adanson (1763) together with Pariana Aubl., a little herbaceous bam-
divided the grass family into nine informal groups, of busoid grass; tribe Oryzeae included Achlaena Griseb.,
which the two major ones, the Poae and Panica, Beckera Fresen. (= Snowdenia C.E. Hubb.), and a
currently correspond, in a nomenclatural sense, to the heterogeneous group with Hydrochloa P. Beauv. (=
tribes Poeae and Paniceae, respectively. Brown (1810) Leersia Sw.), Hygroryza Nees, Leersia, Luziola Juss.,
published a grass classification based on the spikelet Oryza L., Potamophila R. Br., Zizania L. (Ehrhartoi-
structure and the geographical distribution of taxa. deae), Alopecurus L., Cornucopia L. (Pooideae), and
Later on, Brown (1814) recognized two tribes within the Crypsis Aiton (Chloridoideae); and tribe Tristegineae
family: Poeae and Paniceae. BrownÕs concept included included several Paniceae genera: Arthropogon Nees,
in ‘‘Paniceae’’ the genus Panicum among other genera Melinis P. Beauv., Reynaudia Kunth, Rhynchelytrum
now considered within the tribe Andropogoneae. His Nees (= Melinis), and Triscenia Griseb., and also a few
concept was later followed by several authors to representatives of Arundinelleae (such as Arundinella
circumscribe the subfamily Panicoideae (Link, 1827), Raddi), Andropogoneae (Cleistachne Benth.), and
while the name Paniceae was restricted to the actual Thysanolaeneae (Thysanolaena Nees).
tribe (Brown, 1814). Hackel (1887) followed Bentham (1881) with a similar
Bentham (1881) recognized the Series Poaceae and classification, recognizing 12 tribes in the Poaceae, six of
Panicaceae [approximately similar to the concept of which were equivalent to those proposed by Bentham
Brown (1814)], including within Panicaceae genera in for the Series Panicaceae. Nevertheless, HackelÕs
which the spikelets disarticulate from the pedicel below descriptions were more detailed and included, as diag-
the glumes. Bentham (1881) divided Series Panicaceae nostic characters, novel microcharacters of the fruit such
into six tribes: Paniceae, Maydeae, Oryzeae, Tristegi- as hilum and starch grain types. During the first half of
neae, Zoysieae and Andropogoneae. Tribe Paniceae was the 20th century, new anatomical and cytological features
recognized by the indurate fruiting ‘‘glume’’ with four were added as valuable characters for the classification of
major groups included: (i) ‘‘Paniceae proper’’ with an grasses (Avdulov, 1931; Prat, 1932, 1936).
arbitrary number of genera and distinctive characters Pilger (1940) recognized four tribes within the sub-
being very uncertain, although still recognizing within it family Panicoideae: tribe Paniceae with six subtribes
at least 11 genera as follows: Anthaenantia P. Beauv., (Anthephorinae, Boivinellinae, Lecomtellinae, Melinid-
Amphicarpum Kunth, Beckmannia Host (currently in inae, Panicinae and Trachidinae); tribe Andropogoneae
Poeae s.l.), Chaetium Nees, Eriochloa Kunth, Ichnanthus with six subtribes (Andropogoninae, Dimeriinae, Ischa-
P. Beauv., Oplismenus P. Beauv., Paspalum L., Panicum, eminae, Rottboelliinae, Saccharinae and Sorghinae);
Reimaria Humb. and Bonpl. ex Flüggé (= Paspalum), tribe Arthropogoneae; and tribe Maydeae. Of these four
and Setaria; (ii) ‘‘Cenchrus’’ with spikelets having an tribes, Pilger (1954) included tribes Andropogoneae and
Maydeae in subfamily Andropogonoideae, and consid- laeneae. Within the Paniceae, a total of seven subtribes
ered eight tribes in the Panicoideae: of these, Arthro- were accepted: Arthropogoninae, Cenchrinae, Digitarii-
pogoneae and Paniceae were already described in his nae, Melinidinae, Panicinae and Setariinae.
previous work (Pilger, 1940) and accepted as tribes the
following taxa: Anthephoreae, with a single genus,
Anthephora; Boivinelleae, with Boivinella A. Camus, Materials and methods
Cyphochlaena Hack., and Perulifera A. Camus; Lecom-
telleae, with Lecomtella A. Camus; Melinideae, with Taxonomic sampling
Melinis, Rhynchelytrum, and Tricholaena Schrad.; and
tribe Trachyeae, with Trachys Pers. Also, Pilger (1954) We included a total of 155 genera currently consid-
recognized tribe Isachneae in the Panicoideae with ered part of the subfamily Panicoideae, each represented
Coelachne R. Br., Heteranthoecia Stapf, Isachne R. by at least one species (Clayton and Renvoize, 1986;
Br., and Limnopoa C.E. Hubb. Watson and Dallwitz, 1992; Zuloaga et al., 2000;
Brown (1977) divided tribe Paniceae into four sub- Sánchez-Ken and Clark, 2010). Among outgroups we
tribes according to a phylogenetic concept based on leaf included 18 genera of the PACMAD clade (excluding
anatomy and photosynthetic types. Subtribe 1 included Panicoideae) and four genera from the BEP clade,
mostly species with paniculate inflorescences and leaf rooting with Bromus inermis Leyss. The final matrix
cross-sections with Kranz anatomy and a single bundle included 265 taxa; 15 genera were only considered with
sheath (Kranz MS); these plants used the NADP–malic morphology because ndhF sequences were not obtained
enzyme subtype of the C4 photosynthetic pathway (C4 due to lack of voucher material or because vouchers
NADP–ME). BrownÕs subtribe 2 included plants that failed to amplify: Acostia Swallen, Acritochaete Pilg.,
were all non-Kranz and C3; subtribe 3 included species Arthragrostis Lazarides, Baptorhachis Clayton and Ren-
with racemose inflorescences and leaf cross-sections with voize, Eccoptocarpha Launert, Holcolemma Stapf and
Kranz anatomy and a double bundle sheath (Kranz PS), C.E. Hubb., Hubbardia, Hydrothauma Hubbard,
C4 PEP–carboxykinase subtype (PEP–CK); and sub- Lecomtella, Microcalamus Franch., Oryzidium C.E.
tribe 4 included species with paniculate inflorescences Hubb. and Schweick., Streptolophus Hughes, Tarigidia
and leaf cross-sections with Kranz PS, C4 NAD–ME. In Stent, Thyridachne C.E. Hubb., and Trachys. Hubbardia
this work, Brown proposed evolutionary hypotheses heptaneuron Bor has been collected only twice, in spray-
among taxa and related the photosynthetic syndrome to moistened rocks near the Jog Falls on the Sharavati
environmental adaptations. He hypothesized that An- River (Yadav et al., 2010). We were unable to obtain
dropogoneae (all Kranz MS, C4 NADP–ME) were material of this species, which is critically endangered
derived from some ancient Kranz MS Paniceae or but is the subject of an effort at restoration (Yadav
Panicoideae, and predicted two or three independent et al., 2009). A list of the species, voucher material and
origins for the Kranz syndrome within Paniceae. Genbank accession numbers used in the molecular study
Clayton and Renvoize (1986), following in part the is presented in Appendix 1.
classification of Pilger (1954), recognized seven tribes in
Panicoideae, based mainly on spikelet morphology, Morphological data matrix
upper floret texture and leaf anatomical type. The two
major tribes were Paniceae and Andropogoneae; the The morphological data set was modified from the
former was divided into seven subtribes: Arthropogon- matrix published by Zuloaga et al. (2000) with partic-
inae, Cenchrinae, Digitariinae, Melinidinae, Neurachni- ular modifications: two extra characters were added,
nae, Setariinae (= Panicinae) and Spinificinae; while while other changes included adding states for the
Andropogoneae was organized into 11 subtribes. Other expanded outgroup, and changes in the coding scheme
minor tribes within Panicoideae include Arundinelleae, concerning the multistate non-additive characters that
Eriachneae, Hubbardieae, Isachneae and Steyermar- are here coded as additive whenever the states are
kochloeae. considered internested (Lipscomb, 1992). A total of 57
Watson and Dallwitz (1992) treated six tribes in the characters were scored (Appendices 2 and S1). In the
Panicoideae: Andropogoneae, Arundinelleae, Isach- present analysis, terminals were coded at the species
neae, Maydeae, Neurachneae and Paniceae. They placed level if DNA sequence data were available. For the
Steyermarkochloeae and Eriachneae within Arundinoi- terminals where DNA sequence data were not available,
deae, and Hubbardia in Isachneae. we coded the characters at generic level, scoring any
In one of the most recent treatments of the American polymorphic character as such. The characters were
Panicoids, Zuloaga et al. (2007b) recognized eight tribes scored on herbarium material deposited at BAA,
based on morphology and molecular data: Andro- CAMB, K, MO, NSW, PRE, SI and US (abbreviations
pogoneae, Arundinelleae, Centotheceae, Gynerieae, according to Index Herbariorum; B. Theirs, http://
Isachneae, Paniceae, Steyermarkochloeae and Thysano- sweetgum.nybg.org/ih); if vouchers were not available,
we followed the information found in the DELTA reverse strands were sequenced with a minimum overlap
system (Watson and Dallwitz, 1992). of 90% for every taxon. Single-pass sequencing was
performed on each template using selected primers to
DNA extraction, amplification and sequencing complete a bidirectional contig of the full sequence.
Editing and assembling of sequences was conducted
Total DNA was extracted from leaves of plants using Chromas Pro ver. 1.34 (Technelysium Pty, Ltd,
collected in the field and dried in silica gel using Tewantin, Qld, Australia). Quality of sequences was
modified Cetyltrimethylammonium Bromide (CTAB) assessed by visual inspection of the chromatograms.
protocol from Doyle and Doyle (1987). A DNeasy Plant Sequences were manually aligned using BioEdit ver.
Mini Kit (Qiagen, Hilden, Germany) was used when 5.0.9 (Hall, 1999), adding gaps to the matrix. Each base-
fresh material was not available, and herbarium spec- pair gap was treated as missing data, but informative
imens were analysed. indels were considered as presence ⁄ absence characters
A plastid marker, ndhF, was amplified; this gene during the analyses.
encodes a large subunit of one of the proteins that takes
part in photorespiration (the NADPH plastiquinone Phylogenetic analyses
oxidoreductase). This gene is thought to be conservative
within the plant genome although, as an actively We used TNT ver. 1.1 (Goloboff et al., 2008) for
evolving gene of the plastome, its nucleotide-substitu- phylogenetic analyses. We analysed both the combined
tion rate is twice as high as that of rbcL (Seliverstov matrix and the ndhF sequence matrix on its own. For both
et al., 2009). These characteristics make ndhF valuable matrices we used Driven search, finding the minimum
for phylogenetic reconstruction and it has proven to be a length 100 times with default settings for Sectorial
useful tool in different phylogenetic lineages of plants searches and Tree fusing (Goloboff, 1999), all with
(Kim and Jansen, 1995; Catalán et al., 1997; Aliscioni ‘‘Random Seed = 0’’. The resulting trees were submitted
et al., 2003; Sánchez-Ken et al., 2007). The complete to Ratchet (Nixon, 1999) and Drift (both default settings)
ndhF gene (ca. 2000 bp) was amplified using primers until 10 000 trees were found. Since there were no
specified by Olmstead and Sweere (1994), Clark et al. differences among the strict consensus trees calculated
(1995), and Aliscioni et al. (2003). For silica-dried from the original 100 optimal trees and the 10 000 optimal
samples, three pairs of primers were used (5F–972R, trees found during ratchet and drifting, we did not
972F–1666R and 1666F–2110R). For herbarium sam- continue the searches. To improve the resolution of the
ples, five smaller fragments were amplified using 5F– strict consensus tree, we used the ‘‘Tree pruning’’ option
536R, 536F–972R, 972F–1318R, 1318F–1666R and to find taxa that caused more than five nodes to collapse.
1666F–2110R. For non-overlapping fragments, we filled These species were excluded from the final consensus
in missing base pairs with N, although these characters trees. Jackknife values (Farris et al., 1996) were calculated
represent a small proportion of the entire sequence. by 10 000 resampling iterations with a removal probabil-
PCR reactions were performed in 25 lL final volumes ity of 36%, using 10 replicates of Ratchet to find the
with 50–100 ng of template DNA, 0.2 lm of each minimum length once in each replication. As for the
primer, 25 lm dNTP, 5 mm MgCl2, 1· buffer and 1.5 analyses above, all taxa were retained in the matrix during
units of Taq polymerase provided by Invitrogen Life the resampling, but taxa found to cause conflict under the
Technologies. The reaction conditions were: a first tree searches were excluded from the consensus when
period of denaturation at 94 °C for 5 min, followed by calculating the final jackknife frequencies.
35 cycles of denaturation at 94 °C for 30 s, annealing at To explore character evolution within the study
48 °C for 60 s, and extension at 72 °C for 90 s. Final group, we generated two phylograms, one showing the
extension at 72 °C for 6 min terminated the reactions. A amount of character changes of the 20 inflorescence
negative control with no template was included for each characters; the other showing the 31 characters related
series of amplifications to check for contamination. to the spikelet morphology. Both phylograms were
PCR products were run out on a 1% TBE agarose gel generated on the basis of one of the optimal trees from
stained with SYBR Safe DNA gel stain (Invitrogen) and the combined data set.
visualized in a blue light transilluminator. Cleaning of
PCR products was done by Macrogen, Inc. (Seoul,
Korea) using the Montage PCR purification kit from Results
Millipore and following the manufacturerÕs protocol.
Sequencing reactions were also performed by Macrogen ndhF sequence analysis
using the ABI PRISM BigDye Terminator Cycle
Sequencing Kits with AmpliTaq DNA polymerase Sequence alignment was trivial, with five informative
(Applied Biosystems, Seoul, Korea) following the pro- indels between 6 and 24 bp long, as already described by
tocols supplied by the manufacturer. Both forward and Giussani et al. (2001). The aligned sequences included
Fig. 1. Strict consensus tree resulting from analysis of the ndhF sequences. Numbers below the branches indicate jackknife values from the ndhF
sequence analysis; numbers above the branches indicate jackknife values from the combined analyses (ndhF + morphology). Positions of taxa only
included in the combined analysis are marked with black dots. Boxes show an alternative topology resulting from the combined analyses.
Fig. 1. (Continued).
2074 bp, of which 770 were phylogenetically informa- sequences, showed that the subfamily Panicoideae
tive. The final tree length was 4000 steps long. To would be polyphyletic if tribe Isachneae was included.
improve resolution, Stereochlaena cameronii (Stapf) Sánchez-Ken et al. (2007) consequently reinstated the
Pilg. was excluded from the strict consensus tree shown subfamily Micrairoideae including tribes Micraireae,
in Fig. 1 and from the consensus when calculating Isachneae and Eriachneae. We sequenced two additional
jackknife support. species of tribes Isachneae and Eriachneae, Isachne
leersioides Griseb. and Pheidochloa gracilis S.T. Blake,
Combined morphological and molecular analyses respectively. Both taxa grouped within the Micrairoi-
deae clade supported by a strong jackknife value of 99,
The morphological matrix included 57 characters. in agreement with previous results (Duvall et al., 2007;
A nearly collapsed consensus tree was obtained when Sánchez-Ken and Clark, 2007; Sánchez-Ken et al.,
analysing the morphological matrix due to a high degree 2007).
of homoplasy combined with a low character ⁄ taxa ratio. Hubbardia is a monotypic endemic genus from east
The combined morphological and molecular matrix India, which is recognized by having spikelets dorsally
yields optimal trees 4943 steps long. In general, the compressed with two flowers, disarticulation above the
consensus tree from the combined matrix was better glumes, both glumes persistent, and the palea of both
resolved for most clades, although only few clades florets absent. Clayton and Renvoize (1986) and Sán-
showed over 50% jackknife values in the combined data. chez-Ken and Clark (2010) treated this genus in tribe
All supported clades from the combined data matrix were Hubbardieae, subfamily Panicoideae. Although DNA
clades also appearing when the ndhF matrix was analysed sequences were not obtained for Hubbardia in the
on its own. Consequently, we show the consensus tree and present study, the combined analysis clearly indicates
jackknife values found when analysing only the ndhF that the genus belongs to the Micrairoideae clade and is
sequences, and compare those with the values that are sister to Isachne (Fig. 1). All genera of subtribe Micrai-
found when morphology is added to the matrix (Fig. 1). rioideae are characterized by having the upper anthe-
To improve resolution in the jackknife consensus tree cium cartilaginous. The placement of Hubbardia outside
from the combined analysis, all taxa lacking DNA the Panicoideae is consistent with current molecular
sequence data were excluded. Three monotypic genera phylogenetic studies (Sánchez-Ken, unpublished data).
from tropical Africa—Hydrothauma, Oryzidium and
Thyridachne—represented only by morphological data, Subfamily Panicoideae s.l.
were excluded in the strict consensus tree. In some trees,
Hydrothauma is placed as sister to either the Andropogo- Subfamily Panicoideae s.l. (including Panicoi-
neae–Arundinelleae s.s.–Paniceae s.l. clade; the Andro- deae + Centhothecoideae) is well supported with a
pogoneae–Arundinelleae s.s.–Paniceae x = 10 clade strong jackknife value of 96 and 92 in the ndhF and
(Paspaleae); or the Paniceae s.s. x = 9 clade. Thyridachne combined analyses, respectively. Both analyses sup-
is placed within Boivinellinae subclade, as sister to ported 11 out of 12 tribes proposed by Sánchez-Ken and
subclade Neurachninae, or in one of the three basal Clark (2010) within the Panicoideae, because Hub-
nodes within the Paniceae s.s. x = 9 clade. Oryzidium is bardieae was included into Micrairioideae (Fig. 1).
placed as sister to Melinis repens (Willd.) Zizka within Gynerieae and Thysanolaeneae are both monotypic
Melinidinae subclade, or as sister to Reynaudia filiformis tribes, while the other six tribes are monophyletic:
(Spreng. ex Schult.) Kunth. Andropogoneae, Arundinelleae s.s., Centotheceae, Cyp-
erochloeae, Tristachyideae and Zeugiteae. Our analyses
do not support the monophyly of tribes Chasmanthieae,
Discussion Paniceae s.l. and Steyermarkochloeae.
Our phylogeny revealed good resolution within the Tribe Steyermarkochloeae

PACMAD clade, and several major groups were recov-
ered by ndhF in agreement with previous works (Gómez- Tribe Steyermarkochloeae includes two monotypic
Martı́nez and Culham, 2000; Duvall et al., 2001, 2007; genera: Steyermarkochloa Davidse and R.P. Ellis and
Giussani et al., 2001; GPWG, 2001; Sánchez-Ken and Arundoclaytonia Davidse and R.P. Ellis, endemic to
Clark, 2007, 2010; Sánchez-Ken et al., 2007; Bouchenak- Amazonian South America. Taxonomic classification of
Khelladi et al., 2008). this tribe has been contradictory; Davidse and Ellis
(1984, 1987), on the basis of anatomical and morpho-
Subfamily Micrairoideae logical characters, included this tribe under Arundinoi-
deae, while Watson and Dallwitz (1992) suggested its
Recent phylogenetic studies (Duvall et al., 2007; close relation to Bambusoideae. GPWG (2001) and
Sánchez-Ken and Clark, 2007), based on DNA Zuloaga et al. (2003, 2007b) considered tribe Ste-
yermarkochloeae in the Panicoideae, in agreement with basic chromosome number of x = 10. Most of the
Clayton and Renvoize (1986), who mentioned its x = 10 Paniceae are plants of the Americas, with the
resemblance to Hymenachne P. Beauv. exception of Dallwatsonia B.K. Simon from Australia,
Steyermarkochloa angustifolia (Spreng.) Judz. is here and Lecomtella and Baptorhachis from Africa. This
included for the first time in a phylogenetic analysis. clade is clearly related to the Andropogoneae–Arund-
Whether using ndhF sequences alone or in combination inelleae s.s. x = 10 clade, indicating that the basic
with morphology, Steyermarkochloa angustifolia is sister chromosome number may be a synapomorphy. The
to the Tristachyideae clade within the Panicoideae. second clade includes genera with a basic chromosome
However, the branch in the ndhF tree, together with that number of x = 9 and a pantropical distribution.
of Reynaudia filiformis [35 steps long, see below], is one
of the longest in the data matrix, with a total of 45 The American x = 10 Paniceae s.l. (= tribe Paspaleae)
changes. Since the sister clade of Steyermarkochloa
angustifolia is a relatively long branch (38 changes), we Based on the ndhF phylogeny, the American x = 10
excluded the Tristachyideae clade to explore if the clade is moderately supported with jackknife values of
position of Steyermarkochloa angustifolia was affected 80 (58 in the combined analysis) if Reynaudia is
by long-branch attraction (Pol and Siddall, 2001). When considered part of it. If Reynaudia is excluded, the clade
the Tristachyideae clade was left out of the analyses, is strongly supported, with jackknife support values of
Steyermarkochloa angustifolia drifted to several posi- 98 (93 in the combined analysis). Reynaudia filiformis,
tions and appeared related to the Thysanolaena clade or an endemic species from Cuba, is sister to the remaining
the Orthoclada–Chasmanthium clade, but was never x = 10 clade and shares with it a 6 bp insertion.
sister to Arundoclaytonia dissimilis. Davidse and R.P. However, the ndhF sequence of R. filiformis differs
Ellis in several positions from the remainder sequences in
Unfortunately, we failed to obtain a complete ndhF the x = 10 clade, and is one of the longest branches,
sequence of Arundoclaytonia dissimilis, hence we used with 35 changes; the chromosome number is unknown
the partial one obtained by Sánchez-Ken and Clark for this species. Reynaudia differs from other members
(2007). In our combined analysis (ndhF + morphol- of the American x = 10 Paniceae s.l. in several charac-
ogy), Arundoclaytonia is nested within the Chas- ters of the spikelet, such as awned and bilobed glumes,
manthieae clade, closely related to Chasmanthium and and the fertile floret with two stamens. Although it is
Bromuniola, in agreement with previous results reported possible that Reynaudia is part of the American x = 10
by Sánchez-Ken and Clark (2010). While the mono- clade, the taxon remains incertae sedis because of the
phyly of tribe Steyermarkochloeae should be tested moderate jackknife value and the divergent spikelet
further, its inclusion in subfamily Panicoideae is cor- morphology. The genus was previously included in
roborated here. subtribe Arthropogoninae (Clayton and Renvoize, 1986;
Zuloaga et al., 2007b), but it is distinct from Achlaena,
Tribe Paniceae s.l. Altoparadisium Filg., Davidse, Zuloaga and Morrone,
and Arthropogon according to its phylogenetic position
We obtained ndhF sequences for almost all the (see below).
diversity of the tribe Paniceae s.l.; 90% of the currently Excluding Reynaudia, all other American x = 10
recognized genera of the tribe are represented here. Our genera form a highly supported (98 ⁄ 93) monophyletic
molecular sampling is unique in the addition of many group with C3, C4 NADP–ME and intermediate C3–C4
underrepresented genera and Old World entities, prin- genera, and 10 nucleotide synapomorphies; this group is
cipally from Africa and Australia, such as taxa from assigned to tribe Paspaleae. The tribe is divided into
subtribes Cenchrinae, Neurachninae, Setarinae and three major subclades corresponding to the subtribes
Spinificinae. We incorporated in the morphological Arthropogoninae, Otachyriinae and Paspalinae.
matrix all genera recognized within the Paniceae. A
previous phylogenetic study based only on morphology Subtribe Otachyriinae
(Zuloaga et al., 2000) also included a large sample with
more than 100 genera of Paniceae. However, the This subclade, which is concordant with previous
monophyly of the tribe was not tested (Zuloaga et al., groupings (Duvall et al., 2001; Giussani et al., 2001;
2000), as the tree was rooted with Isachne and no species Aliscioni et al., 2003), includes species of Anthaenantia,
of Andropogoneae or other panicoid tribes were Dallwatsonia, Hymenachne, ‘‘Panicum’’ sect. Laxa, Pla-
considered. giantha Renvoize, Otachyrium Nees, and Steinchisma
In agreement with previous molecular studies (Gómez- Raf. The subclade is strongly supported both when
Martı́nez and Culham, 2000; Giussani et al., 2001), our analysing the ndhF data set independently or combined
results showed that tribe Paniceae is paraphyletic and with morphology (jackknife values 93 versus 84).
splits in two major clades, one including taxa with a Dallwatsonia, a monotypic genus endemic to Australia
(Simmon, 1992), is morphologically similar to the plastic molecular synapomorphies, in agreement with
American incertae sedis ‘‘Panicum’’ sect. Laxa. This previous results (Scataglini et al., 2007; Rua et al.,
genus shares with sect. Laxa the presence of a lax 2010). During the past decade, the circumscription of
inflorescence with crowded spikelets disposed unilater- Paspalum has changed to include Thrasya in Paspalum
ally on the branches, and an ellipsoid spikelet, with the subgenus Harpostachys (Denham, 2005; Denham and
lower glume shorter than the spikelet, while the upper Zuloaga, 2007). In order to keep Axonopus and Pasp-
anthecium is membraneous. It is noteworthy that alum both monophyletic, it would be necessary to
Dallwatsonia and species of sect. Laxa share fusoid cells include Centrochloa and Ophiochloa in Axonopus; and
as seen in leaf cross-sections, although these are irreg- Spheneria, Thrasyopsis and Reimarochloa in Paspalum.
ularly distributed in Dallwatsonia. This genus may have When describing Acostia, a monotypic genus from
been introduced recently into Australia, as reported for Ecuador, Swallen (1968) pointed out its resemblance to
Cliffordiochloa parvispiculata B.K. Simon [= Stein- Digitaria Haller and Panicum, distinguishing the genus
chisma laxa (Sw.) Zuloaga] (Simon, 2003) and for by the absence of the lower glume. Webster et al. (1989)
Fasciculochloa sparshottiorum B.K. Simon and C.M. related this genus to Panicum and included Acostia in its
Weiller [= Steinchisma hians (Elliot) Nash] (Simon, synonymy. In our results, based on the combined
1999). analysis, Acostia is not related to Panicum s.s. or to
Digitaria, but is sister to the Axonopus–Centrochloa–
Subtribe Paspalinae Ophiochloa clade, with which it shares a reduced or
absent lower glume, and a cartilaginous upper anthe-
Subclade Paspalinae includes Anthaenantiopsis, Axon- cium.
opus P. Beauv., Centrochloa Swallen, Echinolaena Desv.,
Gerritea Zuloaga, Morrone and Killeen, Hopia, Ichnan- Subtribe Arthropogoninae
thus, Ocellochloa, Ophiochloa Filg., Davidse and Zulo-
aga, Paspalum, Reimarochloa Hitchc., Renvoizea, The relationship of the genera of Arthropogoninae to
Spheneria Kuhlm., Streptostachys Desv., Thrasyopsis other Panicoideae has been controversial. Some of them
Parodi, Panicum validum Mez, and P. tuerckheimii were placed in the tribe Andropogoneae (Kunth, 1833;
Hack. (both ‘‘Panicum’’ species as incertae sedis), most Steudel, 1853–1855); Tristegineae (Bentham, 1881; Hac-
of them having unilateral inflorescences, with C3 or C4 kel, 1887); or Melinidinae (Bews, 1929; Hitchcock, 1936;
NADP–ME photosynthetic pathways. Some genera Roshevitz, 1937), or were grouped in an independent
have a stipe and disarticulate at the base of the upper tribe, Arthropogoneae (Pilger, 1954). Our results place
anthecium, while other genera are characterized by the the Arthropogoneae within the tribe Paspaleae, so their
presence of simple papillae and bicellular microhairs at name becomes subtribe Arthropogoninae; the other
the apex or base of the upper palea. The clade is tribes are distantly related, with tribe Andropogoneae in
moderately supported (jackknife values for ndhF 86; the x = 10 Andropogoneae clade, and tribes Melinid-
combined 82; Fig. 1) and characterized by three molec- inae and Tristegineae (= Melinidinae) as part of the
ular synapomorphies. In the combined analysis, the x = 9 Paniceae s.s. clade.
genera Lecomtella and Baptorhachis are represented by Subclade Arthropogoninae, also called the ‘‘Ambig-
morphological data and fall within subclade Paspalinae, uous clade’’ by Giussani et al. (2001), included several
although the position has little jackknife support. heterogeneous genera with noteworthy variation in
Lecomtella is a monotypic genus endemic to Madagas- morphological, anatomical and physiological charac-
car with an unusual arrangement of spikelets on the ters. This subclade is weakly supported when analysing
branches, having bisexual spikelets at the tips of the the ndhF sequences on their own (jackknife value of 73)
branches and staminate ones below (Clayton and but with very low support when jackknifing the
Renvoize, 1986), and a peculiar winged stipe at the base combined data set (jackknife value of 54); a total of
of the upper floret. Baptorhachis is endemic to Mozam- three molecular synapomorphies define the group.
bique, and has an inflorescence with a solitary raceme Subclade Arthropogoninae includes seven genera with
with broad foliaceous rachis and spikelets with the lower C3 photosynthetic pathway (Apochloa, Canastra, Ho-
glume absent. molepis Chase, Oplismenopsis Parodi, Phanopyrum
The largest genera within subclade Paspalinae, Axon- (Raf.) Nash, Stephostachys and Triscenia) while the
opus and Paspalum are both paraphyletic (Giussani remaining nine genera are C4 NADP–ME (Achlaena,
et al., 2001; Aliscioni, 2002). Centrochloa and Ophio- Altoparadisium, Arthropogon, Coleataenia, Cyphonan-
chloa are nested within Axonopus in a clade supported thus, Keratochlaena, Mesosetum Steud., Oncorachis, and
by 17 molecular synapomorphies, four of which are Tatianyx Zuloaga and Soderstr.). Apochloa, Canastra,
non-homoplastic, in agreement with López and Mor- Coleataenia and Cyphonanthus have been segregated
rone (2010). Reimarochloa, Spheneria and Thrasyopsis recently from Panicum s.l. (Zuloaga et al., 2006, 2010;
are included within the Paspalum clade by three homo- Morrone et al., 2007; Sede et al., 2008), whereas Kera-
tochlaena and Oncorachis (Sede et al., 2009b) were genera, of which Anthephora, Chaetopoa C.E. Hubb.,
segregated from Streptostachys. In the ndhF phylogeny, Chlorocalymma Clayton, Tarigidia and Trachys have
three minor subclades within the subclade Arthropog- involucral structures (the other two genera are repre-
oninae are highly supported and include anatomically sented only by morphological data). These five genera
uniform C4 NADP–ME genera (with the exception of are related to Megaloprotachne C.E. Hubb. and Digi-
Triscenia, a monotypic non-Kranz genus). The Achlaena, taria, which lack involucral bracts. Tarigidia represents
Altoparadisium and Arthropogon subclade (jackknife an intermediate form between Anthephora and Digitaria
value of 100 in both analyses) is supported by 18 (Loxton, 1974; Vega et al., 2010); its resemblance to
nucleotide substitutions; these genera share a hairy callus Anthephora is represented by spicifom and terete inflo-
at the base of the spikelet and the upper anthecium is rescences, spikelets grouped in clusters and surrounded
laterally compressed; Achlaena and Altoparadisium show by several glumes, occasionally fused, which are almost
distinctive Kranz cells. In the second subclade, Colea- as long as the lower lemmas. Similarities between
taenia and Triscenia share five base-pair changes. Digitaria and Tarigidia are mostly due to the character-
Triscenia is a monotypic genus endemic to Cuba, and istic flattened upper lemma with broad, thin margins. In
similar to Coleataenia in its racemose inflorescence and Trachys, the spikelets are associated with bractiform
spikelet morphology; Triscenia differs from Coleataenia involucres, which are formed by reduced spikelets
by having the upper anthecium cartilaginous with flat surrounding the clusters.
margins and non-Kranz anatomy. A third group is a
subclade comprised of three genera, Keratochlaena, Subtribe Cenchrinae
Mesosetum and Tatianyx, that have a caryopsis with a
linear hilum; this subclade is supported by 10 substitu- The ‘‘Bristle clade’’ or subtribe Cenchrinae within
tions (jackknife value of 99 in both analyses). The linear Paniceae s.s. is moderately well supported, with jack-
hilum is, however, a homoplasious character state, being knife values of 76 ⁄ 80 in the ndhF versus combined
also present in Homolepis, Oncorachis and Oplismenopsis analysis. The genera in this subtribe are characterized by
(Arthropogoninae), in Streptostachys (Paspalinae), and setae in the inflorescence, although the setae can be
in Acroceras Stapf and Louisiella C.E. Hubb. and J. modified in various ways. The position and abundance
Léonard (x = 9 Paniceae s.s.). of setae is variable; they can be solitary and present only
at the tips of the branches, or numerous and associated
The Pantropical x = 9 Paniceae (= tribe Paniceae s.s.) with every spikelet, or solitary and associated with only
a few spikelets of the inflorescence. Doust and Kellogg
The Paniceae s.s. clade, most members of which have (2002), in their study of the development of the
a basic chromosome number of x = 9, has a jackknife inflorescence in species of the ‘‘Bristle clade’’, showed
value of 76 for ndhF data alone and 77 when combining that all taxa are quite similar at early stages of
both data sets; the group is distributed in tropical and development; later on, changes in the extent of branch-
subtropical regions of the world. Although the basal ing, the differentiation pattern of primordia, and the
generic relationships are not resolved, there are well elongation of the inflorescence axis together caused the
established groups within the x = 9 clade. marked differences seen among the mature inflorescenc-
es. Among genera here included in the subtribe Cen-
Subtribe Anthephorinae chrinae, several were previously placed in the ‘‘Bristle
clade’’: Cenchrus, Ixophorus Schltdl., Odontelytrum
Traditionally, all genera with highly modified inflo- Hack. (=Cenchrus), Paspalidium Stapf (=Setaria L.),
rescences were grouped in subtribe Cenchrinae sensu Pennisetum (=Cenchrus), Pseudoraphis Griff., Setaria,
Clayton and Renvoize (1986). Our study showed that Spinifex L., Stenotaphrum Trin., Uranthoecium Stapf,
Cenchrinae is not monophyletic, and that modified and Zygochloa S.T. Blake. In addition, we have
inflorescences have evolved independently by non- confirmed the placement here of two taxa without
homologous characters in two different lineages: within bristles: Panicum antidotale Retz. and Zuloagaea bulbosa
genera of the ‘‘Bristle clade’’ (see below), bristles are (Kunth) Bess (Gómez-Martı́nez and Culham, 2000;
homologous to an inflorescence branch (Koch, 1843; Zuloaga et al., 2000; Duvall et al., 2001; Giussani et al.,
Sohns, 1955; Butzin, 1977); in the Anthephorinae 2001; Doust and Kellogg, 2002; Aliscioni et al., 2003;
subclade, the involucral bracts are possibly derived Doust et al., 2007; Donadı́o et al., 2009; Kellogg et al.,
from the lower glume or from sterile spikelets (Reeder, 2009; Chemisquy et al., 2010). The ndhF sequence data
1960; Clayton and Renvoize, 1986). In the present also confirm the placement in this clade of Chamaera-
analysis, the Anthephorinae subclade is supported by 16 phis, Dissochondrus (Hillebr.) Kuntze, Hygrochloa
base changes and an 18 bp long deletion, with a strong Lazarides, Paractaenum P. Beauv., Paratheria, Pseud-
jackknife value of 99 and 88 in the ndhF or the combined ochaetochloa Hitchc., Setariopsis Scribn. and Xerochloa,
analysis, respectively. Anthephorinae includes seven a result consistent with the presence of setae in their
inflorescences. Morphological data in the combined ably variable in vegetative and reproductive characters.
analysis also place Holcolemma and Streptolophus in the Some genera have lax branched inflorescences (e.g.
Cenchrinae. Holcolemma is a genus of four species that Melinis, Tricholaena) whereas others have unilateral
grow in Eastern Africa, Southern Asia and Australia; racemes (e.g. Moorochloa Veldkamp, Urochloa P.
this genus is characterized by a contracted inflorescence, Beauv.). The genus Thuarea has highly specialized
with the terminal spikelet of each branch associated with inflorescences, with one to two persistent bisexual
an inconspicuous seta. Holcolemma is similar to species spikelets in the basal portion of a leafy rachis and
of Setaria, but is distinguished by its winged lower several deciduous staminate spikelets in the distal
palea; the latter character is also a feature of the portion; also, in this genus the rachis, after fertilization,
monotypic Mexican genus Ixophorus. In the combined bends to enclose the fruit in a structure similar to a
analysis, Holcolemma and Ixophorus appear as sister capsule. Within the Melinidinae, the lower glume varies
species. Streptolophus, a monotypic genus from Angola greatly in length, from absent (in Yvesia A. Camus and
with spikelets surrounded by a spiny involucre, is sister Thuarea), to rudimentary or nearly as long as the
to Paratheria prostrata Griseb., which has solitary spikelet; both glumes, the lower lemma and the upper
spikelets subtended by large, solid setae. anthecium are muticous or aristate, the upper anthecium
In addition to Zuloagaea bulbosa and Panicum is smooth or with conspicuous transverse rugosities with
antidotale, our analyses identified four genera of the simple or compound papillae present. Eccoptocarpha, a
clade that also lack setae: Acritochaete (morphology monotypic genus from Africa, Tanzania and Zambia, is
only), Alexfloydia B.K. Simon, Snowdenia, and Whiteo- sister to Moorochloa in the analyses of the combined
chloa C.E. Hubb. Bess et al. (2005) reported that the matrix. This genus has PEP–CK anatomy, and is
inflorescence development of Zuloagaea bulbosa is distinguished morphologically from Moorochloa by
similar to that of Panicum miliaceum L. (type species having the upper anthecium borne on a slender, sinuous
of Panicum s.s.), and that the former species is anom- rachilla internode, and by having a reticulate network of
alous in having lost the synapomorphy of the ‘‘Bristle veins on the upper glume and lower lemma; the latter
clade’’. As in Zuloagaea bulbosa, Panicum antidotale, character was also observed in some species of Urochloa
Whiteochloa capillipes (Benth.) Lazarides, and Alexfloy- (e.g. U. humidicola, U. dictyoneura). In spite of the
dia repens B.K. Simon have spikelets and inflorescences morphological variability found in this clade, two
that are morphologically similar to those of Panicum subclades were recovered. The first subclade, which
s.s.; developmental patterns in these species should be includes Leucophrys Rendle, Melinis, Moorochloa and
studied to determine the basis for the absence of bristles. Tricholaena, is strongly supported: the genera share five
Acritochaete, a monotypic genus of Eastern Africa, base changes and morphological characters, such as
without setae, was thought by Clayton and Renvoize disarticulation at the base of the upper anthecium, and
(1986) to be related to Digitaria; this genus has a small the upper anthecium smooth and shiny. The other
lower glume and a chartaceous–cartilaginous upper subclade, which includes Chaetium, Eriochloa, Mega-
anthecium similar to that in Cenchrus and related thyrsus, Scutachne Hitchc. and Chase, and Urochloa, is
genera; its inclusion in this clade needs to be confirmed also strongly supported: all these genera share seven
by DNA data. In addition, Snowdenia, nested within base changes, although we found no morphological
Cenchrus, has spikelets similar to those present in characters as synapomorphies.
Cenchrus unisetus (Clayton and Renvoize, 1986), and
could represent a distinct case of suppression of setae Subtribe Panicinae
during inflorescence development. The absence of setae
in at least six taxa in the group represents independent Based on our molecular and combined analyses,
losses during evolution. Arthragrostis, Louisiella, Panicum s.s., and Yakirra
Lazarides and R.D. Webster form a well supported
Subtribe Melinidinae clade, the subtribe Panicinae (jackknife values of 65 and
93 in the ndhF and combined analysis, respectively). All
Our analysis confirms previous results in this subtribe these genera are C4 NAD–ME. Arthragrostis, a mono-
(Gómez-Martı́nez and Culham, 2000; Giussani et al., typic genus from Australia, is in a large polytomy
2001; Aliscioni et al., 2003; Christin et al., 2007, 2008; together with Panicum and Louisiella. Arthragrostis is
Vicentini et al., 2008; Salariato et al., 2010). The distinguished by having primary inflorescence branches
Melinidinae includes 12 genera plus Panicum deustum deciduous and upper anthecium stipitate; this stipe is
Thunb., and is weakly supported by jackknife values of formed by a filiform internode of the rachilla without
66 in the ndhF analysis and 62 in the combined analysis. lateral appendages (see discussion of this character
The subtribe is characterized by mainly C4 photosyn- below). Yakirra, a genus with five species that grow in
thesis, with leaf anatomy characteristic of the PEP–CK Myanmar and Australia, is also characterized by the
subtype. The subtribe includes genera that are remark- presence of a homogeneous stipe at the base of the upper
anthecium; this stipe includes two small auricles in its clade includes in our analysis six genera, four of them
distal portion. In the molecular analysis, Yakirra is endemic to Australia: Calyptochloa, Neurachne, Para-
placed as sister to the remaining species of the Panicinae, neurachne and Thyridolepis; while Ancistrachne also
while in the combined analysis it is nested in a clade with extends to south-eastern Asia, the Philippines and
some species of Panicum s.s. and Arthragrostis. Both New Caledonia; and Cleistochloa grows in Australia
Arthragrostis and Yakirra are morphologically similar and New Guinea. The Neurachninae clade is only
to species of Panicum sect. Rudgeana in having an open weakly supported in the molecular analysis, with a
and lax panicle, spikelets dorsiventrally compressed, jackknife value of 63, but is strongly supported in the
similar nervation of glumes and lower lemma, and upper combined analysis (jackknife value of 94). The subtribe
anthecium indurate, glabrous, and with simple or is characterized by having inflorescences with reduced
compound papillae toward the apex of lemma and secondary branches, and spikelets abaxially disposed.
palea (Zuloaga, 1987); moreover, all species of Panicum From an anatomical and physiological point of view,
sect. Rudgeana have a stipitate upper anthecium. the Neurachninae includes four non-Kranz genera:
However, this stipe is heterogeneous in sect. Rudgeana Ancistrachne, Calyptochloa, Cleistochloa and Thyridol-
and homogeneous in both Arthragrostis and Yakirra. A epis) and one Kranz genus of the NADP–ME subtype,
stipitate upper anthecium has originated several times (Paraneurachne); while Neurachne is atypical in includ-
independently within the Paniceae s.l. It has been ing non-Kranz species, Kranz species of the subtype
observed in Echinolaena, Ichnanthus and Paspalum, NADP–ME but with anatomical deviations, and C3 ⁄ C4
and in some species of Dichanthelium (Hitchc. and intermediates such as N. minor S.T. Blake. The NADP–
Chase) Gould, Renvoizea and Ocellochloa. The elonga- ME photosynthetic subtype represents here a derived
tion of the rachilla between the lower and the upper feature from C3, and has evolved independently from
lemma could be part of a strategy for dispersal of the the rest of the Paniceae; the C3 ⁄ C4 intermediate species
caryopsis; Davidse (1987) pointed out that the stipe at appear to be transitional stages. Calyptochloa and
the base of the anthecium in species of Echinolaena, Cleistochloa are closely related genera with several
Ichnanthus and Panicum is an elaiosome, connected with synapomorphies, such as the presence of cleistogenes
dispersal by ants. Based on our results and the similar in leaf axils, secondary inflorescence branches reduced
features observed in Arthragrostis, Panicum s.s. and to a single adaxial spikelet, the lack of a lower palea, and
Yakirra, these three taxa could eventually be considered macrohairs on the upper palea.
in a single genus. Louisiella, the remaining genus in
subtribe Panicinae, is monotypic and grows in western Subtribe Boivinellinae
Africa; this genus is in a strongly supported clade, both
in the molecular and combined analysis, with Panicum Giussani et al. (2001) recognized as the ‘‘Forest shade
elephantipes Nees ex Trin., an American member of sect. clade’’ a group of genera of forest shade habitats:
Dichotomiflora of Panicum s.s. Louisiella shares with Acroceras, Echinochloa P. Beauv., Lasiacis (Griseb.)
P. elephantipes several characters and ecological prefer- Hitchc., Oplismenus P. Beauv., Parodiophyllochloa, and
ences, such as plants floating or decumbent on mud, Pseudechinolaena Stapf. In this study, the clade is
culm internodes spongy, ligules membraneous–ciliate, expanded to include Alloteropsis J. Presl, Amphicarpum
spikelets with the lower glume reduced, upper glume and Kunth, Cyrtococcum Stapf, Cyphochlaena, Entolasia
lower lemma acuminate, longer than the upper anthe- Stapf, Ottochloa Dandy and Microcalamus; all 10 genera
cium. Louisiella differs from P. elephantipes and mem- are therefore taxonomically circumscribed in subtribe
bers of sect. Dichotomiflora in having a caryopsis with a Boivinellinae. Microcalamus is represented only by
linear hilum. A more detailed study of Panicum s.s. is morphological data; nevertheless, preliminary ndhF data
necessary to circumscribe this genus and to establish its for Microcalamus confirms its position in the subtribe
relationships with Arthragrostis, Louisiella and Yakirra. (C. Grennan and M. Duvall, pers. comm.); when
including morphological data in the combined analysis,
Subtribe Neurachninae Microcalamus is sister to Acroceras. The Boivinellinae
was moderately supported by jackknife values of 81 ⁄ 72
Blake (1972) grouped Neurachne, Paraneurachne S.T. when using ndhF sequences or combined data sets,
Blake, and Thyridolepis S.T. Blake in tribe Neurach- respectively; the subtribe is supported by three molec-
neae. Later, Clayton and Renvoize (1986) included these ular changes. Furthermore, several genera, such as
genera in the subtribe Neurachninae within the tribe Acroceras, Amphicarpum, Cyphochlaena, Cyrtococcum,
Paniceae s.l. Our study showed a close relationship of Entolasia, Lasiacis, Microcalamus, Oplismenus, Otto-
these three genera with others previously included in chloa, Parodiophyllochloa and Pseudechinolaena, are C3,
subtribe Setariinae (Clayton and Renvoize, 1986): Anc- prefer the interior or edge of forests, and all have broad
istrachne S.T. Blake, Calyptochloa C.E. Hubb., and leaves. Echinochloa is a C4 NADP–ME genus; its
Cleistochloa C.E. Hubb. As a result, the Neurachninae position, when analysing sequences of phyB, a coding
gene from the nucleus, is unrelated to the Boivinellinae Willis et al. (2000) and Bruhl et al. (2006) restricted the
clade (Vicentini et al., 2008), a fact that might be genus to only one species, Homopholis belsonii C.E.
attributed to hybridization or reticulation events. Hubb., and transferred H. proluta (F. Muell.) R.D.
Finally, Alloteropsis is a heterogeneous genus, includ- Webster and Panicum subxerophilum Domin to the new
ing C4 PEP–CK, C4 NADP–ME, C4 NAD–ME, or Australian genus Walwhalleya Wills and J.J. Bruhl, with
non-Kranz species (Ellis, 1977; Hattersley and Watson, an additional new species, W. pungens (Wills and J.J.
1992; Ibrahim et al., 2009). Bruhl) Wills and J.J. Bruhl. Our study grouped both
genera in a well supported clade with jackknife values of
Subclade A (Dichanthelium—’’Panicum’’ sect. 97 ⁄ 99. This result contradicts that of Willis et al. (2000),
Clavelligerae) who related Walwhalleya to Digitaria, Entolasia, and
species of Panicum s.l. Homopholis and Walwhalleya are
This subclade, previously described by Giussani et al. both physiologically C3 genera and sister to the subclade
(2001) and Aliscioni et al. (2003), includes American that includes the three photosynthetic C4 pathways
species of the genus Dichanthelium and African species (NADP–ME, PEP–CK and NAD–ME) in subtribes
of ‘‘Panicum’’ sect. Clavelligerae. Although the Dichan- Cenchrinae, Melinidinae and Panicinae. This sister
thelium–Clavelligerae subclade is well supported in our relationship is present in the strict consensus tree of
analyses, with 15 bp substitutions and jackknife values both ndhF and the combined analyses, and could
of 99 ⁄ 100, its final position and relationships with other represent a link to a non-Kranz ancestor for the three
Paniceae are uncertain. Species of Dichanthelium (ca. 55) C4 subtribes.
are characterized by having inflorescences with cleistog-
amous and chasmogamous flowers, and many species Character evolution
with foliar dimorphism. Species of ‘‘Panicum’’ sect.
Clavelligerae (ca. 10) share the presence of spikelets with Photosynthetic pathway. In all optimal trees, from both
conspicuous glandular hairs. To confirm relationships the combined data set and the ndhF sequence analyses,
within this subclade, it will be necessary to study more C4 photosynthesis is plesiomorphic for the Paspaleae–
representatives of Dichanthelium and Clavelligerae. Andropogoneae–Arundinelleae s.s. clade, and is fol-
lowed by four reversals to C3. This unambiguous
Subclade B (Poecilostachys–Sacciolepis–Trichanthe- optimization is obtained because the C4 species Rey-
cium–Panicum incertae sedis) naudia is the sister group to the Paspaleae in all optimal
trees. When analysing the combined data set, all three
Subclade B comprises the genera Sacciolepis Nash main clades within the Paspaleae (Arthropogoninae,
and Trichanthecium (Zuloaga et al., 2011), together with Otachyriinae and Paspalinae) are ambiguous concerning
species of Panicum incertae sedis of sections Monticola the plesiomorphic state of the photosynthetic pathway.
and Verrucosa (Aliscioni et al., 2003), and ungrouped For these three clades, the ambiguous optimization is
species of ‘‘Panicum’’. Most of the taxa included here are resolved in favour of a plesiomorphic C4 pathway due to
non-Kranz, have a disarticulation zone at the base of the the position of Reynaudia and the Andropogoneae–
upper anthecium, and have bicellular microhairs at the Arundinelleae s.s. clade (see optimization in Fig. 2). The
apex or all over the anthecium surface. Morphological position of Reynaudia is weakly to moderately sup-
synapomorphies need to be explored for better charac- ported, and remains as incertae sedis (see above). If
terization of all species of the clade. Also, relationships Reynaudia was placed elsewhere in the tree, or was not
among species of this clade and those of subtribe included, the ancestral photosynthetic pathway for the
Boivinellinae should be scrutinized with the addition of Paspaleae–Andropogoneae–Arundinelleae s.s. clade as
other American and African species. The genus Poe- well as the Paniceae s.s. would be unambiguously C3,
cilostachys Hack. is embedded in this clade only when and there would be multiple origins of C4. The latter
using molecular data, with a jackknife value of 81. In the result is consistent with the previous work of Christin
combined analysis, Poecilostachys and Hylebates Chip- et al. (2008) and Vicentini et al. (2008).
pin. are sister taxa, although this subclade is unsup- When analysing the combined data set, Anthephorinae
ported. Both genera occur in forest shade areas and are is basal in the Paniceae s.s. This topology causes an
characterized by their broad leaves and open panicles. ambiguous optimization of the photosynthetic pathway
As a result, the relationship of both genera with other at the root of Paniceae s.s., as well as an ambiguous
Paniceae s.s. is still not resolved. optimization of the entire Paniceae s.l. (Paniceae s.s. +
Paspaleae) + Andropogoneae–Arundinelleae s.s. clade.
Subclade C (Walwhalleya–Homopholis) Note that these two ambiguous optimizations are
resolved in favour of a plesiomorphic C3 photosynthesis
Homopholis C.E. Hubb. is a small genus with two both in Paniceae s.s. and at the base of the Paniceae s.l.–
species endemic to Australia (Webster, 1987). Recently, Andropogoneae–Arundinelleae s.s. clade in some trees
A B
Fig. 2. Character evolution within the subfamily Panicoideae s.l. shown as phylograms for 35 spikelet characters (A) and 17 inflorescence characters
(B) when optimized on one of the optimal trees for the combined data set. Changes in the photosynthetic pathway have been optimized on both
phylograms. Line drawings represent morphological variations in the spikelets (1–13) and quantitatively modified inflorescences in subtribes Paspalinae,
Anthephorinae and Cenchrinae. References: 1, Oplismenopsis najada; 2, Otachyrium versicolor; 3, 15, Hymenachne amplexicaulis; 4, Paspalum arenarium;
5, Paspalum lilloi; 6, Paspalum simplex; 7, Digitaria ciliaris; 8, 20, Lasiacis sorghoidea; 9, Urochloa fusca; 10, Melinis repens; 11, Panicum elephantipes; 12,
25, Setaria parviflora; 13, Cenchrus myosuroides; 14, Coleataenia prionitis; 16, Paspalum densum; 17, Paspalum lachneum; 18, Paspalum ceresia; 19,
Anthephora hermaphrodita; 21, Melinis minutiflora; 22, Urochloa plantaginea; 23, Panicum peladoense; 24, Cenchrus spinifex; 26, Setaria verticilliformis.
when the C3 species Hydrothauma manicatum is included contributing to the branch lengths in these clades or in
in the matrix. In the three possible placements of subtribes Melinidinae and Panicinae (Fig. 2B). Qualita-
H. manicatum (see Results), an ambiguous optimization tive modifications of the inflorescences seem to have
of the photosynthetic pathway is retained only when occurred nearly exclusively within the major C4 XyMS
H. manicatum is placed as sister to the entire Paniceae clades Paspalinae, Anthephorinae and Cenchrinae
s.l.–Andropogoneae–Arundinelleae s.s. clade. Obtaining (Fig. 2B, 16–19, 24–26), the one exception being the
a DNA sample for Hydrothauma could therefore poten- highly modified inflorescences in Thuarea (Melinidinae).
tially add information concerning the plesiomorphic A relationship between C4 XyMS photosynthesis and
photosynthetic pathway within the Paniceae s.s. modified inflorescences is reported here for the first
time. The basis for such a co-evolution is uncertain, but
Morphological characters. Paniceae s.l. includes an enor- the presence of unmodified inflorescences at the base of
mous morphological diversity, mainly recognized as the major C4 XyMS clades suggests that the modifica-
different patterns of spikelet and inflorescence type. tion of the inflorescences in terminal clades would be a
Changes in the spikelets, such as size and nervation delayed response to the change in photosynthetic
of the glumes, as well as texture and ornamentation of pathway (see Giannini and Goloboff, 2010 for a
lemma and palea, were among the most common discussion on delayed responses of phylogenetic corre-
morphological modifications and constituted nearly lated characters).
two-thirds of the coded morphological characters. Illus-
trations of the morphological variability of the spikelets
within the tribes Paspaleae and Paniceae s.s. are shown in Taxonomic considerations
Fig. 2A. Branch lengths in Fig. 2A show that changes in
spikelet characters have occurred in all subtribes. Major Based on our results, we propose changes to the
groups are defined by some changes, such as papillae at classification of Panicoideae to realign the genera of the
the apex of the upper palea in the Panicinae; absence of subfamily with particular emphasis on the tribe Paniceae
lower glume in most species of the Paspalinae (Fig. 2A, 4– s.l. Hubbardia, treated under the monotypic tribe
6); and reduction in number of nerves in the glume in all Hubbardieae in Panicoideae, is now included in the
species of Anthephorinae (Fig. 2A, 7). However, most subfamily Micrairoideae. Tribe Paniceae s.l. must
spikelet characters characterize minor clades within be split into two tribes: Paspaleae, to include the
subtribes, such as awned glumes in Oplismenopsis, Cana- American x = 10 Paniceae genera; and Paniceae s.s.,
stra, Arthropogon, Altoparadisium and Achlaena (Arthro- to include the Pantropical x = 9 Paniceae genera.
pogoninae, Fig. 2A, 1); expanded lower palea in Within this classification we propose a new realignment
Otachyrium, Plagiantha and Steinchisma (Otachyriinae, of subtribes based on molecular and morphological
Fig. 2A, 2); and herbaceous lemma and palea in Hym- synapomorphies.
enachne (Otachyriinae, Fig. 2A, 3). Tribe Paspaleae J. Presl, Reliq. Haenk. 1(4–5): 208.
The inflorescences show fewer qualitative changes 1830. TYPE: Paspalum L., Syst. Nat. (ed. 10) 2: 846,
than the spikelets do within the tribes Paspaleae and 855, 1359. 1759.
Paniceae s.s. In general, most of the variation among the = Lecomtelleae Pilg. ex Potztal, Willdenowia 1: 771.
inflorescences is found in quantitative changes of the 1957. TYPE: Lecomtella A. Camus, Compt. Rend.
lateral branching as well as internode length within Hebd. Séances Acad. Sci. 181: 567. 1925.
the panicles, yielding large, lax, multiflorous to smaller, We propose to reinstate the tribe Paspaleae for the
more compressed, and few flowered inflorescences. species in the Paniceae x = 10 clade. The genera of this
Qualitative modifications of the inflorescences within clade are mostly native to the Americas, with a basic
Paniceae s.l. include abortion of the fertile apex of the chromosome number of x = 10. Ecologically and
main axes and rachises in Paspalinae, Anthephorinae morphologically, the group is remarkably variable and
and Cenchrinae; lack of branching beyond first order; includes species that are non-Kranz, species that are
and modification of the disarticulation zone of the Kranz with the C4 NADP–ME subtype, and species that
diaspores. The plesiomorphic diaspore involves disar- are physiologically intermediate between C3 and C4.
ticulation at the base of the spikelet, such that the Subtribe Arthropogoninae Butzin, Willdenowia 6(3):
caryopsis falls together with the glumes. This disartic- 516. 1972. TYPE: Arthropogon Nees, Fl. Bras. Enum.
ulation has been modified towards more inclusive Pl. 2(1): 319. 1829.
diaspores that articulate at the base of the primary Photosynthetic pathway. C3, C4 NADP–ME.
branches or main axes. For more thorough description The subtribe includes 16 genera and approximately 68
of these characters and their morphological implications American species:
in each clade, see the discussion of the subtribes above. Achlaena (1), Altoparadisium (2), Apochloa (15),
Notably, qualitative inflorescence character changes Arthropogon (3), Canastra (2), Coleataenia (7), Cypho-
are nearly absent in several of the major C3 clades, not nanthus (1), Homolepis (5), Keratochlaena (1), Mesosetum
(25), Oncorachis (2), Oplismenopsis (1), Phanopyrum (1), 1984. TYPE: Paspalidium Stapf, Fl. Trop. Afr. 9: 582.
Stephostachys (1), Tatianyx (1), Triscenia (1). 1920.
Subtribe Otachyriinae Butzin, Willdenowia 6: 182. = Subtribe Spinificinae Ohwi, Acta Phytotax. Geo-
1970. TYPE: Otachyrium Nees, Fl. Bras. Enum. Pl. 2(1): bot. 11(1): 56. 1942. TYPE: Spinifex L., Mant. Pl.: 163,
271 (misprinted as p. 273). 1829. 300. 1771.
Photosynthetic pathway. C3, C4 NADP–ME and C3– = Subtribe Uranthoeciinae Butzin, Willdenowia 6:
C4 intermediates. 182. 1970. TYPE: Uranthoecium Stapf, HookerÕs Icon.
The subtribe includes six genera and incertae sedis Pl. 31: t. 3073. 1916.
species of ‘‘Panicum’’ sect. Laxa; five genera are Amer- = Subtribe Xerochloinae Butzin, Willdenowia 6: 184.
ican, and one endemic to Australia. The subtribe has a 1970. TYPE: Xerochloa R. Br., Prodr. 196. 1810.
total of seven genera and approximately 38 species: = Subtribe Snowdeniinae Butzin, Willdenowia 6(3):
Anthaenantia (4), Dallwatsonia (1), Hymenachne (7), 516. 1972. TYPE: Snowdenia C.E. Hubb., Bull. Misc.
Otachyrium (7), Panicum group Laxa (12), Plagiantha Inform. Kew 1929(1): 30. 1929.
(1), Steinchisma (6). = Subtribe Pseudoraphidinae Keng and Keng f.,
Subtribe Paspalinae Griseb., Spic. Fl. Rumel. 2: 468. Bull. Nanjing Bot. Gard. 1988-1989: 7. 1990. TYPE:
1846. TYPE: Paspalum L., Syst. Nat. (ed. 10) 2: 846, Pseudoraphis Griff., Not. Pl. Asiat. 3: 29. 1851.
855, 1359. 1759. Photosynthetic pathway. C4 NADP–ME.
= Reimarochloinae Caro, Dominguezia 4: 41. 1982. The subtribe includes 26 genera and approximately
TYPE: Reimarochloa Hitchc., Contr. U.S. Natl. Herb. 328 species;
12(6): 198. 1909. Acritochaete (1), Alexfloydia (1), Cenchrus (110),
Photosynthetic pathway. C3 and C4 NADP–ME. Chamaeraphis (1), Dissochondrus (1), Holcolemma (4),
The subtribe includes 19 genera, mainly from Amer- Hygrochloa (2), Ixophorus (1), Panicum antidotale,
ica, with two Old World genera: Lecomtella and Paractaenum (1), Paratheria (2), Plagiosetum (1), Pseud-
Baptorhachis, and approximately 475 species: ochaetochloa (1), Pseudoraphis (6), Setaria (160), Setari-
Acostia (1), Anthaenantiopsis (4), Axonopus (85), opsis (2), Snowdenia (4), Spinifex (4), Stenotaphrum (7),
Baptorhachis (1), Centrochloa (1), Echinolaena (8), Stereochlaena (5), Streptolophus (1), Uranthoecium (1),
Gerritea (1), Hopia (1), Ichnanthus (30), Lecomtella (1), Whiteochloa (5), Xerochloa (4), Zuloagaea (1), Zygo-
Ocellochloa (12), Ophiochloa (2), Panicum s.l. (2), chloa (1).
Paspalum (310), Reimarochloa (3), Renvoizea (10), Subtribe Melinidinae (Hitchc.) Pilg., Nat. Pflanzen-
Spheneria (1), Streptostachys (2), Thrasyopsis (2). fam. (ed. 2) 14e: 95. 1940. Tribe Melinideae Hitchc.,
Tribe Paniceae R. Br., Voy. Terra Austral. 2: 582. U.S.D.A. Bull. (1915-23) 772: 18. 1920. TYPE: Melinis
1814. TYPE: Panicum L., Sp. Pl.: 55. 1753. P. Beauv., Ess. Agrostogr.: 54. 1812.
Subtribe Anthephorinae Benth., J. Linn. Soc., Bot. 19: = Subtribe Tristegininae Harv., Gen. S. Afr. Pl. (ed.
30. 1881. TYPE: Anthephora Schreb., Beschr. Gras. 2: 2): 428. 1869, nom. illeg. TYPE: Tristegis Nees, Horae
105. 1810. Phys. Berol: 47, 54. 1820, nom. illeg. superfl.
= Unranked Digitariastrae Stapf, Fl. Trop. Afr. 9: = Subtribe Brachiariinae Butzin, Willdenowia 6: 189.
12. 1917. Subtribe Digitariinae (Stapf) Butzin, Willden- 1970. TYPE: Brachiaria (Trin.) Griseb., Fl. Ross. (14):
owia 6: 509. 1972. TYPE: Digitaria Haller, Hist. Stirp. 469. 1853.
Helv. 2: 244. 1768, nom. cons. Photosynthetic pathway. C4, PEP–CK.
Photosynthetic pathway. C4 NADP–ME. The subtribe includes 14 genera and approximately
The subtribe includes seven genera and approximately 210 species:
238 species: Chaetium (3), Eccoptocarpha (1), Eriochloa (30),
Anthephora (12), Chaetopoa (2), Chlorocalymma (1), Leucophrys (1), Megathyrsus (3), Melinis (27), Mooro-
Digitaria (220), Megaloprotachne (1), Tarigidia (1), chloa (3), Panicum deustum, Rupichloa (2), Scutachne (2),
Trachys (1). Thuarea (2), Tricholaena (4), Urochloa (130), Yvesia (1).
Subtribe Cenchrinae (Dumort.) Dumort., Anal. Fam. Subtribe Boivinellinae (A. Camus) Pilg., Nat. Pfl.-Syst.
Pl. 64. 1829. Unranked Cenchrinae Dumort., Observ. (ed. 2) 14e: 101. 1940. Tribe Boivinelleae A. Camus, Bull.
Gramin. Belg.: 139.1824. TYPE: Cenchrus L., Sp. Pl. 2: Mus. Hist. Nat. 31: 393. 1925. TYPE: Boivinella A.
1049. 1753. Camus, Bull. Soc. Bot. France 72: 175. 1925.
= Subtribe Setariinae (Dumort.) Dumort., Anal. = Subtribe Microcalaminae Butzin, Willdenowia 6:
Fam. Pl. 64. 1829. Unranked Setariinae Dumort., 189. 1970. TYPE: Microcalamus Franch., J. Bot. (Mo-
Observ. Gramin. Belg.: 137. 1824. TYPE: Setaria P. rot) 3(17): 282, f. b. 1889.
Beauv., Ess. Agrostogr. 51, 178, pl. 13, f. 3. 1812, nom. Photosynthetic pathway. C3, C4 NADP–ME and C3–
cons. and typ. cons. C4 intermediates.
= Subtribe Paspalidinae Keng and Keng f. ex S.L. The subtribe includes 13 genera and approximately
Chen and Y.X. Jin, Acta Phytotax. Sin. 22(6): 475. 129 species:
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of this article, our friend and renown botanist, Dr Poaceae subfamily Pooideae based on chloroplast ndhF gene
Osvaldo Morrone. sequences. Mol. Phylogenet. Evol. 8, 150–166.
This research was supported by ANPCyT (grant nos. Chemisquy, M.A., Giussani, L.M., Scataglini, M.A., Kellogg, E.A.,
32664 and 01286). Field collections were carried out by Morrone, O., 2010. Phylogenetic studies favor the unification of
Pennisetum, Cenchrus and Odontelytrum: a nuclear, chloroplast and
funds awarded by the National Geographic Society (grant
morphological combined analysis. Ann. Bot. 106, 107–130.
nos. 7792-05 and 8365-07) and by the Myndel Botanical Christin, P.A., Salamin, N., Savolainen, V., Duvall, M.R., Besnard,
Foundation. We also thank the staff of the Darwinion, G., 2007. C4 photosynthesis evolved in grasses via parallel adaptive
who kindly helped us throughout, and Mariana Valente genetic changes. Curr. Biol. 17, 1241–1247.
for her assistance in the preparation of figures. Christin, P.A., Besnard, G., Samaritani, E., Duvall, M.R., Hodkinson,
T.R., Savolainen, V., Salamin, N., 2008. Oligocene CO2 decline
promoted C4 photosynthesis in grasses. Curr. Biol. 18, 37–43.
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Supporting Information rone, F499149. Cenchrus ciliaris L., AY029625. Cench-
rus compressus (R. Br.) Morrone, AY029672. Cenchrus
Additional Supporting Information may be found in flaccidus (Griseb.) Morrone, AF499150. Cenchrus myos-
the online version of this article: uroides Kunth, AF499152. Cenchrus pilosus Kunth,
Appendix S1. Morphological matrix used in the AY623746. Cenchrus setigerus Vahl, AF499153. Centot-
analyses. Numbers in the first row represent the char- heca lappacea (L.) Desv., AY847122. Centrochloa sing-
acters described in the Appendix 2, together with their ularis Swallen, Rua 790 (BAA), JN604678. Centropodia
coding. Numbers within brackets show a polymorphism glauca (Nees) Cope, AF251462. Chaetium bromoides
for the particular character of the species involved. (J. Presl) Benth. ex Hemsl., AY029626. Chaetopoa pilosa
Dashes indicate character inapplicable for a taxon. Clayton, Bidgood et al. G-59 (K), JN604679. Chamae-
Please note: Wiley-Blackwell is not responsible for the raphis hordeacea R. Br., Craven 4098 (CAMB),
content or functionality of any supplementary materials JN604680. Chasmanthium curvifolium (Valdés-Reyna,
supplied by the authors. Any queries (other than missing Morden and S.L. Hatch) Wipff and S.D. Jones,
material) should be directed to the corresponding author AY847125. Chasmanthium latifolium (Michx.) H.O.
for the article. Yates, AY029694. Chasmanthium laxum (L.) H.O.
Yates, AY847126. Chionachne koenigii (Spreng.) Thwa- pallens (Sw.) Munro ex Benth., AY029638. Isachne
ites, AF117397. Chlorocalymma cryptacanthum Clayton, arundinacea (Sw.) Griseb., AY847119. Isachne leersio-
Greenway and Kanuri 14210 (MO), JN604681. Chryso- ides Griseb., Zuloaga and Morrone 9572 (SI), JN604692.
pogon fulvus (Spreng.) Chiov., AF117398. Cleistachne Ixophorus unisetus (J. Presl) Schltdl., AY623749. Kar-
sorghoides Benth., AF117400. Cleistochloa rigida (S.T. roochloa purpurea (L. f.) Conert and Türpe, AF251458.
Blake) R.D. Webster, Thompson BIL-160 (BRI), Keratochlaena rigidifolia (Filg., Morrone and Zuloaga)
JN604682. Coleataenia anceps (Michx.) Soreng, Morrone and Zuloaga, EU805492. Lasiacis sorghoidea
AY188455. Coleataenia longifolia (Torr.) Soreng, (Desv. ex Ham.) Hitchc. and Chase, AY029639. Leu-
AY188482. Coleataenia petersonii (Hitchc. and Ekman) cophrys mesocoma (Nees) Rendle, GU594628. Loph-
Soreng, AY188479. Coleataenia prionitis (Nees) Soreng, atherum gracile Brongn., AY847129. Loudetia simplex
AY029652. Coleataenia tenera (Beyr. ex Trin.) Soreng, (Nees) C.E. Hubb., AY847117. Louisiella fluitans C.E.
AY188491. Cymbopogon flexuosus (Nees ex Steud.) Hubb. and J. Léonard, Fay 6160 (MO), JN604693*.
Will.Watson, AF117404. Cyperochloa hirsuta Lazarides Megaloprotachne albescens C.E. Hubb., Hansen 3382
and L. Watson, AY847139. Cyphochlaena madagascar- (MO), JN604694. Megastachya mucronata (Poir.) P.
iensis Hack., Pignal 1146 (MO), JN604683. Cyphonan- Beauv., AY847123. Megathyrsus maximus (Jacq.) B.K.
thus discrepans (Döll) Zuloaga and Morrone, Simon and S.W.L. Jacobs, AY029649. Melinis repens
DQ646392. Cyrtococcum accrescens (Trin.) Stapf, Hu (Willd.) Zizka, AY029675. Merxmuellera macowanii
and But 23191 (MO), JN604684. Dallwatsonia felliana (Stapf) Conert, AF251457. Mesosetum chaseae Luces,
B.K. Simon, Clarkson and Neldner 9872 (BRI), AY029641. Micraira subulifolia F. Muell., AY622316.
JN604685. Danthonia californica Bol., AF251459. Microstegium nudum (Trin.) A. Camus, AF443813.
Danthoniopsis dinteri (Pilg.) C.E. Hubb., AY847116. Mnesithea selloana (Hack.) de Koning and Sosef,
Dichanthelium acuminatum (Sw.) Gould and C.A. Clark, AF117401. Molinia caerulea (L.) Moench, U21995.
AY188485. Dichanthelium clandestinum (L.) Gould, Moorochloa eruciformis (Sm.) Veldkamp, AY188452.
AY188461. Dichanthelium cumbucana (Renvoize) Zulo- Neurachne alopecuroidea R. Br., Kellogg 1029 (MO)
aga, AY188464. Dichanthelium koolauense (H. St. John JN604695. Ocellochloa chapadensis (Swallen) Zuloaga
and Hosaka) C.A. Clark and Gould, AY029627. and Morrone, AY188486. Ocellochloa piauiensis (Swal-
Dichanthelium sabulorum (Lam.) Gould and C.A. Clark, len) Zuloaga and Morrone, AY029656. Ocellochloa
AY029654. Dichanthium aristatum (Poir.) C.E. Hubb., stolonifera (Poir.) Zuloaga and Morrone, AY188488.
AF117409. Digitaria ciliaris (Retz.) Koeler, AY029630. Oncorachis ramosa (Zuloaga and Soderstr.) Morrone
Digitaria didactyla Willd., AM849203. Digitaria radico- and Zuloaga, AY029686. Ophiochloa hydrolithica Filg.,
sa (J. Presl) Miq., AY029628. Digitaria setigera Roth, Davidse and Zuloaga, AY029642. Oplismenopsis najada
AY029629. Dissochondrus biflorus Kuntze ex Hack., (Hack. and Arechav.) Parodi, AY188453. Oplismenus
Wood and Perlaan 1998 (MO), JN604686. Distichlis hirtellus (L.) P. Beauv., AY029644. Orthoclada laxa
spicata (L.) Greene, AF251464. Echinochloa colona (L.) (Rich.) P. Beauv., AY847128. Otachyrium versicolor
Link, AY029631. Echinochloa frumentacea Link, (Döll) Henrard, AY029643. Ottochloa nodosa (Kunth)
AY029632. Echinolaena inflexa (Poir.) Chase, Dandy, Maxwell 89-561 (MO), JN604696. Panicum
AY029633. Elionurus muticus (Spreng.) Kuntze, adenophorum K. Schum., AY188454. Panicum antidotale
AF117410. Entolasia stricta (R. Br.) Hughes, Snow Retz., AY188456. Panicum aquaticum Poir., AY029658.
and Simon 7297 (MO), JN604687. Eragrostis curvula Panicum bergii Arechav., AY188457. Panicum cervica-
(Schrad.) Nees, U21989. Eriachne pulchella Domin, tum Chase, AY188459. Panicum chloroleucum Griseb.,
AY618659. Eriochloa punctata (L.) Desv. ex Ham., AY188460. Panicum claytonii Renvoize, AY188462.
AY029634. Gerritea pseudopetiolata Zuloaga, Morrone Panicum deustum Thunb., GU594631. Panicum dicho-
and Killeen, Morrone et al. 4219 (SI), JN604688. Gyne- tomiflorum Michx., AY188466. Panicum elephantipes
rium sagittatum (Aubl.) P. Beauv., AY847120. Hetero- Nees ex Trin., AY029647. Panicum fauriei Hitchc.,
pogon contortus (L.) P. Beauv. ex Roem. and Schult., AY029650. Panicum hylaeicum Mez, AY188470. Pani-
AF117411. Homolepis glutinosa (Sw.) Zuloaga and cum miliaceum L., AY188472. Panicum millegrana Poir.,
Soderstr., AY029637. Homolepis isocalycia (G. Mey.) AY029660. Panicum mystasipum Zuloaga and Morrone,
Chase, AY029636. Homopholis belsonii C.E. Hubb., AY188474. Panicum nephelophilum Gaudich.,
Blake 13252 (BRI), JN604689*. Hopia obtusa (Kunth) AY029645. Panicum olyroides Kunth, AY188475. Pan-
Zuloaga and Morrone, AY029659. Hygrochloa aquatica icum pedersenii Zuloaga, AY029646. Panicum pilosum
Lazarides, Craven 2379 (CANB), JN604690*. Hylebates Sw., AY188480. Panicum racemosum (P. Beauv.) Spr-
cordatus Chippindall, Davidse 73-2633 (MO), JN604691. eng., AY188481. Panicum repens L., AY188467. Pani-
Hymenachne donacifolia (Raddi) Chase, AY029635. cum rudgei Roem. and Schult., AY029661. Panicum
Hymenachne grumosa (Nees) Zuloaga, AY188468. Hym- sellowii Nees, AY188484. Panicum stramineum Hitchc.
enachne pernambucensis (Spreng.) Zuloaga, AY188478. and Chase, AY188489. Panicum trichanthum Nees,
Hyparrhenia hirta (L.) Stapf, AF117412. Ichnanthus AY188492. Panicum tricholaenoides Steud., AY188493.
Panicum tuerckheimii Hack., AY188494. Panicum vali- Pilg., Hill and Horn 27203 (MO), JN604707. Spinifex
dum Mez, AY188495. Panicum verrucosum Muhl., sericeus R. Br., EF189895. Sporobolus indicus (L.) R.
AY188496. Panicum virgatum L., U21986. Paractaenum Br., U21983. Steinchisma decipiens (Nees ex Trin.) W.V.
novae-hollandiae P. Beauv., Speek 1264 (CANB), Br., AY188499. Steinchisma hians (Elliott) Nash,
JN604697. Paraneurachne muelleri (Hack.) S.T. Blake, AY029685. Steinchisma laxa (Sw.) Zuloaga,
Newbey 10800 (MO), JN604698. Paratheria prostrata AY029655. Steinchisma spathellosa (Döll) Renvoize,
Griseb., Laergaard et al. 16915 (C), JN604699*. Paro- AY188500. Stenotaphrum secundatum (Walter) Kuntze,
diophyllochloa cordovensis (E. Fourn.) Zuloaga and AY029684. Stephostachys mertensii (Roth) Zuloaga and
Morrone, AY188463. Parodiophyllochloa missiona (Ek- Morrone, AY188471. Stereochlaena cameronii (Stapf)
man) Zuloaga and Morrone, AY188473. Parodiophyllo- Pilg., Davidse and Handlos 7214 (MO), JN604708*.
chloa ovulifera (Trin.) Zuloaga and Morrone, Steyermarkochloa angustifolia (Spreng.) Judz., Berry
AY029653. Parodiophyllochloa penicillata (Nees ex et al. 6016 (MO), JN604709. Stipagrostis zeyheri (Nees)
Trin.) Zuloaga and Morrone, AY188477. Paspalum De Winter, AF251455. Streptostachys asperifolia Desv.,
arundinellum Mez, AY029663. Paspalum conjugatum AY029687. Tatianyx arnacites (Trin.) Zuloaga and
P.J. Bergius, AY029669. Paspalum conspersum Schrad., Soderstr., AY029688. Thrasyopsis juerguensii (Hack.)
AY029666. Paspalum foliiforme S. Denham, AY029690. Soderstr. and A.G. Burm., Rua 728 (BAA), JN604711.
Paspalum glaziovii (A.G. Burm.) S. Denham, Thuarea involuta (G. Forst.) R. Br. ex Sm., GU594624.
AY029689. Paspalum haumanii Parodi, AY029664. Thyridolepis mitchelliana (Nees) S.T. Blake, Latz 13500
Paspalum malacophyllum Trin., AY029671. Paspalum (MO), JN604710. Thysanolaena maxima (Roxb.) Kuntze,
paniculatum L., AY029667. Paspalum remotum J. Rémy, U21984. Trichanthecium cyanescens (Lam.) Zuloaga and
AY029668. Paspalum vaginatum Sw., AY029665. Pasp- Morrone, AY188465. Trichanthecium parvifolium (Lam.)
alum virgatum L., AY029670. Phacelurus digitatus (Sib- Zuloaga and Morrone, AY188476. Trichanthecium wett-
th. and Sm.) Griseb., AF117418. Phanopyrum steinii (Hack.) Zuloaga and Morrone, Y188497. Trichola-
gymnocarpon (Elliott) Nash, AY188469. Pheidochloa ena monachne (Trin.) Stapf and C.E. Hubb., FJ486535.
gracilis S.T. Blake, Lazarides 8871 (MO), JN604700. Tripsacum dactyloide, AF117433. Triscenia ovina (L.) L.,
Phragmites australis (Cav.) Trin. ex Steud., U21997. Ekman 989 (MO), JN604712*. Tristachya biseriata Stapf,
Plagiantha tenella Renvoize, AY029674. Plagiosetum AY847118. Uniola paniculata Stapf, AF251463. Urant-
refractum Benth., EU819409. Poecilostachys oplismeno- hoecium truncatum (Maiden and Betche) Stapf, SJ9599.
ides (Hack.) Clayton, Carvalho 4202 (C), JN604701*. Urochloa mutica (Forssk.) T.Q. Nguyen, AY029691.
Pohlidium petiolatum Davidse, Soderstr. and R.P. Ellis, Walwhalleya subxerophila (Domin) Wills and J.J. Bruhl,
AY847130. Pseudechinolaena polystachya (Kunth) Snow and Simon 7316 (MO), JN604713. Whiteochloa
Stapf, AY029676. Pseudochaetochloa australiensis capillipes (Benth.) Lazarides, Risler 1804 (MO),
Hitchc., Simon 3826 (CAMB), JN604702. Pseudoraphis JN604714. Xerochloa laniflora Benth., Latz 11688 (MO),
paradoxa Pilg., EF189892. Reimarochloa acuta (Flüggé) JN604715. Yakirra australiensis (Domin) Lazarides and
Hitchc., Zuloaga and Morrone 9537 (SI), JN604703. R.D. Webster, Latz 12164 (MO), JN604716. Yvesia
Renvoizea trinii (Kunth) Zuloaga and Morrone, madagascariensis A. Camus, GU594636. Zeugites capil-
EU107783. Reynaudia filiformis (Spreng. ex Schult.) laris (Hitchc.) Swallen, AY847133. Zeugites pittieri Hack.,
Kunth, Zuloaga and Morrone 9538 (SI), JN604704. AY632374. Zoysia matrella (L.) Merr., U21975. Zuloa-
Rupichloa acuminata (Renvoize) D. Salariato and Mor- gaea bulbosa (Kunth) Bess, AY029648. Zygochloa parad-
rone, AY029692. Sacciolepis indica (L.) Chase, oxa (R. Br.) S.T. Blake, EF189896.
AY029677. Schizachyrium scoparium (Michx.) Nash,
AF117420. Scutachne dura (Griseb.) Hitchc. and Chase, Appendix 2
GU594616. Setaria barbata (Lam.) Kunth, AF499145.
Setaria geminata (Forssk.) Veldkamp, AY029662. Se- Characters and coding; a description of the characters
taria grisebachii E. Fourn., AF499141. Setaria italica and character states presented as considered in the
(L.) P. Beauv., AF499140. Setaria lachnea (Nees) analyses. Characters 22, 28, 32, 42, 44 and 45 were
Kunth, AY029683. Setaria macrostachya Kunth, coded additive. *, Characters and character states that
AY029678. Setaria palmifolia (J. König) Stapf, differ from Zuloaga et al. (2000).
AY029680. Setaria verticillata (L.) P. Beauv.,
AF499139. Setaria viridis (L.) P. Beauv., U21976. Inflorescence
Setariopsis auriculata (E. Fourn.) Scribn., Cóbar and *Char. 0. Main axis of inflorescences: terminating in a
Garcı´a 1082 (MO), JN604705. Snowdenia petitiana (A. spikelet (0), terminating in a bristle (1), terminating in a
Rich.) C.E. Hubb., Barney 113 (K), JN604706*. Sor- naked point, without a bristle (2).
ghastrum nutans (L.) Nash, AF117421. Spartina pecti- *Char. 1. Rachis: terminating in a spikelet (0), termi-
nata Link, AF251465. Spartochloa scirpoidea (Steud.) nating in a bristle (1), terminating as a foliaceous axis (2),
C.E. Hubb., AY847140. Spheneria kegelii (Müll. Hal.) terminating in a naked point, without a bristle (3).
Char. 2. Foliaceous rachis: absent (0), present (1). Floret

Char. 3. Bracts of the inflorescences: absent (0),
present (1). Char. 30. Lower lemma: muticous (0), awned (1).
Char. 4. Involucral bristles (cauline): absent (0), Char. 31. Apex of the lower lemma: entire (0), bifid (1),
present (1). trilobate (2).
Char. 5. Involucral bracts: absent (0), present (1). Char. 32. Lower palea: absent (0), present, not
Char. 6. Cleistogenes in leaf axils: absent (0), present (1). expanded (1), present and expanded (2).
Char. 7. Apex of the pedicel: truncate (0), oblique (1). *Char. 33. Upper anthecium: dorsiventrally com-
Char. 8. Disarticulation at the base of the spikelet: pressed (0), laterally compressed (1), cylindrical (2).
absent (0), present (1). *Char. 34. Upper anthecium: crustaceous (0), cartilag-
Char. 9. Disarticulation at the base of the primary inous (1), hyaline (2), membraneous to herbaceous (3).
branches: absent (0), present (1). Char. 35. Upper lemma texture: smooth (0), trans-
Char. 10. Disarticulation at the base of the inflores- versely rugose (1).
cence: absent (0), present (1). Char. 36. Upper lemma with basal scars or appendages:
Char. 11. Disarticulation between the lower glume and absent (0), present (1).
lower lemma: absent (0), present (1). Char. 37. Upper lemma: muticous (0), awned (1).
Char. 12. Disarticulation at the base of the pedicel: Char. 38. Upper lemma: not differentiated at the apex
absent (0), present (1). (0), differentiated at the apex (1).
Char. 13. Disarticulation at the node of the main axis: Char. 39. Upper lemma (margins): tucked in onto the
absent (0), present (1). palea (0), lying flat and exposed on the palea (1).
Char. 14. Disarticulation at the base of the upper Char. 40. Upper palea: absent (0), present (1).
anthecium: absent (0), present (1). Char. 41. Upper palea: tightly clasped by the lemma
*Char. 15. Unilateral spikelet: absent (0), present (1). (0), gaping (1).
*Char. 16. Second order branches in inflorescence: *Char. 42. Upper palea: without simple papillae (0),
multiflowered (0), reduced to a single abaxial spikelet with simple papillae (1), with compound papillae (2).
(1), reduced to a single adaxial spikelet (2). *Char. 43. Distribution of papillae in the upper palea:
all over surface (0), at the apex only (1).
Spikelet *Char. 44 Upper palea (presence of microhairs):
without bicellular microhairs (0), with bicellular micro-
*Char. 17. Spikelet shape: not gibbous (0), gibbous (1). hairs at the apex and ⁄ or base (1), with bicellular
*Char. 18. Presence of incomplete florets distal to microhairs all over its surface (2).
fertile florets: absent (0), present (1). *Char. 45. Upper palea (presence of macrohairs):
Char. 19. Breeding system: plants with at least with some without macrohairs (0), with macrohairs at the apex
perfect flowers (0), no perfect flowers, staminate and and ⁄ or base (1), with macrohairs all over its surface (2).
pistillate flowers on the same plant (1), no perfect flowers, Char. 46. Upper palea (apex): straight (0), recurved
staminate and pistillate flowers on different plants (2). (1).
Char. 20. Hairy callus at the base of the spikelet: Char. 47. Stamen (number): three (0), two (1), one (2).
absent (0), present (1). Char. 48. Lodicules : present (0), absent (1).
*Char. 21. Stipe at the base of the upper floret: absent Char. 49. Style base: free (0), fused (1).
(0), present (1). Char. 50. Ovary: glabrous (0), pilose (1).
Glumes Fruit
Char. 22. Lower glume: present (0), vestigial (1), *Char. 51. Caryopsis (hilum): punctiform (0), linear (1).
absent (2).
Char. 23. Lower glume: not saccate (0), saccate (1). Leaf anatomy
Char. 24. Lower glume: muticous (0), awned (1).
Char. 25. Upper glume: present (0), absent (1). *Char. 52. Photosynthetic pathway: C3 XyMS+ (0),
Char. 26. Lower and upper glume: of different size (0), C4 XyMS– (1), C4 XyMS+ (2).
the same size (1). *Char. 53. Chloroplasts on the parenchymatous sheath:
*Char. 27. Upper glume: as long as or longer than the absent (0), centripetal (1), centrifugal (2).
lower lemma (0), 1 ⁄ 2 or less the length of the lower Char. 54. Distinctive Kranz cells: absent (0), present
lemma (1). (1).
*Char. 28. Upper glume: 2 or 4-nerved (0), enerved (1), 1- Char. 55. Fusoid cells: absent (0), present (1).
nerved (2), 1–3 nerved (3), 3–5-nerved (4), 5–11 nerved (5). *Char. 56. Basic chromosome number (x): 9 (0), 10 (1),
Char. 29. Upper glume: muticous (0), awned (1). 11 (2), 12 (3) 13 (4), 5 (5), 6 (6), 7 (7), 17 (8).

Phylogeny of The Paniceae (Poaceae, Panicoideae) Integrating Plastid DNA Sequences and Morphology Into A New Classification

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Phylogeny of The Paniceae (Poaceae, Panicoideae) Integrating Plastid DNA Sequences and Morphology Into A New Classification

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Cladistics

Cladistics (2011) 1–24

Phylogeny of the Paniceae (Poaceae: Panicoideae): integrating plastid

Paniceae is one of the largest tribes of subfamily

Ó The Willi Hennig Society 2011

Our phylogeny revealed good resolution within the Tribe Steyermarkochloeae

Webster, R.D., 1987. The Australian Paniceae (Poaceae). J. Cramer, Appendix 1

Char. 2. Foliaceous rachis: absent (0), present (1). Floret

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