2015 Book FloweringPlantsMonocots

Edited by K.
Kubitzki
Volume XIII
Flowering Plants
Monocots
Poaceae
Elizabeth A. Kellogg
THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
For further volumes see list at the end of the book and:
http://www.springer.com/series/1306
.
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki
XIII
Flowering Plants Monocots
Poaceae

With 96 Figures
Donald Danforth Plant Science Center
St. Louis
Missouri
USA
Series Editor
Prof. Dr. Klaus Kubitzki

Universit€at Hamburg
Biozentrum Klein-Flottbek und Botanischer Garten
22609 Hamburg
Germany
ISBN 978-3-319-15331-5 ISBN 978-3-319-15332-2 (eBook)

DOI 10.1007/978-3-319-15332-2
Springer Cham Heidelberg New York Dordrecht London
Library of Congress Control Number: 2015935331
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To P.F.S.
.
Preface
The goal of this book is to present a state-of-the-art view of grass taxonomy and to
summarize our current understanding of morphological variation in the grasses.
Today, both aspects of agrostology have been engulfed in a flood of molecular
sequence data and an equally large influx of developmental genetic data. I have tried
throughout to incorporate these companion data as they affect our interpretation of
morphological characters and our understanding of phylogeny.
The first sections of the book describe grass morphology, character by character. I
also refer to many studies of developmental genetics that illuminate the genetic basis of
traditional taxonomic characters. Often the data are incomplete, focused on only a
handful of major cereal crops. Where possible, I include information on critical genes
underlying each set of characters. As is conventional in the literature, names of genes
are written in lowercase italics and proteins in uppercase Roman letters. Today, it is
much less common to undertake broad surveys of particular characters across a large
group of organisms than it was in the early 20th century. I hope that, by highlighting
gaps in our knowledge, such survey work can be encouraged in the future.
In terms of taxonomy, this book represents an effort to update the major treat-
ments of grass genera provided by Clayton and Renvoize (Genera Graminum, 1986)
and Watson and Dallwitz (Grass Genera of the World, 1992 onward). In the decades
since those publications, the major clades of grasses have been identified by the Grass
Phylogeny Working Group (2001), and expanded by Sánchez-Ken et al. (2007) and the
Grass Phylogeny Working Group II (2012), leading to the recognition of 12 monophy-
letic subfamilies. Remarkably for such a large family, all but a handful of species (fewer
than ten) are confidently placed in a subfamily. Within the subfamilies, the major
monophyletic tribes have been recognized, and the limits of these are largely stable.
Within the tribes, broad agreement on subtribal limits is emerging, although a number
of genera remain unplaced at this level.
As outlined in more detail in the section Subdivision of the Family, the major
innovation of this book is its phylogenetic approach. The work of Clayton and
Renvoize (1986) and Clayton et al. (2006 onward) arises from the philosophy of the
evolutionary or phenetic school of taxonomy, and Watson and Dallwitz (1992 onward)
also use an explicitly phenetic approach. More recent checklists are based on current
phylogenies (Simon 2007; Simon et al. 2011 onward; Soreng et al. 2012 and onward),
updating the classification frequently as indicated by recent molecular studies. The
classification used here is similar but not identical to those in the checklists, and
includes the rationale for many of the taxonomic decisions.
Remaining phylogenetic and taxonomic problems of the family are at the generic
level. Many genera recognized by Clayton and Renvoize (1986) and currently accepted
by Clayton et al. (2006 onward) and Watson and Dallwitz (1992 onward) are para- or
polyphyletic. This book, like the online checklists, updates those generic limits based
on current molecular phylogenetic studies in an effort to recognize only monophyletic
genera. Nonetheless, current phylogenies leave many loose ends and not all generic
vii
viii Preface
problems can be resolved by current data. Problems outstanding are discussed

throughout the text.
In the formal descriptions I have used several conventions to convey phylogenetic
information. I have attempted to make descriptions more or less hierarchical, so that
character states are only those that apply at a particular level. This means that the
descriptions of the family, subfamilies, tribes and subtribes do not encompass all
possible character states, but only include those that are likely to be synapomorphic
for the clade and/or applicable to the early-diverging taxa. Thus, for example, Chlor-
idoideae are described as having bisexual flowers because dioecy is derived later in the
history of the clade. However, the hierarchical aspect of the descriptions breaks down
frequently because the ancestral state of a clade is often uncertain; plant habit and
ligule structure are two good examples. In these cases, several states are listed.
Taxa that are clearly not monophyletic are indicated in quotes—e.g., “Chloris”, or
“Panicum” s.l., the latter being distinct from Panicum s.s., which is monophyletic.
Putative synapomorphies are indicated in italics. The strength of the evidence for these
varies, so that they should be considered as hypotheses to be tested.
I had initially hoped to avoid many of the arcane grass-specific floral terms, in an
effort to make the entire book more accessible to non-agrostologists. This effort was
not particularly successful, although I had no trouble describing bamboo inflores-
cences without the terms iterauctant and semelauctant. As laid out in the section on
Flower Structure, recent data on the grass floret suggest that it is simply a zygomorphic
monocot flower, and not as peculiar as formerly believed. Accordingly, I have used the
term “flower” instead of “floret”. This is certain to irritate some people, but may make
things clearer to others. As described in the section on Inflorescence Structure, the
terms spike, raceme, and panicle are inaccurate and so are not used. Instead, inflor-
escences are described according to the number of orders of branching and whether
axillary branches proliferate, as in some Andropogoneae.
As noted in the section Subdivision of the Family, phylogenies show that the gross
morphology of grasses is subject to substantial convergence and is not a good guide to
evolutionary history. While many of the well-supported monophyletic groups are
marked by strong synapomorphies, these are often characters of micromorphology
or even genome structure (e.g., chromosome number) and are thus not useful in the
herbarium or in the field. Identification keys are therefore cumbersome and many taxa
are keyed out more than once. The presentation here thus illustrates the tension
between a fully phylogenetic classification and one that is developed for identification
purposes.
In summary, I hope that this book provides food for thought, encouragement for
debate, and an impetus for additional research.
St. Louis, MO Elizabeth A. Kellogg

Acknowledgements
This project has required input from many people and I am grateful to everyone who
helped. The project depended completely on having full access to the incomparable
resources of the Missouri Botanical Garden. For this I am grateful to Peter Raven for
being instrumental in bringing me to St. Louis in the first place, and being a staunch
supporter of me and this work. I also depended on the help of the superb staff,
particularly Jim Solomon for access to the collections and Bob Magill for the Tropicos
database. The entire project required a thoroughly well-curated grass collection, which
is available at the Garden because of the long-time efforts of Gerrit Davidse; I thank
Gerrit for his willingness to be interrupted with questions and for sharing his taxo-
nomic expertise. Ihsan Al-Shehbaz let me share his office for several years and gave me
a place to leave books and provided great friendship. Much of this book was written
while I held the E. Desmond Lee and Family Professorship in Botanical Studies, at the
University of Missouri-St. Louis. That position was designed explicitly to link the
university with the Botanical Garden, and this book is one result of that formal
collaboration.
Many colleagues contributed expertise in their particular taxonomic groups. Neil
Snow and Paul Peterson kindly shared a pre-publication manuscript on Leptochloa and
the Eleusininae, which relieved a number of chloridoid headaches, and Paul also
shared a pre-publication manuscript on placement of a substantial number of chlor-
idoid genera. Neil Snow provided valuable updated information on Disakisperma and
Leptochloa. Rob Soreng shared unpublished data on several pooid taxa and kept me up
to date on changes in the very useful online Catalog of World Grasses. Jordan Teisher
provided valuable comments on Arundinoideae and Micrairoideae, and was able to
correct several errors that had propagated in the literature. I thank Francisco Vazquez
and Mary Barkworth for sharing their considerable expertise in Stipeae. Jimmy Triplett
contributed helpful comments on Arundinarieae, and shared unpublished data on
Pleioblastus. Finally, my many friends and colleagues at the Instituto de Botánica
Darwinion in Buenos Aires shared their incomparable knowledge of the Panicoideae;
I wish to thank particularly my long-time collaborator and director of the Institute,
Fernando Zuloaga, as well as the late Osvaldo Morrone, and Liliana Giussani.
Special thanks are due to Lynn Clark who spent a huge amount of time working
through the treatment of the Bambusoideae, correcting many faux-pas and clarifying
points of morphology. Her input is substantial enough for me to acknowledge her
specifically at the beginning of that section. Furthermore, she went through the
descriptions of grass morphology at the beginning of the book, and made sure that
any statements applied also to the bamboos.
Bryan Simon checked all the taxonomic treatments in detail and carefully com-
pared the numbers of species with those in Grassworld and Grassbase. I thank him in
particular for his patience with my insistence on a phylogenetic classification. I also am
grateful to Maria Vorontsova and David Simpson for their enthusiasm and encourage-
ment for this project. There were many moments during the writing phase when I
ix
x Acknowledgements
wondered how many would read this treatment—the firm support of Bryan, Maria,
and David kept me pushing ahead.
The classification and description of morphology presented here incorporates
much new information generated by my laboratory with steady support from the
National Science Foundation. Former and current graduate students who contributed
data and expertise include Emilie Bess, Paulo Camara, Ken Hiser, John Hodge, Daniel
Layton, Russell Spangler, Jill Preston, Sarah Youngstrom, Cassiano Welker, and Jin-
shun Zhong. In addition, this project has benefited from the careful work of post-
doctoral fellows Sandra Aliscioni, Janet Barber, Hugo Cota, Andrew Doust, Matt Estep,
Pu Huang, Elma Kay, Simon Malcomber, Michael McKain, Roberta Mason-Gamer,
Sarah Mathews, Renata Reinheimer, Jimmy Triplett, Tony Verboom, and Michael
Zanis.
I am indebted to Klaus Kubitzki, the editor of this Springer book series, for the
invitation to undertake this project and I appreciate his patience. Its completion took
far longer than he had hoped or expected—he will surely be pleasantly surprised that
the end of the tunnel has been reached. I also thank Monique T. Delafontaine for her
meticulous copy editing, and Andrea Schlitzberger at Springer (Heidelberg) for guid-
ing this through to publication.
Finally, I heartily thank Peter Stevens who has encouraged this project from the
start, reading the entire manuscript at least once and some parts several times,
commenting on the contents, and providing a steady stream of relevant references.
In addition, he has patiently cooked a very large number of nourishing meals for the
two of us.
I should conclude by saying that, notwithstanding this wonderful input from
colleagues, all taxonomic decisions and any remaining errors in the book are my own.
Contents
Poaceae – General Information

Description of the Family, Vegetative Morphology and Anatomy . . . . . . . . . . . 3
Vegetative Morphology and Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Inflorescence Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Transition to flowering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Bract formation and inflorescence phyllotaxis . . . . . . . . . . . . . . . . . . . . . . . 28
Numbers of orders of branching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Numbers of branches or spikelets at each order of branching . . . . . . . . . . . . 30
Branch angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Axis elongation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Disarticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Flower Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Embryology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Ovule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Megagametophyte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Embryo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Endosperm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Karyology and Genome Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Chromosome number . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Polyploidy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Genome size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Genome mapping, sequencing and conservation of gene order . . . . . . . . . . . 58
GC content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Pollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Morphology and anther structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Pollination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Pollen transfer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Pollen germination and pollen tube growth . . . . . . . . . . . . . . . . . . . . . . . 65
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
xi
xii Contents
Fruit and Seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Unassisted or wind dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Dispersal in time: seed dormancy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Dispersal by animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Role of the awn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Forcible dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Phytochemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Phenolics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Derivatives of amino acids (tryptophan, phenylalanine, tyrosine) . . . . . . . . . 79
Terpenoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Chemicals produced by symbionts or pathogens . . . . . . . . . . . . . . . . . . . . . 82
Phytosiderophores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Distribution, Habitats and Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Distribution and maintenance of grasslands . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Distribution of major clades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Species diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Conservation and response to climate change . . . . . . . . . . . . . . . . . . . . . . . . 89
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Reproductive Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Unisexual vs. bisexual flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Self-pollination and self-incompatibility systems . . . . . . . . . . . . . . . . . . . . . . 94
Cleistogamy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Barriers to interspecific crossing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Asexual reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Fossil Record and Dates of Diversification . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Domestication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Origins of major cereal crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Morphological changes in domestication . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
Genetic and genomic changes in domestication . . . . . . . . . . . . . . . . . . . . . . . 114
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Affinities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Classification of the Poaceae
Subdivision of the Family . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Key to the Subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
I. Subfamily Anomochlooideae Pilg. ex Potztal (1957) . . . . . . . . . . . . . . . . . . . . 131
Key to the Genera of Anomochlooideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Spikelet Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
II. Subfamily Pharoideae L.G. Clark & Judz. (1996) . . . . . . . . . . . . . . . . . . . . . . 135
Key to the Genera of Pharoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Genera of Pharoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Contents xiii
Bistigmatic clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
III. Subfamily Puelioideae L.G. Clark et al. (2000) . . . . . . . . . . . . . . . . . . . . . . . 139
Key to the Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
BEP plus PACMAD clades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
BEP clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
IV. Subfamily Ehrhartoideae Link (1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Key to the Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Ehrhartoideae Incertae Sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
I. Tribe Phyllorachideae C.E. Hubbard (1939) . . . . . . . . . . . . . . . . . . . . . . . . . 144
II. Tribe Ehrharteae Nevski (1937) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
III. Tribe Oryzeae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
III.1. Subtribe Oryzinae Griseb. (1853) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
III.2. Subtribe Zizaniinae Benth. (1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
V. Subfamily Bambusoideae Luerss. (1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Tribes and Subtribes of Bambusoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Key to the Genera of Bambusoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Group I – Herbaceous bamboos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Group II – Branch complement of two or more subequal
branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Group III – Branch complement with one branch only or one dominant
branch and two to many smaller laterals . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
Tribes and Genera of Bambusoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
IV. Tribe Arundinarieae Asch. & Graebn. (1902) . . . . . . . . . . . . . . . . . . . . 159
V. Tribe Bambuseae Kunth ex Dumort. (1829) . . . . . . . . . . . . . . . . . . . . . . 169
Neotropical clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Paleotropical clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
VI. Tribe Olyreae Martinov (1820) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
VI. Subfamily Pooideae Benth. (1861) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Tribes and Subtribes of Pooideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Key to the Genera of Pooideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Group I – Unbranched inflorescences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Group II – Inflorescence axis branched, spikelets without awns . . . . . . . . . 202
Group III – Inflorescence branched, spikelets awned, the awn straight . . . . 204
Group IV – Inflorescence branched, spikelets awned, the awn curved,
twisted or geniculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Tribes and Genera of Pooideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
VII. Tribe Brachyelytreae Ohwi (1941) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
VIII. Tribe Nardeae W. D. J. Koch (1837) . . . . . . . . . . . . . . . . . . . . . . . . . . 208
IX. Tribe Phaenospermateae Renvoize & Clayton (1985) . . . . . . . . . . . . . . . 209
X. Tribe Stipeae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
XI. Tribe Meliceae Rchb. (1828) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
XII. Tribe Diarrheneae C.S. Campb. (1985) . . . . . . . . . . . . . . . . . . . . . . . . . 221
XIII. Tribe Brachypodieae Harz (1880) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
XIV. Tribe Bromeae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
XV. Tribe Triticeae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
XVI. Tribe Poeae R. Br. (1814), s.l . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260
xiv Contents
PACMAD Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

VII. Subfamily Aristidoideae Caro (1982) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
Key to the Genera of Aristidoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268
Genera of Aristidoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
VIII. Subfamily Panicoideae Link (1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
Tribes and Subtribes of Panicoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Key to the Genera of Panicoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Group I – Inflorescence unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Group II – Inflorescence branched; inflorescence or branch axes
disarticulating at the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
Group III – Inflorescence branches unbranched, not disarticulating . . . . . . 278
Group IV – Primary branches of inflorescence branched, although
branches sometimes very short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Tribes and Genera of Panicoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
XVII. Tribe Steyermarkochloeae Davidse & R.P. Ellis (1984) . . . . . . . . . . . . 281
XVIII. Tribe Tristachyideae Sánchez-Ken & L.G. Clark (2010) . . . . . . . . . . 282
XIX. Tribe Chasmanthieae W. V. Br. & B. N. Smith ex Sánchez-Ken & L. G.
Clark (2010) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284
XX. Tribe Gynerieae Sánchez-Ken & L.G. Clark (2001) . . . . . . . . . . . . . . . . 286
XXI. Tribe Centotheceae Ridl. (1907) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
XXII. Tribe Andropogoneae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . 289
XXIII. Tribe Paspaleae J. Presl (1830) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
XXIV. Tribe Paniceae R. Br. in Flinders (1814) . . . . . . . . . . . . . . . . . . . . . . 323
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342
IX. Subfamily Danthonioideae N.P. Barker & H.P. Linder (2001) . . . . . . . . . . . 347
Key to the Genera of Danthonioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
Genera of Danthonioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 352
X. Subfamily Chloridoideae Kunth ex Beilschm. (1833) . . . . . . . . . . . . . . . . . . . 353
Tribes and Subtribes of Chloridoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Key to the Genera of Chloridoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Group I – Inflorescences unbranched, with spikelets sessile or pedicellate;
spike-like panicles may also key here . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Group II – Inflorescence with two or more unbranched branches, either
digitate or spread along the main axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355
Group III – Inflorescence branches themselves branched, at least some
spikelets borne on secondary or higher-order branches, the spikelets
mostly pedicellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 358
Tribes and Genera of Chloridoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
XXV. Tribe Centropodieae P.M. Peterson, N.P. Barker &
H.P. Linder (2011) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
XXVI. Tribe Triraphideae P.M. Petersen (2010) . . . . . . . . . . . . . . . . . . . . . 362
XXVII. Tribe Eragrostideae Stapf (1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . 363
XXVIII. Tribe Zoysieae Benth. (1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
XXIX. Tribe Cynodonteae Dumort. (1824) . . . . . . . . . . . . . . . . . . . . . . . . . 371
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
XI. Subfamily Micrairoideae Pilg. (1956) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Key to the Genera of Micrairoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Genera of Micrairoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
Contents xv
XII. Subfamily Arundinoideae Burmeist. (1837) . . . . . . . . . . . . . . . . . . . . . . . . 403

Key to the Genera of Arundinoidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
Genera of Arundinoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
Poaceae – General Information
Description of the Family, Vegetative Morphology and Anatomy
Poaceae (R. Br.) Barnh. (1895).
Gramineae Juss. (1789).
Rhizomatous perennials, bisexual or monoe- VEGETATIVE MORPHOLOGY AND ANATOMY

cious. Culms herbaceous to somewhat lignified,
erect. Leaf blades broad, with pseudopetioles, Roots
ligules membranous or a fringe of hairs. Inflor-
As in most seed plants, the radicle of the grass
escences branched or unbranched, the floral units
embryo is the first structure to emerge from the
subtended by bracts. Perianth green to brown or
caryopsis at seed germination. Additional roots
absent. Stamens 6; style branches and stigmas
also form at the scutellar node in some taxa; these
3. Pollen monoporate, with an annulus, with
have been called “transitory node roots” by Hos-
channels in the exine, lacking scrobiculi. Ovule
hikawa (1969), who notes that they are present in
1. Embryo lateral, differentiated with clear root
many (but not all) Pooideae and in Ehrharta, but
and shoot meristems enclosed by sheaths (coleo-
absent in rice and in all other grasses investi-
rhiza and coleoptile), several embryonic leaves,
gated. Although the radicle and scutellar node
and a lateral haustorial organ (scutellum). Fruit
roots (together known as seminal roots) are usu-
indehiscent, with one seed, the seed coat fused to
ally described as short lived, they have been
the inner wall of the pericarp, the hilum linear.
found to survive at least 3.5 to 4 months and
Mesophyll with fusoid cells and cells with invagi-
extend to depths of two feet (Weaver and Zink
nated cell walls, midrib complex. Epidermis with
1945). In annuals that have been investigated,
multicellular microhairs, with alternating long
including wheat, plants can survive and flower
and short cells, the short cells developing silica
with the seminal roots alone (Weaver and Zink
bodies. Photosynthetic pathway C3.
1945). Roots subsequently form from the meso-
A full description of Poaceae including all
cotyl and the coleoptile nodes (Hoshikawa 1969),
character variation is lengthy and obscures the
with additional roots arising from subsequent
ancestral conditions for the family that are the
nodes of the main stem and its branches. The
basis for the description above (GPWG 2001).
roots produce an extensive fibrous network
Other familiar characters such as reduction of
(Fig. 1) (Kutschera and Lichtenegger 1982).
the style branches and stigmas to two occurred
Roots may also develop from the lowermost
well after the origin of the family, so are listed in
nodes of the plant, from decumbent stems or
clade or subfamily descriptions below. Modifica-
stolons, and from rhizomes. In the latter case,
tion of the inner perianth to form lodicules may
the roots bind the soil and can stabilize sand
be synapomorphic for the family and have been
dunes. Roots emanating from rhizomes also con-
lost in Anomochlooideae, but it is simpler to
tribute to formation of sod.
assume that the origin of lodicules occurred
The root apical meristem has been studied in
before divergence of Pharoideae and the remain-
detail in maize and rice, and is presumed to be
der of the grasses. The grasses almost certainly
similar in other grasses. The meristem has a closed
originated in shady moist environments; occupa-
organization similar to that in the well-studied
tion of open habitats occurred several times inde-
eudicot Arabidopsis, but unlike the eudicots root
pendently well after the origin of the family.
E.A. Kellogg, Flowering Plants. Monocots, The Families and Genera of Vascular Plants 13, 3
DOI 10.1007/978-3-319-15332-2_1, # Springer International Publishing Switzerland 2015
4 Description of the Family, Vegetative Morphology and Anatomy
Fig. 1. Grass root systems. A Deep rhizomes and roots of Ammophila arenaria. B Shallow rhizomes and roots of Poa
compressa. C Roots from a caespitose species, Deschampsia caespitosa. (From Kutschera and Lichtenegger 1982)
cap initials are wholly separate from the progeni- concentration of ions across the suberized cell
tor cells for other tissues (Coudert et al. 2010). The layers of the root, causing water to flow in to the
quiescent center, at least in maize and rice, xylem (Cao et al. 2012; Holloway-Phillips and
includes hundreds of cells (Hochholdinger et al. Brodribb 2011). High root pressure develops at
2004). Initiation of lateral roots begins with cell night when leaves are not actively transpiring,
divisions in both the pericycle and endodermis, and effectively refills xylem vessels that have
rather than just in the pericycle as in eudicots. embolized during the day. Embolism is a particu-
Cross-sectional anatomy of the mature roots lar problem for some grasses, which fail to close
of grasses is similar to that of other monocots their stomata even when the leaf water potential is
(Clark and Fisher 1987). The root has an epider- quite low. For example, Lolium perenne is able
mis, an exodermis, a cortex of variable width, an to continue gas exchange and photosynthesis
endodermis, pericycle, and polyarch stele. Both even when the leaves have lost over 50 % of
the exodermis and endodermis have suberized their hydraulic conductivity (Holloway-Phillips
cell walls that undergo secondary and even and Brodribb 2011). The ability of xylem to
tertiary thickening, most notably on the inner repair embolisms limits the height of plants in
face of the cells. However, a symplastic pathway general; in bamboos, there is a high correlation
through these walls is maintained by plasmodes- (r2 ¼ 0.81) between root pressure and observed
mata, and some apoplastic flow appears to occur height (Cao et al. 2012).
through the exodermis (Hose et al. 2001). Most Root hairs form just behind the growing
species of grasses have roots with a central pith, point of the root, in vertical files, and hair cells
but a few (generally annuals) have a central xylem alternate with non-hair cells (Clowes 2000). In
vessel; the pericycle and pith may or may not be many species, an epidermal cell divides asymmet-
sclerified, and the extent of tertiary thickening rically to produce a large daughter cell (an atri-
varies in the endodermal cells (Goller 1977). Silica choblast) that will not become a hair and a
may be deposited in endodermal cells (Goller smaller daughter cell (a trichoblast) that will
1977; Hose et al. 2001). Goller (1977) notes that become one. In the grasses, as well as in their
variation in root anatomy correlates with subfa- relatives Restionaceae, Juncaceae, and Cypera-
milial classification, but his Table II suggests that ceae, the trichoblast is the daughter cell that is
many characters may be diagnostic for genera or closest to the apical meristem, whereas it is the
tribes rather than subfamilies. The differences are other way around in other monocots (Clowes
generally quantitative (see summary in Clark and 2000; Dolan and Costa 2001). In rice, however,
Fisher 1987). Enlarged storage roots are rare in the trichoblasts and atrichoblasts do not differ in
the grasses (Clark and Fisher 1987). size immediately after division, but rather
Grasses are known to develop high root pres- undergo differential growth such that the atricho-
sure. This is controlled actively by raising the blasts become larger (Kim and Dolan 2011). In
VEGETATIVE MORPHOLOGY AND ANATOMY 5
other grasses, the trichoblasts and atrichoblasts Grasses also develop associations with fungi.
are not different in size at all (Rothwell 1966; Row Most grasses can develop arbuscular endomycor-
and Reeder 1957). rhizae, associations that can substantially improve
Particularly in dry environments, many uptake of phosphorus. Development of a function-
grasses also develop a rhizosheath, a discrete ing symbiosis is under genetic control, involving
layer of soil particles that is firmly attached to some genes that are common throughout land
the root and fully separable from the surrounding plants and others that appear to be grass specific
soil (Price 1911; Thomas 1921; Wullstein et al. (Yang et al. 2012). Ascomycetes in the family Cla-
1979; Wullstein and Pratt 1981). Rhizosheaths vicipitaceae are generally arthropod pathogens,
have been studied in a handful of crop plants but one clade shifted to form symbiotic associa-
(Duell and Peacock 1985; McCully 1995; St. tions with grasses (Spatafora et al. 2007). A mem-
Aubin et al. 1986; Young 1995) and in a small ber of this clade, Metarhizium robertsii, will invade
number of xerophytic grasses; there appear to be the roots of switchgrass (Panicum virgatum), and
differences between crops and xerophytes, but the stimulate root hair growth.
literature is sparse. The most comprehensive The genetic basis of grass root development is
description of a rhizosheath is for Lyginia bar- only beginning to be explored (Hochholdinger
bata (Restionaceae) (Shane et al. 2011), in which et al. 2004; Hochholdinger and Zimmermann
the sheath appears to be similar to grasses. In 2008; Smith and De Smet 2012). For example, the
maize, the rhizosheath forms about 1 cm behind gene Rootless concerning crown and seminal roots
the root apex and extends 20–30 cm back from (Rtcs) has been cloned from maize, and encodes a
the apex (McCully 1995), whereas in grasses that transcription factor with a Lateral Organ Bound-
normally grow in dry sand, the rhizosheath may aries (LOB) domain; Rtcs controls formation of all
be much longer, up to several meters (Buckley shoot-borne roots, both seminal roots and crown
1982; Price 1911). In roots with both sorts of roots (Majer et al. 2012). Homologues have also
rhizosheaths, root hairs are unusually dense, been cloned from rice, where they appear to have a
and curl around the sand grains of the sheath. In similar function (Smith and De Smet 2012). Other
maize, the root hairs in the region of the rhi- loci affecting lateral root initiation and elongation
zosheath are living (McCully 1995), whereas they have been characterized in both maize and
are persistent and possibly dead in the long rhi- rice (Hochholdinger et al. 2001; Hochholdinger
zosheaths of desert grasses, suggesting that they 1998; Kitomi et al. 2011; Liu et al. 2009). Despite
are no longer taking up water. The rhizosheath of progress cloning genes in rice and maize, the
maize appears only early in development, and is size of the plants makes study of root systems
lost as the epidermis matures and sloughs off. In difficult. Fortunately, the recent development of
contrast, in a mature rhizosheath of a xerophyte, Brachypodium distachyon as a model system will
the hypodermis, exodermis, epidermis and root certainly provide new tools for understanding the
hairs plus sand grains form a layer that is largely controls of root architecture in the grasses (Cho-
impermeable to water. Inside the hypodermis the chois et al. 2012).
outer cortex breaks down, creating a long empty
tube surrounding the inner cortex and stele
Stems
(Buckley 1982; Wullstein and Pratt 1981).
Bacteria are found in the rhizosheath (Goch- The shoot apical meristem has a characteristic
nauer et al. 1989; Wullstein and Pratt 1981) and zonal organization like that of most seed plants;
appear to have several roles. First, they apparently the outer layer (tunica) consists of cells that
secrete polysaccharides that, along with mucilage divide primarily anticlinally, whereas the inner
secreted from the root tip itself, glue the rhi- part consists of cells with less consistent patterns
zosheath together (McCully 1995; Price 1911). of division. Brown et al. (1957) suggested that the
Second, they fix nitrogen (Bergmann et al. 2009; “festucoid” grasses (an informal group that at the
Wullstein et al. 1979), although it is unclear how time included all grasses outside the Panicoideae)
much of this is translocated into the plant. Third, have two tunica layers in the meristem, whereas
they may have antibiotic properties and protect the panicoids have only one. In maize (a pani-
the root from fungi (Shane et al. 2011). coid) the outer L1 layer gives rise to the
epidermis, whereas other tissues are specified by stem, and that the trait is controlled by a single
the inner cells (Jackson 2009). locus (Houshmand et al. 2007). Breeding for a
As the stem matures, cells in the internodes solid stem confers resistance to the wheat stem
stop dividing and differentiate basipetally. This sawfly, suggesting that taxa with solid stems have
leaves a small meristematic region, the interca- enhanced protection against insects. The wheat
lary meristem, at the base of each internode just mutant tiller inhibition (tin) also causes the lower
above the next lowest node. Although the inter- three internodes of the culm to become solid,
calary meristem is a weak spot on the stem, sup- apparently by diverting sucrose away from axil-
port is provided by the surrounding leaf sheaths. lary buds (Kebrom et al. 2012). This result hints
Protoxylem and protophloem are present in the that this taxonomically important character may
meristem, so that vascular continuity is main- reflect a fundamental difference among species in
tained (Clark and Fisher 1987 and references the way carbohydrates are partitioned.
therein). Activity of this meristem allows lodged Stems in most grasses are herbaceous, but
grass stems to right themselves. become woody in members of Bambuseae and
Internodes are generally short in early devel- Arundinarieae (the woody bamboos). In these
opment, and those near ground level often elon- taxa, dense clusters or caps of sclerenchyma
gate little if at all. The timing of internode cells form both externally and internally around
elongation varies between species, but is often the vascular bundles of the stem (Liese 1998). The
similar within major taxa. For example, in Pooi- bundles themselves are closely spaced, resulting
deae, internode elongation is delayed until just in an extremely hard culm. The height of
before flowering. In contrast, in Bambuseae and bamboos is strongly correlated with root pres-
many Panicoideae, internodes elongate appar- sure, which provides the force needed to refill
ently independent of flowering. The timing of embolized xylem vessels (Cao et al. 2012). Other
internode elongation also determines whether large reed-like grasses (e.g., Phragmites, Thysano-
the plant is grazing resistant or not (Branson laena) and the handful of shrubby ones (e.g.,
1953; Holechek et al. 1998); as long as the shoot Cladoraphis spinosa) also develop hard woody
apical meristem is near ground level it cannot be culms, but whether these are histologically and
easily removed by large herbivores. developmentally similar to bamboo stems is
Internally, internodes may be solid through- unknown (GPWG 2001).
out development, or may become hollow. In some Most leaves on a grass plant have a single bud
bamboos, the internodes at the base of the plant in their axils. The buds are under both develop-
may be solid, whereas those at higher nodes are mental and environmental control, and their fate
hollow. The distinction between solid and hollow also depends on where they form on the plant.
internodes is not absolute, however, in that some Axillary buds from the short basal internodes
species have aerenchyma in the internode. Varia- may grow horizontally to form stolons or rhi-
tion in the internal anatomy of the internode may zomes, or may grow more or less vertically to
be taxonomically diagnostic, although it is highly form axillary branches. When the upright
homoplasious (GPWG 2001), and is probably branches occur near ground level they are
most useful at the level of genus and species. known as tillers. As the axillary branch develops,
Most Pooideae tend to have hollow internodes, it may break through its subtending leaf sheath
whereas other subfamilies are more variable (an extravaginal branch) or not (an intravaginal
(Brown et al. 1959a). Clearly the oft-repeated jin- branch). In general, rhizomes and stolons are
gle “sedges have edges, and rushes are round, and formed from extravaginal branches, whereas til-
grasses are hollow right to the ground” is a seri- lers may be either extra- or intravaginal.
ous over-simplification. Pooideae tend to have The extent and nature of basal branching
wider hollows than other taxa, but the ecological controls the overall architecture of the plant.
and evolutionary significance of hollow inter- Grasses that form only tillers develop a clumped
nodes is unknown; Brown et al. (1959a) suggest architecture (i.e., are caespitose), whereas at the
a correlation of hollow internodes with moist other extreme those that form rhizomes or sto-
habitats. Genetic studies in durum wheat show lons are spreading and may be sod-forming. Til-
that having a solid stem is dominant to hollow lers may be geniculate at the base and root from
their nodes so that they are scarcely distinct from Tiller Angle Control1 (TAC1) is a quantitative
short rhizomes. Nonetheless, the growth form of trait locus that also affects tiller angle but the
any particular species is generally reasonably underlying gene has not yet been cloned (Yu
constant. et al. 2007).
The number of tillers is controlled by hor- Branches also form on the stem (culm) itself,
mones, particularly by auxin, strigolactones, and although this is taxon specific. For example,
brassinosteroids, and by carbohydrate levels. branched culms are unknown in the Pooideae,
When auxin transport is inhibited, or when the whereas they are common in Panicoideae (partic-
apical meristem of the plant is removed, the num- ularly Andropogoneae) and Olyreae, and almost
ber of tillers increases and their angle becomes universal in Bambuseae and Arundinarieae.
wider (Li et al. 2007; Xu et al. 2005). Although this Genetic studies (Doust et al. 2004; Doust and
has been demonstrated experimentally only in Kellogg 2006) have shown that the genes that
rice, it is likely that the result is general. Proteins control the formation of culm branches differ
in the strigolactone pathway, such as HIGH TIL- from those that control basal branching (tiller-
LERING DWARF1 and DWARF10, and in the ing). The genetic basis of the trait is thus consis-
brassinosteroid pathway, such as DWARF AND tent with the taxonomic observations.
LOW-TILLERING, also regulate the number of Although the extent of branching – whether
tillers in rice (Arite et al. 2007; Tong et al. 2012; as tillers, rhizomes, or culm branches – is taxon
Zou et al. 2006) and are likely to be involved in specific, it is also controlled by the environment,
other grasses as well. Tiller outgrowth is affected particularly shade and light, and mediated by
by TEOSINTE BRANCHED1 (TB1) and its ortho- auxin, cytokinin and strigolactone, at least in
logues, a cell-cycle regulator that integrates input the crop plants studied (Doust 2007a; McSteen
from hormonal pathways with environmental sig- 2009; Wang and Li 2008b). Many of the cellular
nals (McSteen 2009; Ramsay et al. 2011; Remiger- mechanisms controlling branch formation are
eau et al. 2011). Tiller production is also affected shared among monocots and eudicots, but others
by carbohydrate partitioning; diversion of appear to be grass specific (see citations in Doust
sucrose from axillary buds to the main stem in 2007b).
wheat, as apparently occurs in tiller inhibition Not all leaves on all grasses bear axillary
(tin) mutants, leads to reduced tillering (Kebrom buds. For example, the lower culm nodes of
et al. 2012). some bamboos fail to form buds (Clark and
Tiller angle is also under genetic control and Fisher 1987). In most such cases, whether the
has been investigated extensively in rice, in which bud is specified but simply fails to differentiate,
tiller spreading affects yield and pest resistance or whether the signal for bud formation is never
(Wang and Li 2008a). Plants that spread too transmitted or received, is unknown.
much shade their neighbors and thereby reduce A few woody bamboos (e.g., Chusquea) have
grain production per unit area, whereas those multiple axillary buds, which may be formed by
that are too upright are susceptible to insect supernumary axillary meristems in the axil of a
pests and pathogens because of increased contact single leaf, or may represent a highly compressed
with other plants and higher humidity within the branch complex. These have never been studied
clump. Several proteins have been identified that developmentally.
control tiller angle, including PROSTRATE As in most monocots, the branches produced
GROWTH1 (PROG1) (Jin et al. 2008; Tan et al. by axillary buds bear an adaxial prophyll. This is
2008), LOOSE PLANT ARCHITECTURE 1 generally two-keeled and is particularly promi-
(LPA1) (Wu et al. 2013), LAZY1 (Li et al. 2007), nent in the woody bamboos, where variation in
and PIN-FORMED2 (Chen et al. 2012; Xu et al. its shape is often taxonomically useful.
2005). The latter two proteins regulate auxin The node is complex, both internally and
transport, whereas the mechanism of PROG1 externally. Internally it is marked by a plexus of
influence is unknown. LPA1 affects tiller angle extensively anastomosing vascular tissue that
by controlling the growth of cells on the adaxial forms just above the point of insertion of the
side of the branch; longer cells in that position leaf (Liese 1998; Sharman 1942). Although only
lead to a more spreading tiller (Wu et al. 2013). a handful of grass species have been investigated,
the nodal plexus is a web of transverse vessels in rice by LPA1 (Wu et al. 2013). In general, the
that connect the axial vessels (Pizzolato 2000); the pulvinus lacks sclerenchyma and instead is sup-
overwhelming majority of the latter end at the ported by collenchyma (Paiva and Machado 2003).
nodal plexus with only a tiny percentage extend- In North American grasses, the sheath pulvinus is
ing through (Shane et al. 2000; see also André almost universally present, whereas the nodal pul-
1998). Even in taxa with hollow internodes, the vinus occurs primarily in Panicoideae and Chlor-
node is more or less solid. In the bamboos there is idoideae, and is generally absent in Pooideae
a clear woody wall, the diaphragm (Liese 1998). (Brown et al. 1959b; Clore 2013).
The vascular structure and function of the In a few grasses the lower nodes of the culm
node are described in detail for rice and barley, may be enlarged to form storage organs (Burns
and are presumed to be broadly similar in other 1945). Whereas in a few species the thickened
grasses (Yamaji and Ma 2014). A vascular bundle organs are leaves (e.g., Poa bulbosa), and thus
that will ultimately extend into a leaf can be traced the structure is a true bulb, in other species the
to two nodes below the leaf (i.e., leaf node minus storage organ is the stem and so is properly a
2) where is it relatively small in diameter (called a corm (e.g., Zuloagaea bulbosa, Melica bulbosa,
diffuse vascular bundle by Yamaji and Ma 2014). Arrhenatherum avenaceum var. nodosum, Ehr-
It connects through the next node above (leaf harta capensis and relatives). Species with corms
node minus 1; a transit vascular bundle), and at often occur in areas with low summer rainfall
the leaf node appears enlarged, with an increased (Burns 1945; Verboom et al. 2003). Most bulb- or
number of xylem elements and phloem sieve corm-bearing species are in subfamily Pooideae,
tubes (an enlarged vascular bundle). Thus any but a few are panicoid (Zuloagaea bulbosa; Bess
given node contains diffuse vascular bundles, et al. 2006) or ehrhartoid (Verboom et al. 2003).
transverse vascular bundles and expanded vascu- In terms of life history, the ancestral condition
lar bundles. Xylem transfer cells in the expanded for the grasses is herbaceous, perennial, and rhi-
vascular bundles move solutes to the diffuse vas- zomatous (GPWG 2001), but the annual habit has
cular bundles and thus up to higher nodes in the been derived repeatedly. Humans have particu-
plant. Such a pathway has been demonstrated for larly exploited the annuals (e.g., wheat, maize,
silicon (Si), which accumulates at nodes where it rice), in which much of the photosynthate is accu-
is then distributed among vessels at the nodes mulated in seeds. Multiple genes control the switch
(Yamaji et al. 2008, 2012; Yamaji and Ma 2009). between annual and perennial. As shown in sor-
A similar pathway exists in the phloem, transport- ghum and rice, some of the same loci are involved
ing zinc (Zn) and also the toxic metal cadmium in both species, suggesting that changes of plant
(Cd); both Zn and Cd accumulate in the nodes and habit can occur relatively easily (Hu et al. 2003).
are distributed to developing tissues (Satoh-Naga-
sawa et al. 2012; Yamaguchi et al. 2012). Trans-
Leaves
porters for copper (Cu) and manganese (Mn) are
also located at the nodes but neither mineral accu- Morphology and development
mulates; rather, influx and efflux are tightly con-
trolled to keep levels consistent throughout the As in all seed plants, the shoot in grasses is made
plant (Deng et al. 2013; Yamaji et al. 2013). up of repeating units known as phytomers or phy-
Externally, the node is often marked by a tomeres; each unit consists of a leaf, an internode,
slightly swollen area, the nodal pulvinus, which is and an axillary bud. Whether the internode and
often surrounded by a corresponding area in the bud should be associated with the leaf above or
sheath. This area is flexible and is involved in below is a matter of debate, but however defined
reorienting the stem in response to lodging, the phytomer is repeated over and over in the
although in mature stems it may become lignified growth of the grass shoot (Clark and Fisher 1987).
and lose the capacity to bend (Kaufman et al. 1987). Leaves form on the flanks of the shoot apical
Bending of the pulvinus correlates with changes in meristem. The position of a nascent leaf can be
levels of auxin and gibberellin, which is consistent identified initially by a change in expression of
with the role of these hormones in cell expansion the meristem identity gene knotted1, which is
(Clore 2013; Wolbang et al. 2007), and is regulated switched off in the cluster of cells that will
become the leaf primordium (Jackson et al. 1994). involved in positioning the blade (Foster and
Auxin becomes concentrated in these cells, and Timmermans 2009), and in rice is controlled by
their subsequent divisions lead to the formation the gene LPA1 (Wu et al. 2013). Curiously for such
of a leaf primordium (Reinhardt et al. 2003). The an obvious morphological feature, the wedge-
primordium develops both laterally and in the shaped region has no standard name. It is called
proximo-distal axis to become a broad flat struc- a “leaf joint” in some papers on rice (e.g., Wu et al.
ture that encircles the meristem. In maize, the leaf 2013), whereas is has been called a “dewlap” by
forms from the outer two layers of the shoot some taxonomists (e.g., Martı́nez-y-Pérez et al.
apical meristem, with the L1 layer sometimes 2008; Pohl 1980) and by agronomists working on
contributing to the mesophyll as well as produc- sugarcane (Artschwager 1951), although the term
ing the epidermis (Poethig 1984). About 40 cells is not widely used. Maize geneticists call this
initially contribute to the circumference of the region an auricle. Taxonomists, however, reserve
leaf primordium, and the entire primordium is the term “auricle” for the tiny prongs or hooks
made up of about 200 cells. that extend from the wedge-shaped region, and
Leaf initiation in the grasses (as in all grami- “auricle” is used in this sense here. In this taxo-
noid Poales) is strictly distichous, except in the nomic sense, maize lacks auricles. Bowden (1970)
tiny moss-like grass Micraira. In this species, reports that the dewlaps of Andropogon gayanus
serial sections through the shoot show a phyllo- var. bisquamulatus (Hochst.) Hack. secrete sweet
taxis of 3/8; there is no evidence of twisting of the nectar, but this observation appears not to have
stem or sheaths (Philipson 1935a). Spiral phyllo- been followed up.
taxis has also been reported for the large reed-like Leaf length and width are variable within and
species Arundoclaytonia dissimilis, but this plant between species, and are often taxonomically infor-
has never been studied in detail. Although Jud- mative. Fiorani et al. (2000) found that differences
ziewicz and Soderstrom (1989) cite Page (1951) to in leaf length in species of Poa can be attributed to
suggest that leaf initiation is spiral in Strepto- changes in the rate of growth rather than its dura-
chaeta, Page was in fact referring to the inflores- tion – i.e., leaves grow faster for about the same
cence. The leaves are distichous. length of time, a result extended by Sugiyama
Cell division and expansion initially occur (2005) to several species of C3 grasses. Arredondo
throughout the young leaf primordium, but and Schnyder (2003) also found a correlation
actively dividing meristematic cells become between the size of the meristem at the base of the
increasingly restricted to the base of the young blade and the rate of leaf elongation in eight species
leaf so that the leaf matures from the apex to the of pooid grasses. If this result is generally true, it
base (Sharman 1942). The region of active divi- may point to phylogenetically correlated differ-
sion, termed the “proliferative zone” by Sylvester ences in regulation of the cell cycle.
and Smith (2009), is later divided into two by the Also at the blade-sheath boundary is an adax-
developing ligule and sheath, so that two meris- ial flap of epidermal tissue, the ligule; in most
tems are formed, one each at the base of the blade cases this is small, no more than a couple of
and the base of the sheath. millimeters long, although it may be several cen-
Mature leaves in the grasses generally consist timeters long in some woody bamboos. During
of a distal blade and a proximal sheath. The sides leaf development, the blade-sheath boundary is
of the leaf blade – on either side of the midrib – are defined by a region of increased cell division
controlled developmentally by the NARROW- known as the pre-ligular band; this marks the
SHEATH proteins, which define a lateral composition where the ligule will form (Sharman
partment of the leaf (Nardmann et al. 2004). The 1942; Sylvester et al. 1990). Development of the
region where the blade and sheath join is known ligule has been studied extensively, especially in
as the collar; in some taxa this corresponds to a maize (reviewed by Foster and Timmermans
region of more flexible tissue and less scleren- 2009). Many genes are involved in the proper
chyma (Paiva and Machado 2003). In many positioning and development of the ligule, but
grasses there is a wedge-shaped region on either only a few seem to affect ligules exclusively,
side of the leaf at the collar that can be identified whereas others disrupt the entire structure of
by its color and texture. This region appears to be the leaf.
In a detailed description of ligule develop- the sheath from the culm varies among genera or
ment in Deschampsia, Melica and Phyllostachys, groups of genera.
Philipson (1935b) noted that the ligule appears to Leaf angle is under the control of brassinos-
form from separate domains, a central epidermal teroids (Tong et al. 2012), as well as other plant
one, and lateral extensions of the margins of the hormones whose effects interact with those of the
sheath, thus anticipating by more than 50 years brassinosteroids (Song et al. 2009). However,
the discoveries of the lateral domains in maize other controls of leaf angle operate independently
leaves (Scanlon and Freeling 1997). Ligule mor- of hormonal pathways, affecting cell division in
phology is remarkably variable and is often taxo- and around the collar. These include the genes
nomically informative. Membranous ligules Leaf inclination2 (Zhao et al. 2010), Increased leaf
predominate among members of the BEP clade, angle1 (Ning et al. 2011), and LPA1 (Wu et al.
whereas the PACMAD subfamilies often have 2013).
ligules consisting of a fringe of hairs.
A ligule-like structure, which may be mem-
branous or ciliate, may form on the abaxial side
Leaf epidermis
of the collar region, and is variously known as a
contraligule, pseudoligule, or external ligule; the
In most grasses, the leaf epidermis consists of a
development of this structure has never been
single layer of cells that form in long files parallel
studied. It occurs in some genera such as Puelia
to the proximo-distal axis of the leaf (Fig. 2).
(Puelioideae), Streptogyna, and in most, if not all,
Because the leaf matures from tip to base, these
genera of Bambuseae and Arundinarieae, and can
long files represent a developmental gradient that
be helpful for genus and species identification.
has helped in the investigation and understand-
The base of the leaf blade above the ligule
ing of differentiation. Many of the late cell divi-
may be constricted to form a pseudopetiole. Pres-
sions are asymmetric, with a single cell giving rise
ence of a pseudopetiole is ancestral and synapo-
to two differently sized daughter cells (see Fig. 8A
morphic among the grasses (GPWG 2001). All
in Sylvester et al. 1990). The result of this is a
species of Anomochlooideae, Pharoideae, and
characteristic alternation of long and short cells
Puelioideae have pseudopetioles, as do most spe-
in the mature epidermis. The alternation is not
cies of Bambusoideae. The structure also appears
perfect, and short cells may occur in pairs or files
in some Panicoideae. Development of pseudope-
of up to five. Alternation of long and short cells
tioles has never been studied and it is unknown
occurs in other monocots as well, but only in the
how it affects – or is affected by – the meristem at
cell files that will produce stomata. Thus the
the base of the blade, nor is there information on
asymmetric cell divisions that occur throughout
its contribution to the hydraulic architecture of
the epidermis in the grasses represent a change in
the leaf.
position of a developmental program (Kellogg
In taxa such as Streptogyna (BEP clade, incer-
2000). Long-and-short cell alternation is shared
tae sedis), most bamboos, some Panicoideae (e.g.,
with Joinvillea, one of the close relatives of the
Thysanolaena, tribe Centotheceae; Gerritea, tribe
grasses, and provides one piece of evidence for
Paspaleae), Micrairoideae (Micraira), Arundinoi-
their close relationship (see Affinities); because
deae (Molinia), some species of Ehrharta, in
Ecdeiocolea is leafless, the ancestral condition for
Macrochloa (Stipeae), and Aristida (Aristidoi-
the grass sister group is unknown (Campbell and
deae), an abscission zone forms at the collar and
Kellogg 1987). The epidermis is often structured
the leaf blades disarticulate. The anatomy of the
differently over the veins (costal region) than it is
abscission zone varies among species (R€oser and
between them (intercostal regions), a difference
Heklau 2011). In other species (e.g., in some spe-
that is generally consistent within a species or
cies of Rytidosperma, Danthonioideae), an abscis-
genus (Fig. 2). The genetic basis of the asymmet-
sion zone forms at the base of the sheath and the
ric cell divisions that will produce stomata is
entire leaf is deciduous. In the culm leaves of
becoming increasingly well understood (Abrash
woody bamboos, even if the blade persists for
and Bergmann 2009), but whether this machinery
several seasons, it will ultimately disarticulate
is also activated in other short cells is unknown.
from the sheath. Subsequent disarticulation of
Fig. 2. Leaf epidermis showing files of cells extending mal cells bear papillae. B Monocymbium cerisiiforme
along the proximo-distal axis. A Oxychloris scariosa (Panicoideae), showing bilobate silica bodies in short
(Chloridoideae), showing characteristic saddle-shaped sil- cells over the veins. (From Watson and Dallwitz 1992
ica bodies in short cells over the veins. Intercostal epider- onward)
Epidermal long cells vary only slightly in Epidermal short cells have various fates,
shape, usually being rectangular but sometimes depending on the species, position in the plant,
fusiform. Their longitudinal walls, however, may and position in the leaf (or leaf-like structure such
be sinuous and thus interlocking, or straight; the as the lemma). One common role of short cells is to
latter condition occurs frequently, but not univer- differentiate as silica-containing cells. The accumu-
sally, in Pooideae. The shape of the longitudinal lation of silica in the leaf is a character shared by
wall is determined in part by localization of micro- most of the commelinid monocots, and accumula-
tubules and actin (Frank et al. 2003). In maize, in tion specifically in the epidermis is a character of
which the longitudinal walls are normally sinuous, the Poales (Stevens 2012). Monosilicic acid, Si
mutations in loci known as Brick (because they (OH)4, is produced by weathering of soils, and is
create brick-like cells when mutated) create long taken up by the plant both actively and passively in
cell shapes reminiscent of those occurring in, for a highly regulated process (Yamaji et al. 2008, 2012;
example, species of Poa or Ehrharta. A survey of Yamaji and Ma 2014). Silica is deposited in an
grasses staining for tubulin and actin would help amorphous non-crystalline form as silicon dioxide
determine whether distribution of these cytoskele- (SiO2.nH2O) throughout the plant, particularly in
tal proteins is the primary determinant of taxo- tissues involved in support of the stems and leaves
nomic variation in cell wall morphology. (Isa et al. 2010), but also in the walls of guard cells,
In the grasses, as in most commelinid mono- subsidiary cells, and epidermal papillae (Ueno and
cots (Stevens 2012), the stomata are paracytic – Agaric 2005); the rate and developmental timing of
there are two guard cells, with the stoma oriented accumulation is specific for different cell types
parallel to the long axis of the leaf, and two sub- (Sakai and Sanford 1984). In addition, silica com-
sidiary cells parallel to the guard cells. In stomatal monly accumulates in specialized short epidermal
development, a short cell divides longitudinally cells (silica cells) in which it is deposited initially in
to give two cells and then these divide, again the cell wall and then accumulates centripetally,
longitudinally, to give a set of four, more or less while the cellular contents break down (reviewed
parallel, rectangular cells (Abrash and Bergmann by Prychid et al. 2004). Silica deposition protects
2009). The outer two differentiate into subsidi- the plant from bacterial and fungal pathogens, sup-
aries and the inner two into guard cells. The ports the stems, reduces the uptake of toxic metals,
ancestral and most widespread condition is for and regulates water loss (Isa et al. 2010; Ma and
subsidiaries to be somewhat dome-shaped in sur- Yamaji 2006). In addition, silica rapidly wears
face view, although in some species they are down the mandibles of insect herbivores and
triangular. However, in the pooid clade that reduces digestibility (Massey and Hartley 2009).
includes Poeae, Triticeae, Bromeae and Brachy- Silica accumulation is presumed to be energetically
podium, the subsidiaries have parallel walls, a costly, but its effect on plant biomass varies
condition that is uniquely derived. The subsidi- between species; this variation affects susceptibility
aries overlap the guard cells in all Pooideae that to herbivores and thus competitive interactions
originated after the divergence of Nardeae. (Garbuzov et al. 2011).
Silica may also be a defense against large more or less like a muffin. The nucleus then
mammalian herbivores, although the data on moves up in the cell closer to the outer wall. As
this are less clear. Simpson (1951) famously pro- the outer wall continues to expand, growth
posed that the evolution of hypsodont teeth in becomes asymmetrical, and the cell elongates
equids was driven by a shift to diets of grass, and parallel to the surface of the leaf (Kellogg 1990).
their high concentrations of silica. However, San- In cells that will become prickles, the tip of
son et al. (2007) have recently questioned whether the cell develops a sharp point and silica accu-
silica bodies are in fact hard enough to wear down mulates in the tip (Prat 1932). Silica accumulation
mammalian tooth enamel, as had been suggested appears to occur early before the leaf has fully
(Baker et al. 1959). In addition, Str€omberg (2006) opened and before silica deposition in other cells
showed that hypsodonty appeared well after the (Motomura et al. 2006). Prickles generally point
spread of grasslands, at least in North America, toward the distal end of the structure on which
weakening the hypothesized connection. they occur, but sometimes point toward the prox-
The shape of the silica deposits (silica-bodies imal end (i.e., are retrorse). Presence or absence
or phytoliths) is often characteristic of particular of prickle hairs on particular structures often is a
taxonomic groups. These shapes were originally good field identification character. However, I
described based on their appearance in two know of no study in which this character has
dimensions (Metcalfe 1960; Prychid et al. 2004), been evaluated in a phylogenetic context; intui-
and early attempts to use silica body characteris- tively, it seems as though it should be highly labile
tics in phylogeny reconstruction found that they in evolutionary time. The function of prickle
were highly homoplasious (Kellogg and Campbell hairs is unknown, but could help deter small
1987; Kellogg and Watson 1993). The description herbivores such as slugs and nematodes.
and classification of silica bodies (phytoliths) is Bicellular microhairs are found in all non-
improving due to efforts to describe them in pooid grasses (Johnston and Watson 1976), but
three dimensions rather than two (Piperno 2006; their development has never been investigated.
Piperno and Pearsall 1998). For example, bilobate As their name implies, these trichomes have
silica bodies occur in many subfamilies of grasses only two cells, one apical and one basal, but the
(Fig. 2A, B). However, those of Stipeae are asym- shape of the cells is often characteristic of partic-
metrical in cross section, while those of Panicoi- ular taxa. In the “panicoid type” microhair, both
deae are generally symmetrical. Aristidoid cells are longer than wide, and internal mem-
bilobates tend to have a long thin shaft between branes are not readily visible (Amarasinghe and
the lobes, whereas in other taxa the shaft is short Watson 1988, 1989). In contrast, in the “chlori-
and in bamboos is lacking altogether. So-called doid type” the apical cell is nearly as wide as long.
“oryzoid” bilobates are elongated perpendicular The distinction between the two is not sharp,
to the long axis of the leaf. This form is found in however, and a graph of the length-width ratio
Ehrhartoideae, some bambusoids (Olyreae), and of the apical cells in all microhairs in the family
arundinoids (Eriachne); careful analysis of these, shows that the variation is continuous (GPWG
however, shows that they can in fact be distin- 2001; Kellogg, unpublished observations). Some
guished (Prasad et al. 2011). More detailed chloridoid microhairs contain internal mem-
description of silica bodies will not wholly solve branes in the basal cell and secrete salt
the problem of homoplasy in the character, how- (Liphschitz and Waisel 1974; Marcum 1999; Oi
ever. Individual plants have a range of silica body et al. 2012), but there is no evidence that the
forms (Piperno and Pearsall 1998) and thus char- panicoid hairs are secretory (Amarasinghe and
acterization of silica bodies within a species or Watson 1989). Other chloridoid microhairs, the
genus must be somewhat quantitative. “Enneapogon type” have internal membranes in
Short cells may also form trichomes. These the apical cell, but appear to be non-secretory
may be unicellular (prickles, macrohairs) or (Amarasinghe and Watson 1988, 1989). Lack of
bicellular (microhairs), and may accumulate sil- microhairs is synapomorphic for all Pooideae
ica or not (Prat 1932). Development of prickles after the divergence of Brachyelytrum, Nardus
and macrohairs begins with enlargement of the and Lygeum, but ecological consequences of this
cell. As the trichome develops, the outer wall loss are unknown. Microhairs with more than two
expands, so that the outline of the cell looks cells are reported for Joinvillea (Joinvilleaceae), a
close outgroup of the grasses, and for Streptogyna Xiang et al. 2012), in some cases via affecting the
crinita and several members of Bambusoideae differentiation between the ab- and adaxial sides
(Soderstrom and Judziewicz 1987). of the leaf (Hibara et al. 2009). Comparative stud-
Macrohairs on the leaf blades are inherited ies have yet to be done on most of these proteins,
independently of hairs on other plant parts so it is unclear which components of the bulli-
(Moose et al. 2004), supporting their use as a dif- form specification network may have been
ferent taxonomic character. Macrohairs are often selected to produce the diversity of leaf anatomy
surrounded by a multicellular and slightly raised observed among the grasses.
set of epidermal cells (Prat 1932). These have been Multicellular structures known as “glands”
reported to be secretory (Bowden 1971), with the have been observed in some grasses, particularly
cells containing a variety of sugars and pectic sub- in Danthonioideae (Linder et al. 1990). Two-
stances. However, the number of species investi- celled salt glands have also been described in
gated is tiny, and macrohairs would repay closer Spartina (now part of Sporobolus s.l.; Peterson
investigation (Sylvester and Smith 2009). et al. 2014), in which an epidermal initial cell
In areas around some or all of the veins, expands downward into the mesophyll, and later
adaxial epidermal cells may differentiate as bulli- divides asymmetrically to form a small apical cell
form cells. These cells are enlarged in the ab- (Fahn 1979). Salt glands have also been charac-
adaxial direction, extending into the region nor- terized in Chloris gayana; the density of glands on
mally occupied by mesophyll. They may occur as leaves increases when the plant is grown in higher
fans of cells or in irregular groups, and may or concentrations of salt (Oi et al. 2012). The apical
may not be associated with other colorless meso- cell of the gland has a complex endomembrane
phyll cells, and thus provide a source of taxonom- system and a high number of mitochondria, sug-
ically informative characters. Bulliforms may gesting that salt excretion is energetically costly.
occur on either side of the mid-vein only, or on The epidermis of many grasses is coated with
the sides of lateral veins as well. Some species lack a layer of wax. This is characteristic of leaves on
bulliform cells entirely. juvenile plants in maize and the presence of wax
Bulliform cells can rapidly take up or lose has been used as a marker of the transition from
water. By expanding and contracting, bulliform juvenile to adult morphology (Moose and Sisco
cells are reported to control leaf rolling (Arber 1996). In some taxa, such as Sorghum, the wax
1934; Bidlack and Jansky 2011), but there is sur- forms on the stem and sloughs off as large flakes.
prisingly little evidence to support the hypothesis In some bambusoids, wax forms only on one
that changes in bulliform turgor are actually caus- portion of the leaf, indicating very precise, cell
ative (Arber 1934). In the resurrection species Spor- specific, genetic and developmental control.
obolus stapfianus the outer wall of the bulliforms is
thick and water is lost to the adjacent mesophyll
cells (Dalla Vecchia et al. 1998). While water loss Internal anatomy of leaves
from the bulliforms may lead to leaf rolling in some
species, in S. stapfianus the bulliforms are involved The internal anatomy of grass leaves has been
in maintaining hydration of the mesophyll. In late studied extensively (Brown 1977; Ellis 1976; Met-
development of the leaf, bulliform cells may accu- calfe 1960; Watson and Dallwitz 1992 onward).
mulate silica (Motomura et al. 2004). Most investigations have focused exclusively on
A number of proteins regulate the number of the cross-sectional appearance of the middle por-
bulliform cells in each group. Transcription fac- tion of the adult leaf blade, so comparative data
tors that help specify identity of the ad- and on sections in other planes, on sheaths and on
abaxial sides of the leaf blade determine whether early development are limited. In most species,
bulliforms will develop on the adaxial side (as is the mesophyll cells are not tightly packed and are
most common) or abaxial side, and also regulate relatively homogeneous throughout the leaf.
how many bulliforms form in a cluster (Dai et al. However, some Centotheceae have an adaxial
2007; Zhang et al. 2009; Zou et al. 2011). Various palisade layer. In some early-diverging grasses
enzymes also affect bulliform cell development and in virtually all bamboos the mesophyll is
(Fujino et al. 2008; Hu et al. 2010; Li et al. 2010; interrupted by fusoid cells (Fig. 3A).
Fig. 3. Cross-sections of leaves of selected grasses. A parenchyma sheath, rch radiate chlorenchyma, sg scleren-
Dinochloa macclellandii (Bambusoideae). B Poa sp. chyma girder, st stomatal apparatus, x xylem. (From
(Pooideae). C Bouteloua sp. (Chloridoideae). ac arm cell, GPWG 2001, p. 405, with permission of the Missouri
bc bulliform cell, ch chlorenchyma, fc fusoid cell, is inter- Botanical Press; drawn by M. Kojima)
cellular space, ms mestome sheath, p phloem, ps outer
In all the early-diverging lineages of grasses, and vascular tissue, the fusoid cells develop
mesophyll cells have obvious invaginations of the large vacuoles. As the leaf blade emerges from
cell wall when viewed in cross section; such cells the sheath of the leaf below it, the fusoid cells
are known in the literature as arm cells (Fig. 3A). appear to die and their walls apparently collapse.
Such cell wall invaginations also occur in all Page (1947) then speculates that some force must
bamboos, and in the tribe Oryzeae of subfamily be operating on the fusoid cells to preserve their
Ehrhartoideae (GPWG 2001). Invaginated cell regular shape during development. In long-lived
walls also appear in Phragmites (subfamily Arun- leaves of some bamboos the fusoid cells accumu-
dinoideae), where they constitute a reversal fol- late silica in their lumens (Motomura et al. 2004).
lowing loss at the base of the PACMAD clade. March and Clark (2011) find that shade grown
Longitudinal sections of leaves have revealed leaves of Chusquea, Phyllostachys, and Yushania
additional variation in internal morphology (Sán- all developed fusoid cells, whereas sun-grown
chez-Ken and Clark, unpubl. data). leaves did not. They speculate that perhaps fusoid
Mesophyll cells with invaginated walls are cells are a way to increase light availability inside
apparently uniquely derived in the grasses. They the leaf.
are lacking in Joinvilleaceae. Because Ecdeioco- Grass leaves generally contain sclerenchyma
leaceae are leafless, there are no data for this associated at least with the vascular bundles,
family. However, a similar phenotype appears in although there are often clusters of sclerenchyma
other more distantly related families. For exam- at the leaf margin as well (Fig. 3A, C). The distri-
ple, Restionaceae have cells with invaginated cell bution of sclerenchyma within the leaf is often
walls in their stems, where they are known as peg distinctive and can be helpful in species identifi-
cells (Cutler 1969). cation. Sclerenchyma may extend from the vas-
Fusoid cells are large, rectangular to cigar- cular bundle to the abaxial epidermis, the adaxial
shaped mesophyll cells, and appear curiously epidermis or both, or may be present as a cap
empty in leaf cross sections. They are apparently over the bundle (Ellis 1976).
synapomorphic for the grasses; the plants in The functional consequences of different pat-
which they occur are shade loving and prefer terns of sclerenchyma distribution are largely
moist habitats. They occur only in the early- unknown. Leaves with sclerenchyma girders
diverging lineages, Streptogyna, the Bambusoi- extending from the bundle sheath to both epi-
deae, and a handful of Panicoideae in tribe Pas- dermes are known as “heterobaric”, whereas
paleae, subtribe Arthropogoninae. Superficially those without such girders are called “homoba-
similar cells have been reported in Centotheceae ric”. In heterobaric leaves, such as those in Hor-
(Panicoideae) as well, but they appear to be deum vulgare, the girders, called bundle sheath
laterally expanded bundle sheath cells and are extensions in the physiology literature, provide a
more accurately described as bundle sheath direct hydraulic connection between the vascular
extensions. The function and development of bundle and the epidermis (Buckley et al. 2011)
fusoid cells are unknown and it is not even allowing stomatal movements to respond rapidly
clear whether they are alive or dead, although and easily to water availability. Bundle sheath
some data suggest that they may be dead at extensions also divide the leaf into functional
maturity. compartments that affect the structural and func-
Development of fusoid cells has been tional aspects of the leaf, although this functional
described in some detail in Streptochaeta spicata compartmentation has not been investigated in
(Page 1947). As the primary vascular bundles grasses. Investigations in dicotyledonous trees
begin to differentiate, most mesophyll cells are and shrubs suggest that heterobaric leaves have
still dividing in all three planes of the leaf (ab- lower leaf mass per unit area, more nitrogen per
adaxial, lateral, and proximo-distal). However, unit mass, and have higher photosynthetic capac-
the mesophyll cells adjacent to the vascular tissue ity per unit mass, than homobaric leaves (Lia-
cease to divide in the ab-adaxial and lateral koura et al. 2009). In homobaric leaves, CO2 can
planes and simply enlarge; they continue to diffuse laterally for a distance of several milli-
divide along the proximo-distal axis. While cell meters, whereas the bundle sheath extensions of
division continues in all mesophyll, epidermis a heterobaric leaf effectively prevent such
diffusion (Pieruschka et al. 2008). It seems likely whereas this layer is apparently absent in Arundo
that such relationships will hold in grasses as and Phragmites (Arundinoideae) (Carolin and
well. Jacobs 1973). In mestome sheaths in general the
The phloem of grass leaves contain two dis- outer walls are suberized, minimizing the apo-
tinct sorts of sieve elements (Botha 2013). The plastic pathway for water.
canonical sieve element is thin-walled, and is In C4 plants, the carbon-fixing enzyme
associated with a companion cell. This pair of Rubisco is produced only in the bundle sheath
cells appears to function in a manner similar to cells, rather than in the mesophyll as in C3 species.
that of dicot sieve elements and companion cells, Rubisco is then replaced in the mesophyll by PEP
in which the sieve element is responsible for long carboxylase (Kanai and Edwards 1999; Wang et al.
distance transport, is enucleate a maturity, and is 2011). Atmospheric CO2 is hydrated to carbonic
metabolically dependent on the companion cell, acid and subsequently fixed as a four-carbon com-
to which it is connected by plasmodesmata. The pound oxaloacetate (OAA) in the mesophyll. OAA
other sort of sieve element is thick-walled, is then reduced to either malate or aspartate, and
appears not to be involved in transport, and is is shuttled to the bundle sheath, where it is dec-
symplastically isolated from the thin-walled sieve arboxylated and the carbon is reduced by the
elements; instead it is connected to the vascular normal Calvin Cycle to make three-carbon sugars.
parenchyma of the metaxylem. The function of This process leaves the three-carbon phosphogly-
the thick-walled sieve elements is not known, nor ceric acid, which is returned to the mesophyll.
is their taxonomic distribution. They occur in Because the C4 pathway requires constant
grasses and sedges but whether they appear in shuttling of carbon compounds between meso-
other monocots is unknown (Botha 2013). phyll and bundle sheath, every bundle sheath
The internal anatomy of the leaf varies con- cell is next to and connected directly with a meso-
siderably depending on the photosynthetic path- phyll cell in C4 plants. This is accomplished by
way used. Grasses may use either the conventional increasing the density of venation. In all C4
C3 pathway, or the high efficiency C4 pathway. grasses, the vascular bundles are separated by
Within the family, the C4 photosynthetic pathway no more than two mesophyll cells (Fig. 3C) (Hat-
has arisen multiple times among members of the tersley and Watson 1975). In a few C4 taxa, such
PACMAD clade (Christin et al. 2007, 2008; as Arundinella, the fully developed veins are
Edwards and Smith 2010; GPWG II 2012; Vice- separated by more than two mesophyll cells, but
ntini et al. 2008). C4 is a mechanism for reducing in the place of minor veins are lines of cells that
the amount of oxygen available to ribulose 1,5 appear unusual in cross section and were initially
bisphosphate carboxylase/oxygenase (Rubisco), given the name “distinctive cells” by Tateoka
thus minimizing photorespiration. By reducing (1958). These cells function in carbon reduction,
the energy lost to photorespiration, C4 plants produce RuBisCO (Sinha and Kellogg 1996), and
make better use of light energy; Zhu et al. (2008) appear to represent isolated bundle sheath cells
have calculated that C4 plants may capture 24 % from degenerate minor veins. Photosynthetic
more energy than C3 plants. In addition, C4 plants pathway correlates perfectly with vein spacing.
make more efficient use of water and nitrogen Leaves of all or nearly all genera of the family
(Sage 1999). have been examined anatomically (Watson and
In all C3 grasses, there are two rings of cells Dallwitz 1992 onward), and a substantial majority
around the vascular bundles, the ancestral condi- of these have also been evaluated using stable
tion (Fig. 3A, B). The outer sheath is made up of isotopes (Brown 1977) (C. Osborne, Univ. of
thin-walled parenchyma cells and is known as the Sheffield, pers. comm.); the ratio of 13C to 12C is
parenchymatous sheath, and the inner sheath has a reliable predictor of photosynthetic pathway.
cells with thick walls and is known as the mes- The genetic regulation of vein spacing is not
tome sheath (Carolin and Jacobs 1973); none of understood, and existing data show that there is
the sheath cells have many chloroplasts. The mes- no simple switch between C3 and C4-like spacing.
tome sheath of some species such as rice and the Vein density in oats (C3) could be increased by
pooid grasses has an osmophilic layer in the cell introduction of entire maize (C4) chromosomes 1,
wall in transmission electron microscopy (TEM), 5, or 9, one at a time (Tolley et al. 2012). While
each chromosomal introduction produced a sig- The C4 NADP-ME taxa in Micrairoideae and
nificant reduction in vein spacing, the number of Aristidoideae are anatomically distinctive. In
cells between the veins was unchanged suggesting both subfamilies there is an outer and an inner
that the cells were merely smaller. Addition of bundle sheath. However, the two C4 genera of
several maize chromosomes also increased the Micrairoideae, Eriachne and Pheidochloa, have
size of the bundle sheath cells. so many chloroplasts in the outer sheath that no
C4 species vary in the enzyme used to decar- cytoplasm or vacuole is visible in cross section
boxylate the four-carbon compound in the bun- (see illustrations in Sinha and Kellogg 1996). In
dle sheath. Most grasses use one of several malic Aristida, both the inner and outer sheaths are
enzymes, some of which use NAD as a co-factor carbon reducing and neither has a suberized
and some of which use NADP; these are known as lamella. Carbon reduction occurs primarily in
NAD-ME and NADP-ME respectively. In addi- the inner sheath (Ueno 1992), but carbon may
tion, some species use PEP carboxykinase (PCK) leak out to the outer sheath, where it can also be
as their primary decarboxylating enzyme. captured by Rubisco and reduced (Hattersley and
Although it is common to classify C4 species Browning 1981).
according to their decarboxylating enzyme, All other C4 species occur within subfamily
many species use more than one. For example, Panicoideae, and use NADP-ME as the decarbox-
PCK activity has been detected in maize, even ylating enzyme. This group includes the most well-
though it is generally considered to be an studied of the C4 plants, maize, and its agronomi-
NADP-ME species (Walker et al. 1997). cally important relatives sorghum and sugar cane.
The structure of the bundle sheath cells gen- These species all have a single bundle sheath, which
erally correlates with the primary decarboxylat- develops from the same cell lineages as the ances-
ing enzyme (Hattersley 1987; Prendergast and tral mestome sheath, and like the mestome sheath
Hattersley 1987; Prendergast et al. 1987; Sinha has a suberized cell wall (Dengler et al. 1985). In
and Kellogg 1996). The C4 species that use pre- these NADP-ME species, the chloroplasts in the
dominantly NAD-ME or PCK include all mem- bundle sheath cells generally lack grana. There is
bers of Chloridoideae, subtribes Melinidinae and also a reduction in the amount of proteins such as
Panicinae (Paniceae, Panicoideae), and the genus LHCPII that anchor the photosynthetic apparatus
Stipagrostis (Aristidoideae). These all have the in the thylakoids (Sinha and Kellogg 1996).
ancestral condition of two bundle sheaths, and Radiate mesophyll cells occur in many C4
the inner bundle sheath retains its ancestral taxa, particularly those with the NAD-ME sub-
structure as a thick-walled mestome sheath type. However, radiate mesophyll is not confined
(Fig. 3C) (Brown 1975). The outer parenchyma to C4 species. It is also recorded for members of
sheath, however, produces large numbers of the tribe Isachneae (Micrairoideae), which are
chloroplasts and is the site of carbon reduction. entirely C3. It is an intriguing possibility that
The structure of the outer bundle sheath often Isachneae may have been derived from C4 ances-
correlates with decarboxylating enzyme as well tors, or conversely that the C4 Micrairoideae
(Hattersley and Watson 1992). In NAD-ME spe- already had radiate mesophyll that made origin
cies, the outer walls of the parenchyma sheath of the C4 anatomy easier.
form an even, more or less circular outline and Data on the anatomy of leaf sheaths are scarce.
are suberized, whereas the outer sheath cells in The veins in the sheath may be more widely
PCK species are often irregular or form a trian- spaced than those in the blade, which affects
gular shaped bundle, and are non-suberized their photosynthetic function. The mesophyll
(Dengler and Nelson 1999). The organelles in cells farthest from the veins have C3 physiology
the outer sheath of NAD-ME plants are often in maize, whereas the ones closest to the veins are
clustered along the walls closest to the vascular physiologically C4 (Langdale et al. 1988). The ana-
tissue (centripetal), whereas the organelles in tomical structure at the top of the sheath also
sheaths of PCK species are often clustered along controls the angle of the leaf blade. In rice, reduc-
the walls closest to the mesophyll. These correlation in the amount of sclerenchyma associated
tions are not absolute, however (Prendergast and with the vascular tissue results in leaves that are
Hattersley 1987; Prendergast et al. 1987). less upright than normal (Ning et al. 2011).
Some grass plants go through a heteroblastic sieve tubes in monocotyledonous leaves. Frontiers
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phosphatidylinositol-anchored protein and modu- Rice leaf inclination2, a VIN3-like protein, regulates
lates rice leaf rolling by regulating the formation of leaf angle through modulating cell division of the
bulliform cells. Plant Phys. 159: 1488–1500. collar. Cell Res. 20: 935–947.
Xu, M., Zhu, L., Shou, H.X., Wu, P. 2005. A PIN1 family Zhu, X.G., Long, S.P., Ort, D.R. 2008. What is the maxi-
gene, OsPIN1, involved in auxin-dependent adventi- mum efficiency with which photosynthesis can con-
tious root emergence and tillering in rice. Plant Cell vert solar energy into biomass? Curr. Opin.
Phys. 46: 1674–1681. Biotechnol. 19: 153–159.
Yamaguchi, N., Ishikawa, S., Abe, T., Baba, K., Arao, T., Zou, J.H., Zhang, S.Y., Zhang, W.P., Li, G., Chen, Z.X.,
Terada, Y. 2012. Role of the node in controlling Zhai, W.X., Zhao, X.F., Pan, X.B., Xie, Q., Zhu, L.H.
traffic of cadmium, zinc, and manganese in rice. J. 2006. The rice HIGH-TILLERING DWARF1 encoding
Exp. Bot. 63: 2729–2737. an ortholog of Arabiodpsis MAX3 is required for
Yamaji, N., Ma, J.F. 2009. A transporter at the node negative regulation of the outgrowth of axillary
responsible for intervascular transfer of silicon in buds. Plant J. 48: 687–696.
rice. Plant Cell 21: 2878–2883. Zou, L., Sun, X., Zhang, Z., Liu, P., Wu, J., Tian, C., Qiu, J.,
Yamaji, N., Ma, J.F. 2014. The node, a hub for mineral Lu, T. 2011. Leaf rolling controlled by the homeodo-
nutrient distribution in graminaceous plants. Trends main leucine zipper class IV gene Roc5 in rice. Plant
in Plant Science 19: 556–563. Phys. 156: 1589–1602.
Inflorescence Structure
Inflorescences in Poaceae vary widely in their Asteraceae have panicles of capitula, and lilacs or
form, so widely that conventional terminology many Clusiceae are characterized by panicles of
for describing them is at best inadequate and at cymes (Stevens 2012).
worst, inaccurate. The problems with descriptive The second, and more serious, problem is
terminology have two sources. The first problem that inflorescences in the grasses are more com-
is that grasses bear compound inflorescences, in plex than in most other flowering plants, and that
which the terminal unit is in fact an unbranched often even the term “panicle of spikelets” fails to
cluster of flowers; each of these unbranched clus- describe the morphology precisely (Kellogg 2000;
ters is a tiny spike, or spikelet. In most literature Perreta et al. 2009). Authors rapidly devolve into
on the grasses, the spikelets are considered to be using intricate terminology, such as “a panicle of
analogous to flowers, and the inflorescence is racemes”, “a complex of partial inflorescences
described using terminology borrowed from ending in racemes” (e.g., Watson and Dallwitz
eudicots. Thus, an axis bearing sessile spikelets 1992 onward). These terms respect the complex-
is called a spike (e.g., Triticum, Lolium), as ity of the structures, but make it difficult to com-
though it were an axis bearing sessile flowers. pare parts and thus to make statements about
An axis with spikelets on pedicels is called a inflorescence evolution.
raceme (e.g., Brachypodium), analogous to a One proposed solution is the typology first
raceme in, for example, Arabidopsis, and an axis suggested by Troll (1964) and developed more
with branched branches is a panicle (e.g., Oryza, extensively by Weberling et al. (1993), Hernández
Avena, Panicum s.s., and many others), and so and Rua (1992), Vegetti (1991), and Vegetti and
forth. These are in fact a spike of spikelets, a Anton (1996). In this system, any group of flow-
raceme of spikelets, and a panicle of spikelets ers that is borne on the main axis (i.e., is the
respectively. (Note also that the term pedicel itself product of the inflorescence meristem) is known
is misapplied in the grasses, referring to the stalk as the florescence, whereas axillary branches
of the spikelet rather than the stalk of a flower.) bearing groups of flowers are known as para-
Because of this compound structure, some clades and their flowers called co-florescences.
authors prefer to use the term “synflorescence” Thus, in the grasses, the spikelet at the apex of
for grasses, to distinguish the inflorescence from the main inflorescence axis (if present) is called
non-compound inflorescences (Judziewicz et al. the florescence, and all other spikelets are co-
1999; Stapleton 1997). Here I use the term inflo- florescences. The complex systems of flores-
rescence in part because it is widely used and also cences and co-florescences are then considered
because the term is commonly applied to both to be polytelic synflorescences. If the arrange-
simple and compound structures. The existence ment of florescences and co-florescences on all
of compound inflorescences is not in itself a paraclades is the same, the synflorescence is
problem. Such structures occur in many angios- described as “homogenized”. Loss of the main
perms without causing apparent confusion. For florescence is described as “truncation”. This
example, Apiaceae often bear compound umbels, typological approach may be hard to connect to
26 Inflorescence Structure
known patterns of development; for example, the TRANSITION TO FLOWERING

term “homogenization” does not refer to any
obvious biological process. In addition, there Early in development, the shoot apical meristem
seems to be little biological distinction between produces only leaves, each of which develops a
the florescence and co-florescences (i.e., a spikelet meristem in its axil. After receiving an endoge-
terminating the inflorescence and a spikelet ter- nous (hormonal) or exogenous (light, heat) sig-
minating a lateral branch). However, the effort to nal, the meristem acquires the capacity to
apply a typological description to multiple species produce an inflorescence (Colasanti and Coneva
of grasses has led to many publications in which 2009; Wellmer and Riechmann 2010). In all flow-
adult inflorescence morphology is diagrammed ering plants, the transition to flowering is con-
and described carefully and precisely (e.g., Gasser trolled by a mobile signal, or florigen, known as
and Vegetti 1997; Kern et al. 2008; Perreta et al. Flowering locus T or FT, that is produced in the
2000; Reinheimer et al. 2009; Vegetti and Pensiero leaves and transported to the shoot apical meri-
1999). Whether or not one chooses to apply stem (Zeevaart 2008). In the meristem, FT binds
Weberling’s (1989) typology, the illustrations in to another regulator, FD. Homologues of both FT
these papers provide valuable comparative data, and FD have been cloned from maize and rice,
and can be a major improvement over attempts to although there may be more copies of each in the
stretch the words “panicle” and “raceme”. grasses than in other angiosperms; the function
Another way to view inflorescence morphol- of both sets of proteins appears to be conserved
ogy is to focus on meristems and their fate, begin- (Danilevskaya et al. 2008; Kojima et al. 2002;
ning with the apical meristem of the plant as it Komiya et al. 2008; Lazakis et al. 2011; Muszynski
shifts from producing leaves to producing bracts et al. 2006; Tamaki et al. 2007; Yan et al. 2006).
and higher-order meristems that will ultimately While the FT-FD module is shared among
lead to spikelet and, hence, flower production. In flowering plants, the proteins that regulate it are
most grasses, the inflorescence is easily identified not. The maize protein INDETERMINATE1 (ID1)
as the product of the shoot apical meristem after and its rice homologue RID1 (¼ OsID1) are only
the transition to flowering, and is a discrete struc- found in grasses (Baumel et al. 2002). These pro-
ture that is terminal on the culm. The problemat- teins are genetically upstream of the FT homo-
ical situations that occur in some woody logues, although they appear not to interact
bamboos (Bambuseae and Arundinarieae) and directly with FT-like genes (Lazakis et al. 2011;
some Andropogoneae are addressed in more Wu et al. 2008). Downregulation of ID1 or RID1
detail below. In this work any spikelet-bearing delays flowering or blocks it entirely. Thus, the
axis that is either terminal on the plant or on an internal regulation of flowering in the grasses
axillary branch with a subtending leaf or prophyll proceeds via mechanisms different from those
is called an inflorescence; this terminology fol- in other plants.
lows Clayton (1969). Under this definition, an In the leaves of many grasses, the FT homo-
inflorescence (as used here) is equivalent to a logues are also regulated by proteins similar to
synflorescence as defined by Judziewicz et al. FRUITULL in Arabidopsis. The FUL-like proteins
(1999). Aggregates of such units are simply plus the SEPALLATA (SEP)-like protein PANI-
described but not given particular names. CLE PHYTOMER2 (PAP2) are responsible for
Recent work in developmental genetics has integrating environmental signals such as vernal-
uncovered the effects of many proteins that con- ization and increasing the expression of FT
trol inflorescence development. As we begin to homologues (Preston and Kellogg 2008; Yan
understand the specific gene-level changes that et al. 2003, 2006). Once in the inflorescence,
affect the inflorescence, we can begin to see the the regulatory roles appear reversed, with the
basis for many of the morphological characters FT-FD complex activating Ful-like and pap2
that have been identified by taxonomists. (Kobayashi et al. 2012). This general regulatory
Descriptions that take into account development pathway has been pieced together from work on
and genetic controls allow more accurate assess- several cereal crops; it will be interesting to learn
ment of homology and more precise definition of whether it is indeed universal in the family as it
characters and character states. now appears.
TRANSITION TO FLOWERING 27
Fig. 4. Proliferating axillary branches in tribe Andropo- bracts plus prophylls. Pl + t’, Pl +t”, etc., higher-order
goneae, subfamily Panicoideae (Coelorachis aurita long paracladia with trophotagma. PsZ, short paracladia
(Steud.) A. Camus). Pl, long paracladium; in this text subzone; this corresponds to the terminal inflorescence,
these are simply called inflorescences. Pl + t, long para- which produces spikelet pairs from its apical meristem.
cladium with trophotagma, where trophotagma denotes (From Vegetti 1999)
In many grasses, it is possible to identify bition extends the entire length of the culm below
three zones or parts of the plant in which axillary the inflorescence; culm branches rarely form in
buds have different fates. The lowest zone is this subfamily, so that inflorescences are terminal
described as the “innovation zone” (Vegetti and on either the primary axis or on axes developing
Weberling 1996), in which buds from the lowest from tillers. In some genera of Andropogoneae
nodes on the plant can produce tillers that reiter- (Panicoideae) (e.g., Hyparrhenia, Andropogon,
ate the structure of the primary axis. Above this is Schizachyrium), axillary branches in the paracla-
an area called the “zone of inhibition” in which dial zone proliferate. Each axillary branch itself
axillary buds generally do not elongate at all produces an axillary branch, which produces
(Vegetti and Weberling 1996). The upper part of another axillary branch, leading to multiple
the plant is then the “paracladial zone” (Vegetti spikelet-bearing axes at each node. Each of the
1991; Vegetti and Anton 1996) in which axillary spikelet-bearing axes produces an adaxial pro-
buds can grow out and produce paraclades, phyll and a single leaf, which has a reduced
inflorescences whose structure is indistinguish- blade and a sheath enclosing the part of the axis
able from that of the terminal inflorescence. that bears the spikelets; the internode between the
The basic three-zone pattern is modified in prophyll and the leaf is generally elongated
several subfamilies. In Pooideae the zone of inhi- (Fig. 4). At maturity then it appears that the entire
upper part of the plant has become a diffuse Kokko 1993; Latting 1972; Whipple et al. 2010).
highly bracteate “inflorescence” in which the ter- Even in species in which bracts are not easily visi-
minal units are repeated complex branching ble under the microscope, molecular markers such
structures. Branch proliferation has been studied as TASSELSHEATH1 (Whipple et al. 2010) and
genetically in maize, and is regulated by the pro- TASSELSHEATH4 (Chuck et al. 2010) indicate
tein GRASSY TILLERS1 (GT1) (Wills et al. 2013); that cells have been set aside to form the bract. In
higher expression of gt1 reduced the number of the axil of each suppressed bract is a primary
proliferating branches. branch meristem, which quickly enlarges and
In Bambusoideae, Judziewicz et al. (1999) obscures the bract meristem.
have defined the synflorescence (equivalent to In the transition to flowering, the shoot apical
“inflorescence” in this work) as “the flowering meristem often changes phyllotaxis from the dis-
structure that emerges from the apex of a leafy tichous pattern of leaf production to a spiral
branch as that branch usually grows from a major pattern of inflorescence branch production. This
axis (or the culm apex).” In many woody phyllotactic shift appears to be plesiomorphic in
bamboos, this inflorescence is similar to an inflo- the family (Kellogg et al. 2013). The inflorescence
rescence in any other grass; the inflorescence axis meristem of the grass sister group, Ecdeiocolea
may be unbranched or branched, and the monostachya (Ecdeiocoleaceae) produces bracts
branches may be elongate or not (capitate). Such and flowers arranged in a spiral (Rudall et al.
conventional inflorescences have been described 2005). In subfamily Anomochlooideae Strepto-
as determinate or semelauctant (McClure 1966; chaeta produces primary branches that terminate
Stapleton 1997); neither term is used here. in flowers and appear to be arranged in a spiral
A more complex situation occurs in some of (Sajo et al. 2008); Anomochloa is reported to be
the woody bamboos, in which the transition from primarily distichous (Judziewicz and Soderstrom
vegetative to inflorescence growth appears to be 1989), but data from early inflorescence develop-
more gradual. In these plants, the branch meri- ment are not available. Spiral inflorescence phyl-
stem ceases to produce foliage leaves but instead lotaxis also appears in Pharus (Pharoideae) (Sajo
produces a set of bracts, some of which have buds et al. 2007), in Oryza and Zizania (Ehrhartoideae)
in their axils, and then eventually begins to pro- (Liu et al. 1998; Moncur 1981; Weir and Dale
duce flowers before terminating with a blind 1960), and many Panicoideae (Bess et al. 2005;
rachilla extension. The bracts are called glumes Brown et al. 2006; Doust and Kellogg 2002; Giulini
or gemmiparous bracts; it is not clear whether et al. 2004; Kellogg et al. 2004; Moncur 1981;
they are homologous with the glumes of more Reinheimer et al. 2005) and Chloridoideae (Liu
conventional spikelets. The buds in the bract et al. 2007; Moncur 1981). Distichous inflores-
axils can then grow out, themselves producing cences do occur in Panicoideae (e.g., Urochloa;
bracts, some with buds, then flowers, and then a Reinheimer et al. 2005) and some Andropogo-
rachilla extension. Depending on the species, this neae (Kellogg 2000), but are clearly derived.
can continue for up to five rounds of branching. In contrast to the spiral phyllotaxis in most
Such reiterating structures have been called pseu- subfamilies, the inflorescence branches of all mem-
dospikelets, because they have some aspects of bers of subfamily Pooideae except Brachyelytrum
spikelet identity and some of branch identity, and are two-ranked, although strict distichy (branches
have been described as indeterminate or iterauc- separated by an angle of 180 ) appears to have been
tant (McClure 1966; Stapleton 1997). derived after the divergence of Nardeae (Kellogg
et al. 2013). The two-ranked inflorescence is quite
obvious even at maturity in members of the tribe
BRACT FORMATION AND INFLORESCENCE PHYLLOTAXIS Triticeae such as Hordeum vulgare (barley) and
Triticum aestivum (wheat), but also can be
The inflorescence meristem initiates bracts on its observed in early development in many Poeae
flanks; in many species (e.g., Hordeum vulgare, (Evans 1940; Fraser and Kokko 1993; Latting
Pennisetum glaucum, Trisetum spicatum) these 1972; Moncur 1981). Recently the two-ranked inflo-
are clearly visible in early development, but their rescence has also been demonstrated in Nardeae,
development is suppressed (Evans 1940; Fraser and Stipeae, Diarrheneae, Meliceae, and Brachpodieae
NUMBERS OF ORDERS OF BRANCHING 29
(Kellogg et al. 2013). The failure of the inflorescence development is similar; in both cases the axis is
meristem to shift to spiral phyllotaxis in Pooideae unbranched.
is thus a synapomorphy for the large clade that is Many other taxa produce only primary
sister to Brachyelytrum, and appears to correlate branches. In these genera, each primary branch
with a shift from shaded to open habitats. meristem gives rise to spikelet meristems; again
Mutations in the rice gene Aberrant Panicle the spikelets may be sessile (e.g., Heteranthoecia,
Organization1 (apo1), which encodes an F-box Eleusine) or pedicellate (e.g., Paspalidium, Gym-
protein homologous to UNUSUAL FLORAL nopogon). Finally, the primary branches may
ORGANS (UFO) in Arabidopsis, display altered themselves branch, producing second-, third-,
phyllotaxis (Ikeda et al. 2005, 2007). While the or higher-order branches.
inflorescence of rice normally produces primary The main inflorescence meristem may termi-
branches in a spiral, apo1 mutants produce two- nate in a spikelet (e.g., Avena sativa, Triticum
ranked primary branches. The change in phyllo- spp., Sorghum bicolor) or not (e.g., Setaria italica,
taxis correlates with a change in the shape of the Zea mays), and may abort soon after forming
meristem. It is possible that changes in apo1 primary branches (e.g., Oryza sativa, Urochloa
expression underlie the pooid synapomorphy. plantaginea, Brachiaria decumbens) (Kellogg
Virtually all primary branch meristems pro- et al. 2013; Moncur 1981; Reinheimer et al.
duce secondary meristems in two ranks (Doust 2005). The fate of the inflorescence meristem
and Kellogg 2002). Although the two ranks are often varies among closely related species (see,
often separated by an angle of 180 (i.e., are for example, Kellogg et al. 2013; Reinheimer and
strictly distichous), in many taxa the secondary Vegetti 2008; Reinheimer et al. 2009), suggesting
branches form on one side of the primary branch. that it may be relatively easy to shift in evolution-
For example, in Paspalum, the primary branches ary time, a point also made by Butzin (1979). The
each bear two ranks of secondaries, which always fate of the inflorescence meristem is independent
form on the abaxial side of the primary. The of that of the primary-, secondary-, and higher-
genetic control of this pattern is unknown. In order branches, which may also terminate in spi-
such unilateral primary branches, the branch kelets or not.
axis is often expanded laterally as though it has The genetic control of branching is agrono-
acquired some aspects of leaf identity. This is mically important and thus has been studied
particularly noticeable in some taxa such as extensively in the cereals (Bommert et al. 2005;
Chlorocalymma, in which the broad rachis of Kellogg 2007; Thompson and Hake 2009). Forma-
the primary branch droops around and encloses tion of branch and floral meristems depends on
the secondaries. auxin transport and biosynthesis, as it does in
eudicots. The auxin efflux carrier PINFORMED1
(PIN1) transports auxin in maize and rice as it
NUMBERS OF ORDERS OF BRANCHING does in Arabidopsis thaliana (Gallavotti et al.
2008), and is phosphorylated by BARREN
Grass inflorescences may be unbranched, or may INFLORESCENCE2 (BIF2 ¼ PINOID in Arabi-
produce primary branches only, or may have up dopsis), an interaction that is also conserved
to six or more orders of branching. Whether among flowering plants. However, BIF2 also
branches are produced or not is determined by phosphorylates an unusual basic helix-loop-
the fate of the inflorescence meristem, which var- helix transcription factor, BARREN STALK1 (in
ies between species and genera (reviewed by Kel- maize), which is orthologous to LAX PANICLE in
logg 2000, 2006; Malcomber et al. 2006; Perreta rice, an interaction not reported for dicots (Skir-
et al. 2009). Inflorescence meristems may pro- pan et al. 2008). The expression pattern and pre-
duce spikelet meristems directly on their flanks, sumed function of BA1/LAX is similar among all
and the spikelets may remain sessile (e.g., Triti- grasses, but somewhat different from the patterns
cum, Ctenium), creating a “spike-like” structure, in eudicots (Woods et al. 2011). Because compa-
or the spikelet may develop a pedicel (e.g., Bra- rable data are not available for other monocots, it
chypodium, some Perotis), and the inflorescence is not clear whether the role of BA1/LAX is
thus be “raceme-like”. However, the underlying unique to the grasses or to a larger clade.
Inflorescence maturation, maturation of each (2009), the architecture of the inflorescence is

individual branch, and maturation of flowers determined by the precise developmental timing
within a spikelet may be acropetal or basipetal, of the conversion of a branch meristem to a
and the direction of maturation in different com- spikelet meristem. If the conversion happens
ponents of the inflorescence may vary (Reinhei- early in development, then few higher-order mer-
mer et al. 2009). For example, the inflorescence of istems are produced, whereas if conversion from
Hyparrhenia is made up of two branches, the branch meristem identity to spikelet meristem
main axis and a lateral branch; one matures identity is delayed, then there is an opportunity
acropetally and the other basipetally (Le Roux for producing more branches.
and Kellogg 1999). In species such as Urochloa The number of primary branches varies inde-
plantaginea and Brachiaria decumbens, branch pendently of the number of secondaries, an
initiation appears to be basipetal, and the pri- observation that is supported by extensive
mary branches may initiate below already formed genetic data. The number of primary branches
branches (Reinheimer et al. 2005; St€ ur 1986), is positively correlated with the amount of tran-
suggesting that meristems form de novo. script of the gene Apo1, mentioned above for its
role in control of phyllotaxis. Apo1 is the gene
underlying the QTL Primary Branch Number in
NUMBERS OF BRANCHES OR SPIKELETS AT EACH ORDER OF rice; higher levels of Apo1 transcript lead to
BRANCHING more primary branches and lower levels lead
to fewer (Terao et al. 2009). While APO1 posi-
The range of values for primary, secondary, and tively regulates the number of primary branches,
tertiary branch number varies extensively the proteins DENSE AND ERECT PANICLE1
between species, although individual species (DEP1), OsMADS34 (¼ PANICLE PHYTOMER2,
often have a characteristic and consistent range or PAP2), and ERECT PANICLE3 (also called
of values. The high variability of these characters LARGER PANICLE) restrict the number (Gao
is obvious simply by considering illustrations of et al. 2010; Huang et al. 2009; Kobayashi et al.
closely related species in almost any grass genus. 2010; Li et al. 2011; Piao et al. 2009). The four
There is often species-specific variation in proteins differ, however, in their effect on sec-
whether the inflorescence is dense (with many ondary branches. APO1 positively regulates the
primary branches) or sparse (with few primary number of secondary branches per primary,
branches), whether the primary branches are DEP1 restricts the number, and OsMADS34 and
densely covered with secondaries or not, and EP3 have little effect at all on the number of
whether tertiary and quaternary branches are secondary branches per primary. Three other
common or rare. The few studies that record proteins, ERECT PANICLE2 (EP2) and rice CEN-
comparative data on numbers of primary TRORADIALIS1 and 2 (RCN1 and RCN2) have
branches find a broad range of variation among no effect on the number of primary branches but
species (Spartina, Kern et al. 2008; Melinidae, restrict the number of secondaries (Nakagawa
Reinheimer and Vegetti 2008; Setaria, Doust and et al. 2002; Zhu et al. 2010). Comparative studies
Kellogg 2002). In Spartina (now part of Sporobo- on these genes and proteins are limited, but DEP1
lus s.l.; Peterson et al. 2014), the number of pri- affects the number of primary and secondary
mary branches does not correlate well with the branches, and inflorescence elongation in rice,
phylogeny (Baumel et al. 2002; Fortune et al. wheat and barley, suggesting a general mecha-
2006), indicating that the number of branches is nism (at least in the BEP clade) for branching
easily modified over evolutionary time. control (Huang et al. 2009).
Perhaps not surprisingly, many genes control The number of primary branches also corre-
branch number. Many of these have pleiotropic lates with the number of vascular bundles in the
effects on other aspects of inflorescence develop- peduncle, suggesting that the number is limited
ment, and thus must be responding to a variety of by carbohydrate supply. EP2 and EP3 control
selective pressures. In general, branch number vasculature and sclerenchyma in the inflores-
controls spikelet number, and hence controls cence and thus affect branch number, along
the number of seeds; there is thus a direct effect with whether the inflorescence is upright or
on fitness. As noted by Ikeda-Kawakatsu et al. nodding (Piao et al. 2009; Zhu et al. 2010).
AXIS ELONGATION 31
Many of the proteins that regulate branch mutants of rice, the normally lax inflorescence of
number do so by interacting directly or indirectly rice is borne upright; this appears to be due to
with the cytokinin pathway. The locus Grain increases in the number of vascular bundles, par-
number 1 (Gn1) in rice encodes CYTOKININ ticularly small vascular bundles, and also in scle-
OXIDASE2 (CKX2), which breaks down cytoki- renchyma in the peduncle (Huang et al. 2009;
nin and limits the number of spikelets and pani- Piao et al. 2009; Zhu et al. 2010). Taxonomists
cle branches (Ashikari et al. 2005), as does its have generally not explored this character in a
ortholog in wheat (Zhang et al. 2012). Conversely comparative context, but it would be of interest to
the cytokinin biosynthetic protein LONELY GUY know whether vascular supply and amount of
(LOG) increases the number of branches (Saka- sclerenchyma generally correlate with the width
kibara 2006). Also regulating CKX2 (directly or of the inflorescence.
indirectly) is EP3, which encodes an F-box pro- Branch angle is also controlled by the activity
tein (Piao et al. 2009); levels of EP3 expression of the pulvinus at the base of each primary inflo-
correlate with levels of CKX2 expression (Li et al. rescence branch. The pulvinus is a prominent
2011). Spikelet number per panicle is also swelling that becomes turgid and forces the
affected by RICE OUTERMOST CELL-SPECIFIC branches away from the main axis (Arber 1934;
GENE5 (ROC5) (Zou et al. 2011). Galinat 1959). RA2 and REL2 contribute to nor-
In Andropogoneae, the spikelets are borne in mal cell structure of the pulvinus, whereas BAR-
pairs; one member of the pair is pedicellate REN STALK FASTIGIATE and BRANCH ANGLE
whereas the other is sessile or nearly so. Studies DEFECTIVE1 control the number of cells (Bai
of developmental morphology, developmental et al. 2012; Gallavotti et al. 2010, 2011). The pres-
genetics, and gene expression all indicate that ence, size and vasculature of the pulvinus have
the pedicellate spikelet is terminal on a very received little or no attention in the comparative
short branch, and the sessile spikelet terminates literature.
the sole lateral branch. The proteins RAMOSA1
(RA1), RAMOSA2 (RA2), RAMOSA3 (RA3), and
RAMOSA ENHANCER LOCUS2 (REL2) working AXIS ELONGATION
together prevent formation of more than one
sessile spikelet and are thus responsible for the Elongation of inflorescence axes occurs after
formation of the spikelet pair (Bommert et al. branching is complete, spikelet meristems have
2005; Gallavotti et al. 2008, 2010; Satoh-Nagasawa been specified, and glumes and lemmas are
et al. 2006). The genes encoding RA1 and RA3 are initiated. Elongation is thus developmentally dis-
only present in the PACMAD clade (and thus may tinct from branching. Genetic studies in a few
be synapomorphic) and are completely absent grasses also show that it is controlled by different
from the genomes of rice and Brachypodium genes (Doust 2007; Doust et al. 2004). In Triticum
(Satoh-Nagasawa et al. 2006; Vollbrecht et al. aestivum, an APETALA2-like gene known as Q
2005; Reinheimer and Kellogg, unpublished); affects elongation of the rachis, and was involved
thus, the RA1-RA2-RA3-REL2 module must also in domestication (Simons et al. 2006); higher
be unique to the PACMAD group. However, RA2 levels of Q make the inflorescence more compact
and REL2 are present in all grasses. Therefore, (see also below). In Hordeum vulgare, another
they must bind with different proteins in the BEP APETALA2-like gene, HvAP2, which is distinct
clade since RA1 and RA3 are not available. from Q, also controls rachis elongation and
thereby number of spikelets (Houston et al.
2013). HvAP2 is regulated by microRNA172
BRANCH ANGLE (miR172); when HvAP2 is mutated in a way that
reduces the interaction with miR172, the inflores-
The angle of inflorescence branches, whether cence axis continues initiating spikelets and fails
spreading or upright, varies considerably. The to elongate. The result is a short inflorescence
erect panicle phenotype has been of particular with an abnormally high number of spikelets.
interest in rice because more erect panicles cor- Because of the high HvAP2 levels, spikelet differ-
relate with increased yield. In DEP1, EP2 and EP3 entiation is delayed, and the window for
inflorescence elongation is shortened (Houston The bracteate structure of the spikelet leads to its
et al. 2013). interpretation as a contracted inflorescence axis.
Elongation of each order of axis is controlled However, this is likely to be an over-simplifica-
independently of the others. Because of this, tion; the mutant phenotype of BRANCHED SILK-
inflorescences with the same number of orders LESS1 (BD1) in maize and FRIZZY PANICLE
of branches and the same number of branches at (FZP) in rice hints that a spikelet is not simply
each order can still look quite different at matu- an abnormally short spike (Chuck et al. 2002;
rity depending on which axes elongate. For exam- Komatsu et al. 2003). If the proteins simply kept
ple, the internodes of both the main axis and the the spikelet axis short, one would expect the
primary branch axes may remain close together mutant phenotype to lead to an elongate branch,
in digitate inflorescences such as Digitaria or but with the normal number of flowers. However,
Cynodon, or the main axis may elongate while mutations in bd1/fzp cause the production of
the primary branch axes do not as in Leptochloa additional branches and flowers (see below),
or Bothriochloa, or internodes may elongate on with the precise phenotype differing between the
both orders of branching as in Panicum or Avena. tassel and the ear of maize and the branched
In some taxa such as Urochloa maxima inflorescence of rice; in addition, axillary meris-
(¼Megathyrsus maximus), internode elongation tems are produced in the axils of the glumes in a
is not consistent along an axis. In this case, a pattern reminiscent of a pseudospikelet in
group of short internodes is followed by one bamboos. Bd1/fzp is expressed in the axils of
long one, followed by another cluster of short glumes in all grasses investigated, suggesting
internodes (Reinheimer et al. 2005). At maturity that its normal function is to repress branching
the primary branches of such inflorescences in that position.
appear whorled or verticillate. If the internode In other respects, the spikelet is similar to a
below the secondaries fails elongate, spikelets on flower. The spikelet is a developmentally integrated
secondary branches appear clustered as in Hor- unit and is determinate in the sense that a flower
deum, or the secondaries themselves cluster as in is – a particularly canalized developmental pro-
Agrostis or other Pooideae. In some Pooideae gram. Although the rachilla ends blindly in the
such as Nephelochloa, inflorescence branches spikelets of many species, the spikelet is not an
are described as whorled; in these species the endlessly reiterating structure. Once the “spikelet
primary branches are almost certainly distichous developmental program” is switched on, the axis
as they are in all pooids, although development ends. (Note that the word “determinate” is used in
has never been investigated for this genus. two different ways in the grass developmental liter-
Apparent whorls are created when a large number ature. Among classical morphologists, a determi-
of higher-order branches, each with contracted nate axis is one that ends in a flower. By this
basal internodes, originates from the distichous definition, many if not all grass spikelets would
primaries. have indeterminate axes because the flowers
appear to be lateral. However, among developmen-
tal geneticists, a determinate structure is one with a
SPIKELETS fixed number of parts. By this definition, the spike-
let is determinate because the number of flowers is
Within the spikelet, flowers occur in distichous either fixed or varies within a narrow range.)
phyllotaxis on a short axis, the rachilla (literally, a A spikelet is also similar to a flower in terms
little rachis). At the base of the spikelet below the of expression of genes that are flower-specific in
flowers are two more bracts, known as glumes. other families. In most angiosperms, including
(These give the grass family the classical name the grass outgroups, the proteins SQUAMOSA
Glumiflorae, transliterated to gloomy-florae by and FRUITFULL (FUL), and certain SQUAMOSA
bored agrostology students.) The glumes are PROMOTER BINDING PROTEIN-LIKE (SPL)
defined as bracts with nothing in their axils. proteins are expressed throughout the flower
The grass spikelet is an evolutionary novelty. but generally not in subtending bracts; they are
It shares some characteristics with a branch sys- thus markers for floral identity (Preston and Hile-
tem and some with a flower, but is clearly neither. man 2010; Preston et al. 2012); the SPL proteins
SPIKELETS 33
are direct regulators of FUL/SQUA (Klein et al. Spikelet size varies by more than an order of
1996). In grasses, SQUAMOSA (¼ APETALA1) is magnitude, from minute (1 mm) to relatively
absent, whereas its paralogue FUL is duplicated large (1 to 2 cm), to enormous (e.g., up to 10 cm
to form FUL1 (¼ VRN1) and FUL2 (Preston and in Colanthelia, Bambuseae). Spikelet size is not a
Kellogg 2006); the best characterized of the SPL direct reflection of flower number, in that some of
proteins is TEOSINTE GLUME ARCHITEC- the largest spikelets (e.g., Hesperostipa) have only
TURE. FUL1, FUL2 and TGA are expressed not one flower and some very small spikelets (e.g.,
only in the flowers but also in the glumes (Preston some Eragrostis species) have several. The con-
et al. 2009, 2012; Preston and Kellogg 2007; Wang trols of spikelet size are largely unknown; it is
et al. 2005). Homologues of these same proteins possible that spikelet size is simply a direct cor-
are present only in floral organs of the grass out- relate of grain size. If this is true, then ongoing
group Joinvillea. Thus, the expression domains of research in rice on grain size will likely uncover
FUL1, FUL2, and TGA are expanded in the some of the regulatory genes.
grasses to include the bracts subtending the flow- The number of flowers per spikelet varies
ers; this change in expression is another synapo- among grasses, but is often fixed within a clade.
morphy of the family. For example, members of Ehrhartoideae have
The pseudospikelets of bamboos are peculiar three flowers (only one fertile), Panicoideae
in that they share some aspects of spikelets and have two, and some clades of Pooideae have
some aspects of branches. They are usually inter- only one. The number of flowers in turn is con-
preted as spikelets in which the most proximal trolled by the length of time that the spikelet
glumes have buds in their axils; these buds meristem actively produces lateral organs. In
develop into new branches with new glume-like both rice and maize, spikelet meristem activity
structures in their axils. The bud-bearing glumes is regulated by proteins known as INDETERMI-
are often followed by one or more conventional NATE SPIKELET1 (IDS1) in maize and its para-
glumes without buds before the axis produces a logue SISTER OF IDS (SID) (Chuck et al. 1998)
flower. It would be of interest to determine (or IDS1-like; Malcomber et al. 2006); related
whether bamboos with pseudospikelets have proteins in rice are OsIDS and SUPERNUMER-
alterations in the structure or expression of Bd1/ ARY BRACT (SNB). Mutations in ids1 lead to
Fzp. A mutation in durum wheat (Triticum turgi- production of extra flowers in the spikelet
dum ssp. durum), branched head, also produces a (Chuck et al. 2008), and mutation of both ids1
structure similar to a pseudospikelet, with a sec- and sid together causes production of extra
ondary spikelet forming in the axil of the glume bracts. Likewise mutation of both Osids and snb
(Shitsukawa et al. 2009); this gene has not been lead to increased numbers of rudimentary glumes
cloned, however. (Lee and An 2012). IDS1, SID1, OsIDS, and SNB
Spikelet compression varies in Poaceae, and are all AP2-like proteins, similar to the wheat
generally correlates with phylogeny. Spikelets domestication gene Q (Simons et al. 2006); like
may be dorsiventrally compressed, laterally com- Q they are controlled by a microRNA in the
pressed or terete. In laterally compressed spike- miR172 family (Chuck et al. 2007, 2008). Thus,
lets, the glumes and lemmas are folded around variation in number of flowers per spikelet
their midribs (conduplicate); this compression observed in grasses could be regulated by the
pattern is nearly universal in Ehrhartoideae, precise timing of expression of miR172.
Bambusoideae, Pooideae, Chloridoideae, and Flower number is also specified by genes act-
Danthonioideae. In dorsiventrally compressed ing later in spikelet development. In rice the
spikelets, in contrast, the glumes and lemmas number of flowers is controlled by MULTI-FLO-
are flat to slightly curved, and spread out on RET SPIKELET1 (MFS1) and TONGARI-
either side of the midrib; this pattern appears BOUSHI1 (TOB1), whose effect is obvious after
most obviously in Panicoideae, but also in Phar- specification of glumes and sterile lemmas (Ren
oideae and in the herbaceous bamboos. The third et al. 2013). MFS1 is also an AP2-like protein and
pattern, less common in terms of numbers of is apparently regulated by microRNAs.
species, is terete spikelets, which are found in Glume morphology is often taxonomically
Aristidoideae and Stipeae (Pooideae). informative and forms a standard part of most
species descriptions. Glumes may be shorter than the position of the distalmost flower is less clear.
the adjoining lemma or longer, sometimes being However, SEM data show that a vestige of the
long enough to enclose the entire spikelet. One or rachilla apex is often visible in early development
both glumes may be suppressed in development (see Bess et al. 2005; Doust and Kellogg 2002).
and reduced to a vestigial flap or a ridge. The The rachilla may be variously elongate or
number of veins varies from zero to many, and contracted between the glumes and lemmas. If it
these may be prominent or obscure. The texture is longer between the upper glume and lowest
of the glumes can be similar to that of the lemma, it is often described as a stipe (e.g., in
lemmas, or they can be harder or softer. In Calamovilfa (now part of Sporobolus s.l., Peterson
some taxa (e.g., Mnesithea or Hackelochloa, et al. 2014)). No data are available on the control
Andropogoneae), the glumes are not only hard of rachilla elongation.
but also elaborately sculptured to form warts,
hooks, or other coruscations. The glumes of
most grasses lack awns, but in some species DISARTICULATION
awns are present and can be longer than the
entire spikelet. The products of the genes Q and The grass inflorescence is clearly adapted for seed
Tenacious glumes in wheat control the amount of dispersal (see section on Dispersal below), and
sclerenchyma at the base of the glume (Nalam there is considerable variation in the pattern of
et al. 2007); in wild wheats the glumes are hard disarticulation (Doust et al. 2014); the disarticu-
and tightly enclose the flower and caryopsis, lation pattern is often consistent within clades
whereas in domesticated wheats there is less scle- and so is commonly used as a taxonomic charac-
renchyma and the glumes break easily during ter. Most commonly, an abscission zone forms at
threshing. the base of the flower, so that the caryopsis falls
Maturation of flowers within a spikelet may with the lemma, palea, and a short rachilla seg-
be acropetal, as in Pooideae, Bambusoideae and ment. In such species, the glumes generally
Chloridoideae, or basipetal as in Ehrhartoideae remain on the parent plant. There is also varia-
and Panicoideae s.s.; this maturation pattern is tion in whether the palea and lemma adhere
independently derived but synapomorphic in firmly to the caryopsis or whether the caryopsis
each of the latter two subfamilies. Expression of is ultimately shed. In the other common pattern
the SEPALLATA-like gene, Leafy hull sterile1 (e.g., in virtually all Paniceae and Paspaleae),
(Lhs1/OsMADS1) correlates with the direction of disarticulation is below the glumes, such that
flower maturation (Malcomber and Kellogg 2004; the entire spikelet is shed as a unit; subsequently
Reinheimer et al. 2006), and thus underlies the the rachilla breaks up, releasing the upper flower.
developmental synapomorphies for Ehrhartoi- In many members of Andropogoneae and Triti-
deae and Panicoideae. In the two subfamilies in ceae, and in the genera with narrow “rat-tail”
which the distalmost flower matures first (Ehr- inflorescences (e.g., Hainardia), disarticulation
hartoideae and Panicoideae s.s.), Lhs1 expression occurs at each node of the inflorescence axis so
is confined to the distal flower. In contrast, in that the dispersal unit is the spikelet plus a rachis
spikelets with acropetal maturation, Lhs1 is internode. In a few taxa (e.g., some species of
expressed in all flowers. Aegilops, Eragrostis sessilispica), the entire inflo-
Some authors have interpreted the distalmost rescence forms the dispersal unit, with disarticu-
flower of the spikelet as being terminal, whereas lation at the most proximal node.
others infer that all flowers are lateral on the Given the variation in the location of abscis-
rachis. Evidence is accumulating for the latter sion zones, it is not surprising that abscission is
interpretation. If development is acropetal, the under the control of several independent genes.
more distal flowers of the spikelet are often Most current data come from cereal crops, in
reduced in size, in some cases being little more which disarticulation is called “shattering” and
than a rudiment with a minute lemma, and often is an undesirable trait that was selected against
the rachilla is prolonged beyond the distalmost early in domestication (see Domestication). Thus,
flower. In this sort of spikelet, all flowers are the Q gene in wheat prevents disarticulation of
clearly lateral. When development is basipetal, the rachis, suggesting that its recessive wild-type
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Flower Structure
Except in Anomochloa and Streptochaeta, grass branches are very short to virtually non-existent
flowers are made up of a gynoecium, androecium, and dominated by the feathery stigmatic tissue. It
lodicules, a palea and a lemma (Fig. 5). While the is thus common to count the number of stigmas,
structure and homologies of the gynoecium and rather than the number of stylar branches. In
androecium are similar to those of other angios- several grass subfamilies, the stylar branches
perms, the homologies of the surrounding struc- appear to originate from separate positions at
tures are less obvious and the terminology more the apex of the ovary, as though the style itself
confusing. Accordingly, flowers are described has become subsumed into the ovary. In this case
here beginning with the gynoecium and working the stylar branches are often referred to as “style
outward. bases” or simply “styles”; thus, a distinction is
The grass outgroups have a gynoecium with rarely made between the obviously compound
two or three carpels and the corresponding num- stylar structure in some subfamilies versus the
ber of locules; the ancestral state is almost cer- outgrowth of a (presumably) single carpel in
tainly three. Although the apex of the carpels many others.
forms a transmitting tract, the length of this var- In subfamily Anomochlooideae, Strepto-
ies considerably. Therefore, if a style is defined as chaeta has one style, with three branches each
a structure borne of the fusion of several carpels, ending in a stigma (Judziewicz and Soderstrom
the length of the style is variable. Distal to the 1989; Preston et al. 2009; Sajo et al. 2008). In
style are stylar branches; these are generally contrast, Anomochloa has a single structure that
extremely short. Most of the stylar branch is presumably corresponds to a single stylar branch
receptive and is therefore generally called a and stigma (Sajo et al. 2012). All Pharoideae have
stigma. Joinvillea (Joinvilleaceae) has a three-car- one style with three branches and three stigmas,
pellary, three-locular ovary, with three virtually as does Puelia (Puelioideae), but Guaduella (also
sessile stigmas (Whipple et al. 2007), as does Puelioideae) has only two branches and two stig-
Georgeantha (Edeiocoleaceae) (Rudall et al. mas. Most descendants of the common ancestor
2005); in Ecdeiocolea the number of carpels and of the BEP and PACMAD clades have two stylar
stigmas is reduced to two (Rudall et al. 2005). In branches and thus two stigmas, but three are
both Joinvilleaceae and Ecdeiocoleaceae, there is present in many bamboos. The GPWG (2001)
little tissue that could be called a style. called all descendants of the common ancestor
In Poaceae, the number of locules in the of Puelioideae and the remaining grasses “the
ovary is reduced to one, and is invariable and Bistigmatic clade”, implying that the reduction
synapomorphic for the family. However, the to two stylar branches is synapomorphic; this
structure of the stylar tissue, the number of stylar interpretation requires the three branches in Pue-
branches, and therefore the number of obvious lia and some Bambusoideae to be derived, but it
stigmas, is variable. The distinction between seems equally plausible that stylar branch num-
styles and style branches is rarely made in the ber was not fixed until after the common ancestor
literature on grasses. In general both the style and of the BEP and PACMAD clades. In some species,
40 Flower Structure
factor (AGAMOUS) that specifies both carpel

identity and floral meristem determinacy, the
grasses have two AG-like genes, originating
from a duplication early in grass evolution. This
duplication was followed by divergence in gene
function, such that one copy of the gene (ZAG1 in
maize, OsMADS58 in rice) specifies carpel iden-
tity and the other copy (ZMM2, OsMADS3) is
preferentially expressed in stamens (Dreni et al.
2011; Kramer et al. 2004; Mena et al. 1996; Yama-
guchi et al. 2006); other genes are also involved (e.
g., DROOPING LEAF (DL), a YABBY-class tran-
scription factor, Li et al. 2011; Yamaguchi et al.
2004).
Grass stamens are conventional, and have
Fig. 5. Flowers. A Floral diagram of a grass with three two thecae and four sporangia. The filament in
lodicules and six stamens. B Floral diagram of a grass with Streptochaeta and Anomochloa is centrifixed
two lodicules and three stamens. C Abaxial view of flower (¼ pseudobasifixed), such that the filament is
of Yushania (Arundinarieae) showing pistil with two sty-
lar branches and two stigmas, three stamens, and three completely surrounded by the connective at the
vascularized and ciliolate lodicules; lemma and palea have point of insertion (Arber 1929; Judziewicz and
been removed. (From GPWG 2001, p. 402, with permis- Soderstrom 1989; Sajo et al. 2009, 2012); such
sion from Missouri Botanical Garden Press; A and B anther attachment is often associated with buzz
drawn by M. Kojima, C by D. Friedrick)
pollination. In most other grasses, the filament is
attached to the abaxial side of the anther and
the third stylar branch can sometimes be seen in elongates swiftly at anthesis so that the anther is
scanning electron micrographs as a tiny apical exserted from the flower. Anthers are versatile
flap (Le Roux and Kellogg 1999). Further reduc- and tremble in response to any breeze, dispersing
tions are seen in some species; for example in copious pollen. Wind pollination appears to be
cultivated maize, the two branches form but are derived in the family, rather than ancestral (Givn-
congenitally fused to form the familiar silk. In ish et al. 2010), but this depends on interpretation
subfamily Pooideae, the stylar branches are sepa- of Anomochlooideae as insect pollinated; the lat-
rate in the earliest diverging lineage, Brachyely- ter has not been confirmed by observation but is
trum. In the next lineage, Nardeae, there is a single merely inferred from anther morphology.
structure; it is unclear whether this represents an Anther dehiscence is consistently longitudi-
unbranched style, or a single stylar branch. After nal. Dehiscence is latrorse in Streptochaeta
divergence of Nardeae, the style branches origi- (Arber 1929; Sajo et al. 2009), whereas in Anomo-
nate from a clear style, whereas the stylar chloa it is introrse proximally and latrorse distally
branches are once more apparently independent (Sajo et al. 2012). In contrast, in grass species with
in Diarrheneae plus its large sister clade. three anthers, the abaxial anther is latrorse or
Styles in most grasses are solid, a condition introrse, whereas the lateral anthers are consis-
rarely found in monocots (Rudall et al. 2005; Sajo tently extrorse (Anton and Astegiano 1973).
et al. 2012), although open styles are described in The earliest grasses and their ancestors had
Pharus (Pharoideae) (Sajo et al. 2007). Pollen six stamens, a condition that appears today in
tubes are then transmitted through the “stylar Streptochaeta, Pharoideae, Puelioideae, and in
core” (Arber 1934). Grass stigmas are plumose some Bambusoideae and Ehrhartoideae
and the receptive branches are multicellular, a (Fig. 5A). In a few Bambusoideae, and in Luziola
character shared with the other families of the of Oryzeae, stamen number is higher than six.
graminid Poales. Reduction of stamen number to four has
Developmental genetic controls of the grass occurred independently in Anomochloa and in
gynoecium appear to be grass-specific. Whereas some species of Ehrharta. Most of the grasses,
eudicots have a single MADS-box transcription however, have only three stamens (Fig. 5B); this
Flower Structure 41
number might be synapomorphic but indepen- by the same genetic controls that regulate the
dently derived in Pooideae and the PACMAD inner tepals of non-grass Poales. The downstream
clade, although some Bambusoideae also have genetic pathways have clearly shifted so that
three stamens which may affect character optimi- instead of activating tepal-like development and
zation in the BEP clade. Further reduction to two photosynthesis, the regulatory genes activate lod-
or even one stamen has occurred repeatedly and icule development.
independently, and often is associated with flow- In Streptochaeta, the three bracts outside the
ers that are obligately self-pollinated and fail to stamens appear to be governed by the same regu-
open at maturity. latory genes that control lodicules (Preston et al.
Outside the stamens are two or three small 2009; Whipple et al. 2007); the three bracts thus
structures known as lodicules, which correspond share with lodicules and inner tepals similarity in
to the inner perianth whorl of other monocots position, gene expression and number. In Anom-
(Fig. 5A–C). As the flower nears maturity, the ochloa, the stamens are surrounded by a ring of
lodicules increase turgor pressure and force the fimbriate structures, which appear to be position-
flower open. The lodicules are generally vascular- ally homologous to lodicules; genetic controls of
ized, although in all Pooideae except for Brachye- their development are unknown (Sajo et al. 2012).
lytrum the vasculature has been lost (GPWG Three lodicules are found in Pharoideae, Bambu-
2001; Jirásek and Jozı́fóva 1968). soideae, and some Pooideae (in tribes Phaenos-
The lodicules represent modified inner peri- permateae and Stipeae), although one of the three
anth organs. Transcription factors that specify appears in a whorl outside the other two. The
petal and stamen identity in other angiosperms significance of this is not known. It has been
(the so-called B class genes) have been identified suggested that two of the lodicules might be mod-
and characterized in multiple grasses (Ambrose ified petals, but that the third might be a sepal;
et al. 2000; Nagasawa et al. 2003; Whipple et al. since there are no genetic or developmental data
2004, 2007). These genes are expressed in both on species with three lodicules, this remains an
stamens and lodicules, as would be expected if open question. However, because most monocots
lodicules were highly modified petals. Impor- have two adaxial sepals rather than one, it seems
tantly, Whipple et al. (2007) find that the B-class more likely that the third lodicule is a member of
genes are also expressed in the inner tepals of the inner perianth, despite its position on the
Joinvillea and Elegia in Restionaceae, indicating floral axis.
continuity of information between the grasses Outside the lodicules is an adaxial bract, the
and their close relatives. In addition, when any palea (Fig. 5A, B). Its position outside the lodi-
of these genes is mutated, the stamens are cules suggests that it could correspond to the
replaced by carpel-like structures and the lodi- fusion of the two outer tepals, a suggestion
cules by palea-like structures. In eudicots, made by Stebbins (1956a, b). A more common
mutants in orthologous genes replace stamens interpretation of the palea, however, is that it is a
by carpels and petals by sepals. Thus, the replace- prophyll. Prophylls in monocots are consistently
ment of lodicules by organs in the next outer adaxial, and are generally two keeled; the palea is
whorl would be expected if they were indeed of adaxial and is two-keeled in most (but not all)
petaloid origin. grasses. However, in most monocots including
Previous authors have speculated that lodi- the grass outgroups, prophylls appear on lateral
cules might be staminodes, sepals, or nectaries, branches, but not on pedicels of flowers. Con-
but the evidence does not support this. If lodi- versely, most monocots including the grass out-
cules were staminodial, then in mutants they groups do have a pair of adaxial tepals. In
should be converted to carpels, which does not Ecdeiocolea these are keeled, so one could specu-
happen. If they were sepaloid or nectaries similar late that their fusion might lead to a bicarinate
to those in eudicots, then mutations in petal- structure.
identity genes would not affect them; since they If the palea were derived from fused outer
are affected, sepaloid or nectary origin is not tepals, it would constitute a novel structure in
supported. In summary, the simplest interpreta- the grasses; the origin of the palea maps to the
tion of the data is that the lodicules are governed “spikelet clade”, including all grasses except
42 Flower Structure
Anomochlooideae. This hypothesis is consistent which is thus either a floral bract or a bract-like
with recent studies of gene expression (Preston abaxial sepal.
et al. 2009; Reinheimer and Kellogg 2009) and Much of grass diversity is reflected in mor-
developmental genetics (Li et al. 2011). Palea phological variation in lemmas, and to a lesser
identity is controlled by AGL6-like proteins extent, paleas. Lemmas vary as much in size and
(e.g., MOSAIC FLORAL ORGANS1/OsMADS6 in texture as the glumes (see above). They differ in
rice), which suppress DL; when both are mutated, the number of veins, in their overall shape, in the
the palea is replaced by two lemma-like organs shape of the apex, and the nature of their attach-
(Li et al. 2011). AGL6-like expression in the palea ment to the rachilla (the callus). Indumentum is
appears only in grasses with spikelets but not in common as is ornamentation of various sorts.
Streptochaeta, in which the outer bracts are not The morphology of the cells in the lemma epider-
fused. Palea identity is also controlled by mis is often diagnostic for species (although it is
RETARDED PALEA1 (REP1), a CYCLOIDEA-like an inconvenient field character).
protein related to proteins that in dicots specify Awns may form on lemmas or glumes or
zygomorphy of the perianth (Yuan et al. 2009). In both, and may be terminal, or originate from
rice, the palea margin is controlled by a set of the sinus between two apical lobes, or may appear
proteins (e.g., the MADS-box proteins CHIMERIC attached to the abaxial side; rarely awns form on
FLORAL ORGANS/OsMADS32 and MOSAIC FLO- paleas. Most awned grasses have a single awn on
RAL ORGANS/OsMADS6) different from those the lemma, but lemmas with multiple awns have
controlling the central part of the palea (REP1, evolved several times. The awns may be straight,
MULTI-FLORET SPIKELET1, DEPRESSED or twisted, or geniculate, or both twisted and
PALEA1), which has led some authors to suggest geniculate. Rather surprisingly, awns are major
that the marginal region of the palea is sepaloid, contributors to photosynthesis in the inflores-
whereas the central region is prophyllar (Ren et al. cence and supply an appreciable amount of pho-
2013; Sang et al. 2012); there are no data extending tosynthate to the developing caryopsis, at least in
this particular model to other grasses. In summary, wheat and barley where their role has been stud-
accumulating evidence is pointing to the somewhat ied most extensively (Tambussi et al. 2007).
heterodox idea that the palea corresponds to two Because many wheat cultivars have awned and
fused adaxial tepals. unawned forms, it is possible to make compar-
Subtending the entire flower is a bract-like isons in a near isogenic background. Awns are
structure, the lemma (Fig. 5A, B). The lemma is reported to improve grain yield when water is
similar to a modified leaf. In studies of floral limiting, but appear to have little effect when
mutants and gene expression, the lemma shares water is plentiful (Grundbacher 1963; Motzo and
some aspects of floral (i.e., presumed perianth) Giunta 2002; Tambussi et al. 2007). They also
identity and some of bract identity. It is possible accumulate silica (Grundbacher 1963). Further
that it represents a novel organ type, resulting discussion of the role of awns is under Fruit and
from the fusion of two developmental programs. seed – Dispersal.
In summary, the data suggest a much simpli-
fied interpretation of the grass floret, which can
be interpreted as a highly modified flower, a term References
that will be used throughout this book. The grass
flower is zygomorphic, usually with two or three Ambrose, B.A., Lerner, D.R., Ciceri, P., Padilla, C.M.,
carpels and a single seed, and 3 or 6 stamens. The Yanofsky, M.F., Schmidt, R.J. 2000. Molecular and
inner perianth (“corolla” ¼ lodicules) has two or genetic analyses of the silky1 gene reveal conserva-
tion in floral organ specification between eudicots
three members; when only two are present, they and monocots. Mol. Cell 5: 569–579.
are the two abaxial ones. The corolla serves a Anton, A.M., Astegiano, M.E. 1973. Notas sobre la morfo-
mechanical rather than attractive function. The logia floral de Gramineas. Kurtziana 7: 49–53.
outer perianth (“calyx” ¼ palea) is comprised of Arber, A. 1929. Studies in the Gramineae. VI. 1. Strepto-
chaeta. 2. Anomochloa. 3. Ichnanthus. Ann. Bot. 43:
the two lateral adaxial tepals, which are fused. 35–53.
The abaxial outer tepal is either lost or forms a Arber, A. 1934. The Gramineae: a study of cereal, bamboo,
common primordium with the bract (lemma), and grass. New York: Macmillan.
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Caporali, E., Zhang, D.B., Kater, M.M. 2011. Func- protein, determines spikelet meristem fate and ster-
tional analysis of all AGAMOUS subfamily members ile lemma identity in rice. Plant Phys. 162: 872–884.
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in rice. Development 130: 705–718. DROOPING LEAF regulates carpel specification and
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expression is ancient and palea expression is new. Yuan, Z., Gao, S., Xue, D.W., Luo, D., Li, L.T., Ding, S.Y.,
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Embryology
OVULE et al. (1994). The distinction between hemiana-

tropous and hemicampylotropous is slight, par-
The gynoecium initially forms as a bulge at the ticularly in the absence of a clear funiculus.
top of the floral meristem, which enlarges into a Whatever typology is used to describe the ovule,
dome-like structure. On the abaxial side of the the micropyle always ends up pointing toward the
dome (i.e., the side toward the lemma) a circum- base of the ovary, and the ovule is attached to the
ferential rim forms and grows upward, and soon ovary wall for a substantial portion (often more
encircles the entire gynoecium; this leaves the than ½) of its length.
ovule primordium looking like a ball in a cup, The grass ovule has two integuments. In most
but displaced toward the adaxial side (Le Roux grasses, only the inner integument forms the
and Kellogg 1999; Percival 1921; Zaitchik et al. micropyle (Anton de Triquell 1987; Bhanwra et al.
2000). In other words, the ovule is lateral from 1991; Kam and Maze 1974; Sajo et al. 2008; Ver-
early stages of development. Although not illu- boom et al. 1994) and the ovule is described as
strated, development of the ovule in Sorghum is endostomic; this developmental pattern has been
described in a similar way (Artschwager and suggested to be synapomorphic for Poaceae (Lin-
McGuire 1949). In the early diverging grasses der and Kellogg 1995). In Pharus (Sajo et al. 2007)
Streptochaeta, Anomochloa and Pharus, the and in Panicum miliaceum (Davis 1966), both inte-
ovule is said to initiate basally, but is then dis- guments are reported to form the micropyle, sug-
placed to a lateral position by differential growth gesting a reversal in the character; however, the
of the ovary wall (Sajo et al. 2007, 2008, 2012); figure in Sajo et al. (2007) shows a normal endos-
another interpretation of their photos is that ini- tomic ovule. The inner integument may be more
tiation is lateral. Published descriptions of the than two cells thick near the micropyle (Verboom
grass ovule imply that there is great variation in et al. 1994). There are some reports of taxa in which
its morphology, variously describing it as anatro- the inner integument does not fully enclose the
pous, campylotropous, or amphitropous and nucellus (i.e., a “naked nucellus”), so that a normal
everything in between. Anton de Triquell (1987) narrow micropyle fails to form (Bhanwra et al.
reviews this literature and notes that in fact grass 1991; Choda et al. 1982); the effect of this on fertili-
ovule morphology is quite uniform, and that the zation and ovule development is unknown.
“variation” reflects simple misapplication of In most grasses, the outer integument devel-
ovule typology. The funiculus is virtually absent ops more slowly than the inner one (Bhanwra
and the outer integument often fairly short until et al. 1991; Davis 1966; Maze and Bohm 1973;
late development, so the ovule type is not always Verboom et al. 1994), and often is much shorter
clear; Anton de Triquell (1987) argues that the than the inner integument at anthesis. However,
ovule is best described as hemicampylotropous. both integuments develop simultaneously in
At maturity the ovule is described by Sajo et al. Streptochaeta (Sajo et al. 2008). The outer integu-
(2007, 2008) as hemianatropous, a term also ment generally degenerates after fertilization
applied to a set of arundinoid taxa by Verboom (Bhanwra et al. 1991; Sajo et al. 2007, 2008).
46 Embryology
MEGAGAMETOPHYTE although only 16 are shown in the figure. Prolifera-

tion of the antipodals is not unique to Poaceae. It
The megaspore mother cell differentiates from a also occurs in the Restionaceae (Campbell and
cell immediately inside the nucellar epidermis and Kellogg 1987) but nothing is known about other
no parietal cell forms; occasionally, however, two Poalean families, so it is unclear if it is indepen-
megaspore mother cells are observed (Artschwager dently derived in Restionaceae and Poaceae, or if it
and McGuire 1949). The lack of a parietal cell means is synapomorphic for the entire clade. Proliferation
that ovules are tenuinucellate, but in some taxa of antipodals does not occur in the early diverging
ovules become pseudo-crassinucellate by periclinal grass genera Streptochaeta and Pharus (Sajo et al.
division of the epidermis (Artschwager and 2007, 2008), suggesting that it could be a synapo-
McGuire 1949; Aulbach-Smith and Herr, Jr. 1984; morphy for the BEP plus PACMAD clades. How-
Bhanwra 1988; Bhanwra et al. 1991; Verboom et al. ever, data on this character are too scattered to
1994). The phylogenetic distribution of pseudocras- draw any conclusions about its consistency within
sinucellate ovules is unclear. Bhanwra (1988) sug- major groups.
gests that the condition is common in Panicoideae, The overall shape of the embryo sac varies
but it is also reported in Bambusoideae (Bhanwra among species. In the panicoids, the axis from the
et al. 2001) and Pooideae (Maze and Bohm 1973). egg cell through the antipodals is straight and in
Meiosis I leads to a pair of cells, one of which is line with the proximo-distal axis of the ovule,
closer to the micropyle and the other to the chalazal whereas in the other subfamilies the egg-antipodal
end of the ovule; the latter tends to be larger. At axis is curved so that the antipodal cells appear
Meiosis II, the chalazal cell divides in a plane cross- more or less lateral at maturity (Anton de Triquell
wise to the long axis of the developing megagame- 1987; Bhanwra 1988). As with the preceding char-
tophyte, whereas the micropylar cell may divide acters, the number of taxa sampled is small so it is
either crosswise or longitudinally. These divisions not clear how widely these generalizations apply.
result in a linear (Bhanwra et al. 1991; Wu et al. In subfamily Danthonioideae, the synergids
2011) or T-shaped tetrad of megaspores (Sajo et al. appear to function as haustoria, growing out
2007; Stover 1937), respectively. Variation in tetrad through the micropyle (Philipson and Connor
pattern is reported within a single species (Aul- 1984; Verboom et al. 1994); this character has
bach-Smith and Herr, Jr. 1984), suggesting that been interpreted as synapomorphic for the sub-
the pattern is not of phylogenetic significance and family. Haustorial synergids have been observed
may in fact be random. In some cases, the second in all studied genera of the subfamily (nine of 17
cell division fails entirely in the micropylar member genera, 39 of 281 species), although the character
of the dyad, leading to a triad of cells instead of a appears to be polymorphic in Pentameris.
tetrad (see references in Anton de Triquell 1987, Apomixis occurs in many grasses and is dis-
Bhanwra 1988, Choda et al. 1982); it is not clear cussed under Reproductive systems.
whether this is characteristic of particular species,
or is simply a common but sporadic occurrence.
The megagametophyte is monosporic and EMBRYO
develops from the chalazal megaspore (Anton de
Triquell 1987; Evans and Grossniklaus 2009; Stover The first division of the zygote is transverse; the
1937; Wu et al. 2011). In most species, the megaga- basal cell then divides again transversely, then the
metophyte is of the Polygonum-type, with three apical cell divides vertically or obliquely, and
mitoses followed by cellularization leading to finally the basal cell divides again, giving rise to
seven cells; the central cell contains two nuclei. In the suspensor (Artschwager and McGuire 1949;
most grasses, the antipodal cells continue to divide Jones and Rost 1989). Early embryo development
after cellularization, although the number of extra in the grasses is usually described to be of the
divisions varies (Anton and Cocucci 1984; Evans “asterad” type (Bhanwra 1988; Febulaus and Pull-
and Grossniklaus 2009; Shadowsky 1926; Stover aiah 1994; Sajo et al. 2007), meaning that both the
1937; Wu et al. 2011); Yamaura (1933) reports as basal and the apical cell of the two-celled embryo
many as 300 antipodals in Sasa paniculata, contribute to formation of the embryo proper. In
EMBRYO 47
contrast, both Nardmann and Werr (2009) and no homology to organs in other monocots (“cae-
Jones and Rost (1989) suggest that only the apical nogenetic”), and that the epiblast is an outgrowth
cell contributes to the embryo proper in maize of the coleorhiza. His view of the coleoptile ech-
and rice, respectively, and Taylor and Vasil oes that of Boyd (1931). Weatherwax (1920), in
(1995) report no particular pattern of cell division contrast, suggested that the coleoptile is a leaf.
in Cenchus americanus (¼ Pennisetum glaucum, Recent evidence suggests that the coleoptile
pearl millet). plus scutellum together constitute the cotyledon
The embryo proper enlarges, first becoming (Takacs et al. 2012). The scutellum functions like
club shaped, and then elongating. As the embryo cotyledons in many plants to take up nutrients
elongates, it begins to differentiate into a bipolar from the endosperm during germination.
embryo with shoot and root apical meristems Because the coleoptile develops on the same side
(Takacs et al. 2012). On one side of the embryonic of the shoot apical meristem as the scutellum, it
axis, a broad shield-shaped and peltate structure appears to be the sheathing portion of the cotyle-
forms, the scutellum, which is generally longer donary structure, whereas the scutellum may be
and broader than the embryo itself. As the apex the distal tip (Takacs et al. 2012). The first leaf
of the scutellum begins to elongate, the shoot forms on the opposite side of the embryo, thus
apical meristem is seen initially as a densely establishing the distichous phyllotaxy that will
packed group of cells on one side of the embryo, characterize the plant. This interpretation is sup-
above which a notch forms. Above the notch, a ported by global analyses of gene expression,
bulge of cells forms, and these ultimately enlarge which show high similarity between the coleoptile
to form a ring or collar around the developing and scutelllum, with few genes expressed
meristem (Artschwager and McGuire 1949; Nard- uniquely in the coleoptile (Takacs et al. 2012).
mann and Werr 2009; Taylor and Vasil 1995); this As the first leaf begins to form, the scutel-
ring then elongates into a tube like structure, the lum + coleoptile (i.e., cotyledon) begins to
coleoptile. The primary root meristem is acquire some leaf-like characteristics. A distinct
organized endogenously. Opposite the scutellum set of genes is expressed in the scutellum + cole-
and just above the root meristem, an epidermal optile structure plus the first leaf (Takacs et al.
outgrowth may form; this is the epiblast. In some 2012). The tubular coleoptile apparently has a
grasses, the scutellum elongates basipetally protective function for the developing seedling,
alongside, but separate from the coleorhiza; in but also provides photosynthate (Jackson 2009).
the pooid grasses, however, this elongation Nardmann et al. (2004) show that in maize the
scarcely occurs (Reeder 1957). In the former narrowsheath genes are expressed in the margins
case, when the embryo is viewed in longitudinal of the coleoptile as it begins to encircle the
section there is a clear space between the scutel- nascent meristem. These genes are expressed in
lum and the base of the embryo known as the the margins of all leaf-like organs in the plant.
scutellar cleft. The functional significance of the During embryonic development, a vascular
different scutellar structures, if any, is unknown. trace differentiates to the scutellum and one to
The homologies of the scutellum and the the coleoptile. In some species, these two traces
coleoptile have been the subject of much discus- are separated by a clear internode, known as the
sion, backed, until recently, by remarkably little mesocotyl internode. Brown (1960) felt that the
data. As summarized by Brown (1960), the scu- mesocotyl had no homology to organs in other
tellum has been variously considered a cotyledon, monocots, but that it might be similar to the
a part of a cotyledon, an outgrowth of the radicle, hypocotyl in dicots; this makes little sense if the
the embryonic axis, a leaf sheath, or an aborted scutellum is the cotyledon, since a hypocotyl
bud. The coleoptile, while clearly a sheathing would have to be below the cotyledon, whereas
structure of some sort, has also been considered the mesocotyl is clearly above the scutellum.
a cotyledon or part of it, or two or more fused After formation of the coleoptile, the apical
leaves, or fused cotyledons, or a prophyll, or a meristem produces one or more true leaves
bud. Brown (1960), citing developmental evi- before seed maturation. The epiblast, which is
dence, concludes that the scutellum is a modified so prominent particularly in pooid grasses,
cotyledon, that the coleoptile is a structure with might correspond to a leaf, but no data exist to
48 Embryology
embryo both root and shoot apical meristems

are fully organized before seed maturation; vas-
cular tissue is visible. In evolutionary time, a
heterochronic shift must have occurred such
that embryo development was accelerated relative
to seed maturation, or seed maturation was
delayed relative to embryo development (Kellogg
2000).
Embryo characters have been used to help
determine relationships among grasses and were
among the most consistent in an early cladistic
study (Kellogg and Campbell 1987). In particular,
the scutellar cleft may be present or absent, an
epiblast may be present or absent, the mesocoty-
lar internode may be short or long, and the
embryonic leaf margins may meet or overlap
(Reeder 1957) (Fig. 7). However, the latter char-
acter in particular varies within many higher taxa
and may be less diagnostic than previously
believed. A long mesocotyl internode appears to
be the sole morphological synapomorphy for the
large PACMAD clade (GPWG 2001).
Fig. 6. Bambusa multiplex. Seedling in two stages of

development: left fruit with radicle and plumule just ENDOSPERM
beginning rapid elongation; right fruit with attached
young plant showing branching root, and culm shoot Endosperm formation has been studied exten-
with first foliage leaf. (From McClure 1966)
sively because of its agronomic importance, but
the focus has been on only a handful of taxa. Most
test this hypothesis. In Oryza sativa and Poa data come from maize, with additional observa-
pratensis, the first leaf formed after the coleoptile tions from wheat, barley, rice, sorghum, and
lacks a blade, and only one more embryonic leaf more recently Brachypodium. (An additional
is formed (Sylvester et al. 2001). This presents an problem is that endosperm development in
interesting comparison with members of the wheat is generally presented in transverse sec-
Bambusoideae, in which the first seedling leaf tion, whereas that in maize is presented in longi-
lacks a blade, and the first few internodes of the tudinal section; this makes drawings and
seedling often fail to elongate (Soderstrom 1981) photomicrographs difficult to compare.) Details
(Fig. 6). It is possible that the first leaf after the of endosperm development differ, but it is
coleoptile is bladeless in all members of the BEP unclear whether the differences are specific for
clade; in contrast, in Zea mays (PACMAD clade) entire taxonomic groups or whether they are
the blade is present but reduced. Oryza and Poa unique to the few species studied. Given the
develop only one more embryonic leaf, whereas high fitness consequences of provisioning a
Zea mays produces four more leaves in the seed, selection might have shaped endosperm
embryo. development in different ways in different
The embryo is lateral within the developing lineages. In describing endosperm below, I hope
seed, but the mature embryo structure with its to encourage more comparative studies that
highly differentiated tissues and organs is a syn- might illuminate the consistency of these charac-
apomorphy for the family. In most other mono- ters across many taxonomic groups.
cots, including members of Poales, the mature In all grasses that have been studied, the
embryo is a small cluster of cells, lacking any central cell of the megagametophyte contains
obvious structures. In contrast, in the grass two nuclei, which are unfused until fertilization
ENDOSPERM 49
Fig. 7. Embryos. A Longitudinal section of a pooid of a panicoid embryo apex showing overlapping embry-
embryo showing presence of an epiblast (ep) and coleop- onic leaf margins. D Cross section of a pooid embryo apex
tile (c). B Longitudinal section of a panicoid embryo showing embryonic leaf margins that meet. (From GPWG
showing presence of a scutellum (s), a scutellar tail (st), 2001, p. 402, with permission from Missouri Botanical
and elongated mesocotyl internode (mi). C Cross section Garden Press; drawn by M. Kojima)
(Jones and Rost 1989). One of the two sperm Among other characteristics, the nucellar epider-
fertilizes the central cell to form a triploid mis in wheat breaks down while the aleurone
nucleus, which then begins to divide within differentiates (Percival 1921), whereas the nucel-
hours, undergoing several rounds of division lar epidermis is persistent in Brachypodium, as it
before the zygote divides at all (Artschwager and is in rice (Ellis and Chaffey 1987).
McGuire 1949; Brown and Lemmon 2007; Perci- At the same time that the endosperm is devel-
val 1921). The nuclei divide initially without cyto- oping, the nucellus breaks down. Soon after pol-
kinesis; the preprophase band does not form, nor lination, the cells of the nucellus undergo
do cell walls (Brown and Lemmon 2007; Opano- programmed cell death; the space that the cells
wicz et al. 2011; Sabelli and Larkins 2009a; Leroux had occupied is filled by the developing endo-
et al. 2014). The endosperm nuclei migrate to the sperm (Nguyen et al. 2007).
chalazal end of the embryo sac, and a large central As cellularization proceeds, the endosperm
vacuole develops so that the nuclei are pushed differentiates into four distinct tissues, the aleu-
toward the periphery of the endosperm cell (Opa- rone, the starchy endosperm, transfer cells, and
nowicz et al. 2011; Weatherwax 1930; Leroux et al. the layer surrounding the embryo, called the
2014). After nuclear proliferation, microtubules embryo-surrounding region or ESR (Sabelli and
form, radiating from the surface of each nucleus, Larkins 2009a; Leroux et al. 2014). The latter is
and cell walls are laid down between the nuclei in evanescent, becoming recognizable as soon as
the peripheral layer. Anticlinal walls are then cellularization occurs, but then breaking down
formed between the next layer of nuclei to the (Cossegal et al. 2007; Opsahl-Ferstad et al. 1997;
inside, creating a sort of three-sided box or alve- Weatherwax 1930). The cells in the ESR appear to
olus surrounding each nucleus. The nucleus then be metabolically active and may be involved in
divides and undergoes cytokinesis toward the transfer of sucrose to the developing embryo,
center of the cell, thus moving the layer of par- although there are no symplastic connections
tially enclosed nuclei inward by one. By about 3 to between the two. In maize the ESR cells express
6 days after pollination in the cereals, the entire a specific set of genes (the ESR genes) that may be
central cavity is fully cellularized. Some aspects of involved in signaling and/or pathogen response
developmental timing differ between Brachypo- (Opsahl-Ferstad et al. 1997). Unfortunately for
dium and wheat and may be worth exploring in comparative biologists, the ESR has been
other pooid grasses (Opanowicz et al. 2011). described in detail only in maize. Cytologically
50 Embryology
similar sets of cells have been reported in wheat bohydrates and amino acids from the maternal
(Smart and O’Brien 1983), and were apparently tissue (Sabelli and Larkins 2009a). Although Rost
observed by Percival (1921), and barley (Brown et al. (1990) did not find transfer cells in the fruit
et al. 1994), but not characterized in detail. How- of Briza maxima, it is possible that the stage of
ever, proteins that are specific to the ESR in maize development was too late to observe them.
are apparently not present in rice, which raises It is no exaggeration to say that cereal endo-
questions about the generality of this cell type. sperm feeds the world, and that the cells of the
The outer layer of endosperm cells (in the starchy endosperm are the center of it. Soon after
grasses as well as other angiosperms) is known cellularization is complete, the cells of the starchy
as the aleurone layer. These cells are morphologi- endosperm undergo remarkable alterations of the
cally different from other endosperm cells and are normal cell cycle, in which the chromosome com-
generally cuboidal. The thickness of the aleurone plement of each nucleus undergoes endoredupli-
varies among taxa, although it has not been sur- cation, creating cells with as much as 96 times the
veyed in any systematic fashion; maize and wheat haploid amount of DNA (Kowles and Phillips
have aleurone that is only one cell layer thick 1985; Nguyen et al. 2007; Sabelli and Larkins
whereas there are three layers in barley; the thick- 2009a, 2009b). The ploidy levels of the starchy
ness in rice and Brachypodium is variable (Ellis endosperm vary between crops, with lower ploidy
and Chaffey 1987; Opanowicz et al. 2011; Sabelli in wheat than in other species that have been
and Larkins 2009a). The aleurone responds to studied (Nguyen et al. 2007). Endoreduplication
gibberellin secreted by the developing embryo, begins in the center of the endosperm and extends
by synthesizing enzymes that mobilize starch in gradually to the periphery, a process that in maize
the endosperm. Humans have taken advantage of takes about a week. Endoreduplication generally
this process to create malted barley, the starting does not extend to the aleurone, which remains
point for beer and whiskey. diploid, except in barley (Keown et al. 1977). Dur-
At germination the aleurone secretes ing this process of endoreduplication, the cells
enzymes that break down the macromolecules begin to accumulate starch and proteins.
in the starchy endosperm. As germination pro- Starch granules accumulate in amyloplasts,
ceeds however, the aleurone undergoes pro- under the control of starch synthases, starch
grammed cell death (Nguyen et al. 2007). branching enzymes and starch de-branching
The vascular strand on the adaxial side of the enzymes. A single amyloplast may produce one
ovary projects into the nucellus in some taxa and or several starch granules; if several are produced
is surrounded by a set of cells known as the the granules will coalesce at maturity and appear
nucellar projection. The aleurone surrounding compound, whereas a single large granule will
the nucellar projection is sometimes modified appear simple. Species with simple starch gran-
cytologically. In Triticeae in particular, the endo- ules are found in Panicoideae s.s. and some of the
sperm expands on either side of the vascular early-diverging panicoid tribes (Sánchez-Ken and
strand and nucellar projection so that the latter Clark 2010; Tateoka 1962), whereas those of other
structures end up at the bottom of a deep groove subfamilies tend to be compound. Starch gran-
known as the crease. In contrast to the Triticeae, ules are bimodal in size in Triticeae (Tomlinson
the vascular strand of the ovary is quite short et al. 2003), whereas they are uniform and small
in Cenchrus americanus (¼Pennisetum ameri- in Brachypodium (Opanowicz et al. 2011).
canum, ¼P. glaucum) (Fussell and Dwarte Starch biosynthesis involves a substantial
1980), and scarcely extends past the hilum. number of grass-specific proteins, probably con-
Transfer cells differentiate in the region of the tributing to the high starch content in the endo-
endosperm adjacent to the placenta (Artschwager sperm in the family. Genes encoding major
and McGuire 1949; Charlton et al. 1995; Fussell enzymes of the starch biosynthetic pathway
and Dwarte 1980; Jones and Rost 1989; Rost et al. were all duplicated as part of the whole genome
1984; Weatherwax 1930). They have highly inva- duplication that occurred coincident with the ori-
ginated cell walls and correspondingly extensive gin of the family (Comparot-Moss and Denyer
plasma membranes (Schiel et al. 1984); this 2009; Li et al. 2012). Following duplication, the
increased surface area promotes transfer of car- genes diverged in function, and exhibited an
References 51
accelerated rate of evolution with positive selec- leria, Sphenopholis and Trisetum [the
tion observed on a number of amino acids in the endosperm] remains liquid for years.” The occur-
proteins. For example, the rate-limiting enzyme rence of liquid endosperm is sporadic and does
in starch biosynthesis is ADP-glucose pyropho- not seem to follow phylogenetic lines. However,
sphorylase. This enzyme has distinct forms that this study relied on a subjective assessment of
occur in the cytosol and plastids of plants; endosperm condition; it would be interesting to
whereas the plastid form is common in most assess the protein and lipid composition of the
starch storage organs of most plants, the cytosolic endosperm in a more rigorous way now that
form is apparently unique to grasses (Comparot- appropriate analytical tools are available for
Moss and Denyer 2009) and is expressed predom- doing this. Liquid endosperm is also reported in
inantly in the endosperm (Tetlow et al. 2004). A the bamboo Melocanna (Ramanayake and Weer-
protein kinase known as SnRK1b also appears to awardene 2003), but its chemical composition
be restricted to cereals, and is important in the was not investigated.
process of sucrose transport and storage (Jain
et al. 2008). However, other Poales have not
been studied. References
In addition to starch, the endosperm includes
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Karyology and Genome Structure
CHROMOSOME NUMBER possible, it will only be provided by genomic data

from Anomochlooideae, Pharoideae, and the
The grasses, and in particular the cereals, have grass sister taxa.
long been investigated as models for studies of Base chromosome number is fairly stable in
chromosome number, morphology, dynamics, many clades of grasses and for some is the shared
and evolution (e.g., Avdulov 1931; Stebbins character that provides evidence for monophyly.
1956). The number of chromosomes varies from The core Pooideae (Poeae + Triticeae + Bro-
2n ¼ 4 in species of Zingeria and Colpodium meae) are marked by having their genes arranged
(Kim et al. 2009) to 2n ¼ ca. 266 in Poa litorosa in seven large chromosomes (GPWG 2001; Kel-
(Hair and Beuzenberg 1961). Based on a parsi- logg and Campbell 1987). The Panicoideae are
mony optimization, the chromosome base num- divided into three major clades, corresponding
ber of x ¼ 12 is synapomorphic for the spikelet to chromosome base numbers of x ¼ 10 (Andro-
clade, i.e., all grasses except Anomochlooideae pogoneae and Paspaleae) and x ¼ 9 (Paniceae s.
(GPWG 2001). s.) (Giussani et al. 2001). A base number of
The chromosome number for the common x ¼ 11 is synapomorphic for Olyreae (Bambusoi-
ancestor of the family is uncertain. In the out- deae) and is independently derived and synapo-
groups, n ¼ 18 for Joinvillea plicata (Newell morphic for Aristidoideae (GPWG 2001). Despite
1969) and 2n ¼ ca. 38 (i.e., n ¼ 19) for Ecdeioco- this overall pattern of stability, base chromosome
lea monostachya (Hanson et al. 2005). The hap- number does vary within some genera; examples
loid number in Anomochloa is n ¼ 18 include Phalaris (x ¼ 6 or 7; Baldini 1995), Bra-
(Judziewicz and Soderstrom 1989); this number chypodium (x ¼ 5, 7, 8, 9; Catalán et al. 2012),
combined with those of the immediate outgroups and Cenchrus (Pennisetum) (x ¼ 5, 7, 8, 10, 17;
would suggest that the common ancestor also had Chemisquy et al. 2010, among many others).
n ¼ 18. However, Streptochaeta is n ¼ 11 (Jud- B chromosomes have been reported for over
ziewicz and Soderstrom 1989). Using methods of 200 species of grasses (Hunziker and Stebbins
comparative genomics, estimates of the ancestral 1987). These are supernumerary chromosomes,
chromosome base number are x ¼ 10 (Salse et al. derived from the normal (“A”) chromosome
2008), or x ¼ 7 (Devos 2010); these were postu- complement; they tend to accumulate in the
lated to have undergone allotetraploidization fol- male gametes by non-disjunction at meiosis
lowed by rearrangements, leading ultimately to (Jones and Rees 1982). B chromosomes have
an ancestor with x ¼ 12. This estimate converges been studied in particular detail in maize (Birch-
on that suggested by several authors (Avdulov ler and Han 2009; Jones et al. 2008) and Secale
1931; GPWG 2001; Raven 1975). Stebbins (1985) (Jones and Rees 1967, 1982), among others. Many
suggested a base number of 5, 6 or 7, Mehra et al. aneuploid chromosome numbers reported in the
(1968) and Sharma (1979) argued for x ¼ 6, and literature may be caused by mistaking B chromo-
Flovik (1938) suggested x ¼ 5. If a resolution is somes for A chromosomes.
56 Karyology and Genome Structure
Changes in chromosome number reflect rear- (Spangler et al. 1999; Wilson et al. 1999). One
rangements of existing chromosomes that reduce popular idea is that the ancestor of the tribe had
or increase the number of centromeres. Centro- a 2n number of 10, and thus a haploid number of
meres are generally stable within a species, but five (Celarier 1956; Garber 1950). This idea is sup-
neocentromeres form relatively easily, and con- ported by the handful of species in Andropogo-
versely former centromeres can be lost when chro- neae that exhibit n ¼ 5 (Coix aquatica, and
mosomes combine. The mechanisms by which this members of Sorghum subg. Parasorghum
happens are only beginning to be understood (¼Sarga)); however, these taxa are clearly derived
(Birchler and Han 2009). At the DNA level, a in the phylogeny (Spangler et al. 1999), pointing to
centromere consists of a short repetitive sequence, a secondary reduction in chromosome number.
interspersed with retrotransposons and occasional Garber (1950) and Price et al. (2005) found that
transcribed genes (Fan et al. 2011). The repetitive chromosomes of the x ¼ 5 species of Sorghum
sequence is generally species-specific and may were considerably larger than those of the x ¼ 10
affect the viability of hybrids, but it has not been taxa; the x ¼ 5 sorghums also have a higher 2C
explored in a systematic context. DNA content than the x ¼ 10 species (Price et al.
One apparently common mechanism of 2005). These observations raise the possibility that
reduction in chromosome number has been docu- x ¼ 5 represents an intriguing chromosomal
mented in Aegilops tauschii (Triticeae, Pooideae), fusion event, rather than preservation of the
in which genome architecture has been compared ancestral base chromosome number. One can
to that of rice (Luo et al. 2009). Aegilops chromo- imagine that a burst in retrotransposon activity
some 1 is collinear with rice chromosomes 5 and could have led simultaneously to the abrupt
10. The genes found on rice 10 appear in the increase in genome size seen in this group, as
middle of the Aegilops chromosome, whereas the well as the genome rearrangements that led to
genes of one arm of rice 5 appear to have been five large rather than 10 small chromosomes.
attached at one end of rice chromosome 10, with
the other arm of rice 5 attached at the other end of
rice chromosome 10. It thus appears that an entire POLYPLOIDY
chromosome (rice 10) has been inserted into the
centromeric region of another (rice 5). The gene Allopolyploidy is common in Poaceae, leading
content and order of the inserted chromosome is Hunziker and Stebbins (1987) to claim that “Gra-
maintained, and the two individual arms of the mineae is the only large family in which high
chromosome into which it is inserted become the frequency of polyploids prevails throughout the
distal ends of the new chromosome. The centro- family.” One of the earliest allopolyploids to be
mere of the inserted chromosome remains func- studied in detail was wheat (Kihara 1982). Pasta
tional, whereas the centromere into which it is wheat (Triticum durum) is an allotetraploid
inserted is inactivated. derived from the diploid Triticum urartu and a
Segmental duplications appear to be less species of goatgrass (probably similar to Aegilops
common, but have been documented in members speltoides) (see Domestication). This tetraploid
of the genus Zizania (Oryzeae, Ehrhartoideae), then crossed with another diploid species of goat-
which have 2n ¼ 30 or 34. A genome map of grass (Aegilops tauschii) to produce a sterile
Zizania palustris shows that the base number of hybrid. In experimental hybrids of the two spe-
x ¼ 15 reflects duplication of portions of rice cies, non-reduced gametes are formed at high
chromosomes 1, 4, and 9 (Kennard et al. 2000). frequencies in both the male and female gameto-
These duplicate regions must have somehow phytes, so that self-pollination can lead to fertile
acquired functional centromeres, but the mecha- hexaploids (Matsuoka 2011).
nism involved is unknown. Other similar cases have been demonstrated.
Because of general interest in the ancestry of For example, the polyploid species of Elymus are
cultivated maize, there is considerable literature clearly derived from species of Pseudoroegneria
on chromosomal evolution in its relatives in the and Hordeum, as shown by cytogenetic studies
tribe Andropogoneae. The ancestral base chromo- (Dewey 1984) and molecular sequence data (Kel-
some number (n) for the tribe is most likely 10 logg et al. 1996; Mason-Gamer et al. 2002, 2010a,
GENOME SIZE 57
2010b). The woody bamboos are all polyploid, the smallest diploid genomes reported for Orope-
with most species being tetraploid (Calderón and tium thomaeum (haploid size (1C) ¼ 0.25 pg
Soderstrom 1973; Peng et al. 2013; Stebbins 1971). DNA or ca. 205 Mbp) and the largest for Secale
Switchgrass (Panicum virgatum) is an allotetra- cereale (8.28 pg DNA or ca. 6790 Mbp); polyploid
ploid, and common millet (P. miliaceum) is the genomes are of course commensurately larger.
result of an independent polyploidization event The ancestral size of a 1C genome in the family
(Triplett et al. 2012). The genus Zea is an ancient is estimated to be 1.5–2.6 pg DNA (Caetano-
allotetraploid (Blanc and Wolfe 2004; Schnable Anolles 2005), although this estimate was pub-
et al. 2009; Swigonova et al. 2004). Sugar cane is a lished before the estimation of genome size of
complex polyploid, with an uncertain base chro- the outgroup Ecdeiocolea monostachya at 1 pg
mosome number, although phylogenetic data sug- of DNA (Hanson et al. 2005); this small value
gest that the base number might be 10 (Kellogg might make the ancestral grass estimate slightly
2012). lower. Phylogenetic analyses have identified a
Within every group of Poaceae, it is possible major increase in genome size in the Pooideae,
to identify well-supported clades that are clearly with a particularly dramatic increase in the Triti-
similar morphologically but within which hybri- ceae (Bennetzen and Kellogg 1997; Kellogg and
dization and polyploidy are rampant (“. . . big Bennetzen 2004). The protein-coding genes in the
messy geographically conditioned splodges of grass genomes are largely in the same order, and
divergence as yet unsanitized by extinction variation in genome size thus reflects variation in
. . .”; S. Mayo, pers. comm., referring to Anthur- the space between genes. The “space” is occupied
ium, but could be any of several groups of by retrotransposons, which are small pieces of
grasses). For example, in the tribe Paniceae, the DNA that copy themselves into RNA and then
Cenchrus clade and most likely the Setaria clade are copied back to DNA and reinserted (Devos
are easily distinguished, but within them rela- 2010; SanMiguel et al. 1996). The retrotranspo-
tionships are complex, polyploids are common, sons preferentially reinsert into themselves so
and morphology is highly homoplasious. Other they form a nested arrangement; the ends of ret-
such groups include the Triticeae in subfamily rotransposons are identical (Long Terminal
Pooideae, and the genus Oryza in subfamily Repeats, or LTRs), and can be identified even
Ehrhartoideae. Within the tribe Poeae, it is likely after multiple copies of the transposon are
that subtribes Airinae, Poinae, and Loliinae also inserted between them.
represent such lineages within which reticulate Chromosome numbers do not correlate well
evolution is common, but between which hybri- with genome size. For example, rice has 12 pairs
dization is rare. of chromosomes and the haploid DNA content is
Polyploidy does not correlate with domesti- about 0.5 picograms (pg) of DNA. In contrast,
cation in angiosperms (Hilu 1993), nor is there an barley has only seven pairs of chromosomes but
obvious correlation in the grasses. Of the “Big a haploid DNA content of about 5.55 pg (Kellogg
Three” – maize, wheat and rice – the first two and Bennetzen 2004). Likewise, Sorghum biolor
are polyploid, but rice is diploid. Common millet has 2n ¼ 20 and about 0.83 pg of DNA, but the
and finger millet are tetraploids but pearl millet closely related Sarga angusta has 2n ¼ 10 and
and foxtail millet are diploid. Oats are hexaploid more than twice as much DNA per 1C nucleus
but barley and rye are diploid. Sugar cane is (1.85 pg). A corollary is that two plants could
polyploid but sorghum is diploid. It thus seems have the same total amount of DNA in their
unlikely that polyploidy per se confers any par- genomes, but it might be divided into quite dif-
ticular advantage for domestication. ferent numbers of chromosomes.
Although there has been a suggestion in the
literature that genome size may correlate with
GENOME SIZE effective population size (Lynch and Conery
2003), this correlation has not been found in
The sizes of grass genomes vary considerably Oryza, the only grass genus investigated to date
(Caetano-Anolles 2005; Leitch et al. 2010), with (Ai et al. 2012).
GENOME MAPPING, SEQUENCING AND CONSERVATION OF dium than it is to rice or sorghum (Mayer et al.
GENE ORDER 2011). Likewise foxtail millet and sorghum each
have several species-specific inversions, whereas
The rice genome was the first of the grasses to be the two genomes share a set of panicoid-specific
sequenced in its entirety, and contains ca. 389 inversions and translocations that distinguish
mega base pairs (Mb) of DNA (International them from rice (Bennetzen et al. 2012).
Rice Genome Sequencing Project 2005). Although In addition to translocations and inversions,
compact, it still contains a high proportion smaller scale rearrangements cause periodic vio-
(35 %) of transposable elements, or their remains. lations of the overall collinearity of the genomes.
In addition to that of rice, whole genome Genes are moved from their original locations by
sequences are now available for a bambusoid the action of transposable elements and double
grass, Phyllostachys heterocycla (Peng et al. stranded break repair (Wicker et al. 2011).
2013), four members of Pooideae, Brachypodium Although in most cases these processes simply
distachyon (The International Brachypodium Ini- generate pseudogenes, functional genes do also
tiative 2010), Hordeum vulgare (The Interna- appear from time to time, leading to species-spe-
tional Barley Genome Sequencing Consortium cific or clade-specific gene duplicates. This is an
2012), Triticum urartu (Ling et al. 2013) and obvious source of genetic novelty, which could
Aegilops tauschii (Jia et al. 2013), plus four mem- lead to new phenotypes. Some genes that are
bers of Panicoideae, Setaria italica and S. viridis known to be important for morphological varia-
(Bennetzen et al. 2012), Sorghum bicolor (Pater- tion fall in non-collinear regions of the genome,
son et al. 2009), and Zea mays (Schnable et al. hinting at a link between genomic and morpho-
2009). Although the size and highly repetitive logical change. For example, the barley gene Vrs1,
nature of the wheat genome makes it a particu- which controls development and fertility of the
larly difficult challenge for whole genome lateral spikelets at each node, is in a different
sequencing, details of its structure and sequence genomic position in rice and barley (Pourkheir-
are emerging rapidly (Brenchley et al. 2012). With andish et al. 2006). Barley chromosome 2 is col-
continuing rapid developments in sequencing linear with rice chromosome 4 for all markers
technology, it is likely that many additional surrounding Vrs1, but the Vrs1 orthologue in
grass genomes will be available even by the time rice is on chromosome 7, indicating that the posi-
this book is printed. tion of the gene has changed in evolutionary time.
One of the most stunning results of early As genome sequencing becomes increasingly
genome mapping efforts was the demonstration common, it is likely that structural characteristics
that many of the genes in these genomes are in of genes and genomes will be found to character-
the same order; that is, the genomes are largely ize many clades of grasses. It is also likely that
collinear (Gale and Devos 1998; Moore et al. genes will be identified underlying each of the
1995). For example, a comparison of genome morphological synapomorphies.
regions homologous to chromosome 1H (i.e., 1
Hordeum) in Hordeum, Triticum, Brachypodium,
Oryza, and sorghum showed that about 83 % of GC CONTENT
the genes were in the same order (Mayer et al.
2011). Some grass genomes have a higher frequency of
Despite such gross-level collinearity, how- G-C base pairs than do other plants; in other
ever, the distribution of blocks of genes over words, the relative proportions of A-T and G-C
chromosomes varies and periodic large-scale base pairs are not equal throughout the genome,
genome rearrangements can be identified, many as might be expected if genome composition were
of which correlate with phylogeny (Kellogg 2001). entirely random. The extent of GC bias is most
For example, the three genomes of hexaploid pronounced at the third positions of codons, and
wheat have a number of structural similarities in this third-position GC bias varies among species;
common that are not shared with barley, whereas for example Phyllostachys edulis is about 47 % GC
gene order in barley is more similar to Brachypo- on average, whereas Secale cereale is about 68 %
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Pollen
MORPHOLOGY AND ANTHER STRUCTURE ever, simultaneous microsporogenesis, with cyto-

kinesis occurring only after meiosis II is
Stamen morphology is largely uniform through- complete, has been noted in addition to the suc-
out the Poaceae. Anthers are consistently tetra- cessive type (Sajo et al. 2009, 2012). Mature pollen
sporangiate, although Bhanwra (1988) reports is three-celled, with one vegetative and two sperm
bisporangiate anthers in Bromus unioloides. In nuclei (Nakamura et al. 2010).
all grasses, the anther wall consists of an epider- In most grasses, after microsporogenesis the
mis, an endothecium, and a tapetum. In all microspores form a single layer around the inside
known Poaceae, the tapetum is of the secretory of the anther locule, so that the single pore of each
type, as is found in the close relatives of the microspore/pollen grain is in direct contact with
grasses (Furness and Rudall 1998). Tapetal cells the tapetum (Christensen and Horner, Jr. 1974;
generally have two nuclei, but in some taxa have Kirpes et al. 1996). In contrast, in Pharus lappu-
only one (Anton de Triquell 1987; Nakamura laceus, pollen fills the anther locule and is not
et al. 2010). Grasses in the Bistigmatic clade (i.e., restricted to the peripheral region (Kirpes et al.
all grasses except for Anomochlooideae and 1996); however, Sajo et al. (2007) report periph-
Pharoideae) also have a middle layer of cells eral microsporocytes and centrally located pollen
between the endothecium and the tapetum; the grains. Findings on Streptochaeta and Anomo-
middle layer breaks down early in anther devel- chloa are contradictory in terms of pollen posi-
opment. Presence of the middle layer is synapo- tion. Kirpes et al. (1996) report that Streptochaeta
morphic for the Bistigmatic clade; also pollen is peripheral and Anomochloa is not; in
synapomorphic is the fate of the inner walls of contrast, Sajo et al. (2009, 2012) report exactly the
the endothecial cells, which become fibrous at opposite. The discrepancy may be due to exam-
maturity (Artschwager and McGuire 1949; Bhan- ining different stages of development. It is thus
wra 1988; Furness and Rudall 1998). In Anomo- impossible to determine if peripheral pollen is a
chlooideae and Pharoideae, in contrast, the synapomorphy and if so, at what level in the
anther wall lacks the middle layer entirely and phylogeny. Peripheral pollen is derived indepen-
thus has only three layers of cells (Sajo et al. dently in Cyperaceae (Kirpes et al. 1996).
2007, 2009, 2012). The inner walls of the endothe- Grass pollen is remarkably uniform in mor-
cium fail to become fibrous in these two subfa- phology. Pollen grains are more or less spherical,
milies (Sajo et al. 2007, 2009, 2012). with a single germination pore, and in this
Microsporogenesis in most grasses is succes- respect are similar to those of other monocots.
sive, with cytokinesis taking place after meiosis I; The pore is surrounded by a raised rim, or annu-
this character state is also the most common in lus; within the pore is a circular bit of exine, the
the Poales (Furness et al. 2002) and has recently operculum. On the inside (cytoplasmic side) of
been documented in Ecdeiocolea (Sajo et al. the annulus, the foot layer is thickened and lay-
2009). In Streptochaeta and Anomochloa, how- ered (Christensen and Horner, Jr. 1974; Linder
64 Pollen
and Ferguson 1985; Skvarla et al. 2003). The pore four morphogroups of pollen. While the algo-
forms on the side of the microgametophyte in rithm could accurately assign pollen to a mor-
contact with the tapetum, and this position may phogroup in 77 % of the cases, the groups
be important for proper maturation of the pollen themselves did not correlate with taxonomy or
grain (Christensen and Horner, Jr. 1974). In the phylogeny; for example, groups 1 and 2 each
bambusoid genus Pariana, the thickened foot contained a representative of Pooideae, Chlori-
layer is present, but the annulus and operculum doideae, and Panicoideae. It remains to be seen
are absent (Skvarla et al. 2003). Pollen apertures whether the pollen morphogroups correlate with
similar to those in Poaceae are found in other other aspects of biology such as environment.
graminid Poales, although Flagellariaceae, Join-
villeaceae and Restionaceae may not form an
operculum. The pollen wall of Poaceae is colu-
POLLINATION
mellate, with a clear foot layer and tectum. The
cavities in the pollen wall appear to contain little
Pollen transfer
or no material deposited by the tapetum (Heslop-
Harrison and Heslop-Harrison 1987). Unlike Most grasses are wind-pollinated. The tiny flow-
most other Poales, the grasses lack channels ers, feathery stigmas, versatile anthers, and lack
(scrobiculi) perforating the pollen wall (Linder of nectar are all characteristic of the wind polli-
and Ferguson 1985). The lack of scrobiculi nation syndrome, and anyone who suffers from
appears also to be shared with Ecdeiocoleaceae hay fever is painfully aware of the large amount of
and may indicate a sister relationship between the grass pollen that may be airborne at some times
two families. Within the foot layer and tectum, of year. In addition to the well-documented floral
however, narrow channels are visible in transmis- characteristics, wind-pollinated species often
sion electron micrographs (Linder and Ferguson produce a single seed per flower and tend to
1985); these may be unique for Poaceae. have unisexual flowers (Givnish et al. 2010). The
The outer surface of the pollen grain (tectum) single seed per flower is characteristic of and
is covered with tiny pointed bumps, variously synapomorphic for all grasses, and unisexual
called spinules or granules; these may form clus- flowers occur sporadically throughout the family.
ters or islands in pollen of some species, whereas The great diversity of grass flowers, spikelets
the granules are isolated in others (Andersen and and inflorescences has sparked speculation on
Bertelsen 1972; Christensen and Horner, Jr. 1974; whether there has been selection on the grasses
Watson and Bell 1975). Differences in pollen for pollination efficiency. Discussion in the liter-
morphology between species, genera, or larger ature has focused on the relative importance of
taxonomic groups are subtle. Although various simply intercepting pollen on the windward side
authors have tried to find characters to distin- of the inflorescence versus capturing pollen
guish species or genera, and to distinguish wild through vortices created on the leeward side. In
and cultivated plants, these efforts have met with a detailed study of 20 species of pooid grasses,
limited success. In broad taxonomic surveys of Friedman and Harder (2005) measured traits of
grass pollen morphology, Page (1975) and Wat- inflorescence and flower size. They found a nega-
son and Bell (1975) find few characters to distin- tive correlation between numbers of flowers and
guish major groups. In addition, Watson and Bell flower length. The species that they studied could
(1975) find variation within a single grain that be classified as having large or small flowers, and
mirrors the variation between species. Page dense or open inflorescences; all four combina-
(1975) notes that “[t]he overall taxonomic signif- tions of these variables were observed suggesting
icance of sexine pattern in grass pollen is very that there was no optimum. Pollen size is posi-
limited,” and “there is no obvious correlation [of tively correlated with the number of pollen grains
sexine patterns] with habitat, pollination biology and total volume of pollen per flower, indicating
or taxonomy.” that there is no trade-off between pollen size and
A recent attempt to quantify differences in amount. Pollen size also correlates positively with
pollen grain surface morphology has confirmed stigma length.
previous studies (Mander et al. 2013). A classifi- Cresswell et al. (2010) used both modeling
cation algorithm was developed that identified and experimental approaches to suggest that
Morphology and Anther Structure 65
most pollen is captured on the windward side of pollination efficiency is weak. Pollen may not
grass inflorescences and thus that the formation often be limiting. Pollen may be collected primar-
of vortices is largely irrelevant to pollination. The ily on the windward side of the inflorescence,
one exception that they were able to find in the meaning that the formation of eddies by complex
literature was Setaria parviflora (¼ S. geniculata), architecture may not be relevant. Insect pollina-
in which over a third of the pollen was captured tion may occur but its importance is unknown.
on the leeward side of the inflorescence (Niklas
1987). However, S. parviflora oscillates in the
Pollen germination and pollen tube growth
wind, and thus the leeward side of the inflores-
cence may end up facing the wind as it bends. In Lausser and Dresselhaus (2010) divide pollina-
addition, the sterile branches (bristles) character- tion in grasses into five phases, based on their
istic of the genus could presumably have some investigations with Zea mays. Phase I is germina-
effect on the boundary layer and may thus cause tion, in which pollen grains germinate on the
some of the discrepancy. stigmatic papillae. In describing the details of
For selection to operate on pollination effi- this process, Heslop-Harrison and Heslop-Harri-
ciency, pollen must frequently be limiting in natu- son (1987) provide a thorough description of the
ral habitats; however, data on pollen limitation in ultrastructure and histology of pollen grains and
grasses are sparse. McKone et al. (1998) report that stigmas in the grasses. Grass stigmas are “dry”,
Andropogon gerardii and Sorghastrum nutans are without obvious liquid produced. The stigmatic
not pollen limited. Davis et al. (2004) report that cells have a thin cuticle that is interrupted in
pollen is not limiting in Spartina alterniflora when places; this is covered with a layer of protein
it is growing in dense stands but, when density of and mucilage. When the pollen grain lands on
conspecifics is low, then supplemental pollination the stigmatic surface, water quickly (20–25 sec-
raises seed set several fold. onds in rye) moves osmotically from the stig-
Despite the prevalence of wind pollination in matic cell to the pollen grain, forming a
the family, a few taxa may be insect-pollinated. meniscus. Hydration of the pollen grain then
For example, the anthers in Streptochaeta and proceeds over a period of several minutes. Unlike
Anomochloa are basifixed or centrifixed, rather pollen of many eudicots, grass pollen retains a
than versatile, and the stigmas are not plumose water content of 15–20 %, so it is partially
(Sajo et al. 2009). Both characteristics suggest hydrated when it arrives on a stigma, allowing it
insect pollination rather than wind pollination. to germinate almost instantly (Heslop-Harrison
Insects have been observed on inflorescences of and Heslop-Harrison 1992; Heslop-Harrison
the understory grasses Olyra and Pariana in Bra- et al. 1997; Nepi et al. 2001). A single stigmatic
zil (Soderstrom and Calderón 1971). The most cell can rehydrate four or five pollen grains
common visitors were gall midges and phorid (Heslop-Harrison and Heslop-Harrison 1987).
flies; the insects were observed consuming pollen, As germination proceeds, the pollen tube forms
but evidence for effective pollen transfer is still and protrudes through what had been the pollen
lacking. Huang et al. (2002) report visits of hon- aperture.
eybees to Phyllostachys nidularia, and cite other The grass stigma appears to exert little selec-
references of insect visitors to bamboo flowers. tive control over pollen germination (Heslop-
Adams et al. (1981) observed halictid bees Harrison and Heslop-Harrison 1987). Lausser
visiting Paspalum dilatatum in south-central et al. (2010) applied pollen from Lolium multi-
Oklahoma, and followed up with exclusion florum, Poa nemoralis, Oryza sativa, Zea mays,
experiments; these showed that more pollen was Tripsacum dactyloides, Lilium longiflorum, and
transferred and more seed was set when flowers Arabidopsis thaliana to stigmas of Z. mays and
were exposed to bees plus wind than to wind T. dactyloides; all pollen except that from Lilium
alone. germinated. Other more circumstantial evidence
In summary, data on grass pollination are for the possibility of indiscriminate germination
remarkably sparse considering the abundance of comes from data on wide crosses. Allopolyploidy
grass species. Evidence that the diversity of grass is common in the grasses, as noted in the section
inflorescences has been driven by selection for above on Karyology and genome structure.
66 Pollen
In addition, successful pollination has been tion.) Phase V, gametophytic interactions,

demonstrated between wheat and maize, wheat appears to be the only stage in maize in which
and sorghum, and barley and maize (Laurie and signals are required from the female gametophyte
Bennett 1988), as well as between maize and oats (Lausser and Dresselhaus 2010).
(Riera-Lizarazu et al. 1996). Given these results, it Pollen tube growth in connection with self- or
seems likely that pollen from nearly any grass can cross-compatibility is discussed under Reproduc-
germinate on the stigma of nearly any other grass. tive systems.
Phase II of pollination (sensu Lausser et al.
2010) is stigma invasion, in which the pollen tube
grows into the intercellular spaces in the stig- References
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walls (Cosgrove et al. 1997; Valdivia et al. 2009). example of insect pollination in a temperate grass.
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Fruit and Seed
The fruit of Poaceae is a caryopsis, in which the nucellus are broken down. The pericarp in Pap-
seed coat is firmly attached to the inner wall of the pophorum subbulbosum (subfamily Chloridoi-
ovary. This structure is uniquely derived in the deae) is similar, but the inner integument, outer
family. Within this general category, however, integument and nucellus are all collapsed (Rost
there is variation in the structure of the pericarp et al. 1984).
and the ultimate fate of the integuments and In a number of bamboos and chloridoid
nucellus; in particular the inner integument of grasses, the seed coat is free from the pericarp,
the ovule, which generally constitutes the seed or the two are only weakly adherent. This is a
coat, may be intact or crushed at maturity. Only derived condition within the family. The develop-
a few taxa have been studied in detail, and these ment and cellular structure of such fruits have not
are so widely scattered taxonomically that the been studied, so it is not clear what aspects of
data appear somewhat anecdotal; a broader sur- development are modified, or even if the deriva-
vey would help clarify the generality of the obser- tion of the free seed occurs by the same mecha-
vations. nism in all cases.
In wheat (subfamily Pooideae), the outer and Several bamboo genera have fleshy fruits. In
inner epidermes of the pericarp consist of cells the two that have been studied closely, Dinochloa
with the long axes parallel to the apical-basal axis and Ochlandra, embryo and endosperm develop-
of the grain (Percival 1921). Between them are ment are similar to that of other grasses, but the
one or two layers of parenchyma cells with thick pericarp becomes thick and fleshy (Rudall and
walls plus the “cross layer”, which is made up of Dransfield 1989). The structure of the scutellum
cells with their long axes more or less perpendic- is normal, with the apical cells elongated and
ular to the epidermal cells. The inner epidermis is apparently functioning as transfer cells moving
discontinuous, with the long cells widely sepa- nutrients from the endosperm; Rudall and Drans-
rated from each other; at maturity these are field (1989) speculate that nutrients are trans-
often crushed. The seed coat is the product of ferred from the pericarp to the embryo via the
the inner integument, which is two layered at endosperm. The latter is liquid in the baccate
maturity although these layers are also crushed fruits of the bamboo Melocanna (Ramanayake
in late development, along with the nucellus. In and Weerawardene 2003). Vivipary is also
Sorghum halepense, Setaria italica, and Cenchrus reported in Dinochloa; Dransfield (1981) suggests
americanus (¼ Pennisetum glaucum) (subfamily that this character is correlated with the presence
Panicoideae) the mature pericarp consists of the of the fleshy fruit.
outer epidermis plus crushed and broken frag- The point of attachment of the fruit to the
ments of the inner epidermis; layers in between plant is known, inaccurately, as the hilum. Since
break down and are crushed in the mature fruit the term hilum normally refers to the point of
(Fussell and Dwarte 1980; Harrington and attachment of the seed to the placenta, this use of
Crocker 1923; Narayanaswami 1956). The inner the term in grasses is somewhat misleading, but is
integument is intact but the outer integument and so widespread it will be followed here. It is in the
70 Fruit and Seed
basal part of the pericarp, and corresponds to a nodes, dispersing spikelets or clusters of spikelets
region slightly below the placenta (Narayanas- with an attached rachis internode. Finally, the
wami 1956). Other terms include “hilar orifice” entire inflorescence may be shed as a unit (e.g.,
(Harrington and Crocker 1923), “placental pad” some Aegilops and Agrostis species). Disarticula-
(Rost et al. 1984), and “chalazal pad” (Fussell and tion – and hence dispersal – mode may vary
Dwarte 1980). The latter is misleading as well among species of a single genus (e.g., Muhlenber-
since it is roughly opposite the micropylar end gia), or may be characteristic of a genus or tribe.
of the embryo, not the chalaza. The cells of the Some species have more than one sort of
hilum may accumulate pigments in some species, propagule. For example, Sphenopholis obtusata
such as Cenchrus americanus (¼ Pennisetum disarticulates between the two flowers and also
glaucum) (Fussell and Dwarte 1980) and Sorghum below the glumes so that it has one small dia-
bicolor (Giles et al. 1975), in which the area spore, consisting of only the upper flower, and
becomes known as the “black region”; its forma- one large diaspore consisting of the lower flower
tion indicates the end of carbon assimilation by plus the glumes (Rabinowitz and Rapp 1981).
the grain. Seed weight (and hence propagule weight)
The shape and size of the fruit, embryo, and varies within an inflorescence and thus presum-
hilum are frequently used as taxonomic charac- ably affects dispersal ability as well as seedling
ters and can be diagnostic of large groups. For survival. In Eremopyrum distans, seeds from the
example, a punctate hilum is a putative synapo- proximal flower of the spikelet are larger than
morphy for the tribe Poeae (Pooideae). The size those from distal flowers, have higher seedling
of the embryo varies relative to the size of the survival, and form larger plants that produce
caryopsis, and the caryopsis itself may be long more seeds (Wang et al. 2010). In Leymus arenar-
and linear (e.g., some Stipeae) or nearly spherical ius, seeds are heavier from the upper parts of the
(e.g., some Panicoideae). Shape and size of the inflorescence (the reverse of Eremopyrum), and
fruit are under the control of different genetic in the proximal flowers of the spikelet (same as
programs (Gegas et al. 2010). Eremopyrum) (Greipsson and Davy 1995).
Vegetative propagation is also common,
although in most cases it results in very local
DISPERSAL dispersal. However, in some species that grow
near or in water, fragmentation of the parent
The dispersal unit in the grasses varies consider- plant into separate ramets can lead to water dis-
ably, and reflects the many different plant tissues persal. This has been reported for Arundo donax
in which a disarticulation zone can form (Doust in South Africa, for example (Milton 2004). On
et al. 2014). In some taxa, the fruit is shed sepa- the other hand, a study using microsatellite mar-
rately from the remainder of the spikelet. More kers found little evidence that stands of Phragmi-
commonly, the rachilla disarticulates at each tes australis in a Canadian lake were propagated
node, so that the dispersal unit is the entire vegetatively; rather, most clumps were genetically
flower, including the lemma and palea, plus the distinct, pointing to dispersal and propagation by
attached bit of the rachilla internode. In other seed (Belzile et al. 2010).
species, the rachilla disarticulates above the The great diversity of grass dispersal units
glumes but not between the flowers, so that the suggests that dispersal ability may have contrib-
dispersal unit becomes all the flowers of the uted to species diversity in the family. As evi-
spikelet, often including a cluster of sterile dence for this, Davidse (1987) notes that the
lemmas at the distal end. In most Panicoideae, pistillate spikelets or inflorescences of many dioe-
disarticulation is below the glumes so that the cious species are highly modified with awns or
entire spikelet is shed, a characteristic that often burs that appear to assist in dispersal, whereas
provides a useful field identification character for the staminate spikelets are generally unspecial-
the subfamily. In other taxa, disarticulation is at ized. However, as with most other aspects of grass
the base of the primary branch, which then is ecology, dispersal has rarely been studied in a
shed as a unit. In many Triticeae and Andropo- broad systematic context, nor is there a rich
goneae, the inflorescence axis breaks up at the experimental literature. Instead the literature on
DISPERSAL 71
grass dispersal is buried in studies of community general, as a seed matures, the moisture content of
ecology, range management, invasive species and the seed is reduced, particular sets of proteins
community restoration; any one study is limited accumulate, and the hormone abscisic acid (ABA)
to the grasses of a particular, often limited, com- increases, as does sensitivity of the embryo to its
munity. Most existing studies are observational effects; at the same time gibberellin content and
and correlative, although there are some notable sensitivity are low. In the grasses, this process
exceptions as outlined below. This summary is occurs along with accumulation of starch in the
thus incomplete and the definitive review of the endosperm. The details of seed maturation are
literature on grass dispersal has yet to be written. best worked out in Arabidopsis, which lacks
starchy endosperm and thus may not be compara-
Unassisted or wind dispersal ble to the grasses; the controls of seed maturation
and germination must be investigated indepen-
Dispersal in grasses is assumed to be predomi-
dently in the grasses (Barrero et al. 2010; Gutierrez
nantly passive, with seeds being dropped from
et al. 2007; Sabelli and Larkins 2009). (See also the
the inflorescence (“unassisted dispersal”) and/or
section on Embryology above.)
spread by wind. This assumption is based on the
While many grass seeds germinate immedi-
generally small size of the dispersal units, lack of
ately after they are shed, others require a period
obvious morphological characteristics that might
of after-ripening or are dormant. After-ripening
permit other modes of dispersal, or sometimes
is a short period of time in which the seed is
presence of structures that might reduce the ter-
unable to germinate, but germinability generally
minal velocity of the propagule to keep it in the
is gradually restored by a period of dry storage.
air column longer (Davidse 1987). For example,
Dormancy is the inability to germinate even when
Jurado et al. (1991) classified 48 of 58 Australian
environmental conditions are otherwise favor-
grass species as wind-dispersed or unassisted.
able (Baskin and Baskin 1998; Simpson 1990).
Correlation between morphological cate-
Dormancy is under genetic control, and varies
gories and actual dispersal mechanism has rarely
among species and between plants within a spe-
been tested, so the mechanism is usually inferred
cies (Barrero et al. 2012; Baskin and Baskin 1998;
rather than demonstrated. For example, Thom-
Simpson 1990). Baskin and Baskin (1998) list a
son et al. (2010) found data on dispersal mecha-
sample of 11 non-domesticated species in which
nism for only 15 % of eastern Australian grasses.
fresh mature seeds will germinate immediately;
Studies of grasses in Missouri and in Botswana
these include representatives of the Pooideae,
found that propagule weight and morphology
Chloridoideae and Panicoideae, indicating that
affected terminal velocity (measured as the
lack of dormancy is phylogenetically widespread.
speed with which a propagule fell from a specified
A list of genera and species in which dormancy
height) and ability to be transported laterally
has been evaluated is provided by Simpson
(Ernst et al. 1992; Rabinowitz and Rapp 1981).
(1990); it would be interesting to update this
In general, lighter propagules dispersed more
list, which seems likely to expand.
widely, but distances were rarely more than a
Dormancy in the grasses is not related to
few meters at most (Cheplick 1998).
embryo maturity, unlike the case in some other
In one study, grasses were found to disperse
angiosperms (Baskin and Baskin 1998). In all
soon after flowering, although some exceptions
grasses known, the embryo is mature when the
occurred (Rabinowitz and Rapp 1980). Thus, cool-
seed is shed, and if the embryo is excised from the
season grasses (e.g., Pooideae) tend to disperse in
caryopsis it will almost always germinate (Simp-
summer, whereas warm-season grasses disperse in
son 1990). This observation thus points to the
autumn. The exception was Festuca paradoxa,
endosperm, aleurone, or pericarp as maintaining
which dispersed throughout the growing season.
the dormant state.
The physical constraint of the pericarp appears
Dispersal in time: seed dormancy
unlikely to cause dormancy (Baskin and Baskin
The population dynamics of a plant species are 1998; Simpson 1990). Whereas some angiosperms
controlled in part by the timing of germination have seeds that must be scarified before germina-
and the ability of the seed to persist in the soil. In tion occurs, this does not appear to be an absolute
72 Fruit and Seed
requirement in the grasses. However, puncturing let within the inflorescence and position of the
the pericarp just above the embryo does enhance flower within the spikelet (Simpson 1990). For
the percentage of germination in some species, example, in Agrostis curtisii, Avenula marginata,
apparently by permitting penetration of oxygen and Pseudarrhenatherum longifolium, position of
or additional water (Simpson 1990). the seed in the inflorescence influences dor-
Dormancy is affected by the presence of mancy, and interacts with high temperatures
lemmas, paleas and glumes, which are often part and plant ash to control germination (González-
of the dispersal unit (Barrero et al. 2012; Baskin Rabanal et al. 1994).
and Baskin 1998; Simpson 1990). These struc- The connection between inflorescence archi-
tures appear to limit access of water and light; tecture and dormancy may be indirect and be
when they are removed germination percentage is mediated by seed provisioning. In Eremopyrum
increased. In Brachypodium, for example, dor- distans, seeds from the proximal flower of the
mancy enforced by presence of the lemma and spikelet had a higher proportion with physiologi-
palea may last longer than dormancy of the naked cal dormancy and lower seedling emergence the
caryopsis, but is eventually lost in darkness (Bar- next spring (Wang et al. 2010). This pattern cor-
rero et al. 2012). Although there has been some relates with variation in seed weight mentioned
suggestion that the palea, lemma and glumes above. Howard et al. (2012) show that dormancy
contain inhibitors to germination, there is little is related to starch content of the seed, which is
evidence for this. inversely related to sugar content. Higher sugar
The conditions that break dormancy vary content led to reduced sensitivity of the embryo
widely and do not appear to correlate with phy- to ABA in barley, and hence less dormancy. How-
logeny, hinting that the signaling pathways that ever, the relationship between seed size and dor-
control germination are exquisitely sensitive to mancy is not simple, and varies among species
modification by natural selection in response to (Baskin and Baskin 1998).
particular environments. Presence or absence of Among domesticated species, selection has
light, and the quality of the light, are both major reduced dormancy (Barrero et al. 2010). In some
factors in germination (Simpson 1990). Both blue cases this creates the problem of pre-harvest
light, acting through the cryptochrome system, sprouting, in which the seeds germinate on the
and red/far-red light, acting primarily through plant (Gubler et al. 2005). “Viviparous” mutants
phytochrome B, regulate dormancy in grasses; in maize generally result from defects in the ABA
both photoreceptors in turn regulate ABA pathway, suggesting that the seed maturation
(Gubler et al. 2008; Simpson 1990; Xu et al. 2009). program is disrupted. Loss of dormancy in
Moisture and temperature are also important crops is thus associated with a change in the
for some species, but the particular combination hormonal environment of the developing seed.
of the two is species-specific. While some species
(e.g., winter annuals such as Avena fatua) require
Dispersal by animals
high temperature and low moisture, others
require exactly the opposite (Baskin and Baskin The dispersal units in many grasses have hooks
1998). For most species, these requirements have or barbs on various parts, and these could be
never been dissected at the molecular level. adaptations for dispersal by mammals or birds.
Exposure to smoke breaks dormancy in some Mediterranean annual grasses are reported to be
grasses, an obvious adaptation of plants to fire- transported by mammals in Israel (Shmida and
prone ecosystems. The effects of smoke interact Ellner 1983), and the pooid species Agropyron
with those of light, temperature and moisture cristatum and Stipa krylovii were found to be
content, so their effect on dormancy is not easy transported easily in the fur of sheep and goats
to predict (Turner et al. 2009). in Mongolia (Bl€ass et al. 2010). Diaspores of many
Dormancy is affected by inflorescence archi- species of grasses were found in the hair of bison
tecture, suggesting a possible adaptive role for the in tallgrass prairie in North America; most of
considerable inflorescence diversity in the these did not have any obvious hooks or barbs
grasses. Dormancy and germinability vary within that would attach them to the animals, although
a plant, and are affected by position of the spike- many did have hairs or awns whose function is
DISPERSAL 73
unclear (Rosas et al. 2008). A careful develop- grasses in which the caryopsis is enclosed in a
mental study of the hooks in Pseudechinolaena spikelet with awns or hairs are transported by
by Lucas (1979) showed that they are emergences ants in Arizona deserts (Pulliam and Brand
of the back of the glumes, tipped by a single stiff 1975). Davidse (1987) lists 31 species in 11 genera
macrohair. of Panicoideae in which the rachilla contains oil,
Grass fruits will survive passage through the as indicated by staining with Sudan IV, suggest-
guts of mammals, and can be found germinating ing that they may function as elaiosomes. Strep-
in dung. Somewhat surprisingly, the grasses tostachys has a thickened rachilla that also could
reported to move in this way are predominantly be an elaiosome (Morrone and Zuloaga 1991).
pooids, whose caryopses are enclosed only in a
membranous lemma and palea rather than any
Role of the awn
indurated covering. Introduced annuals of the
genera Briza, Bromus, Vulpia, Lolium and Poa Awns are common in grasses and often form a
constitute a high proportion of seedlings sprout- prominent part of the spikelet. Awns most com-
ing from the dung of African ungulates (Milton monly occur on the lemma, but awned glumes are
2004), and nearly half the seeds found in dung of also found in some taxa. The position of the awn,
bison in a tallgrass prairie were from grasses whether on the lemma or glume, varies; it may
(Rosas et al. 2008). Cosyns et al. (2005) fed seeds form at the apex, or from a sinus, or on the
of Agrostis capillaris, Anthoxanthum odoratum abaxial side (generally called the dorsal side,
and Poa pratensis to sheep, cattle, rabbits, horses, although this usage contradicts the use of the
and donkeys, and compared their germination to word “dorsal” in other plant groups). It may be
that of seeds placed directly on soil or on soil plus straight or twisted; generally if twisted the awn is
dung. Germinability for ingested seeds of all spe- hygroscopic, changing shape according to the
cies was mostly less than 10 % of that of non- amount of available moisture. While it is most
ingested seeds, but was more than zero. Seeds of common to have a single awn per lemma or
Eragrostis lehmanniana can pass through the glume, the lateral veins may be prolonged into
digestive tract of a sheep and still be viable (Fre- awns so that the lemma or glume has three, five or
drickson et al. 1997). While Bl€ass et al. (2010) even more awns.
contend that transport of seeds in the gut of gra- Awns are thought to be adaptations for seed
zers is only of minor importance, such transport dispersal (Elbaum et al. 2007; Garnier and Dajoz
could provide a mechanism for relatively long 2001; Peart 1979, 1981, 1984; Peart and Clifford
distance dispersal. 1987). In addition, awns also affect seed provi-
A few grasses have fleshy dispersal units. The sioning (e.g., Li et al. 2006; Motzo and Giunta
pericarp is fleshy in a few species of bamboo (e.g., 2002), and possibly also drought stress (Abebe
Melocanna, Ferrocalamus), whereas the fleshy et al. 2010).
structures in Lasiacis (Panicoideae) are glumes The role of awns in dispersal has been
and sterile lemmas that become shiny black at explored in a few species (Elbaum et al. 2007;
maturity and accumulate oil in their epidermal Garnier and Dajoz 2001; Peart 1979, 1981, 1984;
layers (Davidse and Morton 1973). Spikelets of Peart and Clifford 1987). Wetting and drying of a
Lasiacis were collected from the stomachs of hygroscopic awn causes the dispersal unit (often
fruit-eating birds, indicating that the birds do a flower or spikelet) to shift position and can even
consume them, and the seeds germinated, indi- propel it several millimeters across the substrate.
cating that the seeds can be dispersed. It is Hygroscopic awns in Themeda triandra move the
unknown whether other panicoid grasses will spikelets along the soil and increase the chance
survive passage through the gut of a bird, but that they will become buried (Sindel et al. 1993).
this could provide a mechanism for long-distance The hygroscopic awn in Macrochloa tenacissima
dispersal. (¼ Stipa tenacissima) causes the diaspore to be
Insect dispersal is rarely reported, but may buried in the soil or in crevices (Sch€oning et al.
simply be difficult to detect. Grasses with smooth 2004). The presence of lateral awns, in addition to
caryopses are easily eaten by birds and rodents, the central one, improves the effect of the hygro-
but are harder for ants to grasp. Conversely, scopic awn in locating suitable microsites (Peart
74 Fruit and Seed
1979). In Avena fatua, flowers within a spikelet glume) has long hairs and in which the callus is
mature from bottom to top (acropetally) and the long and pubescent, a combination of characters
hygroscopic action of the awn on the lowermost described as the “burial syndrome” (Humphreys
flower appears to force the flowers out of the et al. 2011). Long hairs on the dispersal unit have
spikelets. The result is dispersal of seeds of vari- an effect similar to that of hygroscopic awns in
ous sizes and at various stages of maturity (Raju affecting orientation of the propagule relative to
and Ramaswamy 1983). the soil (Peart 1979). Long hairs, whether on the
Presence or absence of an awn affects seed propagule itself or only on the callus, also provide
germination. In a series of experiments of six an anchor so that the radicle can penetrate the
species of grasses with awns, Peart (1979) showed soil more effectively. In the absence of anchoring,
that twisting of the hygroscopic awn placed the the force of radicle expansion can push the anthe-
anthecium into a microsite suitable for seed ger- cium out of the ground and can increase mortal-
mination, such as a crack in the soil; removal of ity (Peart 1981). Comparing one species each of
awns resulted in fewer seeds arriving in suitable Aristida and Microlaena, Peart (1981) found an
microsites. He found an interaction between awn interaction between presence of awns, presence of
type (species) and soil type, with some awn barbs on the callus, and soil type.
morphologies more efficient than others in locat- Although awns appear to provide a selective
ing microsites in different sorts of soils. Sch€oning advantage in species in which they occur, awns
et al. (2004) show that seeds from diaspores with have been lost many times in the evolution of
intact awns have higher germination than those grasses. In Danthonioideae, awns have been lost
with awns removed. In addition, awns make the independently about 27 times, although the exact
seed less susceptible to predation by ants; number depends on analytical assumptions
although the awn breaks off eventually, once the (Humphreys et al. 2011). In that subfamily, loss
diaspore is buried the ants are unable to find it. of awns occurs preferentially in annual lineages,
The experiments of Sch€oning et al. (2004) and suggesting that perhaps awns do not confer an
Peart (1979) are highly informative and deserve advantage in species with an annual life history.
to be repeated on a much larger set of species. Awnless danthonioids tend to have glabrous
The length of the awn varies among closely lemmas and short glabrous calluses, both char-
related species, but within a species awn length is acters that imply that seeds are not buried. Con-
highly heritable, with minimal effect of the envi- versely, awned danthonioids generally have
ronment (Garnier and Dajoz 2001). In Hyparrhe- longer calluses and tufts of hairs on the lemmas.
nia diplandra there was a positive relationship Loss of the various components of the burial
between the length of the twisted hygroscopic syndrome occurs in environments where active
awn and both the proportion of buried seeds and burial is less advantageous, and in which burial
mean seed burial depth (Garnier and Dajoz 2001). can be random.
Fire appears to be a selective force on awn Despite the ecological and agronomic impor-
morphology. Peart (1984) found that species tance of awns, the genetic control of their develop-
lacking an awn were more likely to germinate ment has not been explored extensively. A recent
after a fire. He postulated that the awned species study has identified one gene that controls cell
tended to germinate on or near the surface and division in the lemma of rice and thereby regulates
were thus destroyed by fire, whereas species with- awn formation (Luo et al. 2013). This same study
out appendages were more likely to be buried. In found that awn length and grain length were posi-
contrast, Garnier and Dajoz (2001) found a positively correlated with each other, and inversely
tive relationship between awn length and burial correlated with the number of grains per inflores-
depth, with longer awned species being more cence, providing a genetic basis for the trade-off
deeply buried and more likely to survive intense between fruit size and number.
fires. In general, variability in fire occurrence and
intensity also selects for variation in awn length
Forcible dispersal
(Garnier and Dajoz 2001).
Long twisted awns often occur in species in Forcible dispersal, or ballistochory, is not com-
which the outside of the dispersal unit (lemma or mon in grasses but has been reported for Sucrea
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Phytochemistry
Compared to many other angiosperms, the essarily biased toward compounds that are pro-
grasses are not noted for the diversity of their duced in relatively high amounts. Studies often
secondary compounds. However, as indicated rely on extraction of exudates or separation of
below, this may simply reflect lack of investiga- compounds from whole tissue, followed by frac-
tion. A wide range of secondary compounds is tionation and analysis of compounds by HPLC,
reported, but in general the data come from a GC or other methods (e.g., Bais et al. 2006; Xuan
handful of taxa that are particularly important et al. 2006). While such data provide good evi-
for grazing or field crops; the taxonomic distribu- dence of what the plant can produce under par-
tion of most compounds is thus largely unknown. ticular growth conditions, failure to find a
Secondary compounds may be produced by roots compound in an extract does not mean that the
or by above ground tissue, and production may plant cannot produce it. This inability to interpret
be constitutive, or be triggered by herbivore negative data, combined with the patchy sam-
attack, or be produced in response to the pres- pling, means that it is hard to generalize patterns
ence of particular bacteria or fungi. Of the 2400 to broad taxonomic groups.
species in a database of plants with pest-control More interesting evolutionary comparisons
properties (Grainge and Ahmed 1988), about 50 are possible in the cases where the biosynthetic
are grasses; documentation of the chemicals pro- pathway is known, or can be determined by
duced by these plants and their effects on other genetic approaches, as described below for
organisms varies considerably in quality. Com- momilactones in rice. As more biosynthetic path-
pounds produced in response to pathogens are ways are determined, it will be possible to try to
sometimes called phytoalexins, a term that refers locate orthologous genes in other taxa using
to their inducibility rather than their chemical whole genome sequences; this approach will ulti-
composition. A number of grass species harbor mately provide a better picture of how phyto-
fungal endophytes that produce secondary com- chemical diversification has occurred in the
pounds that serve as plant defense; in this case family. In addition, by comparing normal plants
the defense compound is in fact a product of the with those that have a mutation in a single gene in
fungus rather than the plant. Finally, cell walls in the biosynthetic pathway, it is possible to isolate
the grasses have a distinctive chemical composi- the effects of particular secondary compounds on
tion; whether it is adaptive in any way is the plant and the community of organisms
unknown. The material presented here is not around it. In general, however, these investiga-
intended to be a comprehensive review, but will tions have focused on crops and have yet to be
provide some hint of the phytochemical diversity extended to wild species.
to be explored in the family. Root exudates from grasses may have allelo-
Much of the existing literature on secondary pathic properties (Duke 2007), and allelopathy
compounds in grasses reports on the composi- has been reported in most major cereals (Belz
tion of extracts, which are made up of the end 2007). Allelopathic effects are notoriously hard
products of biosynthetic pathways, and are nec- to demonstrate conclusively, because negative
78 Phytochemistry
plant-plant interactions can be caused by many members of subfamily Pooideae are the most
other direct (e.g., competition) and indirect (e.g., common grass producers of flavonols, the pani-
effects on soil microbes) effects. In addition, such coid Bothriochloa bladhii is the only recorded
studies are necessarily biased toward compounds source of 7,30 ,40 -trihydroxyflavone, and the pani-
that are produced in relatively high quantities coids Arthraxon hispidus and Miscanthus tinctor-
under particular growth conditions, or constitu- ius are the only species known to produce
tively. arthraxin, a derivative of luteolin (Harborne and
For at least two of the biosynthetic pathways Williams 1987). Rice seedlings may also release
described below – those for momilactones in rice 5,7,40 -trihydroxy-30 ,50 -dimethoxyflavone, and 3-
and for avenacins in oats – the genes encoding the isopropyl-5-acetoxycyclohexene-2-one-1 into the
relevant enzymes are clustered in the genome environment; these inhibit the growth of Echino-
(Field and Osbourn 2008; Qi et al. 2004; Xu et al. chloa crus-galli and Cyperus difformis, and are
2012). This is apparently the result of genomic particularly potent in combination (Kong et al.
rearrangement; the genes are not linked in other 2004).
grasses. Such an arrangement has also been found Flavonoids are often found as glycosides, in
for related genes in Arabidopsis, and may point to which the flavonoid molecule is bound to a sugar;
selection for co-regulation of components of the diversity comes from the kind and number of
pathway (Field and Osbourn 2008). In some cases sugars and the nature of the bond between the
also the gene products are all anchored at a par- sugar and the flavonoid (sometimes called the
ticular cellular location, forming metabolic units aglycone). Both C-glycosylflavones and O-glyco-
called metabolons; such localization appears to sides are common in the grasses (Harborne and
increase efficiency of biosynthesis and to reduce Williams 1987), as they are in other families. A
accumulation of potentially toxic intermediates. leaf of barley or wheat may contain several dozen
different C-glycosyl-flavones. Maysin is a C-gly-
cosyl-flavone produced in the stigmas of maize,
PHENOLICS where it provides resistance to corn earworm
(McMullen et al. 2009). In addition to the glyco-
Various secondary compounds are produced by sides, some taxa, particularly in the PACMAD
the phenylpropanoid pathway, beginning with clade, produce flavonoid sulphates, but these are
phenylalanine, and proceeding through steps cat- apparently not found (or not recorded) for mem-
alyzed by phenyalanine ammonia lyase, cinna- bers of the BEP clade.
mate-4-hydroxylase, 4-coumarate: CoA ligase, Anthocyanins, a subgroup of flavonoids, are
and chalcone synthase. Some of the most wide- common in Poaceae, and probably occur in all
spread and best characterized of the products of species. The anthocyanins of maize are well
this pathway are the flavonoids. Flavonoids are known, producing brightly colored kernels and
widespread plant pigments that are water soluble ears; study of these pigments and the enzymes
and generally stored in vacuoles. They exhibit that produce them has provided insight into fun-
highly diverse structures, and confer a variety of damental aspects of gene structure and regula-
benefits on the plant, including disease resis- tion, and famously led to the discovery of
tance, deterrence of herbivory, and protection transposable elements (Comfort 2001). All
against UV damage (Grotewold 2006; Harborne grasses investigated produce the anthocyanin
and Williams 2000). The biosynthetic pathway of cyanidin and its derivatives (chiefly cyanidin 3-
flavonoids is shared among seed plants (Tanaka glucoside), whereas the distribution of delphini-
et al. 2008). Like many plants, grasses produce a din and peonidin in plant extracts is more spo-
variety of flavonoids (Harborne and Williams radic (Escribano-Bailón et al. 2004; Fossen et al.
1987), the most common of which is tricin, 2002). Phlobaphenes, polymers of flavan-4-ols,
found in over 90 % of grass species. Other flavo- form in the floral tissues of maize and sorghum
noids are apparently taxonomically restricted, and accumulate in the developing kernel. The
although it is unclear whether these compounds biosynthetic pathway of these is becoming well
really are specific, or whether they have simply known, and is shown to proceed via luteoforol or
not been detected in other taxa. For example, apiforol (Morohashi et al. 2012).
DERIVATIVES OF AMINO ACIDS (TRYPTOPHAN, PHENYLALANINE, TYROSINE) 79
Other compounds produced by the phenyl- pounds are produced as D-glucosides and are
propanoid pathway are the stilbenes. Sorghum stored in the vacuole; when the cell is damaged,
seedlings produce the stilbene trans-piceid in the glucose moiety is removed, releasing the toxic
response to fungal inoculation (Chong et al. aglucone (Sicker et al. 2000). The ability to pro-
2009). In many plants, duplicate genes for chal- duce these compounds appears to be synapo-
cone synthase have been modified so they instead morphic for the grasses or at least the members
encode stilbene synthase, a protein that creates of the BEP plus PACMAD clades; the few dicots
stilbene, a cyclic compound distinct from chal- that produce them do so using non-orthologous
cone. Thus, stilbene synthase has been derived proteins (Dick et al. 2012). The benzoxazinones
repeatedly from chalcone synthase (Chong et al. are derived from indole, which ultimately comes
2009), creating a novel branch in the phenylpro- from tryptophan, and their biosynthesis is cata-
panoid pathway. In sorghum, stilbene synthase lyzed by a set of genes whose chromosomal loca-
appears to be single copy (Paterson et al. 2009; Yu tions vary among the grasses (Sue et al. 2011).
et al. 2005). The genes necessary for synthesizing benzoxazi-
The cell wall composition of the grasses has nones are missing in cultivated barley, but pres-
attracted considerable attention because several ent in other species of Hordeum (Gr€ un et al.
members of the family (e.g., Panicum virgatum, 2005). DIBOA is exuded by the roots of rye, and
Miscanthus x giganteus, and prairie grasses in appears to have some allelopathic activity (Sicker
general) have been suggested as possible sources et al. 2000).
of cellulosic biofuels. The accumulation of ferulic A variety of indole alkaloids has been
acid and coumaric acid in non-lignified cell walls reported from grasses in subfamily Pooideae,
is a characteristic shared with all other commeli- where they are of particular interest because of
nid monocots such as palms, bananas and gin- their toxicity to livestock. The indole alkaloids
gers. Primary walls include glucurono- dimethyl tryptamine and tyramine-related com-
arabinoxylans, a characteristic shared with Zingi- pounds are common in Phalaris, where they
berales and Commelinales. Presence of (1–3,1-4)- cause toxicity to grazers, particularly sheep
b-D-glucans is shared with the other members of (Edgar 1994). Thus, breeding efforts in Phalaris
the graminid clade. have focused on trying to reduce the concentra-
Another major group of phenolics is the qui- tion of these compounds in the plants. Levels of
nones, structurally similar to ubiquinone, an elec- the toxins vary according to cultivar, time of year,
tron carrier in the photosynthetic electron and moisture conditions (Culvenor et al. 2005).
transport chain. Sorgoleone is a p-benzoquinone Perloline (a diazaphenanthrene alkaloid) and E/
produced in root hairs of S. bicolor and S. hale- Z-thesenine-rhamnoside have been isolated from
pense (Czarnota et al. 2001, 2003). It is thought to Lolium perenne (Grimmett and Waters 1943;
inhibit growth of nearby plants by disrupting Koulman et al. 2008).
electron transport in the mitochondrion and the While some indole alkaloids in Pooideae are
chloroplast by binding to specific components of undesirable because of their effects on mammals,
the electron transport chains (Bais et al. 2006; others are beneficial because of their deterrent
Czarnota et al. 2001). It seems likely that benzo- effects on insects. The indole alkaloid gramine is
quinones are produced by other grasses as well. found in barley and in Phalaris, and has been
investigated in the former species because it con-
fers resistance to aphid attack. Barley cultivars
DERIVATIVES OF AMINO ACIDS (TRYPTOPHAN, differ in production of gramine, and production
PHENYLALANINE, TYROSINE) is also affected by specific aphids and by drought
stress (Larsson et al. 2011). Some of the genes for
Grasses produce benzoxazinones, which appear gramine biosynthesis have been identified (Lars-
to be useful in defending the plant against bacte- son et al. 2006).
ria, fungi, and insects (Sicker et al. 2000); the best Elymus (¼ Elytrigia) repens induces nearby
known of these are DIBOA (2,4-dihydroxy-1,4- plants to become more resistant to insect attack.
benzoxazin-3-one) and DIMBOA (produced by It releases a complex mixture of substances from
adding a methoxyl group to DIBOA). The com- its roots, including carboline (6-hydroxy-1.2.3.4-
80 Phytochemistry
tetrahydro-b-carboline-3-carboxylic acid, an sequestered in the vacuole in epidermal cells of

indole alkaloid). When barley plants are grown the shoot, where it may constitute 30 % of the dry
with E. repens, carboline appears to stimulate weight; tissue damage then releases the enzymes
chemical changes in nearby barley plants to that break it down to produce HCN (Poulton
make them less palatable to aphids (Glinwood 1990). Dhurrin may also be modified to a digluco-
et al. 2003). The aphids do not respond to carbo- side, which accumulates in the apoplast and is
line itself, so must react to a chemical released by excreted in guttation droplets (Selmar et al.
the barley plants. Thus, chemical interactions 1996). Dhurrin is also reported in Sorghastrum
between two grasses affect herbivory indirectly. nutans (Gorz et al. 1979), and in members of the
In the pooid genus Festuca, the non-protein S. bicolor complex plus S. arundinaceum, S. hale-
amino acid m-tyrosine is allelopathic, inhibiting pense, S. propinquum, and S. versicolor (Haskins
germination and growth of other species of and Gorz 1985).
plants. The compound is thought to be derived
from L-phenylalanine (Duke 2007). In the roots
of Festuca rubra and F. arizonica m-tyrosine TERPENOIDS
makes up 33–43 % of root extract by dry weight
and is toxic to a variety of dicots and monocots, Grasses produce a wide variety of terpenoids.
but not to fescues that produce the product (Ber- These compounds provide protection against
tin et al. 2007). F. rubra and F. arizonica are in fungal and bacterial pathogens, and some may
separate clades in the genus Festuca with F. rubra also be allelopathic. In addition, volatile terpe-
in the Aulaxyper group, and F. arizonica classi- noids affect herbivory either directly by deterring
fied in the F. ovina group (Darbyshire and Pavlick insect herbivores, or indirectly by attracting para-
2007; Inda et al. 2008). In contrast, three other sitoids that attack the herbivores in complex tri-
members of the F. ovina group, F. longifolia (¼ F. trophic interactions.
trachyphylla), F. ovina, and F. idahoensis, did not A major group of terpenoids is the diterpe-
produce m-tyrosine and indeed were sensitive to noids, particularly the group known as the lab-
its effects on growth. Thus, m-tyrosine produc- dane-related diterpenes, which protect against
tion apparently does not correlate with phylog- fungal disease (Peters 2006, 2010). These are
eny. The only other species known to produce m- synthesized from (E, E, E)-geranylgeranyl
tyrosine is the dicot Euphorbia myrsinitis, but it is diphosphate, also a precursor of gibberellin;
not clear how many other grasses have been their synthesis is characterized by two sequential
tested. cyclization reactions. Many of the genes encoding
Data are sparse on amino acid derivatives enzymes in this pathway have been duplicated
outside subfamily Pooideae. Rice (subfamily Ehr- and modified repeatedly over the course of plant
hartoideae) produces various compounds evolution leading to several thousand compounds
derived from tryptophan when attacked by larvae that are involved in plant defense (Peters 2006,
of the rice striped stem borer; these include trypt- 2010).
amine and serotonin, the latter a well-known ani- Structurally similar compounds may have
mal hormone (Ishihara et al. 2008). Sorghum different effects. For example, in rice, the lab-
(subfamily Panicoideae) produces the cyanogenic dane-related diterpenoid momilactones are alle-
glycoside dhurrin by modifying the amino acid lopathic, whereas the diterpenoid phytocassanes
tyrosine (Poulton 1990). The biosynthetic path- are not. The proteins that synthesize momilac-
way is short, consisting of two cytochromes P450, tones are encoded by a set of genes that are
a UDP-glycose-glycosyltransferase, and a clustered in the genome (Shimura et al. 2007; Xu
NADPH-cytochrome P450 reductase, which et al. 2012). Rice plants lacking either of two of
together form a membrane-bound biosynthetic the biosynthetic genes were grown with lettuce,
complex or metabolon (Jensen et al. 2011; Nielsen which is unusually sensitive to allelopathic com-
et al. 2008). The complex has been transferred to pounds, as well as the common weed Echinochloa
Arabidopsis, which then produces dhurrin, con- crus-galli (barnyard grass) (Xu et al. 2012). Plants
firming the functional importance of the pathway unable to produce momilactone permitted much
(Tattersall et al. 2001). In the plant, dhurrin is greater growth of lettuce and barnyard grass than
TERPENOIDS 81
did the momilactone producers; however, growth nonatriene, b-caryophyllene, humulene, and
of the mutant rice plants was reduced, perhaps by alpha-terpinolene; these compounds appear to
accumulation of toxic intermediates (Xu et al. repel stem borers that attack many grasses in
2012). Like momilactones, phytocassanes are Africa, as well as crops such as maize and sor-
released from the roots of rice seedlings. How- ghum (Khan et al. 2010). The compounds
ever, these diterpenoids have no allelopathic released constitutively by M. minutiflora resem-
effects on lettuce or Chinese cabbage (Toyomasu ble those released by maize after insect attack;
et al. 2008), suggesting that the compounds may thus, intercropping maize and Melinis provides
be produced in response to other pathogens such protection against stem borers. While compara-
as fungi (Hasegawa et al. 2010; Umemura et al. ble experiments have not been done on grasses,
2003). recent work in Brassicaceae suggests that intra-
Triterpenoids may be metabolized to produce specific variation in defense compounds is likely
saponins, which have antimicrobial activity. to be maintained by contrasting selective regimes
While this is most common in dicots, triterpe- in different environments throughout the range
noid saponins – avenacins – are produced in of a species (Prasad et al. 2012).
epidermal cells of the root tip of species of A few grasses, particularly in Andropogo-
Avena and in Arrhenatherum elatius, both in neae, produce oils of commercial importance.
Aveninae (Osbourn 2003). (Oats also produce Chrysopogon (¼ Vetiveria) zizanioides is the
saponins from steroids, the avenosides, but source of vetiver oil, which forms the basis of
these are produced in the leaves.) Avenacin con- many western perfumes, including the classic
fers resistance to the fungal pathogen Gaeuman- Chanel No. 5 (Joy undated; Maffei 2002). Vetiver
nomyces graminis var. tritici (“take-all”) oil is produced in cells just outside the root endo-
(Papadopoulou et al. 1999). The protein that pro- dermis, cells that are also colonized by bacteria.
duces avenacin, b-amyrin synthase, is produced Vetiver plants grown without bacteria fail to pro-
by a gene that is a recent duplicate of a cycloarte- duce oil, and presence of the bacteria correlates
nol synthase-like gene, which is shared with with expression of terpene synthase in the plant;
wheat; apparently after duplication the protein the bacteria then metabolize the terpenes pro-
function was slightly modified to produce avena- duced by the plant to produce the complex mix
cin (Qi et al. 2004). Avenacin producing genes are of compounds characteristic of vetiver oil (Del
clustered in the oat genome, in a location that is Giudice et al. 2008).
not syntenic with the rice genome (Qi et al. 2004). Cymbopogon species are characterized by
Many terpenoids are volatile. Some grasses their oil production and pleasant scent. Among
release isoprene or monoterpenes to the atmo- other compounds, they produce oil that has a
sphere, but the significance of this is unknown, as high percentage of geraniol, which is commer-
is the taxonomic distribution of these emissions cially valuable for flavoring and for perfume, but
(Gibson 2009). Other volatile compounds are is also antimicrobial and may be useful for pest
released when the plant is attacked by herbivores. control (Chen and Viljoen 2010). C. citratus
While the volatile compounds attract parasitoids, (lemon grass) produces a lemon-scented oil that
they can also attract more herbivores, with differ- is used in cooking; the major components are
ent compounds having distinct effects on partic- geranial and its isomer neral (Rodrı́guez Quinta-
ular suites of insects. For example, rice releases nilla et al. 2012). C. martinii (palmarosa) pro-
(E)- b-caryophyllene constitutively; when the duces an oil that is 65 % geraniol (a
gene producing this compound is silenced, both monoterpene) and 20 % geranyl acetate (Prashar
planthoppers and their parasitoids are reduced, et al. 2003; Rodrı́guez Quintanilla et al. 2012),
presumably because without the signal the plant although the relative proportions of these
becomes effectively invisible (Xiao et al. 2012). changes during development; palmarosa oil inhi-
Rice also produces S-linalool when attacked by bits growth of yeast (Saccharomyces cerevisiae)
brown planthoppers; in transgenic plants that do (Prashar et al. 2003). C. nardus produces mostly
not produce the compound, herbivory increased citronellal, geraniol and citronelol, and C. flexuo-
(Xiao et al. 2012). Melinis minutiflora (molasses sus produces geranial and neral (Rodrı́guez Quin-
grass) releases e-ocimene, 3–4,8-dimethyl-1,3,7- tanilla et al. 2012). Oil from C. validus (giant
82 Phytochemistry
turpentine grass) has as its major constituents 2004). The fungus is thus transmitted maternally
myrcene, (E)- b-ocimene, geraniol, linlol, and through the seed. When the fungus enters its
camphene (Chagonda et al. 2000). sexual phase, it invades inflorescence tissue and
Another member of Andropogoneae, Elio- produces conidia; the inflorescence aborts, a con-
nurus elegans, produces a variety of sesquiter- dition known as “choke”.
penes, the major one being campherenone, Fungal infection is beneficial to the plant in
which constituted more than a third of the essen- that it reduces herbivory by insects (Schardl et al.
tial oils extracted from the roots and shoots 2004). Infected plants are more productive than
(Mevy et al. 2002). E. muticus produces primarily their uninfected neighbors. For example, lolines
geranial, neral, and geranyl acetate (Chagonda (saturated pyrolizidines) produced by fungi
et al. 2000). infecting Lolium cuneatum (darnel) and L. arun-
dinaceum (tall fescue) are apparently not toxic to
mammals, but are as effective as nicotine in
CHEMICALS PRODUCED BY SYMBIONTS OR PATHOGENS deterring insect feeding (Schardl et al. 2007).
Species of Clavicipitaceae that infect plants
In some grasses, toxins are produced by fungi or are derived from ancestors that were pathogens
bacteria living on the plants. While some of these in animals (Spatafora et al. 2007). Thus, the abil-
are well studied, others are only referenced briefly ity to produce pathogenic alkaloids is ancestral in
in the literature and thus their prevalence in nat- the fungi, and the grasses have been able to take
ural ecosystems is unknown. advantage of this capacity in the establishment of
Fungi in the family Clavicipitaceae (Hypo- the symbiosis. The shift to plants appears to have
creales, Ascomycota) are endophytes of grasses occurred only once. Epichloë species have been
(White, Jr. et al. 2003). The most famous of these found in many Pooideae, including the early
is ergot (Claviceps purpurea), which infects rye; diverging Brachyelytrum. Although there is a
alkaloids from the fungus affect the mammalian general pattern of co-speciation of hosts and
nervous system, and in humans cause convul- fungi, some host switching is observed (Schardl
sions and seizures, as well as inducing psychoac- et al. 1997).
tive effects. Other well-studied members of the A number of grasses in Poeae in New Zeal-
family occur in the genus Epichloë (and its ana- and, along with Danthonia caespitosa, can be
morph Neotyphodium) (White, Jr. et al. 2003). infected with the bacterium Clavibacter toxicus
These infect many pooid grasses, including (¼ Corynebacterium spp.), which produces cor-
Lolium cuneatum (darnel), the “tares” of the gos- ynetoxins (tunicaminyluracil) (Edgar 1994). The
pel of Matthew, which is a weed of wheat fields. bacterium is spread by a nematode in the genus
Other Epichloë infections are responsible for Anguina. Livestock that ingest infected leaves
“sleepygrass”, an infestation of Achnatherum suffer neurological damage or death. While this
robustum of western North America, which sort of toxin production has been investigated
causes grazing horses to go to sleep for two or only in an agricultural context, it suggests a pos-
three days, and drunken horse grass (Ach- sibility of complex interactions of vector, patho-
natherum inebrians; Asian); the alkaloid respon- gen, and host ecology.
sible has been identified as lysergic acid amide.
Indolediterpenes are also produced and are
another cause of livestock poisoning, as found PHYTOSIDEROPHORES
in dronkgras (Melica decumbens; South Africa)
and huecú (Poa huecu; Argentina). Other hosts of A number of grasses secrete chelating agents,
clavicipatalean fungi include Echinopogon, and phytosiderophores, which enhance iron uptake
species of Festuca and Poa. The alkaloids have in response to low iron concentrations in the
also been widely used medicinally. soil (Kobayashi and Nishizawa 2012; R€omheld
Fungal hyphae extend throughout the above and Marschner 1986). The phytosiderophore is
ground parts of the plant and infect the develop- secreted from the roots into the soil, where it
ing grain, where they form a thin layer outside the binds Fe3+ in a 1:1 ratio. The complex is then
aleurone and inside the seed coat (Schardl et al. taken up into the roots by a transporter (Murata
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Distribution, Habitats and Conservation
DISTRIBUTION AND MAINTENANCE OF GRASSLANDS ever, the conditions of temperature and precipi-
tation that support grasslands also support
Grasses are found on all continents, including forests; thus, these two axes are insufficient for
Antarctica (Convey 2001), and exhibit remark- determining where grasslands will occur (Bond
able ecological diversity. Species occur in deep 2008; Lehmann et al. 2011). Dynamic global vege-
shade of tropical forests, in full sun in deserts, tation models, which attempt to predict vegeta-
from near the poles to the equator, and from sea tion based on climate and soil characteristics,
level to high altitudes. also fail to predict the extent of grasslands;
Not only are grasses widespread, but they are instead such models suggest that forest should
often ecologically dominant, giving their name to be far more common than it is (Bond et al.
a major ecosystem type, grasslands. The prairies 2005). While these environmental parameters
of North America and the vast grasslands of focus on the water use efficiency of grasses, such
Africa, the annual grasslands of the Mediterra- conditions also create an opportunity for fire.
nean and California, the steppes of Russia, the Bond et al. (2005) argue that grasslands in many
pampas of Argentina, and the great bamboo for- parts of the world are maintained by fire; they cite
ests of South America and Asia all attest to the multiple fire-exclusion studies in which grassland
importance of this one family of plants. gradually shifts to forest or woodland. Lehmann
The definition of grassland varies among et al. (2011) also find that rainfall seasonality is
authors (Gibson 2009). In general, grassland is a important in maintaining savannas.
biome with predominantly herbaceous vegeta- Herbivory also affects the balance between
tion, in which grasses are ecological dominants, grasses and woody vegetation, but its effects are
and in which trees and shrubs are not common. context-dependent (Bond 2008). Whether grazing
Some definitions (e.g., that of White et al. 2000) favors trees or grasses depends primarily on the
include areas such as tundra and shrub lands. It is forage preferences of the herbivores. Herbivory
common to include ecological forces in the defi- also interacts with fire. For example, the extent of
nition, such that a grassland is “maintained by tree cover in the Serengeti is primarily regulated
fire, grazing, drought and/or freezing tempera- by fire (Holdo et al. 2009). After eradication of
tures” (White et al. 2000). Because of the variety rinderpest, a virus that caused high mortality in
of definitions of what constitutes grassland, esti- wildebeest, the wildebeest population expanded;
mates vary as to the percent of the Earth’s land this intensified grazing pressure and reduced
surface covered by grasses, but generally range grass cover, leading to reduction in available
between 30 and 40 % (Gibson 2009; White et al. fuel. Fire frequency dropped and tree cover
2000). increased (Holdo et al. 2009).
Grasslands (excluding bamboo forests) are The grasses became ecologically dominant
one of the major biomes of the world, and were during the Miocene, with the forest to grassland
defined by Whitaker (1975) as areas of relatively transition occurring at different times on differ-
low precipitation and high temperature. How- ent continents (Edwards et al. 2010; Str€omberg
88 Distribution, Habitats and Conservation
2011). The exact drivers of this transition are not efficiency C4 photosynthesis and preference for
clear, but likely included a combination of warm temperatures.
increasing aridity and stronger seasonality, While the grasses ancestrally occupied warm
which in turn led to altered fire regimes and moist habitats, there have been two independent
increased herbivory; the latter two then may radiations into cool temperate zones, the Dantho-
have led to transition to grassland. In all cases, nioideae and the Pooideae (Edwards and Smith
the transition to C4 from C3 grassland came later, 2010). The Pooideae not only tolerate cool win-
roughly between 8 and 3 million years ago ters, but many have a specific requirement for
(Edwards et al. 2010). The causes of the shift to vernalization to trigger flowering (Preston and
C4-dominated ecosystems are poorly understood; Kellogg 2008). The ability of Pooideae to tolerate
atmospheric CO2 concentrations have been low cold correlates with an increased number of genes
enough to favor C4 photosynthesis since the Oli- whose expression is modified in response to low
gocene, ca. 30 million years ago (mya), and there temperatures (Sandve and Fjellheim 2010; Sandve
was no noticeable shift at 8 mya (Pagani et al. et al. 2008) and that show evidence of positive
1999, 2005); therefore CO2 concentrations cannot (adaptive) selection (Vigeland et al. 2013). Pooi-
explain C4 grassland expansion. Changes in pre- deae are the major members of Mediterranean,
cipitation are also insufficient to explain C4 grass- Californian, Russian, Argentine, and New Zeal-
land expansion. Models that introduce fire as a and grasslands. Danthonioideae have developed
major parameter fare better in modeling current cold tolerance independently of the Pooideae.
grassland distributions (Scheiter et al. 2012). While an ability to tolerate cold apparently coin-
The rise of grasslands also affected global cided with the origin of the subfamily, major
cycling of silica (Kidder and Gierlowski-Kordesch clades within it (Chionochloa, Danthonia, and
2005). The dry weight of C3 grasses can be 1–3 % Rytidosperma) have developed even greater toler-
silica, a number that may be higher for C4 species ance for lower temperatures (Humphreys and
(6–7 %) and for those in wetlands (10–15 %). As Linder 2013). The mechanisms of cold tolerance
grasses die or are eaten, the silica is recycled in this group are unknown.
within the grassland, or is released into runoff Studies are just beginning to tease apart abi-
and transported to lakes and oceans, where it is otic and biotic environmental correlates of spe-
available for incorporation into diatoms. Geolog- cies diversity in particular clades. In South Africa,
ical evidence for accumulation of diatomite the C4 subfamilies Aristidoideae, Chloridoideae,
apparently coincides with the earliest expansion and Panicoideae sort out differently along gradi-
of the grasslands in the Miocene (Kidder and ents of moisture, temperature, fire frequency, and
Gierlowski-Kordesch 2005). Herbivory and grazing intensity (Visser et al. 2012). Aristidoi-
subsequent passage through a mammalian or deae are most diverse in hot, dry, disturbed or
insect gut appears to increase the solubility of grazed areas, whereas Chloridoideae reach their
silica and thus enhance silica cycling through highest diversity in areas that are hot but wet,
the ecosystem, providing a potential link between with high grazing pressure but infrequent fire.
grassland formation, the rise of large mammalian Panicoideae are most diverse in areas that are
herbivores, and accumulation of diatoms in warm and mesic, but more species of Paniceae
aquatic environments (Vandevenne et al. 2013). occur with high grazing intensity whereas Andro-
pogoneae are the opposite. Andropogoneae are
also most diverse in areas with high fire fre-
DISTRIBUTION OF MAJOR CLADES quency. It will be of considerable interest to
extend this study to other parts of the world.
Individual clades of grasses have distinct ecologi-
cal preferences, and it is possible that the syna-
pomorphies of many groups are physiological SPECIES DIVERSITY
rather than simply morphological. For example,
the North American and African grasslands are Grasslands often harbor high numbers of species,
largely covered with members of the subfamilies and are commonly listed among the world’s
Panicoideae and Chloridoideae, with their high diversity hotspots. For example, the IUCN lists
CONSERVATION AND RESPONSE TO CLIMATE CHANGE 89
234 Centres of Plant Diversity of which 40 are graphic pattern, but also may be influenced by
grasslands, and the World Wildlife Fund taxonomic artifacts. For example, many mono-
(WWF)-US Global 200 Programme identifies 136 typic or oligotypic segregate genera have been
terrestrial ecoregions as having outstanding described in Africa by Stapf and Nees. This
diversity of which 35 are grasslands (Gibson could indicate remarkable diversification on the
2009). Among 25 biodiversity hotspots identified African continent, or could represent the work of
by Myers and Mittermeier (2000) is the Cape a couple of enthusiastic splitters.
Floristic Province in South Africa, an extraordi-
narily species-rich grassland, with over 1700 spe-
cies of plants. CONSERVATION AND RESPONSE TO CLIMATE CHANGE
Despite their high numbers of species, most
grasslands are characterized by a handful of Grasslands typically have excellent soil and thus
dominants (Coupland 1992, 1993); in other are attractive sites for agriculture. They are also
words, most grasses in most ecosystems are rela- easily converted to other human-dominated land-
tively uncommon and fewer than 10 % of grass scapes such as housing developments and shop-
species make up most of the biomass in major ping malls. As such they are surprisingly
grasslands. For example, in the North American endangered in many parts of the world. Of the
tallgrass prairie, the dominant species are Andro- original tall-grass prairies in North America, less
pogon gerardii, Schizachyrium scoparium, Sor- than 10 % of the original acreage remains (Gibson
ghastrum nutans, and Panicum virgatum. All 2009). While many of the grasslands in the British
four happen to be C4 taxa, but the less common Isles are themselves the result of forest removal by
species are also C4. Thus, the photosynthetic humans, the native species are often replaced by
pathway may be only one aspect of their domi- introduced pasture grasses, which provide better
nance. Grassland dominants are significantly forage for cattle (Gibson 2009). Despite their
clustered phylogenetically (Edwards et al. 2010), ongoing conversion to other uses, only a small
indicating that some clades have characteristics fraction of world grasslands is protected, making
that confer a selective advantage. At the same them a highly endangered ecosystem. Hoekstra
time the characters that make a grass dominant et al. (2005) estimate the percent of the world’s
are not shared by all members of the family. biomes converted to other uses and divide this by
Grazing is known to affect species diversity in the percent of the biome protected to develop a
grasslands, but whether the effect is positive, Conservation Risk Index. Temperate grasslands,
negative or neutral depends on the size of the savannas and shrublands had the highest risk
herbivore and the productivity of the particular index of any of the world’s biomes, in part because
plant community. Moderate grazing by large her- the percentage of protected habitat was so low.
bivores in productive communities increases Grasslands are important buffers against
diversity, at least in part by reducing the cover increasing global CO2. Worldwide, grasslands
of dominant species; in communities with lower store almost as much carbon as forests (34 vs.
productivity, however, grazing generally reduces 39 %) (White et al. 2000). Because of the deep
diversity (Bakker et al. 2006; Gibson 2009). Not roots of many perennial grasses, much of this
surprisingly, over-grazing reduces the number of storage is in the soil, rather than above ground
species and can lead to ecosystem change or even as in forests.
loss of plant cover entirely. The field of range Given the complex and poorly understood set
management focuses on techniques that maxi- of factors maintaining grasslands, the future of
mize forage quality and diversity on a sustainable grasslands in the face of climate change is hard to
basis. Nonetheless, as population pressure predict (Bond and Midgley 2012). In many parts
increases in many parts of the world, the tempta- of the world, grasslands are being taken over by
tion to put too many grazing animals on a piece woody vegetation, a pattern that may be caused
of land will often win out. by changes in land use or increased atmospheric
Clayton and Renvoize (1986) note that grass CO2 or a combination of the two (Bond 2008).
genera rarely occur on more than one particular Changes in the amount and timing of precipita-
continent. This may point to a broad biogeo- tion also affect the presence and structure of
90 Distribution, Habitats and Conservation
grasslands. A particularly detailed description of require elimination of weeds of the same family
the famous North American drought of the 1930s (Young and Allen 1997). Many other grasses (e.g.,
is provided by Gibson (2009), summarizing the Imperata cylindrica, Sorghum halepense) are nox-
work of Weaver (1954); the decade-long series of ious weeds that negatively affect agriculture
droughts altered the structure and dominance throughout the world.
relationships of grasses in the prairies. In contrast
to the studies of Bond (2008) and Bond and
Midgley (2012), based on modeling in warm
areas of the world, and Weaver’s (1954) observa-
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Reproductive Systems
Reproductive systems in the grasses are diverse. Lecomtella, Ixophorus), between spikelets within
This section begins by describing the morpholog- an inflorescence (e.g., Pharus, Hypogynium, Ziza-
ical variation in placement of the stamens and nia), or between inflorescences within a plant (e.
pistils, and then follows with discussion of out- g., Zea, many Olyreae).
crossing, self-pollination and apomixis. Andromonoecy is common, particularly in
the Panicoideae, in which spikelets are conven-
tionally two-flowered, with the lower flower often
UNISEXUAL VS. BISEXUAL FLOWERS producing stamens and the upper one being
bisexual. Additional variations on this theme
Grasses may have unisexual or bisexual flowers. occur in the tribe Andropogoneae, with sex
Because of the distribution of the unisexual ver- expression varying between the flowers in each
sus bisexual condition in the immediate out- of the two spikelets of the spikelet pair. For exam-
groups and early diverging taxa, attempts to ple, in many species of Andropogon, the sessile
determine the ancestral condition for the family spikelet has a sterile lower flower and bisexual
are highly sensitive to analytical method and upper flower, while the pedicellate spikelet has a
assumptions (Malcomber and Kellogg 2006). sterile lower flower and staminate upper one; this
Among the graminid Poales, flowers of Flagellar- produces an inflorescence of bisexual and stami-
iaceae are bisexual, whereas those of Restiona- nate flowers in a 1:1 ratio. In other Andropogo-
ceae, Centrolepidaceae and Anarthriaceae are neae (e.g., Elymandra), the most proximal
unisexual. The two immediate outgroups of the spikelet pairs on the inflorescence consist entirely
grasses, Ecdeiocoleaceae and Joinvilleaceae, have of staminate flowers, thus increasing the ratio of
unisexual and bisexual flowers, respectively. In staminate to bisexual flowers.
the grasses, flowers in Anomochlooideae are In contrast to andromonoecy, gynomonoecy
bisexual, but those in Pharoideae are unisexual, is rare in the grasses as a whole. Connor (1981)
and Puelioideae are bisexual. This distribution of reports only eight genera with this distribution of
character states indicates either that unisexual flower types. In two of these, the bambusoids
flowers have arisen repeatedly, or that reversion Diandrolyra and Piresia, the apparently bisexual
from unisexual to bisexual is fairly easy over flowers are in fact staminate with the gynoecium
evolutionary time, or both. non-functional; thus, the plants are actually mon-
Virtually all possible arrangements of unisex- oecious. In the chloridoid genus Munroa, pistil-
ual flowers on the plant are known in the grasses late flowers are proximal to bisexual ones in the
(Connor 1981, 1987). Monoecy is widespread, spikelets of most species, but in M. squarrosa,
appearing in all Pharoideae, tribe Olyreae of pistillate spikelets occur only on lateral inflores-
Bambusoideae, many members of Chloridoideae cence branches, whereas the terminal spikelet is
and Panicoideae, and some Ehrhartoideae. bisexual (Anton and Hunziker 1978); spikelets of
Among monoecious grasses, sex expression may M. argentina are all bisexual. In Centotheca, Coe-
vary between flowers within a spikelet (e.g., lachne and Heteranthecia, the distal flower of the
94 Reproductive Systems
spikelet is often pistillate whereas proximal flow- tion in the jasmonic acid pathway (Yan et al.
ers are bisexual. 2012). However, Malcomber and Kellogg (2006)
Dioecy is not particularly common in the find that TS2 is expressed throughout the plant
grasses. Although Connor (1987) and Connor including anthers, and is highly conserved in all
et al. (2000) list 22 and 20 dioecious genera grasses, suggesting that it may have been co-
respectively, several are monotypic genera that opted for sex determination only in the pani-
are now placed in synonymy in Distichlis, a coids. In addition, variation in the sequence of
genus in which all species are dioecious, and in the protein does not correlate with the presence
Bouteloua, a well supported clade that includes of unisexual flowers. Thus, its major role is
species with almost every conceivable form of unknown but certainly is more general than sim-
dicliny (Kinney et al. 2007). The two dioecious ply to cause organ abortion. Development of sta-
species formerly segregated as Neeragostis are minate flowers in Bouteloua (Chloridoideae) is
now placed in Eragrostis, in which they are the histologically similar to that in Panicoideae, and
only dioecious members (Ingram and Doyle gynoecial abortion also requires TS2 (Chandra
2004). Following current taxonomy, dioecy thus and Huff 2010); interestingly, infection of stami-
occurs in 14 genera in three subfamilies: Chlor- nate flowers with the fungus Salmacisia (¼ Tille-
idoideae (Allolepis, Bouteloua, Cyclostachya, Dis- tia) buchloëana suppresses TS2 expression and
tichlis, Eragrostis, Jouvea, Scleropogon, Sohnsia), causes pistils to develop. In contrast, develop-
Panicoideae (Gynerium, Pseudochaetochloa, Spi- ment of staminate flowers in Zizania (Ehrhartoi-
nifex, Zygochloa), and Pooideae (Poa, the “Leu- deae) is distinct from the panicoid or chloridoid
copoa grade”). pattern. In Zizania, the stigmatic arms of the
While dioecy is uncommon, gynodioecy gynoecium develop, the ovule differentiates and
(plants either entirely pistillate or with bisexual integuments become visible, and only then does
flowers) is even more rare. In Bouteloua and Poa, growth arrest, accompanied by deposition of a
both dioecious and gynodioecious species are dark-staining substance (Zaitchik et al. 2000).
known. Gynodioecy also appears in Austroderia, Sex expression is also affected by the hor-
Chimaerochloa, and Cortaderia in Danthonio- mone gibberellin. In maize, the gene known as
deae. In Poa and the three danthonioid genera, anther ear1 encodes ent-kaurene synthase, in the
some species seem to be entirely pistillate and gibberellin pathway; when it is mutated the pistil-
apomictic, with staminate plants unknown. late flowers of the ear become bisexual and at the
Plants that appear to be gynodioecious may in same time the number of branches in the tassel is
fact be functionally dioecious; morphologically reduced (Bensen et al. 1995). Bouteloua dacty-
bisexual plants in Cortaderia selloana in Argen- loides (buffalograss) is normally dioecious, but
tina serve primarily as pollen donors, so are func- some plants are trimonoecious, producing stami-
tionally staminate despite having gynoecia nate, pistillate and bisexual flowers. In such
(Connor et al. 2000). Male sterility in Cortaderia plants, sex expression can be manipulated by
is under the control of three loci, but these have applying gibberellin, in which case inflorescences
not been investigated at the molecular level (Con- become entirely staminate (Quinn 2000). Inhibit-
nor and Charlesworth 1989). ing gibberellin with paclobutrazol creates inflor-
Unisexual flowers in the grasses always initi- escences that are entirely pistillate.
ate both gynoecium and androecium; one or the
other of these then aborts during development
(Le Roux and Kellogg 1999). In staminate flowers SELF-POLLINATION AND SELF-INCOMPATIBILITY SYSTEMS
of all panicoids investigated to date, the gynoe-
cium develops until the nucellus is visible and It is surprising, given the ecological dominance of
carpel walls form a ridge surrounding it; at this the grasses, how few studies have been under-
stage cell death occurs in the subepidermal cells taken of their breeding systems. Fewer than 100
and growth ceases. Gynoecial abortion in pani- genera have been characterized as self-compati-
coids is under the control of the proteins TAS- ble or self-incompatible (Campbell et al. 1983;
SELSEED1 (Acosta et al. 2009) and Connor 1979), so self-compatibility is unknown
TASSELSEED2 (DeLong et al. 1993), which func- for the majority of the family. Self-compatibility
CLEISTOGAMY 95
is generally more common among annuals than the Z locus are actually complex loci with at least
perennials; because most cereal crops are domes- two genes (Baumann et al. 2000). In addition, a
ticated annuals, it is not surprising that many of third locus known as T (for third) has also been
these are also self compatible (e.g., barley, wheat, identified recently in three pooid grasses
foxtail millet) (Glémin and Bataillon 2009). (reviewed in Yang et al. 2008). Comparative
Of the grasses that have been studied, many genome mapping is likely ultimately to identify
are capable of both self-pollination and outcross- the specific loci corresponding to S and Z; unfor-
ing. In the few such taxa that have been studied in tunately the pooids in which the genetic work has
detail, however, outcross pollen leads to higher been done have some of the largest genomes in
seed set and germinability than self pollen (Fang the family so genomic approaches will be rela-
et al. 2004; Kettenring et al. 2011). Even though tively slow.
barriers to self-pollination are rarely absolute, Pollen tube growth has been described above
estimated rates of outcrossing are strongly under Pollination (section Pollen). The self-
bimodal, with self-incompatible species generally incompatibility response appears to require an
above 80 % outcrossing and self-compatible ones interaction between the tip of the pollen tube
mostly well under 15 % (Connor 1987; Raduski and the stigma, so occurs after pollen germina-
et al. 2012). tion (Heslop-Harrison and Heslop-Harrison
The genetics of self-incompatibility have 1987). Barriers to self-pollination may occur
been worked out in most detail in Secale cereale quite rapidly (within 90 seconds in Secale cereale;
and seven other Pooideae. Self-incompatibility in Heslop-Harrison 1982) or much later after the
these species is controlled by the pollen (gameto- pollen tube has grown into the style – e.g., in
phytic) alleles of two loci, called S and Z (Hayman Alopecurus pratensis (Shivanna et al. (1982) and
1956; Heslop-Harrison 1982; Lundqvist 1954). Andropogon species (Norrmann and Keeler
Pollen growth will be blocked if the S and Z alleles 2003).
of the pollen both match S and Z alleles in the Many grass species are protogynous, which
stigma. This system has two interesting effects; presumably reduces self-pollination, but as with
first, partial compatibility is possible, and second, self-compatibility, the distribution of this charac-
reciprocal crosses may have different outcomes. ter is poorly documented.
For example, if a plant with the genotype S1S1
Z1Z3 is used as the seed parent and a plant with
S1S2 Z2Z3 is the pollen donor, the possible pollen CLEISTOGAMY
genotypes will be S1Z2, S1Z3, S2Z2, and S2Z3. Of
these, three will be compatible and one incompat- More cleistogamous species are reported for the
ible, resulting in 75 % of the pollen being com- grasses than for any other plant family (Campbell
patible. The reciprocal cross will be 50 % et al. 1983; Culley and Klooster 2007). Cleistog-
compatible. Unlike single locus systems that amy, defined as “self-fertilization in an enclosed
occur in other families, the S-Z system will func- flower”, is widespread in the family, occurring in
tion even if gametes are unreduced and the plant over 300 species and 90 genera distributed among
becomes autopolyploid. In addition to the eight most subfamilies (Campbell et al. 1983; Connor
species in which the genetics are known, another 1987; Culley and Klooster 2007). In many taxa,
eight have a gametophytic self-incompability sys- cleistogamy is caused by failure of the inflores-
tem in which reciprocal crosses have different cence to emerge from the sheath; flower develop-
outcomes; it seems likely that these also have an ment is otherwise normal but self-fertilization
S-Z system (Baumann et al. 2000). occurs because the anthers and stigmas are
Even though the basic genetics of the S-Z never exposed. In some cases, retention of the
system was determined more than half a century inflorescence in the sheath is caused by environ-
ago, neither gene has been cloned and their mental factors (e.g., low temperatures, low soil
molecular identity and mode of action remain moisture), and this is particularly true for the
unknown. Loss of self-incompatibility can occur failure of the terminal inflorescence to emerge.
by mutation of alleles that occur only in the However, failure of axillary inflorescences to
pollen, suggesting that perhaps both the S and emerge is often genetically fixed within a species.
In this case, the enclosed axillary inflorescence the style. The mechanism behind this is
may be morphologically distinct from the unknown. In a series of papers, Evans and Ker-
exserted inflorescences elsewhere on the plant, micle (Evans and Kermicle 2001; Kermicle 2006;
and is known as a cleistogene (Chase 1918). Kermicle and Evans 2005, 2010; Kermicle et al.
Two genera – Amphicarpum and Eremitis – 2006) have identified three genetic loci, Gameto-
produce spikelets underground from positively phyte1 (Ga1), Gametophyte2 (Ga2), and teosinte
geotropic shoots, as well as above ground, from crossing barrier1 (tcb1), which they call pollen-
open panicles (Campbell et al. 1983). In Amphi- pistil cross-compatibility (PPCC) loci. Each locus
carpum the seeds of subterranean spikelets are has three alleles that are found in wild popula-
borne more than 3 cm below the surface (Che- tions of Zea mays (teosinte). One allele of tcb1
plick and Quinn 1988), allowing them to survive and all three alleles of Ga1 and Ga2 are also found
the frequent fires of the New Jersey pine barrens in cultivated maize. One allele creates a block in
where the species is native. Two species of Chloris the pistil but pollen growth is competent, the
and one of Paspalum are also reported to have second allows pollen growth but does not confer
subterranean spikelets. a corresponding block in the pistil, and the third
Cleistogamous flowers have much smaller has neither effect. Incompatibility at any of the
anthers than chasmogamous ones, resulting in three loci will lead to markedly reduced success of
lower pollen to ovule ratios (Campbell et al. pollination. Together the three loci reduce cross-
1983). In plants producing cleistogenes or subter- pollination between maize and teosinte in fields
ranean spikelets, fruits of the cleistogamous spi- where the two subspecies co-occur. These loci are
kelets are heavier than those of chasmogamous thus “speciation loci”, providing an early barrier
spikelets, although their relative success in ger- to gene flow between two partially differentiated
mination and seedling establishment varies taxa. This genetic block to hybridization must
between species (Connor 1987). have arisen quite rapidly since maize was domes-
ticated only about 9,000 years ago or less.
Wheat (Triticum aestivum) can be crossed
BARRIERS TO INTERSPECIFIC CROSSING with rye (Secale cereale), but crossability between
these two species depends on the genotype of the
Crosses between species of grasses are often but seed parent (wheat) at a locus known as Kr1;
not always unsuccessful. Knobloch (1968) pub- because wheat is a hexaploid, there are three Kr
lished a useful compendium of crosses that have loci, on chromosomes 5A, 5B, and 5D (Manick-
been attempted in the grasses. In most cases the avelu et al. 2009). A dominant Kr allele at any
offspring were sterile, but exceptions occurred, locus blocks the growth of rye pollen tubes at the
particularly among polyploid species. For exam- base of the style or in the ovary wall, whereas
ple, Norrmann and Keeler (2003) report produc- recessive alleles at Kr have no effect (Lange and
tion of viable hybrids between Andropogon Wojciechowska 1976). Recessive kr alleles occur
gerardii and A. lateralis, A. gerardii and A. glau- naturally in T. aestivum, suggesting that wide
cophyllus, and A. hypogynus and A. lateralis. The crosses could occur from time to time in nature.
parents of these hybrids differ in chromosome The possibility of wide hybridization has also
number, but are all high polyploid (2n ¼ 60 and been investigated in sorghum. Maize pollen will
above). In general, whether crosses – even very germinate on the stigma of most accessions of
wide crosses – are successful or not depends on Sorghum bicolor, but fails to grow more than a
the particular combination of genomes in the few hundred microns (Laurie and Bennett 1989).
parents and not necessarily on their phylogenetic However, in a landrace known as S. nervosum
proximity (Jones and Pasakinskiene 2005). Nr481, maize pollen will germinate and occasion-
As noted under Pollination (section Pollen), ally will reach the ovule. The ability to permit or
pre-zygotic isolating mechanisms exist in the block maize pollen tube growth is controlled by a
grasses but are not universal and where they single locus known as Iap (Inhibition of alien
exist may still be somewhat porous. The stigma pollen tubes). When the seed parent is homozy-
is often not a barrier to pollen germination, but gous recessive at this locus, maize pollen tube
foreign pollen may be blocked in passage down growth is permitted (Laurie and Bennett 1989).
ASEXUAL REPRODUCTION 97
Price et al. (2006) extended this result by testing duce vegetatively via rhizomes or stolons (see
pollen from Sorghum nitidum, S. macrospermum, discussion on Vegetative Morphology and Anat-
and Sarga angusta (¼ Sorghum angustum) on S. omy). Rhizomatous spread may be relatively
bicolor stigmas. In S. bicolor plants with the dom- slow, but the tillers produced may be quite long-
inant Iap allele, foreign pollen germinated, but lived. Harberd (1961) famously estimated that a
was then blocked at various points in the stigmas colony of Festuca rubra might have been over 400
or style branches such that no fertilization years old, and perhaps as old as 1000 years. Simi-
occurred. In plants that were recessive at Iap, lar dates have been estimated for other clonal
however, a few pollen tubes reached the ovary, grasses (de Witte and St€ocklin 2010). A more
although they grew more slowly than pollen from recent study with molecular markers suggested a
S. bicolor. Some viable embryos were obtained minimum age for a plant of Festuca idahoensis as
from all three interspecific crosses, and S. bicolor at least 60 years (Liston et al. 2003).
x S. macrospermum yielded germinable seed. A few species – notably Poa alpina, P. bul-
Thus, pre-zygotic isolation between species is bosa, Deschampsia alpina, and several species of
not absolute. Festuca – produce onion-like bulbils in place of
Post-zygotic blocks to hybridization may spikelets (Gustafsson 1947). This phenomenon is
occur by disruption of either endosperm or sometimes called pseudovivipary to distinguish it
embryo sac development. Crosses between Sor- from vivipary, a term that should be reserved for
ghum bicolor and S. angustum or S. nitidum pro- germination of the seed before being shed from
duced viable embryos that had to be rescued the parent (see discussion under the section Fruit
because of endosperm failure (Price et al. 2006). and seed). Formation of bulbils is often under
In a series of interspecific crosses between Oryza environmental control (Heide 1989; Youngner
sativa and other diploid species of Oryza, polli- 1960), and is common in cool, wet environments
nation and fertilization were normal (Ishikawa (Lee and Harmer 1980) although P. bulbosa has
et al. 2011). However, subsequent development become a widespread weed even in dry areas. In
of the endosperm was disrupted, and the timing P. alpina, plants that produced bulbils are more
of cellularization was abnormal. Seeds were unvi- common at high elevations and have reduced
able not because of problems with embryo devel- fitness if transplanted to lower sites (Steiner
opment but because of the endosperm. Ishikawa et al. 2012). The bulbils may be dormant (as in
et al. (2011) suggest that the failure of endosperm P. bulbosa) or may germinate immediately (as in
development may be caused by inappropriate F. viviparoidea) (Lee and Harmer 1980).
imprinting of parental genes. Many grasses are capable of producing asex-
Other post-zygotic mechanisms affect the ual seed (apomixis or agamospermy). Although
embryo sac. Recently, Yang et al. (2012) have some authors (e.g., Gustafsson 1947; Richards
identified a set of three linked genes in rice that 2003) use the term apomixis to refer to all
confer intersterility between indica and japonica modes of asexual reproduction, it is more com-
lines. These function as a set of Dobzhansky- mon to use apomixis to refer only to asexual seed
Muller interactors, such that particular combina- production (Ozias-Akins 2006; van Dijk 2009), a
tions of alleles cause abortion of the embryo sac, terminology followed here.
apparently by affecting the endoplasmic reticu- Asexual seed production in the grasses can
lum. As with the incompatibility genes described occur via several mechanisms, all of which can be
above, these confer incompatibility only in par- viewed as heterochronic changes in the normal
ticular combinations, whereas in other combina- reproductive process, such that the normal devel-
tions of alleles embryo sac formation and opmental program is triggered at an abnormal
ultimately seed production proceed normally. time or in inappropriate cells (Grimanelli 2012;
Nygren 1967). In sporophytic apomixis (adventi-
tious embryony), a nucellar (maternal) cell devel-
ASEXUAL REPRODUCTION ops directly into an embryo, bypassing
gametophyte production entirely. In gameto-
Asexual reproduction is common in the grasses. phytic apomixis (apospory or diplospory), devel-
Many species, particularly perennials, can repro- opment of the female gametophyte is modified.
In aposporous apomixis, an unreduced embryo maternal to one paternal. Some grasses (e.g.,
sac forms directly from cells in the nucellus, such Tripsacum) appear to tolerate deviation from
that each cell is diploid rather than haploid. The the 2:1 ratio. Others, such as aposporous Panicoi-
unreduced egg then proceeds to form an embryo deae, achieve the ratio by production of a four-
directly, without fertilization. Meanwhile, the celled embryo sac with a single unreduced central
megaspore mother cell undergoes meiosis as cell. This has twice the chromosome complement
usual, and begins normal embryo sac formation. of the reduced sperm that fertilizes it, resulting in
However, the normal megagametophyte generally a 2:1 ratio of maternal to paternal genomes
breaks down before maturing. Aposporous (Ozias-Akins 2006).
embryo sacs tend to form earlier than the normal Pseudogamy makes apomixis difficult to
one (e.g., Kellogg 1987); it is not clear if they detect because simple bagging experiments will
compete for resources or space, or if other pro- allow seed development, and exclusion of pollen
cesses lead to breakdown of the normal embryo will prevent seed set; such results do not distin-
sac. guish agamospermy from self-compatibility.
In diplosporous apomixis, the megaspore Demonstration of apomixis thus requires either
mother cell never undergoes meiosis at all, but observation of megagametophyte development,
instead divides and forms a single embryo sac by assessment of ploidy levels in tissues of develop-
mitosis. Alternatively, meiosis begins but after ing ovules, or analysis of variation among the
the first division the resulting nuclei fuse together offspring of a single plant (Ozias-Akins 2006;
to form a restitution nucleus. In either case, the Ozias-Akins and van Dijk 2007).
embryo sac is diploid and the egg develops with- Non-pseudogamous apomixis is much less
out fertilization. common than pseudogamous, and in grasses is
Apomixis is relatively common in the grasses, known only in species of Calamagrostis, Corta-
having been reported in as many as 60 non- deria (including Lamprothyrsus), Nardus and Poa
hybrid genera (Czapik 2000; Ozias-Akins 2006), (Connor 1981). It is unclear how endosperm
and has clearly arisen more than once in the development in these species is able to proceed
family (van Dijk and Vijverberg 2005). It is well without fertilization.
known and extensively studied in species of Poa Apomixis requires both formation of unre-
(Pooideae), in Cortaderia (Danthonioideae) and duced gametes and parthenogenesis, processes
in a number of panicoids including Bothriochloa, that are not always linked. Unreduced gametes
Tripsacum, Paspalum, Brachiaria, Urochloa max- occur frequently in grasses and may or may not
ima (¼ Megathyrsus), and Cenchrus (which now be correlated with subsequent parthenogenesis.
includes species formerly placed in Pennisetum). In addition, the tendency to produce unreduced
In the panicoid species, aposporous embryo sacs gametes is not necessarily shared among closely
are four-, rather than eight-celled, and lack anti- related species. For example, Norrmann and Kee-
podals. ler (2003) report evidence for an unreduced egg
It is not clear whether sporophytic apomixis cell from Andropogon lateralis, but found no evi-
occurs in grasses. Asker and Jerling (1992) indi- dence for unreduced gametes in over 1000 plants
cate only gametophytic apomixis, whereas Nau- of A. gerardii.
mova (1993) lists eight species with reported The genetic basis of apomixis is not well
adventitious embryony. Although Yu et al. understood, although significant progress has
(2000) report sporophytic apomixis in Bothrio- been made in recent years (Grimanelli 2012;
chloa and Dichanthium, they do not provide sup- Ozias-Akins and van Dijk 2007). Because apo-
porting evidence. Gametophytic apomixis is by mixis would permit fixation of particular hybrid
far more common in the grasses, with apospory combinations it is of considerable interest to
more common than diplospory (Nygren 1967). agriculture. Crosses of sexual and apomictic spe-
In pseudogamous apomixis, embryo forma- cies have shown that apomixis is often dominant
tion is parthenogenetic but fertilization is still (Harlan and de Wet 1963; Ozias-Akins 2006).
needed for endosperm development. Endosperm Genetic studies with molecular markers and
in turn often requires an appropriate ratio of genomic comparisons have been undertaken in
maternal to paternal genomes, generally two five panicoid genera (Cenchrus, Brachiaria, and
References 99
Urochloa (¼ Panicum maximum = Megathyrsus Apomixis is strongly associated with poly-

maximus) of tribe Paniceae; Paspalum in Paspa- ploidy. Because polyploidy itself is so common
leae; and Tripsacum in Andropogoneae), and one in Poaceae, it is not surprising that apomixis
pooid (Poa, tribe Poeae). All but Tripsacum are should also be common. Nonetheless, it is not
aposporous. Ozias-Akins et al. (2003) have shown certain that polyploidy is absolutely required.
that apomixis is associated with a large block of Some evidence from Paspalum and Hieracium
DNA that is maintained in hemizygous condition (Asteraceae) suggests that formation of apomictic
in polyploid Cenchrus squamulatus (¼ Pennise- embryo sacs can occur in diploids, but its fre-
tum squamulatum) and Cenchrus ciliaris. This quency becomes more common upon chromo-
large region does not recombine and contains some doubling (Ozias-Akins 2006).
regions of highly repetitive DNA. In addition, its
chromosomal location appears to vary over evo-
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1982. The pollen-stigma interaction in the grasses. 3. evolution. Melbourne: CSIRO. pp. 307–315
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Bot. Neerl. 31: 307–319. ment of male flowers in Zizania aquatica (North
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Fossil Record and Dates of Diversification
The fossil record of the grasses has been compre- that the family was widespread and diverse by
hensively reviewed by Thomasson (1987), Jacobs that time (Jacobs et al. 1999; Str€omberg 2011).
et al. (1999), and most recently by Str€omberg Many of these fossils are well preserved and can
(2011), and what follows is based heavily on be identified to modern genera or tribes on the
those papers. Available fossils consist of pollen, basis of overall morphology and epidermal char-
macrofossils (leaf fragments, spikelets, fruits), acters (e.g., Elias 1942; Thomasson 1985). The
and phytoliths (silica bodies). Miocene record clearly shows the expansion of
As noted above, pollen of all members of grassland habitats. The earliest grass fossil with
Poales is similar in being spherical and monopo- Kranz anatomy dates to the mid-Miocene, ca.
rate. In grasses and other graminid Poales, the 12.5 mya, in North America (Nambudiri et al.
pore is surrounded by an annulus. In the past, 1978; Whistler and Burbank 1992).
pollen with this general morphology was placed Earlier macrofossils, from the Eocene and
in the form taxon Monoporites annulatus and Oligocene, are less common and also harder to
often assigned to the grasses; however, the annu- assign taxonomically, as might be expected. A
lus is not synapomorphic and many specimens two-flowered grass spikelet has been found in
assigned to M. annulatus have been re-assigned Eocene deposits of North America, dated to
to other families (Linder and Ferguson 1985). The about 55 mya (Crepet and Feldman 1991).
diagnostic character for grasses is the lack of Although the authors assigned this to Arundinoi-
scrobiculi through the exine, a character that deae based on the broad concept of the subfamily
must be assessed with transmission electron prevalent at the time, the characters only allow
microscopy. M. annulatus from 57 million years assignment to the Bistigmatic (multi-flowered
ago (mya) appears in northern South America spikelet) clade; the spikelet does not match any
and is likely to be Poaceae, whereas earlier Maas- currently known taxon. In the early Eocene of
trichtian records (ca. 70 mya) may or may not western Europe, the London clay flora (56–49
represent grasses (Jacobs et al. 1999). Linder mya) is reported to contain grass leaves and
(1987) notes that the Maastrichtian grains “are inflorescences (Chandler 1964), but detailed
rather inadequately illustrated by drawings” in descriptions of these are not available. The next
van der Hammen (1954); more recent reports earliest macrofossil in North America is from
are also not fully diagnostic (Srivastava 2011). about 34 mya (upper Eocene) in the Florissant
Once grass pollen appears in the record, it pro- flora, assigned to Stipa or a close relative (Mac-
vides little further information on diversification Ginitie 1953; Manchester 2001). Early grass
of the grasses. Grass pollen is simply too uniform macrofossils are not known from South America
to assign to any particular major clade. (Jacobs et al. 1999; Str€omberg 2011), despite the
Grass macrofossils – particularly fruits, early pollen records, nor are there macrofossils
lemmas and glumes – dating from the Miocene reported from Africa before the Miocene. Macro-
(23–5.3 mya) have been found in North and fossils have not been found in Australia (Jacobs
South America, Europe, and Africa, showing et al. 1999; Str€omberg 2011).
104 Fossil Record and Dates of Diversification
Fig. 8. Grass phytoliths from the Eocene to Miocene of lobed phytolith, similar to those in Stipeae. E Crenate
North America. A Rondel, assigned to Bambusoideae, phytolith, similar to those in Pooideae. F Tall, keel-shaped
similar to Chusquea. B Two-lobed phytolith, similar to structure, unlike any known in modern grasses. (From
many in the PACMAD clade; compare to Fig. 2B. C Sad- omberg 2005; copyright 2005, National Academy of
Str€
dle-shaped, similar to modern Chloridoideae. D Two- Sciences, U.S.A.)
The lack of early macrofossils may indicate bracteate structure from the same deposit. It is
absence of grasses, but could also reflect poor interpreted as a spikelet, but lacks diagnostic
preservation. Pollen preserves poorly in semi- characters that would make this interpretation
dry environments where the soil pH is above definitive (Poinar, Jr. 2004, 2011). Because the
about 6.0 (Bryant, Jr. et al. 1994); this may affect Cretaceous date of these fossils makes them
the pollen record of dry grasslands in particular. almost as old as the oldest angiosperm, accepting
(Note, however, that the pollen preservation them as Pooideae would require a major revision
study was done in the American Southwest, of all dates of all land plants, and on that basis
which is drier than many of the world’s grass- their identity as grasses is suspect.
lands.) Likewise, macrofossils are rare in part The most widespread but arguably the hard-
because the open habitats of grasses are not con- est data to interpret come from phytoliths, or
ducive to preservation. fossilized silica bodies (Fig. 8). If these are inter-
A handful of Cretaceous macrofossils have preted as current literature suggests, they will
been suggested to represent grasses, although require revision not only of the history of the
the evidence for this assignment is not strong. grasses, but also the history of all angiosperms.
Programinis laminatus is a leaf fragment embed- Phytoliths vary widely in shape among the
ded in Burmese amber, and is thought to be early grasses and thus provide much more taxonomic
Cretaceous, upper Albian (97–110 mya) (Poinar, information than pollen grains do. They are far
Jr. 2004, 2011). The rounded structures that are better preserved in the fossil record and thus
visible on the leaves are interpreted as silica bod- provide more data than the sparse set of macro-
ies similar to those in the Pooideae, although this fossils. Silica bodies have been described for vir-
identification is far from certain. P. burmitis is a tually all genera of grasses, beginning in the first
Fossil Record and Dates of Diversification 105
part of the 20th century (Metcalfe 1960; Prat 1932; Grass fossils have been used to calibrate the
Watson and Dallwitz 1992 onward), but these grass phylogeny and estimate the timing of major
descriptions were based on the shape in surface diversification events (Bouchenak-Khelladi et al.
view (see also Description of the family, Vegeta- 2010, 2014; Christin et al. 2008, 2014; Jones et al.
tive morphology and anatomy: Leaves: Leaf epi- 2014; Vicentini et al. 2008). Pollen, macrofossils,
dermis). While certain shapes are more common and the majority of the phytolith data point to
in some subfamilies than others (e.g., cross origin of the family about 80 mya, or in the upper
shaped silica bodies in Chloridoideae), the shapes Cretaceous. The divergence of the BEP and PAC-
are rarely diagnostic. In an early phylogenetic MAD clades (crown grasses) then is placed at 50
analysis of the family, Kellogg and Campbell to 60 mya (Bouchenak-Khelladi et al 2010; Jones
(1987) excluded silica bodies entirely because of et al. 2014; Vicentini et al. 2008). Christin et al.
unusually high homoplasy. Recent work on phy- (2014) also estimate the BEP-PACMAD split at
tolith morphology, however, has shown that with 51.2 mya (12.3) or 62.6 (7.6) using two differ-
analysis in three dimensions additional charac- ent dating programs, a set of nuclear genes, and
ters can be retrieved and phytoliths can in some calibrations using only fossils from non-Poaceae.
cases be assigned reliably to particular taxa These calibrations and dates all have the satisfy-
(Piperno 2006; Str€omberg 2004). For example, ing property of placing the origin of the PACMAD
most bambusoid silica bodies are saddle shaped clade and hence the earliest appearance of C4
or bilobate in surface view, but when removed photosynthesis about 32 mya in the Oligocene,
from the leaf tissue and viewed from the side, they corresponding to a drop in atmospheric CO2.
appear thick, with irregular points (Piperno and In contrast to all other data, the Indian phy-
Pearsall 1998), and this morphotype appears to toliths would place the origin of grasses apprecia-
be diagnostic for the subfamily. Nonetheless, bly earlier, around 128 mya (Vicentini et al.
much phytolith data relies on the frequency of 2008); this date corresponds to the earliest fossil
particular morphs in an assemblage, rather than a evidence of monocots, which is about 125 mya
single synapomorphic form (Piperno 2006; (Herendeen and Crane 1995) and thus would
Piperno and Pearsall 1998; Str€omberg 2004). require pushing the monocot-dicot split much
The presumed earliest records of grass phy- earlier. Using this same calibration point for a
toliths are from India. Phytoliths found in a dino- dataset based on whole chloroplast genomes,
saur coprolite suggest that dinosaurs consumed Jones et al. (2014) placed the stem node of Poa-
grass (Prasad et al. 2005), a suggestion at odds ceae at 180 mya. Christin et al. (2014), using
with all other fossil information. Morphological nuclear genes, the phytolith calibration plus
diversity of the phytoliths suggests that the BEP non-Poaceae fossils, placed the BEP-PACMAD
clade had already diversified by the date of the divergence at 82.4 (14.8) or 79.1 (3.0) mya,
coprolite (65–67 mya). Additional phytolith data depending on the dating program used. Dates
from similar fossil formations indicate presence based on such calibrations imply a major slow-
of Ehrhartoideae phytoliths at about that date as down of evolutionary rates in the Poaceae. In
well (Prasad et al. 2011), again a surprisingly early addition, such dates would require revision of
record. Because the Indian subcontinent had no hypotheses regarding C4 evolution because atmo-
connections to other landmasses at the time, spheric CO2 was about 1000 ppm at that time. If
accepting the date of the Indian phytoliths the older calibration and dates are correct, then
would force the conclusion that grass diversifica- lower atmospheric CO2 cannot have been the
tion must have either occurred in India, or have driver of C4 but rather it may have been a change
occurred before the breakup of Gondwana. Phy- in seasonality or aridity (Str€omberg 2011). The
tolith assemblages have also been described by older dates also contradict the suggestion of
Brea et al. (2009) and Zucol et al. (2010) for a mid- Ehleringer et al. (1997), which argues that C4
to late-Eocene formation in Patagonia (ca. 39–41 photosynthesis can only be more efficient than
mya); these include a diverse set of phytoliths C3 when CO2 concentrations are relatively low.
including some that could be assigned to chlor- The older date is also incompatible with estimates
idoids or panicoids. of the evolution of cold tolerance in Pooideae
106 Fossil Record and Dates of Diversification
(Sandve and Fjellheim 2010), which would have Brea, M., Zucol, A.F., Raigemborn, M.S., Matheos, S. 2009.
to have occurred when the climate was unusually Reconstrucción de paleocomunidades arbóreas med-
iante análisis fitolı́ticos en sedimentos del paleoceno
warm. The older date for grass origins also affects superio-eoceno? (formación las flores), Chubut,
estimates of evolutionary rates and dates of other Argentina. BAR Int. Series S1870 9: 91–108.
parts of the angiosperm tree. Therefore, accepting Bryant, Jr., V.M., Holloway, G.R., Jones, J.G., Carlson, D.L.
the date and identity of the Indian phytoliths 1994. Pollen preservation in alkaline soils of the
American Southwest. In: Traverse, A. (ed.) Sedimen-
requires overturning virtually all other data and tation of organic particles. Cambridge: Cambridge
inferences about grass, and indeed monocot, University Press. pp. 47–58
evolution. Chandler, M.E.J. 1964. The lower Tertiary flora of south-
In summary, the age of the grasses and their ern England IV. London: British Museum (Natural
History).
rate of molecular evolution are unknown, Christin, P.-A., Besnard, G., Samaritani, E., Duvall, M.R.,
although most lines of evidence point to the Hodkinson, T.R., Savolainen, V., Salamin, N. 2008.
BEP-PACMAD split having occurred 50–60 mya Oligocene CO2 decline promoted C4 photosynthesis
in grasses. Curr. Biol. 18: 37–43.
in the Paleocene or early Eocene, and the grass Christin, P.-A., Spriggs, E., Osborne, C.P., Str€omberg, C.A.
stem node being dated to the upper Cretaceous. E., Salamin, N., Edwards, E.J. 2014. Molecular dating,
In this scenario, both the earliest C4 origins and evolutionary rates, and the age of the grasses. Syst.
the evolution of pooid cold tolerance originated Biol. 63: 153–165.
Crepet, W.L., Feldman, G.D. 1991. The earliest remains of
around the Eocene-Oligocene boundary. How- grasses in the fossil record. Amer. J. Bot. 78:
ever, these dates, despite being supported by 1010–1014.
many disparate sorts of fossils and dating meth- Ehleringer, J.R., Cerling, T.E., Helliker, B.R. 1997. C4 pho-
ods, are incompatible with phytolith data (Chris- tosynthesis, atmospheric CO2, and climate. Oecolo-
gia 112: 285–299.
tin et al. 2014; Prasad et al. 2011; Str€omberg 2011). Elias, M.K. 1942. Tertiary prairie grasses and other herbs
If the phytolith data are accepted at face value, from the High Plains. Geol. Soc. Amer. Spec. Pap. 41:
then the paleohistory of grasses must be revised 1–176.
by millions of years. Such revisions change esti- Herendeen, P.S., Crane, P.R. 1995. The fossil history of the
monocotyledons. In: Rudall, P.J., Cribb, P.J., Cutler,
mates of molecular evolution in the grasses, from D.F., Humphries, C.J. (eds.) Monocotyledons: sys-
being much more rapid than most monocots to tematics and evolution. Kew: Royal Botanic Gardens.
being much slower; such changes also affect esti- pp. 1–21
mates of dates for many important events in Jacobs, B.F., Kingston, J.D., Jacobs, L.L. 1999. The origin
of grass-dominated ecosystems. Ann. Missouri Bot.
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broken down. Much thus rests on the reliable nomics, molecular evolution, and estimated ages of
lineages from the deep phylogeny of Poaceae. Plant
dating of the Indian phytoliths and the phyloge- Syst. Evol. 300: 1421–1436.
netic distribution of their distinctive shapes. Kellogg, E.A., Campbell, C.S. 1987. Phylogenetic analyses
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Oryzeae, which in turn assumes that costal rows Campbell, C.S., Barkworth, M.E. (eds.) Grass system-
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only in that tribe. This assumption can only be Linder, H.P. 1987. The evolutionary history of the Poales/
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Domestication
“. . . agriculture [is] something the grasses did to people as a way to conquer the trees.” (Pollan 2001), p. xxi
ORIGINS OF MAJOR CEREAL CROPS The ancestry of wheat is complex but well
studied, involving several species with three dis-
Because the grasses are common plants, often tinct genomes, conventionally labeled A, B, and
ecologically dominant and with nutritious seeds, D. Diploid A genome wheats include Triticum
it is not surprising that they were harvested and urartu and T. boeoticum. T. boeoticum was
later domesticated by the earliest agriculturalists. domesticated in what is now southeast Turkey
Archaeological evidence shows that wild grasses about 9000 years before present, to give rise to
were used well before agriculture began (Bettin- einkorn wheat (T. monococcum) (Peng et al.
ger et al. 2010; Kislev et al. 1992; Purugganan and 2011). An A genome wheat similar to modern T.
Fuller 2009), and only gradually did early hunter- urartu crossed with a species of goatgrass similar
gatherer groups shift to farming. Fuller (2007) to modern Aegilops speltoides, which provided
suggests a model, modified from Harris (1989), the B genome, and formed a stable allotetraploid.
that describes gathering of wild food and cultiva- Because A. speltoides is widespread and weedy it
tion of domesticated plants as end-points of a is not surprising that the two species would have
continuum, with intermediate stages of “semi- come into contact. The resulting tetraploid
domestication” that spread over various lengths wheats include T. dicoccoides (¼ T. turgidum
of time in various crops (e.g., Tanno and Will- ssp. dicoccoides), T. dicoccum (¼ T. turgidum
cox 2006). In general, this model shows that ssp. dicoccum), and T. araraticum; the former
domestication was a long slow process, occur- two are known as wild emmer and emmer,
ring over hundreds to thousands of years. As respectively, with T. dicoccum being the domes-
population genetic data have been applied more ticated form of T. dicoccoides. (Note that the
extensively to the question of domestication, taxonomy within Triticum is not stable, with
and as more archaeological sites have been each taxon being named variously at the species
investigated, it has also become clear that or subspecies level. In addition, not all authors
many domesticated species were domesticated recognize all species as distinct. Nomenclature
more than once (Weiss et al. 2006). The best here follows Peng et al. 2011.) Like einkorn,
studied of the cereals are the so-called Big emmer wheat was domesticated in southeast Tur-
Three – wheat, rice, and maize – plus barley, key or the southern Levant or both ca. 9500–9000
all of which have an appreciable archaeological years before present; the wild ancestor, T. dicoc-
record. However, additional data are accumulat- coides is fairly widespread in the region and
ing on many other species that are somewhat authors disagree on the number of domestication
less important for modern agriculture, but that events (Luo et al. 2007; Matsuoka 2011; Peng et al.
have been important historically and regionally. 2011). Triticum durum, now used for making
These data permit some generalizations about pasta, is derived from T. dicoccum. Other inde-
the domestication process as well as the effects pendent, domestication events of einkorn and
of selection on genes and genomes. emmer can be detected in the archaeological
110 Domestication
record but did not persist to modern times (Fuller teosinte, Zea mays ssp. parviglumis. Early intro-
2007; Peng et al. 2011). gression into maize from another teosinte, Zea
Although most of the morphological changes mays ssp. mexicana created some confusion
associated with domestication occurred in the about the precise locality of domestication, but
diploid and tetraploid species einkorn and this has been clarified by population genetic data
emmer, neither is a significant crop today. on maize landraces (van Heerwaarden et al.
Instead they have been supplanted by the allohex- 2011).
aploid, Triticum aestivum. Soon after emmer Domesticated barley (Hordeum vulgare)
domestication, cultivated emmer or a very similar appears in archaeological remains about the
tetraploid with A and B genomes crossed with a same time as einkorn and emmer wheat. In addi-
second species of goatgrass, A. tauschii (with the tion to its center of domestication in the Fertile
D genome), to produce T. aestivum or bread Crescent, a second domestication occurred to the
wheat (Kihara 1944; McFadden and Sears 1946). east of the Zagros Mountains (Morrell and Clegg
This hybridization probably occurred multiple 2007; Saisho and Purugganan 2007). Even within
times in the southwestern Caspian region of pres- the Fertile Crescent, there is evidence for at least
ent-day Iran (Wang et al. 2013). The D genome is two domestication events, based on the genes
responsible for providing the genes that give controlling rachis disarticulation. These two
wheat flour its many desirable properties for bak- loci, Brittlerachis1 (Btr1) and Brittlerachis2
ing (Kaltsikes et al. 1968; Kerber and Tipples (Btr2) are linked on chromosme 3H; the alleles
1969; Nelson et al. 2006). Evidence for the hybri- conferring rachis disarticulation are dominant.
dization events leading to bread wheat was first Loss of function at either locus creates a non-
provided by painstaking cytogenetic studies; shattering phenotype. Barley cultivars that are
Kihara then resynthesized T. aestivum from its mutant at btr1 are phylogenetically distinct from
tetraploid and diploid ancestors (Kihara 1982; plants that are mutant at btr2 (Azhaguvel and
Kihara and Lilienfeld 1949). His results have Komatsuda 2007), based on a phylogeny of a
since been confirmed by genome mapping marker linked to the Btr loci.
(Devos et al. 2008; Gale and Devos 1998) and Foxtail millet (Setaria italica) was domesti-
population genetics (Wang et al. 2013). cated from green millet (S. viridis) in northwest-
Rice was domesticated in Asia about 9000 ern China about 7000–8000 years ago (Hunt et al.
years ago from Oryza rufipogon, but the precise 2008; Le Thierry d’Ennequin et al. 2000; Li and
number of origins in a matter of contention (Kel- Wu 1996). Although archaeological sites in the
logg 2009; Vaughan et al. 2008); population Caucasus hint at a possible second center of
genetic data have been equivocal. Recent evi- domestication, the evidence for this is not firm
dence points to a single origin of O. sativa japon- (Harlan 1992; Hunt et al. 2008). The world’s old-
ica in southern China followed by wide dispersal est recorded noodles, dated at 4000 years before
(Huang et al. 2012; Molina et al. 2011). Extensive present were thought to have been made with
human transport of grains over millennia plus foxtail millet, but recent information questions
continued introgression with wild rice (Oryza this claim and suggests that the noodles could
rufipogon) later led to the origin of O. sativa have been made from wheat or barley, both of
indica (Huang et al. 2012). Oryza glaberrima which were also cultivated in the region at the
was domesticated independently in western time (Ge et al. 2011).
Africa from O. barthii, no later than 300–200 BC Finger millet, Eleusine coracana ssp. cora-
(Sweeney and McCouch 2007; Vaughan et al. cana, is an allotetraploid native to Africa, domes-
2008). Although rice grains as old as 3500 years ticated from E. coracana ssp. africana. The date
before present have been identified at sites in of domestication is uncertain. Grains of E. cora-
northeast Nigeria, they were apparently not cana have been found in Ethiopia from a site with
domesticated (Sweeney and McCouch 2007). a possible date of the third millennium BC, but
Zea mays ssp. mays was domesticated in the the date has not been verified (Hilu et al. 1979).
Balsas River Basin in Mexico, about 9000 years The crop was introduced to India from Africa,
before present (Matsuoka et al. 2002; Piperno where it has developed several characteristics
et al. 2009; van Heerwaarden et al. 2011), from distinct from its African forebears (Dida et al.
ORIGINS OF MAJOR CEREAL CROPS 111
2008; Hilu and de Wet 1976). One genome of E. eals. S. officinarum has hybridized repeatedly
coracana was clearly donated by a parent similar with S. spontaneum, a species that is widespread
to the African species E. indica, but the other in Southeast Asia, New Guinea and nearby
genome is unknown (Neves et al. 2005). islands. In the last century, the two species were
Common (Proso or broomcorn), millet (Pan- deliberately hybridized to give rise to most mod-
icum miliaceum) was domesticated in China in ern cultivars and breeding lines of sugarcane. S.
the same general area as foxtail millet but some- officinarum has a chromosome number of
what earlier, perhaps about 10,000 years before 2n ¼ 80, and often produces unreduced gametes.
present (Lu et al. 2009a). The two species have When it crosses with S. spontaneum (2n ¼ 64), in
similar grains, making interpretation of archaeo- which gametes are generally haploid, the chromo-
logical data difficult, but some details of the some complement of the resulting hybrid is
lemma epidermis differ between them (Lu et al. roughly 80 + 32 chromosomes. Loss of S. sponta-
2009b). Panicum miliaceum is an allopolyploid. neum chromosomes is common, giving rise to a
One genome comes from a clade that includes the range of aneuploid numbers (Piperidis et al.
tetraploids P. bergii and P. stramineum, and the 2009).
diploids P. capillare and P. nepheliphilum, Tef (Eragrostis tef) was domesticated in
whereas the other genome is connected to a Ethiopia a little more than 2000 years before
group of polyploid species placed in Panicum present, from an ancestor similar to E. pilosa
sect. Dichotomiflora (Triplett et al. 2012). How- (D’Andrea 2008; Ingram and Doyle 2003); both
ever, few Asian species have been included in any species are tetraploid. Among the cereals, tef is
phylogenetic analysis to date, so the closest rela- distinctive for its incredibly tiny grains,
tives and diploid progenitors of common millet 0.8–1.5 mm long. This is only slightly longer
remain unknown. than the size of grains in E. pilosa (0.5–1.1 mm),
Pearl millet (Cenchrus americanus ¼ Penni- which makes distinguishing the two difficult
setum glaucum) was domesticated in western (D’Andrea 2008). Selection appears to have
Africa, with the oldest remains discovered in the focused on increasing inflorescence branching
Tilemsi Valley in present-day Mali, about 4500 and producing additional tillers, rather than
years before present (Manning et al. 2011; Oumar increasing size of the grains and density of the
et al. 2008). inflorescence. It is possible that this selection
In contrast to other cereal crops, the domes- strategy results from concerns about lodging
tication history of Sorghum bicolor is poorly (D’Andrea 2008).
known. It appears to have originated in sub- The domestication history of many minor
Saharan Africa from a wild progenitor similar to grains, such as fonio (Digitaria exilis), Guinea
S. bicolor ssp. arundinaceum (Zohary and Hopf millet (Brachiaria deflexa), Kodo millet (Paspa-
2000). Domestication is thought to have occurred lum scrobiculatum), barnyard millet (Echino-
in Ethiopia beginning 6000–5000 years before chloa crus-galli), sauwi millet (Panicum
present, but the resultant landraces are highly sonorum), raishan (Digitaria sp.), and Bromus
diverse both morphologically and genetically mango remains poorly studied. Lines of fonio
(Dillon et al. 2007), reflecting the many disparate and Guinea millet with larger seeds and non-
uses of the crop. shattering inflorescences are reported (National
Sugarcane (Saccharum officinarum) origi- Research Council 1996). In the Chad Basin of
nated in New Guinea, where it appears to have northeastern Nigeria, it is possible to buy in
been derived from the wild species, S. robustum local markets a mixture known as kreb, which
(Artschwager and Brandes 1958; Lebot 1999), consists of small grains from several genera of
presumably by selection for clones with high Paniceae, including Echinochloa, Brachiaria, and
sugar and low fiber contents. Unlike the cereals, Panicum (Klee et al. 2000). Bromus mango was
it is generally propagated vegetatively, leading to apparently cultivated in what is now Chile before
a set of widespread clones with low genetic diver- the Spanish conquest (Planchuelo and Peterson
sity. The specific steps in domestication of this 2000), but is now rare.
species are unknown; in general it is much less Modern oats and rye were domesticated in
well studied than domestication of the major cer- Europe, but were previously cultivated and
112 Domestication
possibly domesticated in several distinct sites in mays ssp. mays (cultivated maize) can be crossed
the Near East; these earlier domesticated strains with its progenitor teosinte (Zea mays ssp. parvi-
have not persisted (Weiss et al. 2006). Rye (Secale glumis) as long as maize is the female parent and
cereale) was domesticated from wild plants of the teosinte the pollen parent. (See section on Repro-
same species, which occur in areas corresponding ductive systems above.) In addition, Zea mays
to present-day Turkey (Khush 1963; Sencer and ssp. mays has crossed with Zea mays ssp. mex-
Hawkes 1980; Zohary and Hopf 2000). In com- icana, somewhat confusing efforts to disentangle
parison to species of Triticum and Hordeum, it is the domestication history of the species (van
more tolerant of cool temperatures and is gener- Heerwaarden et al. 2011). Introgression has been
ally found at higher altitudes (Burger and Ell- demonstrated between finger millet (Eleusine cor-
strand 2005). Domestication appears to have acana ssp. coracana) and its wild progenitor
occurred more than once. A few, possibly domes- (Eleusine coracana ssp. africana) (Hilu and de
ticated, grains appear at a site known as Abu Wet 1976), between pearl millet (Cenchrus amer-
Hureyra, on the Euphrates in current Syria, dat- icanus ¼ Pennisetum glaucum) and its wild
ing from about 12,500 years before present, but ancestor (P. glaucum ssp. monodii) (Mariac
these were mixed with wild grains (Weiss et al. et al. 2006; Oumar et al. 2008), between wild
2006; Willcox 2005). Domesticated grains (Triticum dicoccoides) and cultivated (T. dicoc-
(plump, on tough rachises) have been found cum) emmer wheat (Luo et al. 2007), and between
about 8600 years before present in south Central Oryza sativa and O. rufipogon (Huang et al. 2012).
Anatolia, and ca. 4000 years BP in north central
Anatolia (Weiss et al. 2006).
Cultivated oats first appear in the archaeolog- MORPHOLOGICAL CHANGES IN DOMESTICATION
ical record in Europe in the second millennium
BC (Zohary and Hopf 2000). Large caches of oats Domestication results in a set of well-documen-
have been found in archaeological sites in the ted genetic changes caused by the intense selec-
Near East. While these caches appear to lack the tion pressure created by growing in tilled soil,
characteristic morphology of domesticated oats, annual harrowing, annual harvesting and
the numbers of grains suggest that they were subsequent planting (Harlan 1992). One of the
likely cultivated (Weiss et al. 2006). Zohary and earliest traits to be modified during domestica-
Hopf (2000) speculate that oats were probably tion is grain size (Fuller 2007; Purugganan and
domesticated more or less incidentally. The hexa- Fuller 2009). In part, this appears to be the result
ploid oats in the sativa complex are common and of unconscious selection for plants that produce
widespread weeds in fields of wheat and barley vigorous seedlings and that can tolerate the
throughout the Mediterranean; it is thus likely increased planting depth of tilled soil (Purugga-
that they were harvested along with other grains nan and Fuller 2009). Changes in grain size
and were thus selected alongside them. Cultivated occurred rapidly in cultivated wheat, barley and
oat (Avena sativa) is a hexaploid, with genomes rye, with larger grains appearing in the archaeo-
labeled A, C, and D. The A genome appears to logical record over a period of about 500 years
have come from the maternal parent, as shown by (Fuller 2007). The change of grain size in rice is
sequences of chloroplast genes (Peng et al. 2010). confounded with changes in shape, but in general
Based on phylogenetic analysis, the maternal par- the change in grain size appears to have been
ent was similar to the diploid A. wiestii, a parent somewhat slower in rice (Fuller 2007).
also shared by the tetraploid A. murphyi, and the Another early change is loss of disarticulation
hexaploids A. sterilis and A. occidentalis. Chloro- (shattering). (See section on Inflorescence struc-
plast data suggest that the hexaploid wild oats (A. ture: Disarticulation.) Whether this is selected for
fatua) were derived from an independent allopo- depends on the mode of harvest. If seeds are
lyploidization event. collected by shaking the plants and collecting
Most domesticated species are crossable with the falling seeds in a basket, then shattering will
their wild ancestors, and the wild ancestors often be retained. This is seen in North American wild
persist as weeds in cultivated fields (Harlan 1992). rice (Zizania), which has been harvested by
For example, Setaria italica has been crossed with Native Americans for centuries by moving
its progenitor, S. viridis (Wang et al. 1998). Zea through the stands of wild rice in boats and
MORPHOLOGICAL CHANGES IN DOMESTICATION 113
beating the plants so that the seeds fall in to the (Clark et al. 2006; Doebley et al. 1997; Zhou et al.
boat. In contrast, if harvesting is done with a 2011). Superficially similar phenotypic changes
sickle, the selection pressure is reversed to favor occurred during domestication of foxtail millet
plants that retain their seed. A plant that lacks (Setaria italica) and pearl millet (Cenchrus amer-
shattering is heavily, if not completely, dependent icanus ¼ Pennisetum glaucum), but quantitative
on humans for propagation and dispersal, so lack genetic studies find that tb1 was only a minor
of shattering is often taken as diagnostic of a player in those transitions (Doust et al. 2004;
domesticated plant. Several of the genetic loci Remigereau et al. 2011). In both maize and
controlling shattering have been cloned in rice pearl millet, the axillary buds that would form
(Konishi et al. 2006; Li et al. 2006; Zhou et al. tillers are well developed but fail to elongate
2012), and one in wheat (Simons et al. 2006). (Remigereau et al. 2011). In foxtail millet, in
Whether these same loci were selected in other contrast, axillary buds fail to initiate at all, con-
domesticated grasses is unknown. The Btr1 and sistent with a different locus of selection (Doust
Btr2 loci of barley map to chromosome 3 (known et al. 2004).
as 3H, for chromosome 3 in Hordeum); there are In pooid and ehrhartoid grasses, tillering is
two brittle rachis loci in emmer wheat, Br1 and common, with each tiller ending in an inflores-
Br2 that also map to the homoeologous chromo- cence; in contrast culm branching is rare or
somes (3A and 3B) (Matsuoka 2011). Whether the absent. In domestication of crops in these sub-
Hordeum and wheat loci are orthologous is families, the extent of tillering appears not to have
unknown and will await cloning of one or more been a target of selection (Doust 2007). In rice
of the genes (Li and Gill 2006). and wheat the extent of tillering does not differ in
Whereas development of larger grains a consistent way between wild and domesticated
occurred before loss of shattering in wheat, barley plants (Remigereau et al. 2011). Wild finger millet
and rice (Fuller 2007; Purugganan and Fuller (Eleusine coracana ssp. africana) develops more
2009), the order was reversed in pearl millet tillers than cultivated (E. coracana ssp. coracana),
(Fuller 2007). While loss of shattering could in but cultivated plants have more axillary branches
principle have occurred in less than 100 years, than wild ones (Dida et al. 2008).
such rapid genetic change requires intense (albeit The process of sowing and harvesting selects
unconscious) selection imposed by harvesting automatically for plants that germinate promptly
semi-ripe grains with a sickle or by uprooting and synchronously so dormancy and often day-
(Hillman and Davies 1990). Rapid selection also length requirements tend to be lost in domesti-
requires sowing seed in new sites each year to cated species (Harlan et al. 1973). In wild oats,
avoid interbreeding with unselected plants. The einkorn and emmer wheat, spikelets generally
archaeological record, in contrast, indicates that have two flowers; the proximal one is larger and
the transition to non-shattering lines occurred non-dormant whereas the distal one is dormant.
over about 1500 years in wheat and 2000 years Dormancy is lost during domestication. In addi-
in barley, suggesting that wild populations tion to loss of dormancy (dispersal in time),
continued to mix with domesticated ones and domestication also often correlates with loss of
that selection for non-shattering grains was rela- dispersal structures, such as awns in wheat
tively weak (Fuller 2007). (Fuller 2007) and reduction or loss of awns in
Domestication also affects tillering and culm rice (Sweeney and McCouch 2007).
branching, although both the phenotype and the In cultivated rice, most lines have white or
underlying genes vary among domesticated brownish pericarp, representing modification of
grasses. In the domestication of maize, tillering the ancestral red color. Pericarp color is regulated
was suppressed so that cultivated maize gener- by the Rc locus, a bHLH transcription factor
ally has a single culm, in contrast to its highly (Sweeney et al. 2006). The Rc allele in cultivated
branched wild ancestor, teosinte. The change in rice is non-functional, having a 14 bp deletion
architecture was caused by selection on regula- that abolishes DNA binding. The allele appears
tion of the gene teosinte branched1 so that it is to have arisen in a japonica rice and then spread
more strongly expressed in the axillary buds near via introgression throughout almost all cultivated
the base of the plant, preventing bud outgrowth lines (Sweeney et al. 2006).
114 Domestication
In both maize and barley, human selection led still attached. Grains require drying to be broken
to inflorescences that are not merely larger, but free from the surrounding floral bracts (Peng
that have changes in architecture. Wild barley has et al. 2011). Mutations in the genes Q and Tena-
three spikelets per node, only one of which is fer- cious glumes led to free-threshing varieties, in
tile. The clusters of three spikelets form on alter- which the glumes break easily (Nalam et al.
nate sides of the inflorescence such that there is one 2006, 2007; Simons et al. 2006). (See above, Inflo-
row of fertile spikelets on either side; this is known rescence structure: Spikelets.) In addition to con-
as two-row barley. Some modern cultivars retain ferring the free threshing characteristic, the
this trait, but more often selection has fixed muta- dominant (domesticated) Q allele makes the
tions that cause the sterile spikelets to become inflorescence shorter and broader, and the rachis
fertile. This is six-row barley, with three fertile tough and non-disarticulating.
spikelets in clusters on alternate sides of the rachis.
Fertility of the lateral spikelets is controlled by at
least two loci. One, a homeodomain-leucine zipper GENETIC AND GENOMIC CHANGES IN DOMESTICATION
(HD-ZIP I) transcription factor, Vrs1, has been
mutated several times independently in the history The effects of domestication on the genome are
of barley (Komatsuda et al. 2007). A second locus, the subject of much interest and active study. In
INT-C is an ortholog of Tb1, the gene that affects general, domestication creates a population bot-
tillering in maize (Ramsay et al. 2011). Mutations tleneck, which leads to reduced genetic diversity
in INT-C create a phenotype that is intermediate (Doebley et al. 2006; Glémin and Bataillon 2009).
between two-rowed and six-rowed barley; the lat- In cultivated durum and bread wheat, diversity
eral spikelets are partially fertile, and are better has been sharply reduced from wild strains, with
developed than those in two-rowed lines but smal- only 16 % and 31 % of wild diversity respectively
ler than those in six-rowed lines. (Haudry et al. 2007). However, these estimates are
The most dramatic and most famous of the based on a modest number of accessions and
domesticated phenotypes is the cob or ear of markers so should probably be expanded and
maize (corn). This is a botanical monstrosity verified. Foxtail millet retains about 45 % of the
that bears little resemblance to the small pistillate genetic diversity of its wild ancestor green millet
inflorescence of the other subspecies of Zea mays (Wang et al. 2010), and pearl millet retains
and other species of Zea. The wild type inflores- 67–74 % of the wild diversity (Clotault et al.
cence of teosinte has distichous spikelets. At 2012; Oumar et al. 2008). In contrast, landraces
maturity the outer glume plus the associated of maize harbor almost as much variation as is
rachis internode form a rock-hard structure that found in teosinte (83 %; Hufford et al. 2012).
encloses the grain, and the inflorescence rachis Levels of diversity vary among the three gen-
disarticulates at the nodes as in other Andropo- omes of hexaploid wheat (Akhunov et al. 2010).
goneae. The cultivated inflorescence (cob or ear), The A and B genomes, which were inherited from
in contrast, has spikelets in many rows; the spike- emmer wheat, are more diverse than the D genome,
lets are naked, and at maturity the grain grows and single nucleotide polymorphisms (SNPs) in the
well beyond the flimsy glumes, lemmas and A and B genomes are spread throughout the chro-
paleas. The rachis in maize is firm, with no vestige mosomes. The simplest explanation for this pattern
of disarticulation. This complex set of morpho- is past and continuing gene flow between wild and
logical changes is under the control of a single cultivated emmer before polyploidy with Aegilops
major locus, TEOSINTE GLUME ARCHITEC- tauschii, and continuing gene flow between wild
TURE1, which is a transcription factor that nor- emmer and cultivated wheat. The latter two form
mally regulates spikelet development (Preston fertile pentaploid hybrids in nature (Kihara 1982).
et al. 2012; Wang et al. 2005). (See also Inflores- In contrast, SNP diversity in the D genome is very
cence structure: Spikelets, above.) low, confirming previous studies suggesting only a
In wild wheat and barley, as well as in the so- handful of hybridization events leading to the hexa-
called “hulled wheats” – einkorn, emmer, and the ploid T. aestivum (Akhunov et al. 2010). In T.
hexaploid Triticum spelta (spelt) – grains are aestivum, loci related to domestication appear to
dispersed with the glumes, lemma, and palea be clustered in only about 16 regions of the genome
GENETIC AND GENOMIC CHANGES IN DOMESTICATION 115
(Peng et al. 2011). However, because genes in wheat two of these lines are hybridized, heterosis leads
tend to be clustered in general, this clustering may to high-yielding plants that are the basis of cur-
reflect a peculiarity of the wheat genome, rather rent maize production (Duvick 2001). The heter-
than a generalization about domestication. otic groups have become increasingly
Reduction of diversity is especially obvious in differentiated from each other over time; the
regions of the genome harboring genes that are genome of each shows evidence of directional
directly selected. In such regions, genetic diver- selection but no strong selective sweeps (van
sity in the gene under selection is reduced to near Heerwaarden et al. 2012).
zero, as would be expected if a single allele is With the increased availability of molecular
chosen for propagation. During the selection pro- markers and DNA sequence data, it has become
cess, the region of the genome closely linked to possible to identify many of the genes that were
the selected locus is effectively pulled along, so fixed during the domestication process. Although
that an entire piece of the genome shows reduced the sample size remains small, most such genes
diversity. Because selection effectively sweeps appear to be transcriptional regulators, consis-
away variation in the selected region, such a pat- tent with the hypothesis that modification of
tern is taken as evidence of a “selective sweep”. such genes is responsible for much morphologi-
Now that it is possible to sequence portions of the cal change (Doebley et al. 2006). “Domestication
genome of many members of a population, it is genes” include tb1 and tga1 in maize, vrs1 in
possible to scan the genome to look for regions of barley and prog1 in rice, all of which control
reduced variation and to infer past selection. plant or inflorescence architecture, sh1, sh4, and
Such a study in Zea mays estimated that 2–4 % shat1 in rice plus Q in wheat, all of which control
of the genome had been selected during domesti- shattering, and Rc in rice, which affects pericarp
cation and subsequent breeding (Wright et al. color (Glémin and Bataillon 2009). These loci
2005), and similar work in rice finds evidence of have been discussed above and in the sections
55 regions that have undergone selective sweeps on Inflorescence structure and on Fruit and
(Huang et al. 2012). seed. While these nine loci are all transcriptional
Recent studies have been structured to disen- regulators, GIF1 in rice is a cell-wall invertase, an
tangle the early genomic effects of domestication enzyme that controls carbohydrate partitioning
and the much later changes created during 20th and hence grain filling (Wang et al. 2008).
century crop improvement. For example, Hufford In contrast, genes that have been selected
et al. (2012) find that 1–2 % of the annotated during crop improvement (post-domestication)
genes in maize appear to have been selected dur- are much more diverse and include many genes
ing domestication, with a comparable number encoding parts of biosynthetic pathways. This
selected during improvement. While the two pattern also would be expected if crop improve-
sets of loci overlap somewhat, in general domes- ment focuses on such processes as starch biosyn-
tication targeted different genomic regions than thesis (Doebley et al. 2006). Crop improvement
crop improvement did. This is not particularly genes include those controlling kernel color and
surprising, since the process of domestication endosperm composition in maize (c1, r1, sh2, su1,
reduced variation at selected loci, so there was y1) and in rice (waxy), plant height (sd1 in rice,
little for subsequent selection to act on. Compar- rht in wheat), flowering time in rice (hd1, hd6),
ison of gene expression between teosinte and and response to vernalization in wheat (vrn1,
maize finds that some of the same loci identified vrn2). Mutations in waxy affect starch biosynthe-
by population genetic studies exhibit differential sis; by preventing amylose production they lead
expression patterns between the wild and to sticky rice grains that are used for special
cultivated lines, consistent with the hypothesis culinary purposes. Sticky rice varieties exhibit a
of modification through selection (Swanson- broad region (260 kb) of reduced diversity
Wagner et al. 2012). around the waxy locus, indicating strong selec-
The process of maize breeding has led to the tion (Olsen et al. 2006). Kernel color in maize
development of three “heterotic groups”, known varies, with yellow corn being caused by a domi-
as Iowa Stiff Stalk Synthetic (SS), Non-Stiff Stalk nant allele at Y1, which encodes phytoene
(NS), and Iodent (IDT). When plants from any synthase. Breeders have selected for yellow corn
116 Domestication
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from the gene (Palaisa et al. 2006). Doebley, J.F., Gaut, B.S., Smith, B.D. 2006. The molecular
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Doust, A. 2007. Architectural evolution and its implica-
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Affinities
Poaceae are members of the commelinid clade of widely distributed taxonomically. The clade of
monocots (Barrett et al. 2012; Givnish et al. 2010; Commelinales, Zingiberales, and Poales may be
Stevens 2012). Commelinids share the unique about 120 million years old (Janssen and Bremer
characteristic of accumulating ferulic and couma- 2004), unless dating of the Indian phytoliths
ric acids in the walls of unlignified cells; these proves to be correct, in which case it would be
compounds makes the cell walls fluoresce under substantially older (see discussion above under
ultraviolet light. All members of the clade, includ- Fossil record and dates of diversification).
ing Poaceae, produce silica (SiO2) bodies in their The order Poales includes 17 families all of
leaves. Stomata have two subsidiary cells with the which accumulate silica in the epidermis and
long axis parallel to the stomate (paracytic sto- have endosperm in which multiple nuclear divi-
mata), or may have an additional two short sub- sions occur before cell walls are formed (nuclear
sidiaries at each end of the stomate, endosperm) (Stevens 2012). The embryo is tiny in
perpendicular to the long axis (tetracytic). Inflor- all but the grasses. The order is also defined by
escences are bracteate. Embryos of the commeli- molecular characters and sequence data; the
nids are short and broad, but this character has mitochondrial gene, succinate dehydrogenase 3
been modified in the Poaceae, as described below. (sdh3) is absent from all sampled families
The commelinid clade includes Dasypogona- (Adams and Palmer 2003). Many members are
ceae, Arecales (palms, family Arecaceae), plus the wind-pollinated and many have reduced, green-
orders Commelinales, Zingiberales (gingers and ish flowers borne in condensed bracteate inflor-
bananas), and Poales. The latter three orders escences. The term spikelet is applied to the tiny
probably form a monophyletic group, with Poales spike-like inflorescences of many members of
sister to a clade of Commelinales plus Zingiber- Cyperaceae, as well as Poaceae and other poalean
ales. However, analyses of whole plastome data families. However, “spikelets” are clearly not
sets find that monophyly depends on methods of homologous throughout the order and the term
data analysis and the particular genomic regions is only useful within the context of a particular
used (Barrett et al. 2012). Non-sequence-based family. The crown group node for the order is
characters support monophyly of the three estimated to date to about 113 mya (Janssen and
orders, which share a unique chemical composi- Bremer 2004), or 109–106 mya (Leebens-Mack
tion of the primary cell wall (glucurono-arabi- et al. 2005), or 109–108 mya (Magallón and Cas-
noxylans), and stomata with parallel cell tillo 2009), or 99 mya (Magallón and Castillo
divisions, characters that are retained throughout 2009) (all studies ignoring the dates of the Indian
the grasses. Members of Commelinales, Zingiber- phytoliths).
ales, and Poales also produce abundant starchy Within Poales is a clade comprising seven
endosperm, a characteristic that is thus quite families (Anarthriaceae, Restionaceae, Centrole-
ancient. Starchy endosperm is strongly associated pidaceae, Flagellariaceae, Joinvilleaceae, Ecdeio-
with the Poaceae and their overwhelming eco- coleaceae, and Poaceae). In much of the literature
nomic importance, but in fact is much more before 2000, the name Poales was applied only to
122 Affinities
this group of families, and hence had a more ceae share a unique set of nested inversions in
restricted application than it does currently the chloroplast genome (Doyle et al. 1992; Miche-
(Campbell and Kellogg 1987; Kellogg and Linder langeli et al. 2003).
1995). This clade is now sometimes called the The sister group of Poaceae is either Join-
Poales s.s., or the graminid clade (Angiosperm villeaceae, based on morphological characters
Phylogeny Group 2009). The group is clearly (Campbell and Kellogg 1987; Kellogg 2000; Kel-
monophyletic, whether defined by molecular or logg and Linder 1995), Ecdeiocoleaceae, based on
morphological characters. It includes plants with sequences of whole plastomes (Givnish et al.
rhizomes, and distichous (two-ranked) sheathing 2010), or Ecdeiocoleaceae plus Joinvilleaceae (M.
leaves; flowers are small, with inconspicuous McKain, unpublished). Joinvilleaceae include a
tepals, and are often imperfect. In the anther, single genus, Joinvillea, which is distributed in
the endothecial wall has girdle-like thickenings; southeast Asia and the Pacific Islands. Ecdeioco-
the pollen is monoporate and annulate, and has leaceae include two genera, Ecdeiocolea and Geor-
tiny channels known as scrobiculi penetrating the geantha, both native to southwestern Australia.
wall. In the gynoecium of most taxa, the stigmas Pollen characters and leaf epidermal characters
are plumose, and the receptive cells are on multi- conflict in terms of which family is sister to Poa-
cellular branches. Each carpel has a single ortho- ceae. Both Poaceae and Ecdeiocoleaceae have lost
tropous, pendulous ovule, and reduction to a scrobiculi in the pollen wall; this shared loss is a
single ovule per fruit is common. The graminid potential synapomorphy linking the two families.
Poales also share chemical characteristics, includ- However, both Joinvilleaceae and Poaceae have
ing the ability to produce flavones, a primary cell alternating long and short cells in the leaf epider-
wall that includes (1–3,1-4)-b-D-glucans, and mis, a character used by Campbell and Kellogg
sieve tube plastids with cuneate crystals. (1987), Kellogg (2000), and Kellogg and Linder
Although past literature has inferred that the (1995) to suggest a sister taxon relationship of the
entire graminid clade is wind pollinated, Givnish two families. Mature plants of Ecdeiocolea lack
et al. (2010) infer that wind pollination has been leaves so the character cannot be scored in that
independently derived in Restionaceae + Centro- genus; however, alternating long and short cells
lepidaceae, Ecdeiocoleaceae, and in the grasses are not observed in juvenile Ecdeiocolea or
descended from the common ancestor of Puelioi- mature Georgeantha (Givnish et al. 2010 citing
deae and the BEP + PACMAD clades. They sug- Briggs, pers. comm.). Additional morphological
gest that the ancestral condition for the graminid comparisons of Joinvilleaceae, Ecdeiocoleaceae,
Poales is animal pollination, as postulated for and Poaceae are provided in the sections on inflo-
Flagellariaceae, Joinvilleaceae, Anomochlooideae rescence and floral morphology.
and Pharoideae. However, this inference is based
largely on the androecial and stigmatic morphol-
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sequence phylogeny and evolution of Poales. Ann. Michelangeli, F.A., Davis, J.I., Stevenson, D.W. 2003. Phy-
Missouri Bot. Gard. 97: 584–616. logenetic relationships among Poaceae and related
Janssen, T., Bremer, K. 2004. The age of major monocot families as inferred from morphology, inversions in
groups inferred from 800+ rbcL sequences. Bot. J. the plastid genome, and sequence data from the
Linn. Soc. 146: 385–398. mitochondrial and plastid genomes. Amer. J. Bot.
Kellogg, E.A. 2000. The grasses: a case study in macroevo- 90: 93–106.
lution. Ann. Rev. Ecol. Syst. 31: 217–238. Stevens, P.F. 2012. Angiosperm phylogeny website. Ver-
Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In: sion 12, July 2012 [and more or less continuously
Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. updated since]. 2001 onward.
Classification of the Poaceae
Subdivision of the Family
The Poaceae are one of the few large plant and few subtribes have been in use long enough
families that currently enjoy a fully phylogenetic to be widely accepted.
classification, based on a phylogeny generated This treatment of the grasses relies heavily on
largely from DNA sequence data, but supported the tremendous synthetic work of Clayton and
by extensive morphological data (Fig. 9). All Renvoize (1986), Watson and Dallwitz (1992
named subfamilies, tribes, and subtribes are onward), and Clayton et al. (2006 onward), and
strictly monophyletic (i.e., including all the des- would have been completely impossible without
cendants of a given ancestor). Of the ca. 11,000 those earlier works. Although I have included as
species in the family, all can be placed in one much recent information as possible, many
of the early-diverging subfamilies, or in the BEP generic limits recognized here are based on the
or PACMAD clades, using either molecular or previous work. This is particularly true for genera
morphological data or both. Subfamily assign- of tropical Asia and Africa, many of which have
ment is uncertain for fewer than ten species, and yet to be incorporated in the recent wave of
a large majority of species can be placed into molecular phylogenetic studies. The major inno-
tribes and subtribes. Over 1/3 of the species in vation of this book is the phylogenetic approach.
the family now have some sort of molecular data The work of Clayton and Renvoize (1986) and
available, and the remainder exhibit enough mor- Clayton et al. (2006 onward) arises from the phi-
phological synapomorphies to place them confi- losophy of the evolutionary or phenetic school of
dently in clades identified by combined molecules taxonomy, and Watson and Dallwitz (1992
and morphology. onward) use an explicitly phenetic approach.
The subfamilial classification used here fol- More recent checklists use a phylogenetic
lows that of the GPWG (2001), updated by Sán- approach (Simon 2007; Simon et al. 2011 onward;
chez-Ken et al. (2007) and further confirmed by Soreng et al. 2012 and onward), updating the
the GPWG II (2012). The GPWG (2001) aimed classification frequently as indicated by recent
to maximize nomenclatural stability. Although molecular studies. The classification used here is
there are many possible ways to convert a phy- likewise phylogenetic, and is thus similar to those
logeny into a hierarchical classification, the in the recent checklists.
GPWG (2001) chose the combination of groups Many aspects of the classification presented
that retained the most similarity with previous here are congruent with the Classification of
classifications. Recent studies have also defined World Grasses (CWG; Soreng et al. 2012 and
the limits of many of the tribes of grasses; these onward). Both this classification and that of the
are cited under the appropriate subfamily and CWG are based on the same underlying data and
most are widely accepted. I have chosen not to are designed to be phylogenetic. The classification
recognize tribes that are fully redundant with presented here uses fewer ranks (e.g., supertribes
subfamilies (e.g., Phareae in Pharoideae). Delim- are not used), and also avoids monogeneric tribes
itation of subtribes remains a work in progress, or subtribes whenever possible. Therefore, the
128 Subdivision of the Family
Fig. 9. Phylogeny of the tribes of Poaceae, based on the family unplaced in a subfamily, although it clearly
GPWG II (2012). Subfamily names in bold. 1denotes taxa belongs the BEP clade. Dashed line Uncertain position.
not placed by GPWG II. Streptogyna is the only genus of
classification is less finely subdivided than that in forces a choice between narrowly defined subtribes
the CWG (Soreng et al. 2012 and onward). In addi- that are morphologically diagnosable but contain
tion, the delimitation of subtribes sometimes dif- only a single genus versus broader subtribes that
fers, notably in the Pooideae and Chloridoideae. provide information on relationships among
The structure of the phylogeny in those subfamilies genera but that have no morphological
KEY TO THE SUBFAMILIES 129
synapomorphy. The CWG has chosen the former how morphological evolution probably happened
option, creating a subtribal classification that pro- – a stepwise accretion of characteristics that in
vides little information about relationships. I have aggregate permitted a major radiation of several
chosen the alternative, naming more inclusive thousand species in temperate habitats. However,
clades as subtribes; the cost of this decision is that it is inconvenient for easy identification and gen-
the included genera rarely share morphological eration of keys. In the keys throughout this book,
characters. These differences reflect personal pre- therefore, taxa frequently have to be keyed multi-
ferences in ranking and do not affect the underly- ple times, and the leads contain weasel words
ing circumscription of groups. such as “generally” and “usually”.
The grass family includes about 30 % mono- A second challenge for taxonomy in the
typic genera. These are almost always easily rec- grasses is the nature of the morphological char-
ognizable species, with clear morphological acters themselves. The characters most useful for
synapomorphies, and are generally segregates delimiting taxonomic groups are often difficult or
from larger genera. Being monotypic, their impossible to use for identification. This dilemma
generic classification conveys no phylogenetic first became obvious in the 1930s, when Avdulov
information, reflecting only degree of difference. showed that chromosome number – a totally use-
Some of these genera are undoubtedly indepen- less field character – varied in a way that corre-
dent lineages, but others have been found to be lated well with some gross morphological
derived from within well-known genera. I have characters. Additional studies on leaf anatomy
returned the latter to their “parent” genera where were influential in refining the classification,
molecular and/or morphological data support the and most recently molecular data have provided
combinations, but I have hesitated to make clear support for large groups based on other
changes where shared derived characters (molec- (micromorphological) characters. This illustrates
ular or morphological) are lacking. the remarkable ability of natural selection to
The phylogenetic approach used here does shape the gross morphology of the grasses, but
affect many familiar names of species. In an also leads to difficulties in writing keys. Again the
explicit comparison of names in GrassBase (Clay- reader will have a chance to experience for him-
ton et al. 2006 onward) with those in a recent or herself this double-edged sword. The key to the
phylogenetic checklist (Simon 2007; Simon et al. subfamilies below is a case in point.
2011 onward), generic names were changed for an
estimated 10–20 % of species (Vorontsova and
Simon 2012).
KEY TO THE SUBFAMILIES
Despite the advantages of a phylogenetic clas-
sification, the grass classification illustrates one 1. Pseudopetioles present 2
serious disadvantage. The groups shown by 1. Leaves without pseudopetioles 11
molecular data to be monophyletic often do not 2 (1). Conventional spikelets absent
have obvious morphological characters by which I. Anomochlooideae
they can be reliably diagnosed. The subfamily 2. Conventional spikelets present 3
3 (2). Leaf blades resupinate, major veins at an oblique
Pooideae is a case in point. Most species in the angle to the midrib 4
subfamily lack culm branches, lack microhairs, 3. Leaf blades not resupinate, major veins parallel to the
have parallel-sided subsidiary cells in the sto- midrib 5
mata, have laterally compressed spikelets, disar- 4 (3). Stigmas three; leaf blades with microhairs; inflores-
ticulate above the glumes, have a chromosome cence branches and spikelets covered with hooked
hairs II. Pharoideae
complement based on seven pairs of large chro- 4. Stigmas two; leaf blades without microhairs; inflores-
mosomes, and have inflorescences in which the cence branches and spikelets smooth
primary branches initiate in two ranks rather VI. Pooideae (Phaenosperma)
than a spiral. When these characters are mapped 5 (3). Culms woody, usually well over 1 m tall; leaves
on the phylogeny, however, none actually appears dimorphic, the culm leaves with prominent sheaths
and blades reduced or absent, the foliage leaves with
at the origin of the subfamily, and thus none is broad blades
synapomorphic or fully diagnostic. They become V. Bambusoideae (Bambuseae and Arundinarieae)
handy generalizations, but not infallible guides to 5. Culms herbaceous; if lignified and over 1 m tall, then
history. The pattern tells us a great deal about the leaves not dimorphic 6
130 Subdivision of the Family
6 (5). Inflorescence axis flattened, leaf-like, with the spi- 19 (16). Glumes as long as the flowers or longer
kelets borne on one side IX. Danthonioideae
IV. Ehrhartoideae (Phyllorachideae) 19. Glumes generally shorter than the flowers 20
6. Inflorescence axis not flattened 7 20 (19)1. Plants C4, with veins separated by no more than
7 (6). Spikelets with exactly two flowers, the distal one two mesophyll cells
bisexual and the proximal one staminate or sterile; X. Chloridoideae
rachilla extension lacking 20. Plants C3, with veins separated by more than two
VIII. Panicoideae (Paspaleae, Paniceae, Andropogo- mesophyll cells
neae) XII. Arundinoideae plus several Panicoideae (Cen-
7. Spikelets with 1 or more flowers; if exactly two, then not totheca, Megastachya, Cyperochloa, Spartochloa, Chas-
as above 8 manthium)
8 (7). Stamens 6 III. Puelioideae
8. Stamens 2 or 3 (numerous in Pariana), not 6 9
9 (8). Spikelets with 1 flower, unisexual
V. Bambusoideae (Olyreae) References
9. Spikelets with more than 1 flower, generally bisexual 10
10 (9). Style branches and stigmas growing into long Clayton, W.D., Renvoize, S.A. 1986. Genera graminum:
tangled tendrils that harden at maturity grasses of the world. London: Her Majesty’s Statio-
8. Streptogyna (BEP clade Incertae sedis) nery Office.
10. Style branches and stigmas not as above Clayton, W.D., Harman, K.T., Williamson, H. 2006
VIII. Panicoideae (Chasmanthieae and Centotheceae) onward. GrassBase - the online world grass flora.
11 (1). Spikelets with exactly two flowers; rachilla exten- http://www.kew.org/data/grasses-db.html
sion lacking 12 GPWG 2001. Grass Phylogeny Working Group. Phylogeny
11. Spikelets with 1 to several flowers, if two then rachilla and subfamilial classification of the Poaceae. Ann.
extension present 14 Missouri Bot. Gard. 88: 373–457.
12 (11). Upper flower of spikelet bisexual, the proximal GPWG II 2012. Grass Phylogeny Working Group II. New
one staminate or sterile grass phylogeny resolves deep evolutionary relation-
VIII. Panicoideae (Paspaleae, Paniceae, Andropogo- ships and discovers C4 origins. New Phytol. 193:
neae) 304–312.
12. Both flowers of spikelet fertile 13 Sánchez-Ken, J.G., Clark, L.G., Kellogg, E.A., Kay, E.E.
13 (12). Plants several meters tall, reed-like; dioecious 2007. Reinstatement and emendation of subfamily
Micrairoideae (Poaceae). Syst. Bot. 32: 71–80.
Gynerium
Simon, B.K. 2007. GrassWorld: Interactive key and
13. Plants less than 1 m tall, often tiny; flowers bisexual information system of world grasses. Kew Bull. 62:
XI. Micrairoideae 475–484.
14 (11). Ligule membranous 15 Simon, B.K., Clayton, W.D., Harman, K.T., Vorontsova,
14. Ligule a fringe of hairs or a fringed membrane 16 M.S., Brake, L., Healy, D., Alfonso, Y. 2011 onward.
15 (14).1 Lodicules membranous; plants lacking two- GrassWorld. http://grassworld.myspecies.info/
celled microhairs; stomatal subsidiary cells with paral- Soreng, R.J., Davidse, G., Peterson, P.M., Zuloaga, F.O.,
lel sides; leaf veins widely spaced; plants C3 Judziewicz, E.J., Filgueiras, T.S., Morrone, O.,
VI. Pooideae Romaschenko, K. 2012 and onward. A world-wide
15. Lodicules fleshy; plants with two-celled microhairs; phylogenetic classification of Poaceae (Gramineae).
stomatal subsidiary cells dome shaped or triangular; http://www.tropicos.org/docs/meso/CLASSIFICATION
leaf veins closely spaced; plants C4 X. Chloridoideae %20OF%20world%20grasses%202013%20Oct%2029.
16 (14). Spikelets with one fertile flower 17 htm
16. Spikelets with more than one flower 19 Vorontsova, M.S., Simon, B.K. 2012. Updating classifica-
17 (16). Lemma terminating in three awns, their bases tions to reflect monophyly: 10 to 20 percent of spe-
fused to form a column VII. Aristidoideae cies names change in Poaceae. Taxon 61: 735–746.
17. Lemma awnless or with a single awn; if more than one Watson, L., Dallwitz, M.J. 1992 onward. The grass genera
awn then these not fused at their bases 18 of the world: descriptions, illustrations, identifica-
18 (17). Fertile flower subtended by two sterile flowers, tion, and information retrieval; including synonyms,
morphology, anatomy, physiology, phytochemistry,
these often reduced to bract-like structures cytology, classification, pathogens, world and local
IV. Ehrhartoideae distribution, and references, 23rd April 2010 edn.
18. Fertile flower without proximal sterile flowers Wallingford: CAB International. http://delta-intkey.
X. Chloridoideae com/grass/
1
Use of macromorphological characters to distinguish these subfamilies would require developing a key to the level of genera
or small clades. In terms of overall plant structure and spikelet morphology, Chloridoideae are quite similar to several other
subfamilies.
I. Subfamily Anomochlooideae Pilg. ex Potztal (1957)
Rhizomatous perennials, herbaceous. Ligule relationship in phylogenetic trees is an artifact.

fringed or lacking. Leaves with a pulvinus at the Nonetheless, subsequent phylogenetic analyses
distal end of the pseudopetiole, with broad continue to place them as sister taxa (GPWG II
blades. Inflorescence made up of units that defy 2012).
conventional descriptions, generally known as Although each genus in this subfamily has
“spikelet equivalents,” more or less equivalent to been placed in its own tribe, division of such a
a single flower, bisexual. The spikelet equivalents small subfamily seems unnecessary.
sessile, bracteate or not. Additional bract-like
structures arranged in unusual phyllotaxis, but
probably representing highly modified perianth. KEY TO THE GENERA OF ANOMOCHLOOIDEAE
Caryopsis with a hard endosperm, with the
hilum long but inconspicuous, the embryo with 1. Leaves basal, with long pseudopetioles; stamens sur-
or without an epiblast, with a scutellar cleft, with- rounded by a fringe like structure outside of which are
two bracts; bract closest to the floral organs leaf-like,
out a mesocotyl internode, embryonic leaf mar-
with sheath and blade 1. Anomochloa
gins overlapping or not. 1. Leaves borne along the culm, with short pseudope-
Tropical America. Two genera and four spe- tioles; bracts subtending the flower 11; bracts closest
cies. to the floral organs without obvious sheath and blade,
Judziewicz and Soderstrom (1989) suggest but one with a long coiled awn 2. Streptochaeta
that evidence for monophyly of the Anomo- 1. Anomochloa Brongniart Fig. 10
chlooideae is provided by their remarkably large Anomochloa Brongniart, Ann. Sci. Nat. Bot. III, 16: 368
microhairs (75–150 m long), which have a con- (1851).
striction part way up the basal cell, by their dis-
tinctive midrib structure, in which the midrib Leaves broad, looking more like a member of the
projects above the surface of the blade on both Zingiberales than a grass. Ligule a fringe of hairs.
the ab- and adaxial sides, and by their silica Pulvinus at both ends of the pseudopetiole. Inflo-
bodies elongated perpendicular to the proximo- rescence complex, spike-like; each branch sub-
distal axis of the leaf. However, because they tended by an enveloping bract. Flowers
compared Anomochloa and Streptochaeta only subtended by two additional bracts that are sepa-
to the Bambusoideae and select Oryzeae, it is rated by an internode, the upper bract extended
not clear whether these characters are truly syna- into a narrow curved leaf blade. Stamens 4, sur-
pomorphic. The GPWG (2001) questioned rounded by a ring of cilia. Stigma 1. Embryo with
whether any synapomorphies link the two genera, an epiblast, leaf margins meeting. 2n ¼ 36.
and raised the possibility that their sister group One sp., A. marantoidea Brongn., Bahia,
Brazil.
132 I. Subfamily Anomochlooideae Pilg. ex Potztal (1957)
This is one of the most curious of the grasses,

based on its phylogenetic position and its odd
morphology. It is rare and nearly extinct in the
wild and difficult to maintain in cultivation. It has
been the subject of several careful developmental
and anatomical studies, which describe in detail
the nature of the floral parts and subtending
bracts (Judziewicz et al. 1999; Judziewicz and
Soderstrom 1989; Sajo et al. 2012). Establishment
of homology with spikelets of other grasses is
difficult and likely to remain controversial.
2. Streptochaeta Nees Figs. 11a, 11b

Streptochaeta Nees, Fl. Bras. Enum. Pl. 2: 536 (1829).
Leaves with auricles; ligule absent. Inflorescence

spike-like, with spirally arranged, contracted lat-
eral branches, each bearing 11 spirally arranged
bracts with varying morphologies, the sixth bract
ending in a long coiled awn. Awns becoming
entangled at maturity. Embryo without an epi-
blast, leaf margins overlapping. 2n ¼ 22.
Three spp., Central and South America.
The development and possible homologies of
the “spikelet equivalents” are discussed in detail
by Preston et al. (2009) and Sajo et al. (2008).
SPIKELET CLADE
All remaining subfamilies have flowers arranged

in spikelets, tiny spike-like structures subtended
by two bracts (the glumes) (GPWG 2001). See
discussion under Inflorescence structure.
teoles, the bract between bracteoles and axis removed. D

Caryopsis, view from the adaxial side showing linear hilum.
E Portion of flower; upper bract partially dissected away,
showing ovary surrounded by ring of cilia and with anoma-
lous bract found in one spikelet, stamens removed. F Detail
of bracts surrounding the flower, deciduous lower bract and
Fig. 10. Anomochlooideae. Anomochloa marantoidea. A upper bract separated by an internode, the blade-like exten-
Portion of flowering part of the plant, showing the lowest and sion fallen from the latter. G Longitudinal section of flower,
next to lowest bracts subtending spikelet equivalents. B Base lower bract removed, with thickened internode supporting
of inflorescence branch, viewed from the adaxial side, sub- upper bract, ring of cilia, stamens, and gynoecium. (Rep-
tending bract removed. C Basal portion of primary axis, rinted from Judziewicz and Soderstrom 1989, courtesy of
showing small, lower (left) and large, upper (right) brac- Smithsonian Institution; drawn by A.R. Tangerini)
References 133
Fig. 11a. Anomochlooideae. Streptochaeta spicata subsp. Fig. 11b. Anomochlooideae. Streptochaeta spicata subsp.
spicata. A Habit, showing disarticulated spikelet equiva- spicata, showing details of spikelet equivalent. A Portion
lents hanging by awns from the apex of the rachis (note of rachis with pedicel of spikelet-equivalent. B Spikelet
lateral branch of culm terminating in a secondary inflo- equivalent, showing long awn on bract VI. C–G Bracts I–V
rescence). B Base of plant showing stout sympodial rhi- at the base of the spikelet equivalent, showing their vari-
zomes. C Summit of leaf sheath and base of blade showing able venation and shape. H Saccate base of bract VI,
prominent sheath auricle. D Opposite side of summit of showing the internode between bract V and bract VI. I
leaf sheath and base of blade with smaller sheath auricle Bracts VII and VIII. J–L Bracts X–XII. M Androecium
and turgid pulvinus. (Reprinted from Judziewicz and showing short, elongating stamen tube. N Gynoecium. O
Soderstrom 1989, courtesy of Smithsonian Institution; Caryopsis, view from adaxial side, showing linear hilum. P
drawn by G.B. Threlkeld). Same, from abaxial side, showing persistent style and
small, basal embryo. (Reprinted from Judziewicz and
Soderstrom 1989, courtesy of Smithsonian Institution;
drawn by G.B. Threlkeld)
References
GPWG 2001. Grass Phylogeny Working Group. Phylogeny Preston, J.C., Christensen, A., Malcomber, S.T., Kellogg, E.
and subfamilial classification of the Poaceae. Ann. A. 2009. MADS-box gene expression and implica-
Missouri Bot. Gard. 88: 373–457. tions for developmental origins of the grass spikelet.
GPWG II 2012. Grass Phylogeny Working Group II. New Amer. J. Bot. 96: 1419–1429.
grass phylogeny resolves deep evolutionary relation- Sajo, M.G., Longhi-Wagner, H.M., Rudall, P.J. 2008.
ships and discovers C4 origins. New Phytol. 193: Reproductive morphology of the early-divergent
304–312. grass Streptochaeta and its bearing on the homolo-
Judziewicz, E.J., Soderstrom, T.R. 1989. Morphological, gies of the grass spikelet. Plant Syst. Evol. 275:
anatomical, and taxonomic studies in Anomochloa 245–255.
and Streptochaeta (Poaceae: Bambusoideae). Smith- Sajo, M.G., Pabón-Mora, N., Jardim, J., Stevenson, D.W.,
sonian Contr. Bot. 68: 1–52. Rudall, P.J. 2012. Homologies of the flower and inflo-
Judziewicz, E.J., Clark, L.G., Londoño, X., Stern, M.J. 1999. rescence in the early-divergent grass Anomochloa
American bamboos. Washington, DC: Smithsonian (Poaceae). Amer. J. Bot. 99: 614–628.
Institution Press.
II. Subfamily Pharoideae L.G. Clark & Judz. (1996)
Rhizomatous perennials, herbaceous. Leaf blades onward). However, it appears not to have pharoid
resupinate, with pseudopetiole; veins running leaf anatomy, and may not have the resupinate
obliquely from midrib to margin. Inflorescence leaf blades that characterize the rest of the sub-
branched, with hooked hairs on the branches family (Judziewicz and Clark 2007). Following the
and the spikelets. Spikelets unisexual, with one suggestion of Judziewicz and Clark (2007), it is
flower per spikelet, staminate and pistillate spike- placed in Ehrhartoideae.
lets paired, the staminate one pedicellate and the
pistillate one sessile. Glumes 2. Lemmas tubular
or inflated. Lodicules generally absent, or present KEY TO THE GENERA OF PHAROIDEAE
in staminate flowers only. Caryopsis with hard
endosperm, with the hilum extending the full 1. Lemma margins fused, the resulting structure inflated 2
length of the caryopsis, the embryo with an epi- 1. Lemma margins free; if fused, the structure not inflated
blast, with or without a scutellar tail, without a 4. Pharus
2 (1). Ligule membranous, inflorescence disarticulating
mesocotyl internode, the embryonic leaf margins from plant 5. Scrotochloa
overlapping. Inner bundle sheath of veins with 2. Ligule a fringed membrane, inflorescence rachis tough
multiple layers of cells. Intercostal epidermis 3. Leptaspis
with files of long fibers alternating with normal
files of intercostal long cells.
Three genera and 12 species. GENERA OF PHAROIDEAE
Both Scrotochloa and Pharus have a fragile
panicle, which could be a synapomorphy linking 3. Leptaspis R. Br.
the two genera; however, utriculate female
lemmas are present in Leptaspis and Scrotochloa, Leptaspis R. Br., Prodr. Fl. Nov. Holl.: 211 (1810).
and could be a synapomorphy for that pair. This
character conflict prevents determining which Ligule a fringed membrane. Inflorescence
characters are synapomorphic. Virtually all branches not disarticulating. Female lemmas
molecular data on this subfamily come from urceolate, the margins fused, becoming indurate.
Pharus, which has been studied extensively. Lodicules present or absent. 2n ¼ 24.
Good comparative data on Leptaspis and Scroto- Three spp., Old World tropics.
chloa would be particularly valuable. The embry-
onic characters have only been sampled for 4. Pharus P. Browne Fig. 12
Pharus, but are presumed to apply to the whole Pharus P. Browne, Civ. Nat. Hist. Jamaica: 344 (1756).
subfamily. The presence of epidermal fibers is
unusual and may be unique in Poaceae (Soder- Ligule membranous, a fringed membrane, or a
strom et al. 1987). The genus Suddia has been fringe of hairs. Inflorescence branches or the
placed in this subfamily by Clayton and Renvoize whole inflorescence breaking away from the
(1986) and Watson and Dallwitz (1992 and plant. Female lemmas cylindrical, in some species
136 II. Subfamily Pharoideae L.G. Clark & Judz. (1996)
with a tiny beak; covered with hooked hairs. Lodi-

cules sometimes present in staminate flowers.
2n ¼ 24.
Seven spp., Central and South America, West
Indies.
The margins of the lemma are fused in one
species of Pharus.
5. Scrotochloa Judziewicz
Scrotochloa Judziewicz, Phytologia 56: 299–304 (1984).
Ligule membranous, pubescent. Inflorescence

branched, falling entire. Female lemmas urceo-
late, the margins fused, becoming indurate, cov-
ered with hooked hairs. Lodicules absent.
Caryopsis with a groove on the side of the
hilum. Invaginations of mesophyll cell walls
obscure.
Two spp., Ceylon, SE Asia, New Guinea, Sol-
omon Is.
Watson and Dallwitz (1992 onward) report
that this genus has silica bodies with a raised
prickly belt around the equator; this characteris-
tic is not known elsewhere in the grasses.
BISTIGMATIC CLADE
Stigmas two. Orders of stigmatic branching two.

Spikelets with multiple flowers. Anther walls with
a middle layer that breaks down during develop-
ment; the inner walls of endothecial cells becom-
ing fibrous at maturity.
All remaining subfamilies are members of a
large group in which the characters listed above
are synapomorphic (GPWG 2001). Stigma num-
ber reverses to three in some species of Puelia,
and in the Bambusoideae. Flower number per
spikelet has reversed to one in multiple lineages.
Fig. 12. Pharoideae. Pharus glaber. A Flowering plant. B

Leaf base showing pseudopetiole. C Pistillate spikelet
(left) with uncinate trichomes, and staminate spikelet
(right). (From Barkworth et al. 2007, drawn by S. Long)
References 137
Judziewicz, E.J., Clark, L.G. 2007. Classification and bio-

References geography of New World grasses: Anomochlooideae,
Pharoideae, Ehrhartoideae, and Bambusoideae. Aliso
Barkworth, M.E., Capels, K.M., Long, S., Anderton, L.K., 23: 303–314.
Piep, M.B. (eds.) 2007. Flora of North America north Soderstrom, T.R., Ellis, R.P., Judziewicz, E.J. 1987. The
of Mexico. Vol. 24. Magnoliophyta: Commelinidae Phareae and Streptogyneae (Poaceae) of Sri Lanka.
(in part): Poaceae, part 1. New York: Oxford Univer- Smithsonian Contr. Bot. 65: 1–27.
sity Press. Watson, L., Dallwitz, M.J. 1992 onward. The grass genera
Clayton, W.D., Renvoize, S.A. 1986. Genera graminum: of the world: descriptions, illustrations, identifica-
grasses of the world. London: Her Majesty’s Statio- tion, and information retrieval; including synonyms,
nery Office. morphology, anatomy, physiology, phytochemistry,
GPWG 2001. Grass Phylogeny Working Group. Phylogeny cytology, classification, pathogens, world and local
and subfamilial classification of the Poaceae. Ann. distribution, and references, 23rd April 2010 edn.
Missouri Bot. Gard. 88: 373–457. Wallingford: CAB International. http://delta-intkey.
com/grass/
III. Subfamily Puelioideae L.G. Clark et al. (2000)
Rhizomatous perennials, herbaceous, the culms 7. Puelia Franch.

hollow. Leaves with broad blades, with pseudo- Puelia Franch., Bull Mens. Soc. Linn. Paris 1: 674 (1887).
petioles, without auricles, the ligule a fringed
Abaxial ligule present. Spikelets with 2 or 3 (up to
membrane. Inflorescences branched or
7) glumes; distal flower pistillate, proximal 3 to 6
unbranched, sometimes with minute bracts sub-
flowers staminate. Pistillate flower on a long
tending the branches. Spikelets with multiple
internode and subtended by a broad outgrowth
flowers, the proximal several staminate or sterile,
of the rachilla. Filaments of stamens fused. Peri-
the distal ones pistillate or bisexual. Stigmas 2 or
carp free from seed coat. 2n ¼ 24.
3. Caryopsis with a long, linear hilum, the embryo
Five spp., western Africa.
small, its detailed structure unknown.
Two genera and eleven species; tropical Africa.
This subfamily is poorly known. Material is
difficult to acquire and the plants are rarely if ever BEP PLUS PACMAD CLADES
found in cultivation. Presence of proximal stami-
nate or neuter flowers in the spikelet may be Leaves lacking a pseudopetiole. Lodicules 2,
uniquely derived for this group, but the character stamens 3. Mesophyll without fusoid cells.
appears in several other grass taxa, most notably These clades include all the remaining sub-
in the Panicoideae (GPWG 2001). Although each families and thus the overwhelming majority of
genus has been placed in its own tribe, division of the species of the family. The characters listed
such a small subfamily seems unnecessary. were suggested by the GPWG (2001) as possible
synapomorphies for this clade, although all of
them reverse elsewhere in the tree. In addition,
KEY TO THE GENERA the GPWG (2001) suggested that loss of invagi-
nated cells walls might be synapomorphic. How-
1. Abaxial ligule absent; spikelets with short rachilla inter- ever, optimization of this character on recent
nodes, more than six flowers, the distalmost 5–12 trees is ambiguous. These characters will need
bisexual 6. Guaduella to be optimized on the most recent grass phylog-
1. Abaxial ligule present; spikelets with long rachilla inter-
nodes; flowers six or fewer, the distalmost one pistil- eny (GPWG II 2012).
late 7. Puelia
6. Guaduella Franch. Fig. 13
BEP CLADE
Guaduella Franch., Bull. Mens. Soc. Linn. Paris 1: 676
(1887).
This clade has no known morphological synapo-
Spikelets with two to four glumes; distal 5 to 12 morphies.
flowers bisexual, proximal 1 to 3 staminate. Stig- The BEP clade was first identified by Clark
mas 2. et al. (1995) (as the BOP clade), and includes
Six spp., western Africa. Bambusoideae, Ehrhartoideae, and Pooideae.
140 III. Subfamily Puelioideae L.G. Clark et al. (2000)
Early phylogenetic analyses provided little sup-

port for the group (e.g., GPWG 2001; Kellogg and
Linder 1995), but recent data show that the group
is monophyletic (Bouchenak-Khelladi et al. 2008;
Edwards and Smith 2010; GPWG II 2012; Vice-
ntini et al. 2008). The amphi-Atlantic genus Strep-
togyna has been difficult to place, and its variable
position in phylogenies has contributed to low
statistical support for the BEP clade and for rela-
tionships within it. All possible relationships of
the three subfamilies have been suggested, but
increasing support is gathering for the topology
[(Ehrhartoideae) (Bambusoideae, Pooideae)]
(GPWG II 2012; Zhao et al. 2013). The position
of Streptogyna remains ambiguous but it may be
sister to Ehrhartoideae.
BEP clade incertae sedis:
8. Streptogyna P. Beauv. Fig. 14
Streptogyna P. Beauv., Ess. Agrost. 80 (1812).
Rhizomatous or caespitose perennials. Leaf blades

with a pseudopetiole, deciduous, the ligule mem-
branous, abaxial ligule present. Inflorescence nar-
row, branched. Spikelets with several flowers. Distal
flowers reduced. Rachilla breaking up between flow-
ers, the internode developing into a hook. Lemma
awned. Lodicules 3. Stamens 2. Style branches and
stigmas 2 or 3, extending and twisting together at
maturity, becoming hard. Caryopsis with a long
Fig. 13. Puelioideae. Guaduella oblonga. A Rhizome. B linear hilum. Mesophyll cells with or without inva-
Flowering culm. C Portion of spikelet. D Lower glume. E ginated cell walls, mesophyll with fusoid cells.
Upper glume. F Lemma and palea. G Lemma. H and I 2n ¼ 24.
Palea. J Flower. K Lodicule and two stamens. (From Clay- Two spp., tropical America, Africa, Sri Lanka,
ton 1972, drawn by S. Ross-Craig)
India.
BEP CLADE 141
Fig. 14. Incertae sedis. Streptogyna crinita. A Base of lodicule at left and others the anterior lodicules. L
plant showing sympodial habit of rhizomes. B Leaf por- Androecium of 2 stamens with ribbon-like filaments and
tion of mature culms terminating in young inflorescence. gynoecium showing the hairy ovary and style and 2 stig-
C Mature inflorescence. D Ligular region, showing inner mas. M Section of stigma showing retrorse scabrosity. N
and outer ligules. E Young spikelets. F Glumes. G Spikelet Caryopsis, showing small embryo at base and persistent
without glumes. H Section of lemma awn showing hairy apex of ovary. O Caryopsis, showing narrow, elon-
antrorse scabrosity. I Base of lower fertile flower and gated hilum, and persistent hairy apex of ovary. (Rep-
succeeding rachilla segment. J Palea of fertile flower with rinted from Soderstrom et al. 1987, courtesy of
succeeding rachilla segment. K Lodicules, the posterior Smithsonian Institution; drawn by A.R. Tangerini)
142 III. Subfamily Puelioideae L.G. Clark et al. (2000)
GPWG II 2012. Grass Phylogeny Working Group II. New

References grass phylogeny resolves deep evolutionary relation-
ships and discovers C4 origins. New Phytol. 193:
Bouchenak-Khelladi, Y., Salamin, N., Savolainen, V., For- 304–312.
est, F., van der Bank, M., Chase, M.W., Hodkinson, T. Kellogg, E.A., Linder, H.P. 1995. Phylogeny of Poales. In:
R. 2008. Large multi-gene phylogenetic trees of the Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J.
grasses (Poaceae): progress towards complete tribal (eds.) Monocotyledons: systematics and evolution.
and generic taxon sampling. Mol. Phyl. Evol. 47: Kew: Royal Botanic Gardens. pp. 511–542
488–505. Soderstrom, T.R., Ellis, R.P., Judziewicz, E.J. 1987. The
Clark, L.G., Zhang, W., Wendel, J.F. 1995. A phylogeny of Phareae and Streptogyneae (Poaceae) of Sri Lanka.
the grass family (Poaceae) based on ndhF sequence Smithsonian Contr. Bot. 65: 1–27.
data. Syst. Bot. 20: 436–460. Vicentini, A., Barber, J.C., Giussani, L.M., Aliscioni, S.S.,
Clayton, W.D. 1972. Gramineae. In: Hepper, F.N. (ed.) Kellogg, E.A. 2008. Multiple coincident origins of C4
Flora of West Tropical Africa, 2nd edn. Vol. 3, part photosynthesis in the Mid- to Late Miocene. Global
2. London: Crown Agents. p. 361 Change Biol. 14: 2963–2977.
Edwards, E.J., Smith, S.A. 2010. Phylogenetic analyses Zhao, L., Zhang, N., Ma, P.-F., Liu, Q., Li, D.-Z., Guo, Z.-H.
reveal the shady history of C4 grasses. Proc. Natl. 2013. Phylogenomic analyses of nuclear genes reveal
Acad. Sci. USA 107: 2532–2537. the evolutionary relationship within the BEP clade
GPWG 2001. Grass Phylogeny Working Group. Phylogeny and the evidence of positive selection in Poaceae.
and subfamilial classification of the Poaceae. Ann. PLoS ONE 8: e64642.
Missouri Bot. Gard. 88: 373–457.
IV. Subfamily Ehrhartoideae Link (1827)
Rhizomatous, stoloniferous, or caespitose peren- flowers in some taxa. Depending on the species,
nials, herbaceous or with a woody caudex. Ligule the rachilla may be elongated. In species in which
membranous, a fringed membrane, or a fringe of the glumes and sterile lemmas are missing (e.g.,
hairs. Inflorescences branched or unbranched, Hygroryza, Chikusiochloa), the one fertile flower
the spikelets pedicellate. Spikelets with glumes then appears on a long stipe. Maturation of the
0 or 2, with the two proximal flowers sterile and spikelet is basipetal, and ectopic expression of
often reduced, only the distal flower fertile. Flow- some floral identity genes affects the sterile
ers bisexual or unisexual. Lemma with or without lemmas but not the true glumes (Komatsu et al.
an awn. Lodicules 2. Caryopsis with hard endo- 2003; Malcomber and Kellogg 2004, 2006;
sperm, with a long-linear hilum, the embryo with Malcomber et al. 2006).
an epiblast, with a scutellar cleft, without a meso- Silica bodies of this subfamily are distinc-
cotyl internode, with the embryonic leaf margins tive and have been called “oryzoid” (Metcalfe
overlapping. Fusoid cells absent or present, 1960). They occur primarily over the veins, and
mesophyll cells with or without invaginated cell are elongated transverse to the proximo-distal
walls, midrib simple or complex. Silica bodies in axis of the leaf. In addition, more conventional
leaves transverse to the proximo-distal axis. Pho- bilobate silica bodies occur parallel to the prox-
tosynthetic pathway C3. Worldwide distribution. imo-distal axis. Most members of the subfamily
Fifteen genera and 112 spp. are perennial and this is likely the ancestral
Recent literature on this subfamily is sum- state, with the annual habit having arisen sev-
marized by Kellogg (2009); little is known about eral times independently (Kellogg 2009). All
Phyllorachideae, whereas Oryzeae have been Oryzeae and the early-diverging Ehrharteae are
studied in great depth (see below). The spikelet plants of moist habitats, suggesting that this is
structure is unique to the subfamily, and has led the ancestral environment. Oryzeae and Ehrhar-
to some misleading terminology in the literature. teae are thought to have diverged about 35
In some genera, particularly Oryza, the two million years ago, during a period of global
glumes are so reduced that they are either ignored cooling and a reduction in atmospheric CO2
or called “rudimentary glumes”, while the (Vicentini et al. 2008), although data on phyto-
lemmas of the two sterile flowers are inaccurately lith assemblages in India suggest that the tribe
called “empty glumes”. Molecular genetic studies might be twice as old (Prasad et al. 2011) (see
show that the sterile lemmas in fact express some section on Fossil record). The common ances-
genes that prevent them from developing like the tor of Ehrhartoideae may have been native to
glumes and other genes that prevent them from what is now the Old World with several inde-
developing like the fertile lemma (Ren et al. pendent dispersal events to North America
2013); thus, the sterile lemmas do share aspects (Kellogg 2009; Tang et al. 2010).
of both lemma and glume identity and constitute The chromosome base number of most spe-
a novel organ. The situation is further compli- cies of Ehrhartoideae is x ¼ 12, although Zizania
cated by loss of one or more glumes and sterile is unusual in having 2n ¼ 30 or 34.
144 IV. Subfamily Ehrhartoideae Link (1827)
KEY TO THE GENERA Plants 1–3 m tall, with spongy culms. Leaves up to
1 m long, the ligule membranous, with a pseudo-
1. Inflorescence axis terete or angled, but not broad and petiole, the blades sagittate. Inflorescence
leaf-like 2 branches verticillate. Glumes lacking. Mesophyll
1. Inflorescence axis broad, flattened, leaf-like, enclosing
the primary branches and spikelets 14 cells without invaginated cell walls. Lemma with
2 (1). Plant scrambling; apices of some leaves extended prominent veins, the apex acute. Stamens 4.
into long tendrils 16. Prosphytochloa One sp., S. sagittifolia Renvoize, wet sites in Sudan.
2. Plant habit various, but not scrambling and leaf apices Suddia has been placed in Pharoideae by
not tendril-like 3 Clayton and Renvoize (1986) and Watson and
3 (2). Spikelets with two well developed glumes, two sterile
lemmas, and one bisexual flower 12. Ehrharta Dallwitz (1992 and onward). However, it appears
3. Spikelets with glumes minute or absent, sterile lemmas not to have pharoid leaf anatomy, and may not
absent or reduced to small subulate structures, and have the resupinate leaf blades that characterize
one bisexual or unisexual flower 4 the rest of the subfamily (Judziewicz and Clark
4 (3). Spikelets with two sterile lemmas, these often subulate 5 2007). Following the suggestion of Judziewicz and
4. Spikelets lacking sterile lemmas, thus consisting of a
single flower with its lemma and palea 8 Clark (2007), it is placed incertae sedis in Ehrhar-
5 (4). Leaf blades with a pseudopetiole 14. Maltebrunia toideae pending more morphological and molec-
5. Leaf blades without a pseudopetiole 6 ular data.
6 (5). Distal part of lemma with an attenuate beak
20. Rhynchoryza
6. Distal part of lemma awned or not, but not with a beak 7
7 (6). Fertile lemma and palea hardened, with rough I. TRIBE PHYLLORACHIDEAE C.E. Hubbard
tuberculate outgrowths, keeled 15. Oryza (1939)
7. Fertile lemma and palea chartaceous or membranous,
not keeled 19. Potamophila
8 (4). Fertile flower on a long stipe 9 Monoecious perennials with hard culms. Leaf
8. Fertile flower not on a stipe 10 blade cordate to sagittate, with a pseudopetiole,
9 (8). Leaf blades elliptic and floating; plants of India and the ligule a ciliolate membrane. Inflorescence axis
Sri Lanka 18. Hygroryza flattened, broad and leaf-like, enclosing the short
9. Leaf blades linear; plants of southeast Asia
17. Chikusichloa primary branches; primary branches each bearing
10 (8). Spikelets strongly laterally compressed, keeled; a condensed cluster of 1 to 4 spikelets. Spikelets
flowers bisexual 13. Leersia with two flowers, the distal one unisexual, the
10. Spikelets terete or only weakly compressed, not proximal one sterile. Glumes distinct, the lower
keeled; flowers unisexual 11 one subulate and the upper oblong. Mesophyll
11 (10). Lemma unawned 12
11. Lemma awned 13 cells with invaginated cell walls.
12 (11). Inflorescence branches verticillate; spikelets Two genera and three species.
bisexual; plants of Sudan 9. Suddia Phyllorachideae are placed in Ehrhartoideae
12. Inflorescence branches not verticillate; spikelets uni- based on morphology plus a DNA sequence of the
sexual; plants of the New World 22. Luziola rpl16 intron for Humbertochloa greenwayii
13 (11). Fruit ovoid, an achene, the seed coat free from the
pericarp; staminate and pistillate spikelets on the same (Zhang 2000). Additional information on this
inflorescence branches 23. Zizaniopsis pair of genera would be valuable.
13. Fruit linear, a caryopsis, the seed coat fused to the
pericarp; staminate and pistillate spikelets separate in 10. Humbertochloa A. Camus & Stapf
the inflorescence 21. Zizania
14 (1). Staminate and pistillate spikelets on the same Humbertochloa A. Camus & Stapf, Bull. Soc. Bot. France
inflorescence branch 11. Phyllorachis 81: 467 (1934).
14. Staminate and pistillate spikelets in separate inflores-
cences 10. Humbertochloa Inflorescences unisexual. Lower lemma of pistil-
late spikelet rounded; upper lemma tuberculate.
Two spp., Tanzania and Madagascar.
EHRHARTOIDEAE INCERTAE SEDIS 11. Phyllorachis Trimen. Fig. 15

Phyllorachis Trimen., J. Bot. 17: 353 (1879).
9. Suddia Renvoize
Primary branches of the inflorescence each with
Suddia Renvoize, Kew Bull. 39: 455 (1984). one pistillate spikelet and two or three staminate
II. TRIBE EHRHARTEAE Nevski (1937) 145
spikelets. Lower lemma of pistillate spikelet

grooved; upper lemma smooth. 2n ¼ 24.
One sp., P. sagittata Trimen, equatorial Africa.
II. TRIBE EHRHARTEAE Nevski (1937)
Annuals or perennials, rhizomatous, stoloniferous,

or caespitose, sometimes suffrutescent. Leaves
with or without auricles, the ligule membranous
or a fringed membrane. Inflorescence branched or
not. Spikelets with one fertile flower, subtended by
two empty lemmas plus two glumes. Stamens 2, 3,
4, or 6. Epiblast absent. Mesophyll cells with or
without invaginated cell walls; fusoid cells absent.
Vascular system of the midrib simple.
One genus and thirty-seven species.
12. Ehrharta Thunb. Fig. 16

Ehrharta Thunb., Vet. Acad. Handl. Stockholm 40: 217
(1779); phylog. (Verboom et al. 2003).
Tetrarrhena R. Br., Prodr. Fl. Nov. Holl. 209 (1810).
Microlaena R. Br., Prodr. Fl. Nov. Holl. 210 (1810).
Petriella Zotov, Trans. Roy. Soc. New Zealand 73: 235 (1943).
Zotovia Edgar & Connor, New Zealand J. Bot. 36: 569 (1998).
Characters of the tribe. 2n ¼ 24, 48.
Thirty-seven spp., southern Africa, Southeast
Asia, New Zealand.
Recognition of this group as a single genus is
supported by the work of Verboom et al. (2003),
who found that Microlaena is polyphyletic, and
that recognition of Tetrarrhena and Zotovia make
Ehrharta paraphyletic. Stamen number is
reduced to four in the clade that includes species
formerly in Tetrarrhena plus Ehrharta multinoda
(as Microlaena polynoda) and E. stipoides
(as M. stipoides). Stamen number is reduced to
three in the clade of E. delicatula and E. triandra,
and independently to two in the clade that
Fig. 15. Ehrhartoideae-Phyllorhachideae. Phyllorhachis includes E. colensoi (¼ Zotovia colensoi),
sagittata. A Culm with axillary and terminal inflores- E. thomsonii (¼ Z. thomsonii), E. diplax (¼ M.
cences. B Detail from underside of leaf blade showing
venation. C Base of leaf blade. D Raceme showing male avenacea), E. tasmanica (¼ M. tasmanica) and E.
and female spikelets, secondary rachis (a), vestigial spike- avenacea. The lack of fusoid cells and the simple
let (b) and lower glume of female spikelet (c). E, F Sec- midrib anatomy might be synapomorphic; under
ondary rachis, front and back views, showing vestigial
spikelets at a and b. G Axillary inflorescence, showing
this interpretation, these two characters would
primary (a) and secondary (b) rachis. H Staminate spike- also be independently derived in the Pooideae.
let on secondary rachis. I Pistillate spikelet. J Upper Ehrharta species growing in the dry winter-
flower of female spikelet. K Dorsal view of lower lemma rainfall habitats of South Africa represent a rela-
from female spikelet. (From Clayton 1970)
tively recent (possibly late Miocene) radiation
(Verboom et al. 2003). The early-diverging mem-

bers of the group occupied regions of year-round
rainfall, estimated at 689–1,400 mm per year. E.
stipoides is being investigated as a possible
drought-tolerant crop for human consumption
(Shapter et al. 2013).
III. TRIBE ORYZEAE Dumort. (1824)
Annuals or perennials. Leaf blades with or without

a pseudopetiole, the ligule membranous. Spikelets
with 1 to 3 flowers, one fertile, the other 0 to 2 sterile
and reduced to lemmas. Glumes reduced or absent.
Vascular system of the midrib with at least two
vascular bundles, often more, with bundles placed
ad- and abaxially, and often with air spaces.
Eleven genera and seventy species.
A robust and well-sampled molecular phylog-
eny for the tribe shows that all genera are mono-
phyletic (Tang et al. 2010). Previous data linking
Prosphytochloa and Potamophila were based on
misidentification of the Prosphytochloa specimen.
A new specimen of Prosphytochloa plus an acces-
sion of Maltebrunia shows that the two are sister
taxa that together are sister to Leersia. Data on
leaf anatomy relies on the comprehensive work of
Tateoka (1963); vascular structure of the midrib
in Luziola and Hygroryza is simple, similar to
that in Ehrharteae.
All members of Oryzeae grow in damp areas,
many occupy flooded sites, and still others (e.g.,
Hygroryza) are truly aquatic. Although this tribe
scarcely needs dividing into subtribes, the names
are available and correspond to well-supported
clades, and so are provided here.
III.1. SUBTRIBE ORYZINAE Griseb. (1853)

This subtribe has no diagnostic morphological
characters. The leaves may be auriculate or not.
Spikelets are laterally compressed in all genera,
but this character may be ancestral. Fusoid cells
are lacking in the mesophyll of all genera except
for Maltebrunia. Loss of fusoid cells appears to be
a derived character here, although they were lost
in parallel in Ehrharta.
Fig. 16. Ehrhartoideae-Ehrharteae. Ehrharta erecta.

A Habit. B Node with leaf sheath and ligule. C Spikelet.
D Florets. (From Barkworth et al. 2007, drawn by
H. Paźkirková)
III. TRIBE ORYZEAE Dumort. (1824) 147
arsenic from the soil, making it potentially useful

for phytoremediation (Ampiah-Bonney et al. 2007).
14. Maltebrunia Kunth

Maltebrunia Kunth, Rév. Gram. 1: 6 (1829).
Perennials, rhizomatous or not. Leaf blade with a
pseudopetiole, with or without obvious cross
veins. Inflorescence branches branched. Spikelets
with two sterile and one fertile flowers. Margins
of the lemma clasping the palea.
Four spp., Gabon, Tanzania, Madagascar.
15. Oryza L.
Oryza L., Sp. Pl.: 333 (1753); phylog.: Vaughan et al.
(2005).
Porteresia Tateoka, Bull. Natl. Sci. Mus. 8: 406 (1965).
Annuals or perennials, caespitose or rhizomatous.
Culm nodes spongy to solid. Ligule sometimes
ciliolate. Inflorescence branches themselves
branched. Spikelets with two sterile and one fertile
flower, disarticulating below the sterile lemmas
and above the true glumes. Fertile lemma coria-
ceous, asymmetrically keeled, the margins inter-
locking with the margins of the palea, awned or
not. Epidermal cells of the lemma and palea pro-
ducing tuberculate outgrowths. 2n ¼ 24, 48.
Twenty spp., tropical and subtropical world-
wide.
The phylogeny of this group has been
Fig. 17. Ehrhartoideae-Oryzeae-Oryziinae. Leersia dre- resolved by a large genomic data set (Zou et al.
panothrix. A Habit. B Ligule. C Spikelet. D Lemma. E 2008). See also Kellogg (2009) for discussion.
Flower and lodicules separated. F Caryopsis. G–J Spikelets
from other species: G L. tisserantii. H L. friesii. I L. denu- 16. Prosphytochloa Schweick.
data. J L. hexandra. (From Clayton 1970)
Prosphytochloa Schweick., Z€
uchter 31: 193 (1961).
Scandent perennials. Apex of distal leaf blades
13. Leersia Sw. Fig. 17 extended into a long tendril. Inflorescence
branches branched. Spikelets with two sterile
Leersia Sw., Prodr. Veg. Ind. Occ. 21 (1788).
flowers and one fertile.
Annuals or perennials, caespitose or rhizoma- One sp., P. prehensilis (Nees) Schweick.,
tous. Culm nodes spongy to solid. Inflorescence South Africa.
branches themselves branched. Spikelets with
one flower, sterile lemmas absent. Fertile lemma
III.2. SUBTRIBE ZIZANIINAE Benth. (1881)
asymmetrically keeled, awned or not, the margins
inrolled. Stamens 1, 2, 3 or 6. Scutellar cleft This subtribe lacks a clear diagnostic character,
absent. 2n ¼ 24, 48, 60. although some labile characters may be synapo-
Eighteen spp., tropical and warm temperate. morphic depending on their optimization on the
Loss of sterile lemmas is a synapomorphy for phylogeny.
Leersia (Kellogg 2009). Parallel losses have occurred All members of the clade have branched
in some members of subtribe Zizaniinae. Leersia inflorescence branches except for a few species
oryzoides is reported to take up large quantities of of Luziola, and all except for Luziola have fusoid
18. Hygroryza Nees Fig. 18

Hygroryza Nees, Edinb. New Phil. J. 15: 380 (1833).
Prostrate stoloniferous perennials; aquatic.
Culms spongy. Leaf-blades floating, with inflated
sheaths. Spikelets laterally compressed. Sterile
lemmas absent. Flower on a long pedicel.
Lemma awned. 2n ¼ 24.
One sp., H. aristata (Retz.) Nees ex Wright &
Arn., eastern Asia.
19. Potamophila R. Br.

Potamophila R. Br., Prodr. Fl. Nov. Holl: 211 (1810);
phylog.: Abedinia et al. (1998).
Rhizomatous perennials. Spikelets unisexual or
bisexual, these intermixed in the inflorescence.
Sterile lemmas present. Scutellar cleft absent, the
embryonic leaf margins non-overlapping. 2n ¼ 24.
One sp., P. parviflora R. Br., Australia.
P. parviflora may be a diplosporous apomict,
based on lack of overall genetic diversity and
irregular microgametogenesis and megagameto-
genesis (Wheeler et al. 2001).
20. Rhynchoryza Baillon

Rhynchoryza Baillon, Hist. Pl. 12: 291 (1893).
Perennials, 2–3 m tall. Leaves coriaceous, with
auricles. Pedicels of spikelets cuneate. Sterile
lemmas forming a short involucre at the base of
the fertile flower. Fertile lemma with involute
Fig. 18. Ehrhartoideae-Oryzeae-Zizaniinae. Hygroryza margins, the apex extended to a long aerenchy-
aristata. A Habit. B Spikelet. C Palea. D Stamen. (From matous beak and ultimately to an awn.
Koyama 1987) One sp., R. subulata (Nees) Baill., Paraguay to
Argentina.
cells in the mesophyll. According to the phylog- 21. Zizania L.
eny presented by Tang et al. (2010) and the char-
acter distribution shown by Kellogg (2009), Zizania L., Sp. Pl.: 991 (1753); tax.: Terrell et al. (1997).
sterile lemmas have been lost twice in this clade, Aquatic annuals or perennials, caespitose, with
and unisexual flowers have originated twice. or without stolons, 1–3 m tall. Culms spongy.
Spikelets unisexual, staminate spikelets on
17. Chikusichloa Koidzumi lower panicle branches, pistillate ones on upper
Chikusichloa Koidzumi, Bot. Mag. Tokyo 39: 23 (1925). branches. Sterile lemmas absent. Lemmas awned.
2n ¼ 30, 34.
Caespitose perennials. Ligule sometimes ciliolate. Five spp., Eastern Asia and North America.
Spikelets on a long slender pedicel. Glumes absent;
sterile lemmas absent. Stamen 1. Mesophyll cells LUZIOLA + ZIZANIOPSIS CLADE:
without invaginated cell walls. 2n ¼ 24. Sterile lemmas absent. Flowers unisexual.
Three spp., Southeast Asia to Japan. Pericarp free from the seed coat, brittle.
The stamen number clearly represents a This origin of unisexual flowers appears to be
reduction from the ancestral number of 6. independent from that in Zizania.
References 149
22. Luziola Juss. Clayton, W.D., Renvoize, S.A. 1986. Genera graminum:
grasses of the world. London: Her Majesty’s Statio-
Luziola Juss., Gen. Pl. 33 (1789); tax.: Martı́nez-y-Pérez nery Office.
et al. (2008). Judziewicz, E.J., Clark, L.G. 2007. Classification and bio-
Hydrochloa P. Beauv., Ess. Agrostogr. 135, 165, 182 geography of New World grasses: Anomochlooideae,
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V. Subfamily Bambusoideae Luerss. (1893)
This treatment of Bambusoideae includes exten- culm leaves, and gregarious flowering, all of
sive and detailed contributions from Lynn G. which appear to be uniquely derived (Zhang and
Clark (Iowa State University). All remaining Clark 2000). However, molecular phylogenies
errors and misinterpretations are mine. based on nuclear genes indicate that the history
of Bambusoideae is more complex and intriguing
than previously thought (Triplett et al. 2014).
Plants perennial. Culms woody or herba- Arundinarieae (the temperate woody bamboos)
ceous, caespitose or rhizomatous, the nodes flat are tetraploids with genomes designated A and B.
or swollen. Leaves pseudopetiolate, often with Bambuseae (the tropical woody bamboos) are
obvious cross veins, strongly dimorphic in tetraploid and hexaploid with genomes desig-
woody taxa, with culm leaves distinctly different nated C, D and E. Olyreae are predominantly
from leaves on lateral branches (“foliage leaves”), diploid with a genome designated H. Thus, the
the culm leaves with prominent sheaths and Arundinarieae and Bambuseae are the products
reduced blades, the foliage leaves pseudopetio- of separate polyploidization events. Remarkably,
late. Ligule membranous, with or without a fringe however, the B genome of the temperate bamboos
of hairs. Outer ligule present or absent. Inflores- is most closely related to the C genome of the
cences branched or unbranched, bearing spike- tropical ones; in other words, the temperate and
lets or pseudospikelets. Glumes 1 to several, or tropical species have one genome donor in
lacking. Lodicules 3, with extensive vascular tis- common. Chloroplast data place Olyreae sister
sue. Stamens 3 or 6. Ovary with 2 or 3 stigmas. to Bambuseae (Bouchenak-Khelladi et al. 2008;
Hilum linear, as long as the caryopsis. Epiblast Burke et al. 2012; Kelchner and Bamboo Phylog-
present, scutellar cleft present, mesocotyl inter- eny Group 2013; Kelchner and Clark 1997;
node absent, embryonic leaf margins overlap- Sungkaew et al. 2009), although monophyly of
ping. Lamina of first seedling leaf absent. the woody bamboos cannot be rejected statisti-
Mesophyll with fusoid cells, with arm cells with cally (Kelchner and Bamboo Phylogeny Group
asymmetric invaginations in the cell walls. C3. 2013). The data of Triplett et al. (2014) indicate
One hundred sixteen genera, 1441 spp. that woodiness is derived within Bambusoideae,
Phylogenies of Bambusoideae consistently but it is unclear if it evolved once or if each
show strong support for three tribes, Bambuseae, independent polyploidization event correlates
Olyreae, and Arundinarieae, as recognized here with an independent origin of woodiness.
(Bouchenak-Khelladi et al. 2008; Burke et al. Many traditionally recognized genera in
2012; Clark et al. 2007; Kelchner and Bamboo Bambusoideae have proven to be polyphyletic
Phylogeny Group 2013; Kelchner and Clark with the addition of molecular data (see refer-
1997; Sungkaew et al. 2009; Zhang and Clark ences below), indicating that the commonly
2000). Bambuseae and Arundinarieae were long used characters are in fact homoplasious and
thought to be sisters based on the shared char- are not reliable indicators of relationship. Such
acters of woody culms, differentiated foliage and characters include rhizome structure, presence or
152 V. Subfamily Bambusoideae Luerss. (1893)
absence of pseudospikelets, numbers and

arrangement of culm buds and branches, and
inflorescence architecture. In addition, homo-
ploid hybridization has been documented and
appears to be far more common than previously
thought possible (Takahashi et al. 1994; Triplett
and Clark 2010; Triplett et al. 2010, 2014). Several
common “genera” appear to be simply early-gen-
eration (possibly F1) hybrids that combine the
characters of their parent genera. If hybridization
is common, then introgression, particularly of
chloroplasts, may also occur frequently. In this
scenario, the chloroplast phylogenies in the liter-
ature may thus be misleading about relationships
(see, for example, Yang et al. 2013). An additional
confounding factor is the slow rate of molecular
evolution, presumably caused by the very long
generation time of most bamboos. Thus, even
very well sampled molecular phylogenies such
as those of Triplett and Clark (2010) and Zeng
Fig. 19. Bambusoid rhizomes. A Pachymorph (sympo-
et al. (2010) often provide only weak hints of dial) rhizome (Bambusa beechyana). B Leptomorph
phylogenetic history. Finally, all woody bamboos (monopodial) rhizome (Arundinaria amabilis). (From
appear to be polyploid (e.g., Stebbins 1971, 1985; McClure 1966)
Triplett et al. 2014). Not surprisingly, the first
genome sequence of a woody bamboo shows a many collections are sterile, and taxonomists
whole genome duplication event, estimated to have had to develop ways to identify species
have occurred 7–12 mya (Peng et al. 2013), without access to the often-diagnostic inflores-
although it is unclear if the methods used account cences and flowers. As noted by Wong (2004),
for the very slow mutation rate. More studies like “Attempts to identify bamboos using older
that of Triplett et al. (2014) will be needed to accounts can turn out to be frustrating because
address the effects of hybridization and poly- they may not always include adequate descriptions
ploidy on bamboo evolution and hybridization. of the most accessible parts of a bamboo plant . . .”.
A detailed account of bamboo morphology is This is something of an understatement.
provided by Stapleton (1997) and also on the web McClure (1966) drew attention to rhizome
(http://www.bamboo-identification.co.uk/index/ morphology in woody bamboos. In some taxa,
html and http://www.eeob.iastate.edu/research/ the apical meristem of the rhizome is persistent
bamboo/characters.html). In general, morpho- and never turns upward to form a culm; all above
logical work on the subfamily has been uneven, ground culms are formed from meristems that
with some taxa receiving detailed and compre- are axillary on the rhizomes. Such rhizomes are
hensive study and others only being described in known as leptomorph, or monopodial (Fig. 19B).
a fairly superficial way. Numerous good field In other taxa, the apical meristem of the rhizome
characters – most notably the rhizome type and eventually will form a culm, so that the growth of
plant architecture, number of branches per mid- the plant is sympodial as is common in most
culm node, and color of hairs and leaf blades – grasses (Fig. 19A). Sympodial rhizomes are
are not readily available on herbarium specimens. known as pachymorph, a term that reflects exten-
An additional complication is the lack of flower- sive variation in thickness from one internode to
ing material. The bamboos are well known for the next. As the rhizome initiates from a lateral
cyclical flowering, with many species flowering meristem, the first few internodes are slender and
at intervals of tens to over one hundred years lack axillary buds, although they may produce
(Janzen 1976; McClure 1966; Ramanayake and adventitious roots. These first internodes, which
Weerawardene 2003), although sporadic (off- connect the newly formed rhizome to the parent
cycle) flowering is also known. This means that axis, are known as the neck. In some cases, the neck
V. Subfamily Bambusoideae Luerss. (1893) 153
may be one or occasionally several meters long, appears to be blurred, as though the meristem
and will have many nodes. Subsequent internodes goes through a slow transition from producing
are noticeably wider in diameter, produce axillary leaves to producing variously reduced and mod-
buds and often numerous roots throughout the ified bracts, to ultimately producing grass flowers
internode, and are asymmetric so that they curve with lemmas and paleas. In such plants, it is
upward toward the soil surface. After the meri- common for inflorescence branches to be sub-
stem extends above ground and forms the culm, tended by bracts and prophylls. In addition,
internode diameter is again reduced although not there may be glume-like structures proximal to
nearly to the extent seen in the neck. The result is the flowers; these glume-like structures (so-called
a plant with curved swollen rhizomes, connected “gemmiparous bracts”) have buds in their axils.
to their parent axes by narrow internodes. Any bracts distal to the gemmiparous bracts and
Rhizome morphology affects the arrange- proximal to the flowers are then known as
ment of culms. A plant with pachymorph rhi- glumes. Thus, plants with gemmiparous bracts
zomes with short necks will be clump forming. and no additional bracts below the flowers are
Conversely, a plant with either leptomorph rhi- described as lacking glumes, although this
zomes or pachymorph rhizomes with long necks description assumes that a glume is necessarily
will produce more widely spaced culms. a sterile structure. After the flowers are formed,
The leaves of many bamboos bear fimbriae or the buds in the axils of the glume-like structures
oral setae, elongated structures up to several cm then grow out and reiterate the production of
long, at the summit of the sheath of culm and/or glumes and flowers, and the reiterated branchlets
foliage leaves, on the auricles, and sometimes (“rachillas”, although the homology is not clear)
around the ligule. Most commonly these fimbriae may themselves produce more branchlets and
are slender, terete or slightly flattened, less than flowers. This complex, reiterated structure is
0.1 mm wide, and look like particularly long known as a pseudospikelet. Inflorescences with
hairs. A few taxa, however, have much more pseudospikelets are known as iterauctant, in con-
robust structures, 0.2–1.2 mm wide, occurring in trast to those with conventional spikelets, which
the ligular area and often fused to the abaxial side are described as semelauctant. I have avoided
of the ligule. In most of the literature (e.g., Jud- these terms here, preferring to describe the mor-
ziewicz et al. 1999) both sorts of structures are phology using more familiar – if more lengthy –
called fimbriae, although other authors call both descriptors. Presence or absence of pseudospike-
oral setae. Clark and Cortés R. (2004) note the lets has never been analyzed in a phylogenetic
distinction between the two, based largely on context but the taxonomic distribution of the
morphology but also position, and argue that structure suggests that it is highly homoplasious,
the slender structures should be called fimbriae having been gained and/or lost many times in the
whereas the more robust ligular ones should be history of the subfamily.
called oral setae. By this definition, in New World The tribal and subtribal classification pre-
bamboos, oral setae occur only in Bambuseae sented here follows that of the Bamboo Phylogeny
subtribe Guaduinae, in Eremocaulon, Olmeca Group (BPG) (2012), which in turn is based on
and Otatea. Clearly, this distinction needs to be current phylogenetic studies (Kelchner and Bam-
considered for Old World bamboos as well. How- boo Phylogeny Group 2013). The BPG (2012)
ever, because most bamboo taxonomists use the presents a comprehensive table of these studies
terms fimbriae and oral setae interchangeably, in and their main conclusions, so only the most
this account all such structures will be called recent are referenced here. Information for New
fimbriae. World genera of Bambusoideae relies heavily on
Bamboo inflorescences are described in detail Judziewicz et al. (1999), which should be con-
in the section on Inflorescence structure. While sulted for more detailed information. The book
many genera have conventional inflorescences is elegantly illustrated and the taxa are described
with conventional grass spikelets like other much more completely than is attempted here.
grasses, others have inflorescences that are com- Information on Old World bamboos comes from
plex and difficult to describe using standard ter- various sources, but most notably the Flora of
minology. In this latter group, the distinction China (Li et al. 2006). Although I have attempted
between vegetative and inflorescence structures to provide parallel descriptions, information is
lacking on many important characters, particu- 3 (2). Culms bearing thorns and/or root spines at the
larly for Old World taxa. This paucity of descrip- nodes 4
3. Thorns and root spines absent 9
tive material combined with lack of specimens or 4 (3). Thorns formed from modified branches 5
incomplete specimens makes the bamboos the 4. Root spines formed from prop roots, especially at lower
group of grasses for which morphological data nodes 6
are most urgently needed. 5 (4). Culms with white hairs below the nodes; culm leaves
with blade margins continuous with the sheath
margins; palea keels winged; plants of the New World
73. Guadua
TRIBES AND SUBTRIBES OF BAMBUSOIDEAE 5. Culms lacking white hairs at the nodes; culm leaves
with the margins distinct from the sheath margins;
IV. Tribe Arundinarieae palea keels lacking wings; plants of the Old World
Clade I. Bergbambos 97. Bambusa
Clade II. Oldeania 6 (4). Plants of the New World; spikelets with one fertile
Clade III. Chimonocalamus flower 55. Chusquea
Clade IV. Shibataea clade 6. Plants of the Old World; spikelets with several fertile
Clade V. Phyllostachys clade flowers 7
Clade VI. Arundinaria clade 7 (6). Rhizomes leptomorph 34. Chimonobambusa
Clade VII. Thamnocalamus 7. Rhizomes pachymorph 8
Clade VIII. “Indocalamus” wilsonii 8 (7). Rhizomes with short necks, the culms clumped
Clade IX. Gaoligongshania 26. Chimonocalamus
Clade X. Indocalamus s.s. 8. Rhizomes with long necks, the culms solitary
Clade XI. “Ampelocalamus” calcareus 25. Oldeania
Clade XII. Kuruna 9 (3). Culms unbranched 10
V. Tribe Bambuseae 9. Culms branched 11
Neotropical clade 10 (9). Inflorescence branches with subtending bracts;
V.1. Subtribe Chusqueinae spikelets with a rachilla extension; plants of New Cale-
V.2. Subtribe Arthrostylidiinae donia 79. Greslania
V.3. Subtribe Guaduinae 10. Inflorescence branches lacking subtending bracts; spi-
Paleotropical clade kelets lacking a rachilla extension; plants of the New
V.4. Subtribe Bambusinae World
V.5. Subtribe Melocanninae 55. Chusquea subg. Platonia, subg. Magnifoliae
V.6. Subtribe Hickeliinae 11 (9). Foliage leaves with an adaxially projecting midrib;
V.7. Subtribe Racemobambosinae plants of Madagascar, Africa, and Réunion [subtribe
VI. Tribe Olyreae Hickeliinae] 12
VI.1. Subtribe Buergersiochloinae 11. Foliage leaves with the midrib not projecting on the
VI.2. Subtribe Parianinae adaxial side; distribution various, but generally not
VI.3. Subtribe Olyrinae Madagascar or Réunion 19
12 (11). Young shoots covered with black hairs 13
12. Young shoots not covered with black hairs 14
13 (12). Inflorescences dense, capitate, the branches
branched, lateral branches arranged in tight fans
KEY TO THE GENERA OF BAMBUSOIDEAE 112. Cathariostachys
13. Inflorescence unbranched, surrounded by a spathe,
with 1 to 4 spikelets, the spatheate units clustered
The following key should be considered provi- 113. Decaryochloa
sional, especially as regards Old World taxa. 14 (12). Culm leaf sheaths lacking auricles; glumes 1 to 5
Many of the genera are polyphyletic. Others are 15
poorly known, with incomplete character infor- 14. Culm leaf sheaths with auricles; glumes 5 or 6 18
mation in the literature and few available speci- 15 (14). Glumes 1 or 2 16
15. Glumes 3, 4 or 5 17
mens. 16 (14). Spikelets laterally compressed, keeled; style
1. Culms herbaceous; abaxial ligule absent persistent in fruit 115. Hitchcockella
Group I (Olyreae) 16. Spikelets terete, lacking keels; style not persistent
1. Culms woody; abaxial ligule present 2 117. Perrierbambus
2 (1). Culms normally leafless, the plants clump forming 17 (15). Foliage leaves lacking auricles; ovary apex
and rush-like; internodes with cross partitions; near pubescent 114. Hickelia
Rio de Janeiro, Brazil 56. Glaziophyton 17. Foliage leaves with prominent auricles; ovary glabrous
2. Culms normally leafy, the plants clump forming or not; 118. Sirochloa
internodes lacking partitions; distribution various 3 18 (14). Lemma with few veins 116. Nastus s.s.
KEY TO THE GENERA OF BAMBUSOIDEAE 155
18. Lemma with 29 to 41 veins 119. Valiha 12. Pistillate flower symmetrical, generally ellipsoid or
19 (11). Branch complement of two or more subequal fusoid, compressed very little if at all 13
branches Group II 13 (12). Glumes of pistillate spikelets with thickened mar-
19. Branch complement with one branch only or one gins, extending into attenuate apices, these twisting at
dominant branch and two to many smaller laterals maturity 14
Group III 13. Glumes without thickened margins 15
14 (13). Sexes in the same inflorescence, pistillate spike-
lets above, staminate below 135. Sucrea
Group I – Herbaceous bamboos 14. Sexes in separate inflorescences, the terminal inflores-
1. Pistillate lemma and palea leathery, but not hardened; cence staminate and branched, the axillary ones pistil-
lemma with an awn 5–30 mm long; plants of New late and unbranched 136. Raddia
Guinea 121. Buergersiochloa 15 (13). Culms dimorphic on the same plant, with upright
1. Pistillate lemma and palea generally hardened; lemmas culms producing leaves with broad blades and decum-
generally unawned, if awned then the awn less than bent culms producing leaves with blades reduced or
15 mm; plants of the New World 2 lacking; inflorescences commonly but not exclusively
2 (1). Long slender fimbriae present at the top of the borne on the decumbent culms 133. Piresia
sheaths, near the sheath/blade junction; staminate 15. Culms not dimorphic; inflorescences borne on leafy
spikelets in a whorl 3 culms 16
2. Fimbriae absent; staminate spikelets not whorled 4 16 (15). Inflorescence with one distal pistillate spikelet,
3 (2). Leaves with the top of the sheath variously the proximal spikelets staminate 17
decorated with translucent swellings (“lunar marks”); 16. Inflorescences various, often with pistillate and
pistillate spikelets one per inflorescence node staminate spikelets mixed, but not with a single distal
123. Pariana pistillate spikelet 19
3. Leaves without lunar marks; pistillate spikelets one per 17 (16). Spikelets clustered, with many male spikelets
inflorescence, terminal 122. Eremitis surrounding a single terminal female one; the cluster
4 (2). Upper leaf blade inverted, forming a canopy over deciduous; 2 to 5 spikelet clusters on a zigzag axis
the single terminal inflorescence 5 128. Froesiochloa
4. Leaf blades not inverted, not covering the inflorescence 17. Spikelets not clustered 18
6 18 (17). Inflorescences produced from apical and axillary
5 (4). Bases of glumes of pistillate spikelets fused; glumes meristems 125. Arberella
of staminate spikelets tiny, membranous 18. Inflorescences produced only from apical meristems
140. Diandrolyra 134. Reitzia
5. Bases of glumes of pistillate spikelets not fused; glumes 19 (16). Disarticulation of female spikelet above the
of staminate spikelets absent 141. Maclurolyra glumes 126. Cryptochloa
6 (4). Plants producing swollen corms at base of stems 7 19. Disarticulation of female spikelet below the glumes 20
6. Corms absent 9 20 (19). Each inflorescence with few flowers; male and
7 (6). Inflorescences with several branches, each terminat- female flowers in separate inflorescences
ing in a pistillate spikelet, with one or two staminate 132. Raddiella
spikelets below; plants of northern South America 20. Each inflorescence with many flowers; male and
139. Rehia female flowers mixed in the same inflorescence
7. Inflorescences with only two branches, one with stami- 131. Parodiolyra
nate spikelets and the other with pistillate; plants of
Cuba 8 Group II – Branch complement of two or more
8 (7). Leaf blades with tapering bases; flowering culms subequal branches
longer than vegetative ones; pistillate flower longer
than glumes; stamens 3 137. Mniochloa 1. Mid-culm branches apparently paired, with one main
8. Leaf blades with truncate bases; flowering culms shorter branch and one lateral, the two becoming subequal
than vegetative ones; pistillate flower shorter than 40. Phyllostachys
glumes; stamens 2 138. Piresiella 1. Mid-culm branches more than two 2
9 (6). Pistillate lemma awned 10 2 (1). Mid-culm nodes with branches in a fan-shaped
9. Pistillate lemma awnless, or if awned then the awn pattern 3
much shorter than the body of the lemma 11 2. Mid-culm nodes with branching pattern not fan-shaped
10 (9). Awn up to 15 mm long, longer than the body of the 6
lemma; leaf blades deciduous 127. Ekmanochloa 3 (2). Culm internodes alternating in length between long
10. Awn <2 mm long, shorter than the lemma; leaf blades and short; bud prophylls cordate; culm leaf blades
persistent on distal leaves 124. Agnesia erect; spikelets in clusters of 2 or 3 65. Didymogonyx
11 (9). Plants over 50 cm tall, often over 1 m; forming 3. Culm internodes not alternating in length; bud pro-
large clumps 130. “Olyra” phylls triangular; culm leaf blades reflexed; spikelets
11. Plants always less than 1 m tall, generally less than not clustered 4
50 cm 12 4 (3). Foliage leaves dimorphic, the blades of those on the
12 (11). Pistillate flower shaped like a helmet or tooth, main culm much larger than the ones on the lateral
laterally compressed 129. Lithachne branches; leaf sheaths with fused fimbriae creating a
“ruffled” margin; inflorescence branches branched 17. Culm leaves with sheaths lacking a dark girdle
58. Actinocladum between the node and the base of the sheath 18
4. Foliage leaves not dimorphic, leaf sheaths with fimbriae 18 (17). Plants of the New World 19
not fused, the margin not appearing ruffled; inflores- 18. Plants of the Old World 23
cence generally unbranched 5 19 (18). Culms scandent, vine-like; pseudopetioles
5 (4). Blades of culm leaves reflexed, the base narrower strongly reflexed; foliage leaves with one wide
than the distal portion of the sheath; culms often with marginal sector appearing bright green, with no or
mottled and/or scabrous internodes 68. Merostachys reduced wax 20
5. Blades of culm leaves erect, the base about the same 19. Culms erect; pseudopetioles not reflexed; foliage
width as the distal portion of the sheath; culm leaves without a bright green marginal sector 21
internodes generally not mottled or scabrous 20 (19). Flowers spindle-shaped, the lemmas
70. Rhipidocladum sharp-pointed 62. Atractantha
6 (2). Branches as many as 80 per node, in 2 or 3 rows, 20. Flowers not spindle-shaped, the lemmas blunt or acute
appearing verticillate 35. Drepanostachyum but not sharp-pointed 61. Athroostachys
6. Branches fewer than 80 per node, in 1 row or cluster, 21 (19). Plants of the southeastern U.S.; rhizomes
not verticillate 7 leptomorph 45. Arundinaria
7 (6). First internode long, the upper ones very short and 21. Plants of Central or South America; rhizomes
the leaves densely overlapping; branches only one per pachymorph 22
node, but appearing subequal because of very short 22 (21). Plants of Mexico, Central America and Colombia;
internodes; plants of the Guyana Highlands rhizomes with a long neck 75. Otatea
69. Myriocladus 22. Plants of southeastern Brazil; rhizomes with a short
7. First and second internodes roughly similar lengths, the neck 57. Cambajuva
upper ones not noticeably short with the leaves 23 (18). Rhizomes leptomorph 24
overlapping; branches more than one from the same 23. Rhizomes pachymorph 31
node; plants not of the Guyana Highlands 8 24 (23). Flowers borne in conventional spikelets; spikelets
8 (7). Mid-culm branches from multiple buds 9 generally pedicellate; inflorescence with bracts
8. Mid-culm branches from a single bud 14 reduced or lacking 25
9 (8). Culm leaf blade less than 1 cm long, narrow to 24. Flowers borne in pseudospikelets; pseudospikelets
acicular or nearly absent 34. Chimonobambusa generally sessile; inflorescence with well-developed
9. Culm leaf blade more than 1 cm long, flat 10 bracts 30
10. Buds at a single node obviously dimorphic or 25 (24). Culm internodes flattened above the nodes
trimorphic; culm leaves lacking auricles or fimbriae 39. Oligostachyum
55. Chusquea subg. Rettbergia, Euchusquea clade 25. Culm internodes more or less terete, or slightly
10. Buds not clearly dimorphic or trimorphic, although grooved above the nodes 26
sometimes varying somewhat in size; culm leaves 26 (25). Branches 1–3 per node 27
generally with auricles and/or fimbriae 11 26. Branches 3–7 per node 28
11 (10). Culms climbing or scrambling; flowers in 27 (26). Culm branches profusely branched; culm buds
pseudospikelets 91. Holttumochloa always open at front 42. Sarocalamus
11. Culms erect; flowers in conventional spikelets 12 27. Culm branches generally with few or no secondary
12 (11). Shoots fragrant; plants of Asia branches; culm buds initially closed at front
26. Chimonocalamus 29. Pseudosasa
12. Shoots not fragrant; plants of South America 13 28 (26). Stamens 6 31. Acidosasa
13 (12). Plants <2 m tall; foliage leaves with white 28. Stamens 3 29
fimbriae 67. Filgueirasia 29 (28). Culm leaves with the sheath persistent, the blade
13. Plants over 2 m tall; foliage leaves with green or brown reflexed 41. Pleioblastus
fimbriae 71. Apoclada 29. Culm leaves with the sheath deciduous, the blade erect
14 (8). Each lateral branch bearing only 1 or 2 leaves 15 33. Bashania
14. Each lateral branch bearing more than 2 leaves 16 30 (24). Stamens 6 46. Indosasa
15 (14). Branches 4 or 5 per node at mid-culm; sheaths of 30. Stamens 3 43. Sinobambusa
foliage leaves contiguous with branches; terminal leaf 31 (23). Rhizomes with elongated necks 32
lacking a sheath 30. Shibataea 31. Rhizomes with short necks 35
15. Branches 7 to 12 per node at mid-culm; sheaths of 32 (31). Inflorescence branches not subtended by bracts
foliage leaves not contiguous with branches; terminal 44. Yushania
leaf with a sheath 28. Gelidocalamus 32. Inflorescence branches subtended by bracts 33
16 (14). Culm nodes surrounded by a dense ring of 33 (32). Pericarp crustaceous 108. Pseudostachyum
spreading hairs 92. Kinabaluchloa 33. Pericarp fleshy 34
16. Culm nodes glabrous, or hairs not spreading 17 34 (33). Inflorescence a dense head 110. Stapletonia
17 (16). Culm leaves with sheaths with a dark girdle 34. Inflorescence elongate, branching 105. Melocanna
between the node and the base of the sheath 35 (31). Style elongate, hollow 36
93. Dinochloa 35. Style absent or short, or not hollow 39
KEY TO THE GENERA OF BAMBUSOIDEAE 157
36 (35). Palea with 2 awns; stamen filaments fused 5. Foliage leaves with blades erect; fimbriae, if present, not
106. Neohouzeaua curly 6
36. Palea unawned; stamen filaments free 37 6 (5). Culms with a band of white hairs below the nodes;
37 (36). Culm leaf sheaths flat; blade of culm leaf rolled; flowers in pseudospikelets
pseudospikelets with several flowers 73. Guadua (non-thorny forms)
111. Teinostachyum 6. Culms lacking a band of white hairs below the nodes;
37. Culm leaf sheaths with 1 or 2 strong undulations flowers in conventional spikelets 7
toward the apex; blade of culm leaf flat; pseudospike- 7 (6). Culm leaves with sheaths persistent, indurate;
lets with one fertile flower 38 lemma and palea fused to each other at the base
38 (37). Glumes absent 109. Schizostachyum 74. Olmeca
38. Glumes 2 or 3, awned 103. Cephalostachyum 7. Culm leaves with sheaths deciduous (or if persistent not
39 (35). Flowers borne in conventional spikelets 40 indurate); lemma and palea not fused 8
39. Flowers borne in pseudospikelets 45 8 (7) Spikelets with 4 glumes and one flower 55. Chusquea
40 (39). Spikelets with 2 to 8 fertile flowers; glumes 2 or 3 (few specimens of this genus will key here as most
41 have multiple buds and branches)
40. Spikelets with 1 fertile flower; glumes 2, 5 or 6 43 8. Spikelets with 2 glumes and several flowers 9
41 (40). Mid-culm branches 5 to 7 per node 42 9 (8). Foliage leaves with green or brown fimbriae, the
41. Mid-culm branches many per node 90. Vietnamosasa blades with prominent tessellate venation; plants of
(keyed here provisionally assuming that the branch southeastern Brazil 57. Cambajuva
complement arises from a single bud) 9. Foliage leaves with white fimbriae, the blades without
42. (41). Culm leaves with a narrow girdle, ca. 1 mm wide; prominent tessellate venation; plants of cerrado, Brazil
inflorescence one-sided, with bracts at all nodes 67. Filgueirasia (few specimens of this genus will key
36. Fargesia here as most have multiple buds and branches)
42. Culm leaves lacking a girdle; inflorescence open, lack- 10 (4). Rhizomes leptomorph 11
ing bracts 54. Kuruna 10. Rhizomes pachymorph 14
43 (40). Glumes 5 or 6 11 (10). Plants less than 2 m tall 12
81. Not “Nastus” (southeast Asian material) 11. Plants generally over 2 m tall 13
43. Glumes 2 44 12 (11). Culms and axis of inflorescence covered with
44 (43). Culm leaves with sheaths persistent; plants of white powdery wax; culm leaf sheaths longer than the
Africa 24. Bergbambos internodes 48. Sasamorpha
44. Culm leaves with sheaths deciduous; plants of Thai- 12. Culm and axis lacking white powdery wax; culm leaf
land 89. Temochloa sheaths shorter than the internodes 47. Sasa
45 (39). Stamens with filaments fused 13 (11). Internodes with a ring of yellow-brown hairs
87. Pseudoxytenanthera below the node; fruit dry 38. “not Indocalamus”
45. Stamens with filaments free 46 13. Internodes with a ring of white hairs below the node;
46 (45). Flowers with palea longer than the lemma fruit fleshy 27. Ferrocalamus
86. Pseudobambusa 14 (10). Culms erect to arching 15
46. Flowers with palea shorter than the lemma 47 14. Culms scrambling 18
47 (46). Rachilla nodes not disarticulating 15 (14). Rhizome necks elongate, the culms solitary
98. Dendrocalamus 44. Yushania
47 Rachilla nodes disarticulating 97. Bambusa 15. Rhizome necks short, the culms clumped 16
16 (15). Flowers in conventional spikelets; glumes 2;
plants of New Caledonia 79. Greslania
Group III – Branch complement with one branch 16. Flowers in pseudospikelets; glumes several; plants of
only or one dominant branch and two to many temperate to tropical Asia 17
smaller laterals 17 (16). Inflorescence a cluster of pseudospikelets; pseu-
dospikelets with 3 or 4 flowers 86. Pseudobambusa
1. Culm leaves not differentiated from foliage leaves; leaf 17. Inflorescence elongate, branched; pseudospikelets
sheaths densely overlapping 69. Myriocladus with 1 flower 106. Neohouzeaua
1. Culm leaves morphologically distinct from foliage 18 (14). Flowers in conventional spikelets
leaves; leaf sheaths generally not overlapping 2 51. Gaoligongshania
2 (1). Mid-culm branches no more than one per node 3 18. Flowers in pseudospikelets 19
2. Mid-culm branches more than one per node 22 19 (18). Culm nodes with a shelf-like extension; culm leaf
3 (2). Culm internodes flattened or grooved sheaths with horn-like auricles 78. Fimbribambusa
29. Pseudosasa 19. Culm nodes lacking a shelf-like extension; culm leaf
3. Culm internodes terete 4 sheath with auricles not horn-like, or lacking 20
4 (3). Plants of the New World 5 20. (19). Pseudospikelets with one flower
4. Plants of the Old World 10 106. Neohouzeaua
5 (4). Foliage leaves with pseudopetioles reflexed, the 20. Pseudospikelets with more than one flower 21
blades thus appearing to droop; fimbriae curly 21 (20). Base of culm sheath smooth; plants of southern
63. Aulonemia China and Vietnam 76. Bonia
21. Base of culm sheath wrinkled; plants of northern 37 (36). Foliage leaves with green or brown fimbriae, the
Australia 94. Mullerochloa blades with prominent tessellate venation; plants of
22 (2). Culms with multiple buds, usually dimorphic, per southeastern Brazil 57. Cambajuva
node 55. Chusquea 37. Foliage leaves with white fimbriae, the blades without
22. Culms with a single bud per node 23 prominent tessellate venation; plants of cerrado, Brazil
23 (22). Culm leaves with a corky crest at the junction of 67. Filgueirasia
the sheath and the girdle 24 38 (36). Flowers spindle-shaped, the lemmas
23. Culm leaves lacking a corky crest at the junction of the sharp-pointed 62. Atractantha
sheath and the girdle 25 38. Flowers not spindle-shaped, the lemmas blunt or acute
24 (23). Corky crest not bumpy; culms delicate; spikelets but not sharp pointed 39
conventional, with 2 or 3 glumes and few to many 39 (38). Culm leaf blades erect 40
flowers, the flowers not sharp-pointed 64. Colanthelia 39. Culm leaf blades divergent or reflexed 41
24. Corky crest bumpy, tumor-like; pseudospikelets with 40 (39). Culm leaves lacking a dark girdle at the base;
one glume, one prophyll, and one flower, the flower inflorescence with conventional spikelets, the axis
slender, indurate, sharp-pointed, needle-like often zigzag; fruit dry; plants of Central and South
62. Atractantha America, but not extending to Bahia
25 (23). Blade of culm leaf a tiny acicular extension of the 60. Arthrostylidium
sheath, or short and awl-like 26 40. Culm leaves with a dark girdle at the base;
25. Blade of culm leaf broad and flat, not acicular, inflorescence with pseudospikelets, the axis straight;
generally not an extension of the sheath 28 fruit fleshy; plants of Bahia, Brazil 59. Alvimia
26 (25). Branches 2 or 3 per node; plants of Africa 41 (39). Culm and foliage leaf sheaths with long straight
83. Oreobambos fimbriae over 2 cm long; stamens 6 66. Elytrostachys
26. Branches 5 or more per node; plants of Asia 27 41. Culm and foliage leaf sheaths with shorter, curly
27. Secondary branches not pendulous fimbriae; stamens 3 63. Aulonemia
82. Neomicrocalamus 42 (35). Culm leaf sheaths with auricles 43
27. Secondary branches long and pendulous 42. Culm leaf sheaths without auricles 73
49. Thamnocalamus 43 (42). Auricles low, rim-like 44
28 (25). Culms generally broadly spaced, rhizomes 43. Auricles various, but not rim-like 46
leptomorph, or pachymorph with elongate necks 29 44 (43). Culms with a thick band of silvery white hairs
28. Culms clumped, rhizomes pachymorph with short below each node; plants of the Malay Peninsula
necks 35 100. Maclurochloa
29 (28). Foliage leaf blades with a strongly excentric 44. Culms lacking silvery white hairs; plants of various
midrib; plants of the New World 72. Eremocaulon parts of Asia 45
29. Foliage leaf blades with midrib central or only slightly 45 (44). Flowers in conventional spikelets
off center; plants of the Old World 30 120. Racemobambos
30 (29). Inflorescence branches subtended by well-devel- 45. Flowers in pseudospikelets 99. Gigantochloa
oped bracts and prophylls; stamens 6 46. Indosasa 46 (43). Culms erect 47
30. Inflorescence branches with bracts and prophylls 46. Culms scrambling 59
reduced or absent; stamens 3 31 47 (46). Foliage leaves without fimbriae 48
31 (30). Culm internodes glabrous below the nodes 32 47. Foliage leaves with fimbriae 53
31. Culm internodes with a ring of hairs below the nodes 48 (47). Stamens numerous; fruit fleshy; style elongate,
34 hollow 107. Ochlandra
32 (31). Rhizomes pachymorph but with long necks 48. Stamens 6; fruit generally dry; style short, not hollow
44. Yushania 49
32. Rhizomes leptomorph 33 49 (48). Flowers in conventional spikelets
33 (32). Ridge prominent above node; branch buds on a 81. Not “Nastus” (southeast Asian material)
promontory; secondary branches profuse; culm buds 49. Flowers in pseudospikelets 50
open at the front 42. Sarocalamus 50 (49). Palea divided at least 1/3 of its length
33. Ridge above node lacking; promontory lacking; 102. Thyrsostachys
secondary branches lacking; culm buds initially closed 50. Palea undivided, or merely notched 51
at the front 29. Pseudosasa 51 (50). Clusters of pseudospikelets in the inflorescence
34 (31). Culm internodes with yellow brown hairs below subtended by a two-keeled prophyll; rachilla generally
the nodes; fruit dry 52. Indocalamus disarticulating between the flowers 97. Bambusa
34. Culm internodes with white hairs below the nodes; 51. Clusters of pseudospikelets in the inflorescence sub-
fruit fleshy 27. Ferrocalamus tended by a one-keeled prophyll; rachilla generally not
35 (28). Midrib with a single vascular bundle; plants of the disarticulating 52
New World 36 52 (51). Stamens with filaments free 98. Dendrocalamus
35. Midrib with multiple vascular bundles; plants 52. Stamens with filaments fused 99. Gigantochloa
generally of the Old World 42 53 (47). Style elongate, hollow; ovary glabrous 54
36 (35). Plants less than 2 m tall, erect 37 53. Style short, not hollow; ovary generally pubescent 56
36. Plants tall, erect or scandent 38 54 (53). Inflorescence densely capitate 104. Davidsea
TRIBES AND GENERA OF BAMBUSOIDEAE 159
54. Inflorescence elongate 55 70. Style short; ovary often pubescent at least at the
55 (54). Pseudospikelets with one flower; palea awned apex 87. Pseudoxytenanthera
106. Neohouzeaua 71 (70). Palea longer than the lemma, the keels winged;
55. Pseudospikelets with several flowers; palea lacking style short, not hollow 96. Sphaerobambos
awns 111. Teinostachyum 71. Palea shorter than the lemma, the keels without wings;
56 (53). Stamens with filaments fused style elongate, hollow 72
87. Pseudoxytenanthera 72 (71). Stamens with filaments fused; pericarp dry
56. Stamens with filaments free 57 106. Neohouzeaua
57 (56). Pseudospikelets with 1 flower 77. Cyrtochloa 72. Stamens with filaments free; pericarp fleshy
57. Pseudospikelets with 2 or more fertile flowers 58 111. Teinostachyum
58 (57). Culm leaves lacking a wrinkled zone at the base 73 (42). Inflorescence branches not subtended by bracts
97. Bambusa and prophylls; spikelets pedicellate 74
58. Culm leaves with a wrinkled zone at the base 73. Inflorescence branches subtended by bracts and pro-
95. Neololeba phylls; spikelets sessile 75
59 (46). Nodes crested with a narrow shelf-like projection 74 (73). Rhizomes with elongated necks; spikelets with
60 2 or more fertile flowers 44. Yushania
59. Nodes various, but not with a shelf-like projection 61 74. Rhizomes with short necks; spikelets with 1 fertile
60 (59). Culm leaf with blade spreading, the sheath with flower 37. Himalayacalamus
stiff horn-like auricles; flowers in pseudospikelets, 75 (73). Palea divided at least 1/3 of its length
these in clusters of more than 25 per inflorescence 102. Thyrsostachys
node 78. Fimbribambusa 75. Palea undivided, or merely notched 76
60. Culm leaf with blade erect, the sheath with rounded 76 (75). Clusters of pseudospikelets in the inflorescence
auricles; flowers in conventional spikelets, these in subtended by a two-keeled prophyll; rachilla generally
clusters of 2 or 3 per inflorescence node disarticulating between the flowers 77
88. Temburongia 76. Clusters of pseudospikelets in the inflorescence sub-
61 (59). Culm leaf sheaths with a wrinkled base 62 tended by a one-keeled prophyll; rachilla generally not
61. Culm leaf sheaths with the base smooth 65 disarticulating 78
62 (61). Foliage leaf base strongly asymmetrical, with one 77 (76). Culms erect; basal part of culm internode not
side broad and rounded and the other narrow and swollen 97. Bambusa
straight; spikelets with one flower 77. Cyrtochloa 77. Culms scrambling; basal part of culm internode
62. Foliage leaf base symmetrical; spikelets with more swollen 94. Mullerochloa
than one flower 63 78 (76). Stamens with filaments free 98. Dendrocalamus
63 (62). Culm leaf blades reflexed; spikelets with 2 flowers 78. Stamens with filaments fused 79
80. Melocalamus 79 (78). Inflorescence with clusters of pseudospikelets at
63. Culm leaf blades erect; spikelets with more than the nodes; culms often with yellow stripes; plants of
2 flowers 64 India and southeast Asia 99. Gigantochloa
64 (63). Leaf auricles leathery, crescent shaped; palea apex 79. Inflorescence a capitate cluster of pseudospikelets at
without hooked projections; lodicules 0; plants of the end of each lateral branch; culms not yellow
Mindanao, Sulawesi, Maluku, New Guinea, Solomon striped; plants of Africa 84. Oxytenanthera
Islands, Queensland 95. Neololeba
64. Leaf auricles papery, ovate to rounded; palea apex
with two hooked projections; lodicules 3; plants of TRIBES AND GENERA OF BAMBUSOIDEAE
Peninsular Malaysia, Singapore 101. Soejatmia
65 (61). Inflorescence with conventional spikelets 66
65. Inflorescence with pseudospikelets 68 IV. TRIBE ARUNDINARIEAE Asch. & Graebn.
66 (65). Spikelets with one flower (1902)
81. Not “Nastus” (southeast Asian material)
66. Spikelets with more than one flower 67 Rhizomes generally leptomorph. Culms woody,
67 (66). Spikelets on long pedicels 32. Ampelocalamus generally hollow. Branch development basipetal.
67. Spikelets sessile 120. Racemobambos Foliage leaves with an outer ligule. Spikelets lat-
68 (65). Inflorescence a single dense cluster of pseudos- erally compressed. 2n ¼ 48. Midrib complex.
pikelets 69 Twenty-six genera, 533 species.
68. Inflorescence elongate, with clusters of pseudospike-
lets at the nodes 70 This tribe includes all the temperate woody
69 (68). Blade of culm leaf reflexed 104. Davidsea bamboos, and is monophyletic in all molecular
69. Blade of culm leaf erect 85. Parabambusa phylogenies (Bamboo Phylogeny Group 2012;
70 (68). Style elongate; ovary glabrous 71 Burke et al. 2012; Ma et al. 2014; Zhang et al.
2012, Triplett et al. 2014). Leptomorph rhizomes CLADE I. BERGBAMBOS

may be a synapomorphy for this clade, but if so
then the character has apparently reversed in 24. Bergbambos Stapleton
some taxa. Culms of members of Arundinarieae
reach their full height without initiating branches Bergbambos Stapleton, Phytokeys 25: 99 (2013); Soder-
strom and Ellis (1982), morphol., anat.
and produce only culm leaves; later the culms
lignify and produce branches and foliage leaves,
Culms erect or slightly arching. Rhizomes pachy-
beginning with the top of the plant and proceed-
morph, with short necks, with air canals. Branches
ing basipetally (Bamboo Phylogeny Group 2012).
subequal, 5 to 7, in a horizontal line at each node,
Rather than recognize subtribes in Arundi-
from a single bud; nodes not prominent. Culm
narieae, I have followed the work of Triplett and
leaves with persistent sheaths, without auricles,
Clark (2010), Zeng et al. (2010), and Zhang et al.
the blade erect to reflexed. Foliage leaf blade
(2012) and simply listed major clades. Even with
with fimbriae. Inflorescence unbranched. Spike-
large amounts of molecular data, relationships
lets with short pedicels, with 1 fertile flower.
within the tribe are poorly resolved (Burke et al.
Glumes 2. Lodicules 3. Stamens 3. Stigmas 3.
2012; Ma et al. 2014). Most genera are para- or
One sp., B. tessellata (Nees) Stapleton, moun-
polyphyletic, and generic limits are thus highly
tains of South Africa.
likely to change in the near future. Like rhizome
This species is placed in an uncertain position
morphology, characteristics of inflorescence
in molecular analyses (Triplett and Clark 2010;
structure are highly homoplasious on the molec-
Zeng et al. 2010; Zhang et al. 2012), and may be in
ular phylogeny as is leaf epidermal micromor-
a clade sister to all other Arundinarieae. It was
phology (Zhang et al. 2014). Because these were
formerly placed in Thamnocalamus but is clearly
the central characters on which bamboo classifi-
misclassified there. Bergbambos may be sister to
cation was based, it is not surprising that generic
the Sri Lankan Kuruna (Attigala et al. 2014), or to
limits are in extensive need of revision.
Indocalamus wilsonii (Zeng et al. 2010).
Another confounding factor is hybridization.
While the irregular flowering of the bamboos was
thought to minimize opportunities for hybridiza-
tion, crossing has now been clearly documented; CLADE II. OLDEANIA
it appears that there are few reproductive barriers
in this group other than differences in phenology 25. Oldeania Stapleton
(Triplett and Clark 2010; Zhang et al. 2012). The
hybrid taxa tend to combine the morphological Oldeania Stapleton, Phytokeys 25: 100 (2013).
characteristics of the parents and have thus often
been named as new genera. Hybrid genera Culms 15 (20) m tall, erect to nodding. Rhi-
include Hibanobambusa, a hybrid between Sasa zomes pachymorph, with long necks. Aerial
and Phyllostachys (Triplett and Clark 2010; Tri- roots common, particularly at lower nodes,
plett et al. 2014); Pseudosasa japonica, a hybrid spiny. Branches 3 to 5, subequal, the culm slightly
between Sasamorpha and Pleioblastus (Triplett grooved above them, with a well-developed ridge
and Clark 2010; Triplett et al. 2014); Sasaella, a above the node. Culm leaves with sheaths decid-
hybrid between Sasa and Pleioblastus (Triplett uous. Inflorescence branches branched, the
and Clark 2010, and references therein; Triplett branches subtended by bracts, remnants of
et al. 2014); Semiarundinaria (including Brachys- sheaths or hairs. Spikelets with several flowers.
tachyum), a hybrid between Phyllostachys and Empty glumes 2, fertile glumes 4–8. Lodicules 3.
Pleioblastus (Triplett et al. 2010, 2014). These Stamens 3. Stigmas 2.
are not described here. Pseudosasa japonica is Species one, O. alpina (K. Schum.) Stapleton,
the type of P. subg. Pseudosasa, which also con- mountains of Africa.
tains P. owatarii; nothing is known of the origin O. alpina was originally described in Yushania
of the latter. Pseudosasa subg. Sinicae may be but is clearly unrelated to the Asian members of
non-hybrid and is described below in Clade IV, that genus (Zeng et al. 2010). Yushania ambosi-
the Shibataea clade. trensis and possibly all Madagascan Arundinaria
species may also belong to Oldeania, based on

phylogenetic results from Triplett and Clark
(2010).
CLADE III. CHIMONOCALAMUS
26. Chimonocalamus J. R. Xue & T. P. Yi Fig. 20
Chimonocalamus J. R. Xue & T. P. Yi, Acta Bot. Yunnan 1:

75 (1979).
Culms erect, the shoots fragrant. Rhizomes short,

pachymorph. Aerial roots common, particularly
at lower nodes, spiny. Branches 3 to 5, in a hori-
zontal line, subequal or with 3 dominant
branches. Culm leaves with deciduous sheaths,
auriculate or not, with fimbriae, the blade erect
or reflexed. Foliage leaves without auricles, with
or without fimbriae, without prominent cross
veins. Inflorescence branches branched, without
bracts. Spikelets with 4 to 12 flowers. Glumes 2.
Lodicules 3. Stamens 3. Stigmas 2.
Eleven spp., southwest China, eastern Hima-
layas, Burma.
Chimonocalamus is generally monophyletic
in molecular analyses (Triplett and Clark 2010;
Yang et al. 2013; Zhang et al. 2012), but some
species of Chimonocalamus have acquired chlor-
oplasts from species in the Phyllostachys clade,
indicating intergeneric hybridization (Yang et al.
2013). This probably explains the apparent poly-
phyly of the genus in the phylogeny of Zeng et al. Fig. 20. Bambusoideae-Arundinarieae. A–L Chimonoca-
(2010). lamus delicatus. A Portion of culm showing branching
pattern and spiny adventitious roots. B Flowering branch-
let. C Spikelet. D Portion of rachilla internode. E Lower
glume. F Upper glume. G Lemma. H Abaxial view of palea.
CLADE IV. SHIBATAEA CLADE I Adaxial view of palea. J Lodicules. K Abaxial and lateral
views of stamen. L Pistil. M–O C. pallens. M Portion of
culm. N Leafy branchlet. O Abaxial and adaxial views of
Rhizomes leptomorph. Lodicules 3. culm sheath. (From Wu et al. 2007, p. 133, with permission
from the Missouri Botanical Garden Press, St. Louis, and
Science Press, Beijing)
Five genera, 37+ species.
The Shibataea clade is monophyletic in phyloge- latter phylogeny is poorly supported and could be
nies based on plastid genes (Triplett and Clark simply uninformative about the existence of the
2010; Zeng et al. 2010) and could represent a clade.
recircumscribed Shibataeinae. The traditional 27. Ferrocalamus J. R. Xue & Keng f.
Shibataeinae are polyphyletic. In addition to the
genera listed below, three species of “Indocala- Ferrocalamus J. R. Xue & Keng f., J. Bamboo Res. 1: 3
mus” fall in this group (Zeng et al. 2010). The (1982).
clade has no obvious morphological synapomor-
phy, and is not seen the nuclear gene (GBSSI) Culms erect or scrambling, plants shrubby. Inter-
phylogeny of Zhang et al. (2012). However, the nodes terete, with white hairs below the nodes.
Branches 1 per node, or 3 to 5 per node higher up phyletic group that will probably be recognized as
on the plant. Culm leaves with persistent sheaths, a new genus. “Pseudosasa” as conventionally cir-
with or without auricles and fimbriae, the blades cumscribed is polyphyletic (Zeng et al. 2010).
reflexed. Foliage leaves with sheaths lacking auri-
cles, with fimbriae, the blades with distinct trans- 30. Shibataea Makino ex Nakai
verse veins. Inflorescence branches branched,
flexuous. Spikelets with 3 to 10 flowers. Glumes Shibataea Makino ex Nakai, J. Jap. Bot. 9: 83 (1933).
2. Stamens 3. Style 1, stigmas 2. Caryopsis with
fleshy pericarp. 2n ¼ 48. Culms erect, the plants less than 1 m tall. Culms
Two spp., China. flattened above branches. Branches 3 to 5 per node,
This genus is monophyletic in the analyses of more or less equal, without additional orders of
Triplett and Clark (2010) and Zeng et al. (2010). branching. Culm leaves with deciduous sheaths,
lacking auricles, the blade slender, reflexed. Foliage
28. “Gelidocalamus” T. H. Wen leaves with sheaths narrow, similar to petioles, con-
tiguous with the branchlets, the ligule long and
“Gelidocalamus” T. H. Wen, J. Bamboo Res. 1: 21 (1982). involute, the blade with distinct transverse veins;
terminal leaf of each branch lacking a sheath. Inflo-
Culms erect, plants shrubby, caespitose. Branches rescence a fascicle of pseudospikelets, each with a
7 to 12 (to 20) per node, without additional orders spathe-like prophyll and up to 3 bracts, with or
of branching. Culm leaves with persistent without buds in their axils. Pseudospikelets with
sheaths, with or without auricles, the blades 2 to 7 flowers, only the lower 1 or 2 fertile. Stamens
erect. Foliage leaves generally 1 per branch, 3. Style 1, stigmas 3. 2n ¼ 48.
apparently without auricles or fimbriae, the Seven spp., China and Japan.
blades with distinct transverse veins. Inflores- This genus is monophyletic in molecular phy-
cence branches branched. Spikelets with 3 to 5 logenies (Triplett and Clark 2010; Zeng et al.
flowers. Glumes 2. Stamens 3. Stigmas 2. Fruit 2010).
globose, beaked.
Nine spp., China.
“Gelidocalamus” is polyphyletic in molecular CLADE V. PHYLLOSTACHYS CLADE, sensu Zeng et al.
analyses (Zeng et al. 2010; Zhang et al. 2012), with (2010)
some species perhaps related parts of the poly-
phyletic “Indocalamus”. Lodicules 3. Stamens generally 3. Style 1.
Fourteen genera, 392 species.
29. “Pseudosasa” Makino ex Nakai
In addition to the genera listed below, several
“Pseudosasa” Makino ex Nakai, J. Arnold Arbor. 6: 150 species of “Pseudosasa” and “Sasa” fall in this
(1925), p.p. clade. While the clade is retrieved in phylogenies
based on plastid genes, it is not seen the nuclear
Culms erect or nodding; internodes terete or gene (GBSSI) phylogeny of Zhang et al. (2012);
grooved. Branches 1 to 3 per node. Culm leaves however, the latter phylogeny is poorly supported
with sheaths deciduous to persistent, auricles and and could be simply uninformative about the
fimbriae present or absent, the blade erect to existence of the clade.
reflexed. Foliage leaves with sheaths persistent,
auricles and fimbriae present or absent, the
31. “Acidosasa” C. D. Chu & C. S. Chao ex Keng f.
blades with distinct transverse veins. Inflores-
cence unbranched or branched, with the spikelets “Acidosasa” C. D. Chu & C. S. Chao ex Keng f., J. Bamboo
pedicellate on a central axis. Spikelets with 3 to 30 Res. 1: 31 (1982).
flowers. Glumes 2. Stamens 3. Stigmas 3. Metasasa W. T. Lin, Acta Phytotax. Sin. 26: 145 (1988).
Nineteen spp., China, Japan, Taiwan, Korea.
Triplett and Clark (2010) indicate that the Chi- Culms erect, shrubby to arborescent. Rhizomes
nese species of Pseudosasa and Sasa form a mono- leptomorph, elongate. Branches 3, or more at
higher nodes. Culm leaves with deciduous without fimbriae, the blades tessellate with obvi-
sheaths, with or without auricles, the auricles ous cross veins. Inflorescence unbranched or
bearing fimbriae, the blade erect or reflexed. with sparsely branched branches, the spikelets
Foliage leaves without auricles or fimbriae, the pedicellate on a central axis, subtended by
blades with distinct transverse veins. Inflores- reduced sheaths or rings of hairs. Spikelets with
cence unbranched or branched, without bracts, two glumes, the glumes mucronate, with 4 to 14
the spikelets pedicellate on a central axis. Spike- flowers. Stamens sometimes up to 5. Stigmas 2 to
lets elongate, with several to many flowers. 3. Ovary glabrous.
Glumes 2 to 4. Stamens 6. Stigmas 3. Two spp., China.
Eleven spp., southern China, Vietnam. “Bashania” is sometimes considered a subge-
This genus is polyphyletic in the analyses of nus of Arundinaria, but the two genera are clearly
Zeng et al. (2010). A. purpurea is supported as not related. “Bashania” is polyphyletic in the
being sister to Arundinaria gigantea in the phy- analyses of Zeng et al. (2010).
logenomic study of Burke et al. (2012), but other
species of Acidosasa were not included.
34. Chimonobambusa Makino
32. “Ampelocalamus” S. L. Chen, T. H. Wen & G. Y. Chimonobambusa Makino, Bot. Mag. (Tokyo) 28: 153
Sheng (1914); tax.: Stapleton (1994b).
Oreocalamus Keng, Sunyatsenia 4: 146 (1940).
“Ampelocalamus” S. L. Chen, T. H. Wen & G. Y. Sheng, Qiongzhuea J. R. Xue & T. P. Yi, Acta Bot. Yunnan. 2:
Acta Phytotax. Sin. 19: 332 (1981); tax.: Stapleton (1994b). 91–92 (1980).
Menstruocalamus T. P. Yi, J. Bamboo Res. 11: 38 (1992).
Culms clumped, pendulous above. Rhizomes
pachymorph. Branches several per node, 1 often Culms erect, plants shrubby. Rhizomes lepto-
dominant. Culm leaves with deciduous sheaths, morph, elongate. Culm internodes often grooved,
auricles present, fimbriae generally present, the the basal nodes often with prop roots modified to
blade reflexed. Foliage leaves with sheaths with form a ring of spines. Branches 3 (7), more or
auricles, with prominent spreading fimbriae, the less equal. Culms leaves with deciduous sheaths,
blades without obvious cross veins. Inflorescence auricles tiny or absent, the blade much reduced
pendulous, unbranched, or with sparse branched (less than 1 cm long), often simply an acicular
branches, with few or no bracts, the spikelets on point. Foliage leaves with or without auricles
long pedicels borne in clusters at the nodes. Spi- and fimbriae. Inflorescence unbranched, or the
kelets with 2 to 7 fertile flowers plus 1 reduced branches branched, subtended by bracts with
flower distally. Glumes 2. Stigmas 2 or 3. Ovary buds in their axils. Pseudospikelets with several
glabrous, without an appendage. to many flowers. Glumes 1 to 3, often with a bud
Thirteen spp., central Himalayas to southern in the axil of one. Stigmas 2 or 3. Fruit nut-like.
China. Thirty-seven spp., China and Japan.
This genus is polyphyletic in the analyses of The analyses of Triplett and Clark (2010)
Zeng et al. (2010) and Yang et al. (2013), although provide no evidence to accept or reject the mono-
the latter study shows that most species form a phyly of this genus. Zhang et al. (2012) and Peng
clade, with only “A”. calcareus originating inde- et al. (2008) present molecular data to support
pendently; see Clade XI below. synonymy of Qiongzhuea and Menstruocalamus
under Chimonobambusa.
33. “Bashania” Keng f. & T.P. Yi
35. “Drepanostachyum” Keng f.
“Bashania” Keng f. & T.P. Yi, J. Bamboo Res. 1: 171
(1982). “Drepanostachyum” Keng f., J. Bamboo Res. 2: 15 (1983);
tax., Stapleton (1994b).
Culms erect. Rhizomes leptomorph. Branches 3
to 7 per node, from a single bud. Culm leaves Plants shrubby, caespitose. Rhizomes short,
deciduous, the sheaths without auricles, with fim- pachymorph. Branches up to 80 per node, in 2 or
briae, the blade erect. Foliage leaves auriculate, 3 rows, all more or less equal. Culm leaves with
sheaths deciduous, adaxially scabrous, without short. Branches up to 25 per node, the central one
auricles, with or without fimbriae, the blade dominant, from a single bud. Culm leaves with
reflexed. Foliage leaves lacking auricles or fim- sheaths deciduous, leaving a narrow rim, without
briae, without transverse veins. Inflorescence auricles or fimbriae, the blade reflexed. Foliage
branches branched, the branches subtended by leaves with sheaths with or without auricles and
reduced bracts or rings of hairs; basal internodes fimbriae, with transverse veins obscure or absent.
of the inflorescence short so that branches appear Inflorescence unbranched or branched, the
fascicled. Spikelets conventional, with 2 to 6 fer- branches sometimes subtended by rings of
tile flowers plus one reduced flower distally. hairs; basal internodes of the inflorescence short
Glumes 2. Stigmas 2. Ovary glabrous. so that branches appear fascicled. Spikelets con-
Ten spp., Himalayan regions of Bhutan, ventional, with 1 (or 2) fertile flowers proximally,
China, India, and Nepal. 1 reduced flower distally. Glumes 2. Stigmas 2.
“Drepanostachyum” is not monophyletic in Eight spp., Bhutan, China, India, Nepal.
the chloroplast phylogeny of Triplett and Clark “Himalayacalamus” is not monophyletic in
(2010). The nuclear gene phylogeny of Zhang the chloroplast phylogeny of Triplett and Clark
et al. (2012) suggests that it may be derived within (2010). The nuclear gene phylogeny of Zhang
“Himalayacalamus”. et al. (2012) suggests that it may be paraphyletic,
with “Drepanostachyum” derived from within it,
but the tree topology is not well enough sup-
36. “Fargesia” Franchet ported to defend combining the two.
“Fargesia” Franchet, Bull. Mens. Soc. Linn. Paris 2: 1067
(1893). 38. Not “Indocalamus” Nakai
Sinarundinaria Nakai, J. Jap. Bot. 11: 1 (1935).
Borinda Stapleton, Edinburgh J. Bot. 51(2): 284 (1994). Not “Indocalamus” Nakai, J. Arnold Arbor. 6: 148 (1925).
Culms erect. Rhizomes pachymorph, short. Culms nodding, clumped, the plants shrubby,
Branches 7 to numerous per node, becoming generally less than 2 (3) m tall. Rhizomes lepto-
deflexed, subequal. Culm leaves with sheaths per- morph. Internodes terete, with a dense yellow-
sistent or deciduous, leaving a short rim, with or brown ring of hairs below the nodes. Branches
without auricles, with or without fimbriae, the blade one per node, about the same thickness as the
reflexed or erect. Foliage leaves with or without culm. Culm leaves with persistent sheaths, gener-
auricles, with or without fimbriae. Inflorescences
ally with auricles and fimbriae, the blade gener-
unbranched, or the branches branched, generally
subtended by spathe-like sheaths, the spikelets ally recurved. Foliage leaves with sheaths without
borne on one side of the axis or branch. Spikelets auricles, with fimbriae, the blades with distinct
with several flowers. Glumes generally 2. Lemma transverse veins. Inflorescence unbranched or
apex mucronate to awned. Style branches some- branched, the branches subtended by small
times 2, stigmas 2 or 3. bracts. Spikelets with 3 to more than 12 flowers.
Ninety spp., east and southeast Asia. Glumes 2 or 3. Style 1 to nearly absent, stigmas
“Fargesia” is not monophyletic in the chloro- 2 (3). 2n ¼ 48.
plast phylogenies of Triplett and Clark (2010) or Twenty-one spp., one in Japan, all others in
Zeng et al. (2010). The inclusion of Sinarundinaria China.
and Borinda in Fargesia follows the recommenda- “Indocalamus” is polyphyletic (Zeng et al.
tion of the Bamboo Phylogeny Group (2012). 2010; Zhang et al. 2012). The former authors
included 21 of the 23 named species, which fall
37. “Himalayacalamus” Keng f. in virtually every clade of Arundinarieae. Most
members of the “genus” belong here in the Phyl-
“Himalayacalamus” Keng f., J. Bamboo Res. 2: 23 (1983); lostachys clade, but the type (I. sinicus (Hance)
tax., Stapleton (1994b). Nakai) and I. wilsonii are in an uncertain position
outside other major clades in the tribe. The
Culms nodding or pendulous, plants shrubby to description above refers to the majority of the
more or less arborescent. Rhizomes pachymorph,
species and will need to be emended as the genus The analyses of Triplett and Clark (2010)
is redefined. provide no evidence to accept or reject the mono-
phyly of “Phyllostachys”, but it is not monophy-
39. “Oligostachyum” Z. P. Wang & G. H. Ye letic in the analyses of Zeng et al. (2010).
“Oligostachyum” Z. P. Wang & G. H. Ye, J. Nanjing Univ., 41. “Pleioblastus” Nakai

Nat. Sci. 1982: 95 (1982).
Clavinodum T. H. Wen, J. Bamboo Res. 3: 23 (1984). “Pleioblastus” Nakai, section Amari S.L. Chen & G.Y.
Sheng, pro parte.
Culms erect, plants shrubby to arborescent. Rhi- Nipponocalamus Nakai, J. Jap. Bot. 18: 350 (1942).
zomes leptomorph, elongate. Branches 3 (to 7) Polyanthus C. H. Hu, J. Bamboo Res. 10(3): 28–30 (1991).
per node, subequal, from a single bud. Culm
leaves with sheaths generally deciduous, with or Culms erect, sometimes less than 1 m tall.
without auricles and fimbriae, the blades erect or Rhizomes leptomorph. Internodes flattened or
reflexed. Foliage leaves with or without auricles grooved. Branches 3 to 9 per node, subequal,
and fimbriae, with distinct transverse veins. Inflo- from a single bud, producing extensive secondary
rescence terminal on lateral branches, with 2 or 3 branches. Culm leaves with sheaths persistent,
(6) spikelets pedicellate on a slender axis. Spi- with or without auricles, with or without fimbriae,
kelets with several to many flowers, the proximal the blade reflexed. Foliage leaves with or without
ones sterile. Glumes (1) 3 (5), forming a morpho- auricles, with or without fimbriae, the ligules nota-
logical transition to the proximal sterile lemmas. bly long. Inflorescence unbranched or with
Stamens sometimes 4 or 5. Stigmas 3. branched branches. Spikelets with 2 glumes.
Fifteen spp., China. Stamens sometimes up to 5. Stigmas 2 or 3.
“Oligostachyum” is not monophyletic in the Ca. forty spp., Japan, China.
analyses of Zeng et al. (2010). “Pleioblastus” in its conventional sense is not
monophyletic (Zeng et al. 2010). The Japanese
species form a well-supported clade (Zeng et al.
40. “Phyllostachys” Siebold & Zucc.
2010; J. Triplett, pers. comm.), but the relation-
“Phyllostachys” Siebold & Zucc., Abh. Math.-Phys. Cl. ship of these to the various Chinese species is less
K€onigl. Bayer. Akad. Wiss. 3: 745, pl. 5, f. 3 (1843) nom. clear. At least some Chinese species likely belong
conserv. in the genus, but others will need to be trans-
ferred elsewhere. J. Triplett (pers. comm.) notes
Culms erect, the plants arborescent or shrubby. that this genus includes species with rhizomes
Rhizomes leptomorph, elongate. Culms flattened that are leptomorph (monopodial) but short,
above branches. Branches 2 (to 3) at mid-culm thick and tillering, as well as species with more
nodes, unequal, with higher-order branches; conventional elongate leptomorph rhizomes.
internode at base of branch complement short, These two types of rhizome correspond to major
failing to elongate. Culm leaves with sheaths phylogenetic subdivisions in the genus.
deciduous, with auricles and fimbriae, the
blade erect or recurved. Foliage leaves with or 42. Sarocalamus Stapleton
without auricles and fimbriae, the blades with
distinct transverse veins, generally pilose on the Sarocalamus Stapleton, Novon 14: 346. (2004).
proximal portion of the abaxial side. Inflores-
cence of unbranched branches, arranged in brac- Culms erect to nodding, the internodes terete. Rhi-
teate fascicles, the most proximal bract often zomes leptomorph. Branches 1 to 3 per node, erect,
subtending a bud, the next 2 to 6 more distal branching profusely, basal internodes elongate.
bracts scale-like and the 2 to 7 distalmost Culm leaves with sheaths deciduous or persistent,
spathe-like. Pseudospikelets with 2 to 6 fertile with or without fimbriae and auricles, the blade
flowers plus one reduced flower distally. Glumes erect or reflexed. Foliage leaves with or without
0 or 1 (3). Stigmas (1 to) 3. Fruit with an abaxial auricles and fimbriae. Inflorescence unbranched,
groove. 2n ¼ 48. or the branches branched. Spikelets with 4 to 10
Fifty-one spp., Asia. flowers. Glumes 1 or 2. Stigmas 2 or 3.
Three spp., Bhutan, China, NE India, Nepal. Yushania is polyphyletic in the analyses of
Triplett and Clark (2010), with the species from
alpine regions in Africa (Y. alpina) and Madagas-
43. “Sinobambusa” Makino ex Nakai
car (Y. ambositrensis) forming a clade distinct
“Sinobambusa” Makino ex Nakai, J. Arnold Arbor. 6: 152 from the other species of Yushania sampled.
(1925). However, if the African alpine bamboos (now in
Oldeania) are excluded then the genus may be
Culms erect to nodding, the plants arborescent or monophyletic. This description is based largely
shrubby. Rhizomes leptomorph, elongated. Inter- on Chinese material. The type species, Y. niita-
nodes grooved. Branches 3 (5 to 7), more or less kayamensis also falls here, based on the phylog-
equal. Culm leaves with deciduous sheaths, with eny of Triplett and Clark (2010). The observation
auricles and fimbriae. Foliage leaves with or with- of root spines is from Wong (2004).
out auricles and fimbriae. Inflorescence bracteate,
unbranched, bearing 1 to 3 pseudospikelets, each
subtended by a prophyll and 2 or more bracts, the CLADE VI. ARUNDINARIA CLADE
more distal bracts often bearing buds in their
axils; buds developing into additional spikelets. Rhizomes leptomorph, elongate.
Pseudospikelets with as many as 50 flowers. Stig-
mas 2 or 3. Five genera, 126 spp.
Ten spp., China, Taiwan, Vietnam, intro-
duced to Japan. This clade has no obvious morphological synap-
The distinction between “Sinobambusa” and omorphy.
“Indosasa” is based largely on stamen number, Some species formerly known as Pleioblastus
with fewer than 6 (generally 3) in the former fall in this clade, but have not yet been placed in a
versus 6 in the latter. “Sinobambusa” is not new genus. Numbers of species are based on
monophyletic in the analyses of Zeng et al. (2010). estimates from Triplett and Clark (2010).
44. Yushania Keng f. 45. Arundinaria Michx.
Arundinaria Michx., Fl. Bor.-Amer. 1: 73 (1803); phylog.:

Yushania Keng f., Acta Phytotax. Sin. 6: 355 (1957); tax.:
Triplett et al. (2010).
Stapleton (1994c).
Burmabambus Keng f., J. Bamboo Res. 1(2): 173 (1982).
Butania Keng f., J. Bamboo Res. 1(2): 42 (1982). Culms erect, caespitose or not. Branches 1 to 3
Monospatha W.T. Lin, J. Bamboo Res. 13(4): 1–2 (1994). per node, subequal, from a single bud, the
branches often extensively rebranching. One to
Culms erect, plants shrubby; lower nodes some- five internodes at the base of each branch com-
times developing root spines. Rhizomes pachy- plement failing to elongate. Culm leaves with
morph, with elongated necks. Branch number sheaths persistent or deciduous, with auricles,
varying from 1 at most basal nodes to 9 to 11 the blades erect or reflexed. Foliage leaves with
(to 45) at upper nodes, more or less equal in size, auricles and fimbriae, the blades with fine trans-
or the central one larger, from a single bud. Culm verse veins. Inflorescence unbranched, or the
leaves with sheaths persistent or deciduous, with branches branched, with the spikelets pedicellate.
or without auricles, with or without fimbriae, the Spikelets with 6 to 12 flowers. Glumes 1 or 2.
blade erect or reflexed. Foliage leaves with Stamens 3. Style reduced or absent, stigmas 3.
sheaths with or without auricles, with or without 2n ¼ 48.
fimbriae, with transverse veins. Inflorescence Three spp., southeastern US.
branches branched, flexuous, not subtended by This circumscription of Arundinaria follows
bracts. Spikelets with several to many flowers, the Triplett et al. (2010). Arundinaria subgenera
distalmost flower reduced. Glumes 2. Stigmas 2. Bashania and Sarocalamus fall in the Phyllosta-
Seventy-seven spp., Africa, Madagascar, East chys clade and are here recognized as distinct
and southeast Asia. genera. Species previously classified as
Arundinaria in Sri Lanka are unrelated to Arun- 48. Sasamorpha Nakai

dinaria s.s. and are now placed in the genus
Kuruna (Attigala et al. 2014). Sasamorpha Nakai, J. Fac. Sci. Hokkaido Univ. V, Bot. 26:
180 (1931).
46. “Indosasa” McClure Shrubby plants less than 1.5 m tall. Culms and
axis of inflorescence white powdery. Internodes
“Indosasa” McClure, Lingnan Univ. Sci. Bull. 9: 28 (1940). terete, nodes flat. Branches 1 per node. Culm
leaves with persistent sheaths; auricles and fim-
Culms erect to nodding. Branches 3 per node, briae lacking. Foliage leaves with sheaths lacking
more or less equal or the central one larger than auricles and fimbriae, with blades with distinct
the others, the internode grooved above the transverse veins. Inflorescence branched or
branches. Culm leaves with sheaths deciduous, unbranched, with tiny bracts. Spikelets with 4 to
with or without auricles and fimbriae, the blade 8 flowers. Glumes 2. Lodicules 3. Stamens 6. Style
erect or reflexed. Foliage leaves with or without 1, stigmas 3.
auricles and fimbriae, the blades with distinct Five spp., China, Japan, Korea, Eastern Rus-
transverse veins. Inflorescence unbranched, brac- sia.
teate, branches with prophylls. Pseudospikelets Sasamorpha was segregated from Sasa on the
sessile, with many flowers. Glumes 2. Lodicules basis of the white powdery wax on the culms, and
3. Stamens 6. Style 1, stigmas 3. Fruit beaked. the non-auriculate culm leaves. Also the culm leaf
Fifteen spp., China and Viet Nam. sheaths are longer than the internodes, whereas
“Indosasa” is not monophyletic in the ana- they are shorter in Sasa.
lyses of Zeng et al. (2010).
47. Sasa Makino & Shibata, s.s. CLADE VII. THAMNOCALAMUS Munro
Sasa Makino & Shibata, Bot. Mag. (Tokyo) 15: 18 (1901), 49. Thamnocalamus Munro
s.s.
Thamnocalamus Munro, Trans. Linn. Soc. London 26: 33
Plants shrubby, less than 2 m tall, culms erect or (1868); tax.: Stapleton (1994c); phylog.: Guo and Li (2004).
sometimes leaning. Internodes terete, nodes
prominent. Branches 1 per node. Culm leaves Plants shrubby, culms erect to nodding. Rhizomes
with persistent sheaths, with or without auricles pachymorph. Branches about 5 per node, with one
and fimbriae, the blades erect or reflexed. Foliage somewhat larger than the others; branchlets long
leaves with or without auricles and fimbriae, the and pendulous. Culm leaves with sheaths decidu-
blades with distinct transverse veins. Inflores- ous, without auricles, with fimbriae, the blade awl-
cence branched or unbranched, with tiny bracts. shaped, erect. Foliage leaf sheaths without auricles
Spikelets with 4 to 8 flowers. Glumes 2. Lodicules or fimbriae, the blades with distinct transverse
3. Stamens 6. Style 1, stigmas 3. veins. Inflorescence branches branched, the basal
Forty spp., Japan, Korea, eastern China and branches with bracts. Spikelets with 1 to many
eastern Russia. flowers, the distalmost flower reduced. Glumes 2.
“Sasa” as conventionally circumscribed is Stigmas 3.
polyphyletic (Triplett and Clark 2010; Zeng et al. Two spp., China and Himalayas.
2010), with species falling in both the Arundi- Thamnocalamus s.l. is not monophyletic in
naria clade and the Shibataea clade. The type the chloroplast phylogeny of Triplett and Clark
species, S. veitchii (Carrière) Rehder, falls into (2010), but may form a clade when Bergbambos
the Sasa subgroup in the Arundinaria clade (Tri- (Clade I) is excluded. Triplett and Clark (2010)
plett and Clark 2010), so this clade constitutes place Thamnocalamus s.s in the Phyllostachys
Sasa s.s. The placement of the named hybrid clade, whereas Zeng et al. (2010) place it in its
Sasella in this clade shows that the maternal par- own clade.
ent of Sasella was a species of Sasa.
CLADE VIII. “INDOCALAMUS” WILSONII (Rendle) C. S. Culms 1–4 m tall, nodding, clumped. Internodes
Chao & C. D. Chu terete, glabrous. Rhizomes leptomorph. Branch
generally 1 per node, nearly as thick as the
50. “Indocalamus” wilsonii (Rendle) C. S. Chao & culm. Culm leaves persistent, auricles absent,
C. D. Chu fimbriae present, blade erect to spreading. Foliage
leaves without auricles, with fimbriae, the blades
“Indocalamus” wilsonii (Rendle) C. S. Chao & C. D. Chu, with obvious cross veins. Inflorescence branches
J. Nanjing Techn. Forest Prod. 1981: 43 (1981). branched. Spikelets with 3 or 4 flowers. Glumes 2.
Stamen number unknown. Stigmas 2.
Culms less than 1 m tall. Culm leaves without One sp., I. sinicus (Hance) Nakai, China.
auricles or fimbriae, the blade recurved. Foliage Indocalamus sinicus is unrelated to other spe-
leaves with sheaths drying orange-red, without cies of “Indocalamus”, which fall in the Phyllos-
auricles or fimbriae, the blades wavy when dry, tachys clade (Zeng et al. 2010). Because this is the
with obvious cross-veins. Inflorescence branches type species, the remaining species will ultimately
branched. Spikelets with 3 to 7 flowers. Glumes 2. need to be reclassified.
Style reduced or absent, stigmas 3.
One sp., I. wilsonii (Rendle) C. S. Chao & C. D.
Chu, China.
This species is sister to Bergbambos, but with- CLADE XI. “AMPELOCALAMUS” CALCAREUS
out support, in the analyses of Zeng et al. (2010).
53. “Ampelocalamus” calcareus C.D. Chu & C.S.
It is unrelated to other species of “Indocalamus”, Chao
nearly all of which are members of the Phyllosta-
chys clade. “Ampelocalamus” calcareus C.D. Chu & C.S. Chao, Acta
Phytotaxonomica Sinica 21: 204–206.
“A”. calcareus is in a position unrelated to other

CLADE IX. GAOLIGONGSHANIA D.Z. Li, Hsueh & N.H. Xia
species of Ampelocalamus in the phylogenies of
51. Gaoligongshania D.Z. Li, Hsueh & N.H. Xia Yang et al. (2013) and Ma et al. (2014), and is thus
placed in Clade XI of the Arundinarieae; in the
Gaoligongshania D.Z. Li, Hsueh & N.H. Xia, Acta Phyto- latter phylogeny it is sister to all other species of
tax. Sin. 33: 598 (1995). the tribe. The species is poorly known and is not
given a full description here, nor is it placed in the
Culms scrambling or epiphytic. Rhizomes pachy- key pending more comprehensive analysis.
morph. Branches 1 per node, similar in thickness
to the culm. Culm leaves with persistent leathery
sheaths, with auricles, the auricles bearing fim-
CLADE XII. KURUNA Attigala, Kaththriarachchi & L.G.
briae, the blade recurved. Foliage leaf sheaths
Clark
with fimbriae, the blades with transverse veins.
Inflorescence branches branched, sparse, without 54. Kuruna Attigala, Kaththriarachchi & L.G. Clark
bracts. Spikelets pedicellate, with 4 to 10 flowers,
the distal one reduced. Glumes 2. Lemma with a Kuruna Attigala, Kaththriarachchi & L.G. Clark, Phyto-
long awn. Lodicules 3. Stamens 3. Style 1, stigmas taxa 174: 199 (2014).
3. Ovary stalked.
One sp., G. megalothyrsa (Hand.-Mazz.) D.Z. Culms erect to scandent, the internodes terete,
Li, J.R. Xue & N.H. Xia, China. flattened or sulcate. Rhizomes pachymorph,
with short necks. Branches several per node,
from a single bud. Culm leaves hispid abaxially
with fimbriae, without auricles, the blades
CLADE X. INDOCALAMUS Nakai, s.s.
reflexed. Foliage leaf sheaths with fimbriae,
52. Indocalamus Nakai erect. Inflorescences branched or unbranched,
open, the branches lacking subtending bracts.
Indocalamus Nakai, s.s. , J. Arnold Arbor. 6: 148 (1925). Glumes 2, fertile flowers (1) 2 to 6. Palea with an
acute apex or with two sharp points. Stigmas

2 or 3.
Five spp., Sri Lanka.
V. TRIBE BAMBUSEAE Kunth ex Dumort. (1829)

Culms woody. Rhizomes pachymorph. Branch
development acropetal or bidirectional. Outer lig-
ule (contraligule) present. Spikelets laterally com-
pressed, disarticulating above the glumes. Midrib
complex. 2n ¼ (20) 40, (44), 46, 48, 70, 72.
Sixty-three genera, 784 spp.
All Bambuseae have pachymorph rhizomes,
except for Chusquea sect. Chusquea, in which the
rhizomes are amphimorph, presumably a derived
character. As in Arundinarieae, culms of mem-
bers of Bambuseae reach their full height without
initiating branches and produce only culm leaves;
later the culms lignify and produce branches and
foliage leaves, beginning with the base or middle
of the plant and proceeding acropetally or bidi-
rectionally (Bamboo Phylogeny Group 2012).
Height is strongly correlated with root pressure,
which can refill embolized xylem vessels (Cao
et al. 2012). Pseudospikelets are common among
members of Bambuseae, but do not occur in all
genera. The outer ligule also occurs in Strepto-
gyna (BEP clade, incertae sedis). Phylogenetic
analysis of the chloroplast genome shows that Fig. 21. Bambusoideae-Bambuseae-Chusqueinae. Chus-
Bambuseae can be divided into a neotropical quea latifolia. A Branch complement with foliage leaves.
B Inflorescence. C Spikelet. D Culm leaf. E Bud comple-
and a paleotropical group, each of which contains ment. F Node with branches emerging infravaginally; g
several well-supported subclades, recognized as girdle, a persistent section of the leaf sheath encircling the
subtribes (Kelchner and Bamboo Phylogeny culm. (From Clark 1989)
Group 2013). Monophyly of the neotropical
clade is not strongly supported, however. Phylo- absent, the sheaths lacking fimbriae or auricles,
genies of nuclear genes also identify neotropical the blades stiff and upright, rarely reflexed. Ligule
and paleotropical clades, with the C1 genome of with obvious veins. Foliage leaves lacking fim-
the neotropical group most closely related to the briae or auricles. Inflorescence branches
B genome of Arundinarieae (Triplett et al. 2014). branched, sometimes capitate. Spikelets with one
flower, rachilla extension lacking. Glumes four,
NEOTROPICAL CLADE the proximal two sometimes reduced, with or
V.1. SUBTRIBE CHUSQUEINAE Bews (1929) without awns. Lodicules 3. Stamens 3. Stigmas 2.
Stomatal subsidiary cells each with at least 2 papil-
Neurolepidinae Soderstr. & R. P. Ellis in Soderstr. et al., lae.
Grass Syst. Evol.: 238 (1987). One genus, 160 species.
Culms erect to scandent or scrambling, some- 55. Chusquea Kunth Fig. 21

times hanging, shrubby or arborescent. Rhizomes
pachymorph. Culms generally solid, unbranched, Chusquea Kunth, J. Phys. Chim. Hist. Nat. Arts 95: 151
with one bud per node; prop roots sometimes (1822); phylog.: Kelchner and Clark (1997), Fisher et al.
forming root spines. Culm leaves present or (2009, 2014).
Rettbergia Raddi, Agrostogr. Bras. 17–18, t. 1, f. 1 (1823). of the abaxial side with no or reduced wax. Lodi-
Neurolepis Meisn., Pl. Vasc. Gen. 1: 426 (1843). cules 3. Stamens 3. Stigmas 2. Leaf mesophyll with
Swallenochloa McClure, Smithsonian Contr. Bot. 9: 106, f.
43–45 (1973).
sclerenchyma fibers in intercostal regions; midrib
with a single vascular bundle. 2n ¼ 40.
Characters as for the subtribe. 2n ¼ 20, 40, 44, 48 Fifteen genera, 172 spp.
(excluding counts listed as doubtful by Judzie- The sclerenchyma fibers in the intercostal
wicz et al. 1999). regions are unusual in Poaceae. Although the
One hundred sixty spp., Central and South narrow waxless strip on the leaves is apparently
America, West Indies. synapomorphic for Arthrostylidiinae, the charac-
Chusquea is made up of four major clades, ter is also found in other bamboos, particularly
Chusquea subgenus Platonia Fisher & L.G. Clark, Melocanninae (Tyrell et al. 2012). Most, if not all,
C. subgenus Magnifoliae L.G. Clark & Fisher, C. other members of Bambuseae have a complex
subgenus Rettbergia (Raddi) L.G. Clark, and the midrib, consisting of several vascular bundles.
Euchusquea clade, which includes subgenus The phylogeny and considerable morpholog-
Chusquea and subgenus Swallenochloa, both of ical diversity in Arthrostylidiinae are described in
which are polyphyletic (Fisher et al. 2009, 2014). detail by Tyrell et al. (2012). The subtribe is mor-
The Euchusquea clade contains most of the spe- phologically diverse, with variation between
cies of the genus. Species in subgenera Platonia genera in patterns of culm internode elongation,
and Magnifoliae were formerly placed in the position of culm leaf blades (erect to reflexed),
genus Neurolepis, but this “genus” is clearly para- the shape of the prophylls (cordate or triangular),
phyletic; its former generic characters (lack of the number and patterning of branches at each
aerial branching, ligules with obvious veins, and node (one to several dominant ones, branches
a base chromosome number of x ¼ 12) could sometimes in a fan-shaped pattern), inflorescence
possibly be synapomorphies for Chusquea s.l. architecture (branched or unbranched, capitate
The presence of multiple buds at each node, or not), presence of pseudospikelets or conven-
with the buds dimorphic or trimorphic, is syna- tional spikelets, seed coat separable from the
pomorphic for Chusquea s.s. (i.e., subgenus pericarp or not, and pericarp fleshy or not.
Rettbergia plus Euchusquea).
GLAZIOPHYTON + CAMBAJUVA
ARTHROSTYLIDIINAE + GUADUINAE
Inflorescence branches erect, appressed to the
Arthrostylidiinae and Guaduinae are sister taxa, rachis, foliage leaves erect, strongly tessellate,
and share refractive papillae on leaf epidermal fusoid cells in the leaf mesophyll lacking.
cells (Guala et al. 2000; Kelchner and Clark 1997; Glaziophyton and Cambajuva form a clade in
Ruiz-Sánchez et al. 2008; Sungkaew et al. 2009; molecular phylogenies that is sister to all other
Zhang and Clark 2000). Although refractive members of the subtribe (Tyrell et al. 2012). Erect
papillae on the leaf epidermis are likely synapo- leaf blades also occur in some species of Aulone-
morphic for this pair of subtribes, they have ori- mia and Myriocladus but appear to be indepen-
ginated apparently in parallel in Melocanninae dently derived. Likewise tessellation of the
(Bamboo Phylogeny Group 2012). abaxial surface of the leaf occurs in high-altitude
species of both the latter genera; it is possible that
this vein pattern is related to low temperature
(Viana et al. 2013).
V.2. SUBTRIBE ARTHROSTYLIDIINAE Bews (1929)
56. Glaziophyton Franch.
Clump forming species with short thick (pachy-
morph) rhizomes. Branch buds one per node. Glaziophyton Franch., J. Bot. (Morot) 3: 277 (1889).
Prophylls triangular. Foliage leaves with well-
developed, curly or wavy fimbriae at the collar. Culms erect, with chambered pith. First internode
Wax distributed unequally over the foliage leaf very long, subsequent ones very short. Branches
blade surface, leaving one narrow marginal sector generally absent. Culm leaves rare, when present
the sheath without auricles or fimbriae, the apex near their summit by the collar, the blades
of the blade involute, forming a sharp point. spreading to reflexed. Foliage leaves on terminal
Inflorescence branches branched, each subtended branches broader than those below and often
by a basal bract, and with a prophyll. Spikelets reflexed. Inflorescence branches branched or
with 1 sterile flower and 2 fertile flowers. Glumes not, the spikelets on long stalks. Spikelets with
2. many flowers. Glumes 2, their apices acuminate.
One sp., G. mirabile Franch., Rio de Janeiro, Lemma acuminate or with a short beak. Fruit
Brazil. with pericarp free from seed coat.
One sp., A. verticillatum (Nees) Soderstrom,
57. Cambajuva P. L. Viana, L. G. Clark & Filg. Brazil and adjacent Bolivia.
Actinocladum is strongly supported by
Cambajuva P. L. Viana, L. G. Clark & Filg., Syst. Bot. 38: molecular data as sister to Merostachys (Tyrell
98 (2013). et al. 2012). Both genera have caryopses in
which the seed coat is free from the pericarp
Culms erect, the internodes hollow. Culm and fan-shaped clusters of branches. The latter
leaves deciduous, the sheath apex with fimbriae, character also appears independently in Rhipido-
the blade erect. Branches 1 to 7 per node from a cladum and some species of Arthrostylidium.
single bud, usually on a promontory. Foliage
leaves with sheaths bearing erect undulate to 59. Alvimia C. E. Calderón ex Soderstr. & Londoño
crispate fimbriae, the blade apex sharp-pointed.
Inflorescence branches branched, lacking sub- Alvimia C. E. Calderón ex Soderstr. & Londoño, Amer. J.
tending bracts and prophylls. Spikelets with 1 Bot. 75: 833 (1988).
sterile flower, 2 to 4 fertile flowers, and 1 distal
rudimentary flower. Glumes 2. Glumes and Culms scrambling and hanging from vegetation.
lemmas with awns. Lodicules 3. Anthers 3. Stig- Branches several to 100 or more, 1 branch domi-
mas 2. Leaf epidermis lacking refractive papillae. nant, from a single bud on a promontory just
One sp., C. ulei (Hack.) P.L. Viana, L.G. Clark above the node. Culm leaves with sheaths with a
& Filg., southern Brazil. dark line at the base, with or without fimbriae, the
Cambajuva was described to accommodate blades erect. Foliage leaves with dark pseudope-
Aulonemia ulei (Hack.) McClure, which is mor- tioles. Inflorescence branches branched. Pseu-
phologically and phylogenetically distinct from dospikelets long and narrow, with numerous
other species of Aulonemia. flowers. Lemma with a narrow tip. Stamens 2.
Fruit fleshy, falling with lemma, palea and rachis
Remaining species of the subtribe:
internode.
Reflexed pseudopetioles, the leaves thus appearing to
Three spp., Bahia, Brazil.
droop. This character is reversed in Filgueirasia.
Alvimia is strongly supported by molecular
Tyrell et al. (2012) recover three clades among
data as sister to Atractantha (Tyrell et al. 2012).
the remaining species: I. Arthrostylidium, Rhipido-
cladum, Didymogonyx, and Elytrostachys; II. Aulo-
nemia and Colanthelia, plus two species of 60. “Arthrostylidium” Rupr.
Arthrostylidium; III. Actinocladum, Merostachys,
Alvimia, Atractantha, and Filgueirasia. Although “Arthrostylidium” Rupr., Bambuseae 27 (1839); tax.: Jud-
ziewicz and Clark (1993).
these clades are moderately well supported by chlo-
roplast sequence data, none has a diagnostic mor-
Culms scandent at maturity. Branches 3 to many,
phological character.
with one dominant, the branches from a single
58. Actinocladum McClure ex Soderstr. bud on a promontory just above the node. Culm
leaf sheaths without fimbriae, the blades erect.
Actinocladum McClure ex Soderstr., Amer. J. Bot. 68: 1201 Foliage leaf sheaths with fimbriae. Inflorescences
(1981). unbranched, with a zigzag rachis in some species.
Spikelets with 1 to several flowers. Glumes 1 to 3.
Culms erect. Branch complements fan-shaped. Lemma with margins clasping the rachis.
Culm leaves with sheaths with ruffle-like fimbriae Thirty-two spp., Central and South America.
The fimbriae on the foliage leaves may be 63. “Aulonemia” Goudot

inconspicuous. This genus is polyphyletic in the
analyses of Tyrell et al. (2012). “Aulonemia” Goudot, Ann. Sci. Nat., Bot. III, 5: 75 (1846).
Matudacalamus F. Maek., J. Jap. Bot. 36(10): 345 (1961).
61. Athroostachys Benth. Culms erect or scandent. Mid-culm nodes with

one bud, producing one dominant branch nearly
Athroostachys Benth., Gen. Pl. 3: 1208 (1883).
as large as the culm. Culm leaf sheaths with fim-
briae, blades reflexed to spreading, deciduous.
Culms scandent and vine-like. Branches several
Foliage leaf sheaths often purple spotted. Inflo-
per node, all about the same diameter. Culm leaf
rescence branches branched. Spikelets with 2 or 3
sheaths persistent or deciduous, the blades
glumes and several flowers. Lemma with the apex
reflexed, the sheath with a proximal dark-colored,
obtuse, with a brief mucro or arista. 2n ¼ 40.
thickened rim, with fimbriae at the apex, without
Forty spp., montane regions of Central and
auricles. Foliage leaf sheaths with fimbriae, with-
South America.
out auricles. Inflorescence dense and capitate, the
“Aulonemia” as traditionally circumscribed
branches branched but not elongate. Spikelets
is polyphyletic. One species has been removed
with 1 flower plus a small rudimentary one.
to Cambajuva (Viana et al. 2013), and two other
Glume 1, deciduous, the apex acuminate, awn-
species were transferred to Olmeca (Ruiz-Sánchez
like.
et al. 2011a). The remaining species form a para-
One sp., A. capitata (Hook.) Benth., coastal
phyletic group from which Colanthelia is derived
Brazil.
(Tyrell et al. 2012). The Aulonemia/Colanthelia
Molecular data are unavailable for Athroosta-
clade is strongly supported.
chys. Tyrell et al. (2012) speculate that it may be
A few species of “Aulonemia”, including the
closely related to Alvimia and Atractantha.
type, produce several smaller lateral branches in
addition to the dominant one.
62. Atractantha McClure
Atractantha McClure, Smithsonian Contr. Bot. 9: 42 64. Colanthelia McClure & E. W. Sm.
(1973); tax.: Judziewicz (1992); Santos-Gonçalves et al.
(2011). Colanthelia McClure & E. W. Sm., Smithsonian Contr.
Bot. 9: 77 (1973).
Culms scandent, vine-like, or pendent. Branches
three per node, rebranching, with one branch Culms erect or scandent. Branches several per
dominant, from a single bud on a promontory. node, with one branch dominant, divergent,
Culm leaves deciduous, with a pseudopetiole, the from a single bud. Culm leaves often with a dark
sheaths with an obvious skirt of downward point- girdle between the node and the base of the sheath,
ing fimbriae, with a bumpy, tumor-like, corky with a corky crest at the junction of the girdle and
crest at the base of the sheath, the blades gener- the base of the sheath. Foliage leaves with sheaths
ally horizontal to reflexed. Inflorescence sometimes auriculate, the blades deciduous. Inflo-
unbranched, or branched and capitate. Pseudos- rescence unbranched or with branched branches.
pikelets with 1 glume, 1 prophyll, and 1 flower, Spikelets slender, with few to many flowers.
the flower slender, indurate, sharp-pointed, nee- Glumes 2 or 3. Stamens 3. Stigmas 2.
dle-like. Seven spp., southern Brazil and northern
Six spp., coastal Brazil, Amazonas, and south- Argentina.
ern Venezuela. See comments under “Aulonemia”.
Atractantha is strongly supported by molec-
ular data as sister to Alvimia (Tyrell et al. 2012). 65. Didymogonyx (L.G. Clark & Londoño) C.D. Tyr-
The culm leaf blades are small and erect in A. rell, L.G. Clark & Londoño
radiata McClure, an exception to the general state
for Atractantha. Didymogonyx (L.G. Clark & Londoño) C.D. Tyrrell, L.G.
Clark & Londoño, Mol. Phylog. Evol. 65: 146 (2012).
Rhipidocladum sect. Didymogonyx L. G. Clark & Lon- Culms scandent at maturity, often drooping from
doño, Amer. J. Bot. 78: 1271 (1991). trees. Mid-culm branches in fan-shaped clusters,
from a single bud. Culm leaves with sheaths bear-
Culms erect below, arching or scrambling above, ing fimbriae. Inflorescences unbranched, one-
with alternating long and short internodes, the sided, the spikelets crowded, sessile or on short
long ones hollow, the short ones solid or pithy. pedicels. Spikelets with 1 flower, with a rachilla
Prophylls cordate. Culm leaves with sheaths with
auricles and fimbriae. Foliage leaves with sheath
fimbriate, the fimbriae with bases slightly flat-
tened, auricles lacking. Inflorescence contracted,
branched or unbranched. Spikelets in clusters of
2 or 3. Glumes 3 (5), sterile lemmas 1 or 2,
flowers 2 to 12. Mesophyll with sclerenchyma
associated with the bulliform cells. Leaf epider-
mal long cells with papillae overarching the sto-
mata.
Two spp., Colombia and Venezuela.
66. Elytrostachys McClure
Elytrostachys McClure, J. Wash. Acad. Sci. 32: 173 (1942).
Culms scandent, often drooping from trees.

Branches several per node, with one branch dom-
inant, the branches from a single bud. Culm
leaves with sheaths with straight fimbriae, the
fimbriae 2–8 cm long. Foliage leaf sheaths with
or without auricles. Inflorescences on lateral
shoots, dense and capitate. Pseudospikelets each
subtended by a prophyll; each branch with 1 or
2 flowers subtended by 2 glumes. Stamens 6.
Ovary apex pubescent.
Two spp., Central and South America.
67. Filgueirasia Guala
Filgueirasia Guala, Bamboo Sci. Cult. 17: 2 (2003).
Culms erect, <2 m tall. Mid-culm nodes produc-

ing 1 to 15 buds in a line above the node. Culm
leaves with sheaths glabrous on both surfaces, the
blades erect. Foliage leaves with white fimbriae,
the blades erect. Inflorescence branches one- Fig. 22. Bambusoideae-Bambuseae-Arthrostylidiinae.
sided. Spikelets with 3 to 15 flowers. Fruit beaked. Merostachys sp. (A, L) and M. speciosa (B–K). A Seedling
plant, showing basal caespitose clump habit and pachy-
Two spp., Brazilian cerrado. morph rhizomes, with young culm shoot at left. B Culm
sheath from mid-culm node. C Flowering branch, termi-
nating in secund spicate inflorescence. D Leaf sheath
68. Merostachys Spreng. Fig. 22 showing ligule, base of oral setae, petiole, and base of
leaf blade. E A single spikelet of the inflorescence. F
Merostachys Spreng., Syst. Veg. 1: 132, 249 (1824); tax., Palea. G Bristle-like prolongation of rachilla in two
Sendulsky and Soderstrom (1984). aspects. H Lemma. I Stamen. J Gynoecium. K Lodicule
complement. L Seed after having produced a seedling
plant. (From McClure 1973)
extension. Glumes 2, shorter than the flower. triangular, deciduous, the sheaths lacking auri-
Lemma clasping the palea at the base only. cles or fimbriae, the blade confluent with the
Forty-eight spp., Central and South America. sheath. Inflorescence unbranched, the spikelets
Merostachys is strongly supported by molec- often borne on one side. Spikelets with several
ular data as sister to Actinocladum (Tyrell et al. flowers. Glumes 2 or 3 (to 5). Lemma apex obtuse
2012). Both genera have caryopses in which the to acute, awned or not.
seed coat is free from the pericarp and fan-shaped Fifteen spp., Central to South America.
clusters of branches. The latter character also
appears independently in Rhipidocladum and
some species of Arthrostylidium. Species of Mer- V.3. SUBTRIBE GUADUINAE Soderstr. & R. P. Ellis (1987)
ostachys occasionally have two to several flowers
per spikelet. Culm leaves often with the blade margins conflu-
ent with the sheath margins. Culm leaf sheath and
69. Myriocladus Swallen foliage leaf sheath generally with fimbriae. Both
adaxial and abaxial surfaces of foliage leaf blades
Myriocladus Swallen, Fieldiana, Bot. 28: 34 (1951); rev.: with stomata, the adaxial stomata with overarch-
Judziewicz (1998). ing papillae. Lodicules 3. 2n ¼ 46 or 48.
Five genera, 44 species.
Culms erect, the internodes often notably unequal Species with elongated rhizome necks are
in length. Branches 1 per node, but appearing common but not universal in this subtribe; they
clustered as internodes fail to elongate. Prophylls occur in Eremocaulon, Guadua, Olmeca and Ota-
cordate. Culm leaves not differentiated from tea.
foliage leaves. Leaves densely overlapping, with
the sheaths thickened distally, the blades erect to
spreading, deciduous. Inflorescence terminal, 71. Apoclada McClure
unbranched, or the branches branched. Spikelets
Apoclada McClure, Fl. Il. Catarin. 1 (Gram.-Supl.): 57
with 1 to 10 flowers. Glumes generally 2, with or (1967); rev., Guala II (1995).
without awns.
Twelve spp., Guayana Highlands of Vene- Culms erect, clump forming. Mid-culm nodes
zuela, Brazil and Guyana. with 2 to 5 buds. Branches as many as the number
The pattern of internode elongation in Myr- of buds, sometimes rebranching. Culm leaves
iocladus varies. In two species, internodes are with sheaths deciduous, without auricles, with
approximately equal in length. In other species fimbriae, with blades erect. Foliage leaves with
the basal internode is long whereas the upper fimbriae. Inflorescence with few branches. Spike-
ones are scarcely elongate at all, and in still others lets with 7 to 28 flowers. Stamens 3. Stigmas 2.
long internodes are followed by short ones in an Ovary with an enlarged apex, glabrous.
alternating pattern. Molecular data are not avail- One sp., A. simplex McClure & L.B. Smith,
able for Myriocladus, so its placement is uncer- southeastern Brazil.
tain. Tyrell et al. (2012) speculate that it may
belong in a clade with Glaziophyton and Camba-
juva. The cordate prophylls are shared with Didy- 72. Eremocaulon Soderstr. & Londoño
mogonyx, but may not be synapomorphic.
Eremocaulon Soderstr. & Londoño, Amer. J. Bot. 74: 37
(1987); rev., Londoño and Clark (2002).
70. “Rhipidocladum” McClure Criciuma Soderstr. & Londoño, Amer. J. Bot. 74: 35
(1987).
“Rhipidocladum” McClure, Smithsonian Contr. Bot. 9:
101 (1973); tax., Clark and Londoño (1991). Plants strongly rhizomatous, producing culms at
1 to 2 m intervals. Branches more than 1 per
Culms weak, erect below, scandent or clambering node, the central one dominant. Culm leaves
above. Mid-culm branches numerous, in a fan- leathery, with obvious auricles, with fimbriae,
shaped cluster, from a single bud. Culm leaves the blade becoming reflexed. Foliage leaves with
fimbriae, with or without auricles, the midrib

strongly excentric. Inflorescence sparsely
branched. Pseudospikelets elongate, with 1 to 13
(17) flowers, with 0 to 3 proximal bracts bearing
buds in their axils, and 0 to 3 additional bracts
(glumes) lacking buds. Stamens 6. Stigmas 2 or 3.
Four spp., Bahia, Brazil.
73. Guadua Kunth Fig. 23
Guadua Kunth, J. Phys. Chim. Hist. Nat. Arts 95: 150

(1822).
Culms arborescent to scandent, usually with a

band of white hairs below the nodes, and often
above as well. Branches one per node, the second-
ary branches generally modified into thorns. Culm
leaves deciduous, with sheaths contiguous with
the blades, fimbriae sometimes absent. Inflores-
cences of several pseudospikelets, each with 1
bract and 1 prophyll, 1 to several bracts with
buds in their axils, zero to several glumes, and
several flowers. Palea keels winged. Stamens 6.
Stigmas 3. Fruit dry. 2n ¼ 46, 48.
Twenty-seven spp., Mexico, Central and
South America.
Although most of the members of this genus
have dry fruits, that of G. sarcocarpa Londoño is
fleshy. Fig. 23. Bambusoideae-Bambuseae-Guaduinae. Guadua
ciliata. A Portion of stem showing culm leaf with erect
blades. B Branches bearing pseudospikelets. C Apex of
74. Olmeca Soderstr. sheath of foliage leaf, with pseudopetiole and short ligule.
D Culm leaf with triangular blade, blade margins contin-
Olmeca Soderstr., Phytologia 51: 161 (1982); Ruiz-Sán- uous with the sheath. E Node with a single bud, with white
chez et al. (2011a), rev., phylog. hairs above and below the node, the remains of the leaf
sheath forming a rim. (From Davidse et al. 2004, with
permission from the Missouri Botanical Garden Press;
Culms solitary, the rhizome necks elongate. drawn by Bruno Manara)
Nodes with one branch, the branch not itself
branching; branches at mid-culm with a diameter
between 0.5 and 1.0 of that of the culm. Culm designate a type. The name was subsequently
leaves with sheaths persistent, indurate, the validated in 1982 as noted above.
blades erect or reflexed. Spikelets with 2 glumes
and several flowers. Lemma glabrous or with long
75. Otatea (McClure & E. W. Smith) C. Calderón &
hairs at the apex. Lemma and palea fused to each Soderstr.
other at the base. Stamens 3. Stigmas 2 or 3. Fruit
fleshy or dry. Otatea (McClure & E. W. Smith) C. Calderón & Soderstr.;
Five spp., southern Mexico, Honduras. Ruiz-Sánchez et al. (2008, 2011b), rev., phylog.
The rhizome necks are 3–9 m long in one
clade (2 species), but 4–30 cm in the other (3 Culms erect. Mid-culm nodes with one branch
species) (Ruiz-Sánchez et al. 2011a). Soderstrom bud, producing three principal ascending
provided a description and diagnosis of Olmeca branches, diameter of the branches at mid-culm
in 1981 (Amer. J. Bot. 68: 1362), but failed to less than 1/4 that of the culm. Culm and foliage
leaves lacking auricles, with fimbriae (oral setae). chloa and several smaller segregates. Intergeneric
Inflorescence branches branched. Spikelets with hybridization had already been suggested by Goh
multiple flowers. Stamens 3. Stigmas 2 or 3. et al. (2010), and documented by Goh et al.
Eight spp., Mexico, Central America and (2011). A small population of hybrids between
Colombia. Dendrocalamus pendulus Ridl. and Gigantochloa
scortechinii Gamble was found growing near both
PALEOTROPICAL CLADE parental species, in an area that had been dis-
turbed by clearing associated with highway con-
Data from the chloroplast genome strongly sup- struction (Wong and Low 2011). The hybrid
port this clade as monophyletic, but there is no plants flowered but were apparently sterile and
obvious morphological synapomorphy other there was no evidence of F2 plants.
than the biogeography (Bamboo Phylogeny
Group 2012; Kelchner and Bamboo Phylogeny
76. Bonia Balansa
Group 2013). All species may be hexaploid or
higher. The polyploid history of the group is Bonia Balansa, J. Bot. (Morot) 4: 29 (1890); tax.: Xia
complex, involving ancestors that were closely (1996).
related to the ancestor of Arundinarieae (Triplett Monocladus L.C. Chia, H.L. Fung & Y.L. Yang, Acta Phy-
et al. 2014). Melocanninae are clearly sister to a totax. Sin. 26(3): 212 (1988).
clade of the other three subtribes.
Culms scrambling. Branches 1 per node, about
the same thickness as the culm. Culm leaves with
sheaths persistent, with prominent auricles, the
V.4. SUBTRIBE BAMBUSINAE J. Presl (1830)
auricles bearing fimbriae or not, the blade erect
Dendrocalaminae Benth. (1881). or reflexed. Foliage leaves with or without falcate
auricles, with fimbriae. Inflorescence
Culm nodes not swollen. Flowers arranged in unbranched, subtended by a bract and prophyll;
conventional spikelets or pseudospikelets, these distal to the prophyll are 2 or 3 bracts with buds
clustered, with each cluster subtended by bracts. in their axils, and distal to these a 1 cm rachilla
Stamens 6. Style 1 or reduced. Stigmas 3. internode. Pseudospikelets with 3 to 9 flowers;
2n ¼ (48) 70, 72. distal flowers under-developed. Lodicules 3.
Twenty-five genera, 264 spp. Ovary glabrous.
This subtribe has no obvious morphological Five spp., southern China, Vietnam.
synapomorphy. The apex of the ovary of Bambu- The bracts in this genus are sometimes called
sinae often has a different texture and color from “glumes”, but have buds in their axils and are
the body of the ovary itself. The apical tissue is thus gemmiparous bracts.
variously called an umbo, an appendage, or sim-
ply a fleshy outgrowth. The only detailed descrip- 77. Cyrtochloa S. Dransf.
tion of the structure is presented by Holttum
(1956), who shows sketches of sections of ovaries Cyrtochloa S. Dransf., Kew Bull. 53: 861 (1998).
and fruits of a few genera. He describes the out-
growth at the apex of the ovary as similar to a Culms erect or scandent. Branches 3 to many per
style base in Bambusa and Gigantochloa, whereas node, with 1 branch dominant. Culm leaves with
the structure is hollow in Oxytenanthera. I do not sheaths persistent, wrinkled at the base, with
know of any investigation of its development, auricles, the blades ovate or triangular and cor-
anatomy, or function. date. Foliage leaves with auricles absent or falcate,
Within Bambuseae, generic limits are most fimbriae sparse; the blades with strongly asym-
problematical in Bambusinae. The most compre- metric bases. Inflorescence with capitate clusters
hensive phylogeny to date is that of Goh et al. of spikelets at the nodes, the clusters separated by
(2013), which finds evidence for monophyly of a internodes. Pseudospikelets with 1 fertile flower.
group of climbing species and a complex history Glumes 2 to several, the apex acute to acuminate,
of apparent hybridization and/or lineage sorting apex of the upper one sometimes mucronate.
in the genera Bambusa, Dendrocalamus, Giganto- Lemma wrapped around the palea. Lodicules
2 or 3. Ovary with a long appendage, the apex cles, with or without fimbriae, with blades
pubescent. Caryopsis with the pericarp fleshy, or reflexed. Foliage leaf sheaths with or without
free from the seed coat and breakable. auricles, with or without fimbriae. Inflorescence
Five spp., Tropical Asia. unbranched, bearing small clusters of pseudospi-
kelets at the nodes. Pseudospikelets with 1 or
78. Fimbribambusa Widjaja 2 flowers, the proximal one sterile or fertile, the
distal one fertile. Glumes 2, the glumes and
Fimbribambusa Widjaja, Reinwardtia 11: 80 (1997). lemmas orbicular to oblate. Lodicules 3. Stigmas
sometimes 2. Ovary with or without a rounded
Culms scrambling. Nodes with a narrow shelf-like projection at the apex, glabrous. Fruit spherical,
extension. Branches 1 per node, or 5 to 10 per the pericarp fleshy.
node with 1 dominant. Culm leaf sheaths with Five spp., Asia.
stiff linear, horn-like auricles, with fimbriae, the
blades spreading to reflexed. Foliage leaf sheaths 81. Not “Nastus” Juss.
with auricles, with fimbriae. Inflorescences borne
on leafless branches, the pseudospikelets in clus- Not “Nastus” Juss., Gen Pl. 34 (1789).
ters of few to many at the nodes, the clusters
subtended by 2 prophylls. Pseudospikelets with Culms erect or scandent. Lateral branches
2 or 3 flowers. Lodicules 2 or 3. branched, 3 to many, subequal or one or two
Two spp., East Java, New Guinea, dominant, generally arising at the node, but in
Fimbribambusa was segregated from Bam- one species just below the node. Culm leaves with
busa based on the shelf-like extension at the sheaths deciduous or persistent, covered with
nodes, the lodicules being entire, and the ovary black hairs, with auricles and fimbriae. Foliage
glabrous. Fimbribambusa is similar to Tembur- leaves with or without auricles, with or without
ongia but the latter has only 2 or 3 spikelets per fimbriae. Inflorescence unbranched or the
node. branches branched. Spikelets conventional, with
1 fertile flower; rachilla extension present.
79. Greslania Balansa Glumes 5 or 6. Lodicules 3. Ovary with or without
an enlarged apex, the apex pubescent or not.
Greslania Balansa., Bull. Soc. Bot. France 19: 319 (1873). Two spp., Asia, Pacific.
Nastus s.s. belongs in Hickeliinae and is
Culms erect. Branches 1 per node or absent. Culm native to Madagascar and Réunion. The southeast
leaves similar to foliage leaves, the sheaths lack- Asian species fall here and are unrelated to the
ing auricles, fimbriae present or absent. Inflores- rest of the genus. The blades of N. schlechteri
cence branches extensively branched, with (Pilger) Holttum have obvious papillae on the
subtending bracts. Spikelets with 1 flower plus a adaxial surface, visible under low magnification.
rachilla extension. Glumes 2. Lodicules 3. Ovary
glabrous. Fruit with pericarp easily removed from
82. Neomicrocalamus Keng f.
the seed coat.
Four spp., New Caledonia. Neomicrocalamus Keng f., J. Bamboo Res. 2: 10 (1983);
Notes on the morphology and phenology of tax.: Stapleton (1994b).
two of the species are provided by Dransfield
(2002). Culms scandent. Branches many per node, the
central one dominant. Culm leaves with sheaths
80. Melocalamus Benth. deciduous, leaving a rim, auricles absent, the
blade a tiny acicular extension of the sheath.
Melocalamus Benth., Gen. Pl. 3: 1212 (1883). Foliage leaves small, without auricles or fimbriae,
the apex acicular. Inflorescence with primary
Culms scandent. Branches many per node, with branches only, the spikelets sessile, subtended
one branch dominant. Culm leaves with sheaths by bracts and prophylls, but these not bearing
persistent, with a wrinkled basal zone, with auri- axillary buds. Spikelets conventional, with several
flowers, the distal ones reduced. Glumes 1 or 85. Parabambusa Widjaja

lacking. Lodicules 3. Ovary apex pubescent.
Five spp., southern Asia. Parabambusa Widjaja, Reinwardtia 11: 121 (1997).
Neomicrocalamus s.l. is sister to Bonia in the Pinga Widjaja, Reinwardtia 11: 123–124 (1997).
phylogeny of Yang et al. (2008).
Culms scandent, sometimes producing aerial
roots at the nodes. Branches 2 to 5, with 1 branch
83. Oreobambos K. Schum. Fig. 24 dominant, the dominant branch branching. Culm
leaves with sheaths deciduous, leaving a persis-
Oreobambos K. Schum., Notizbl. K€onigl. Bot. Gart. Berlin
1: 178 (1896). tent rim, with auricles and fimbriae, the blade
erect. Foliage leaves with sheaths with erect auri-
Culms scandent or drooping. Branches 2 or 3 per cles and fimbriae, the blades deciduous. Inflores-
node, more or less in a line, one branch domi- cence a cluster of up to 10 pseudospikelets, the
nant. Culm leaves without auricles, with fimbriae, cluster subtended by a bract and a prophyll.
the blade acicular, short. Foliage leaves without Pseudospikelets with 5 to 12 flowers. Glumes 2.
fimbriae, the blades without prominent cross Lemma apex acuminate. Lodicules absent. Ovary
veins. Inflorescence a fascicle of pseudospikelets, glabrous.
Two spp., New Guinea.
surrounded by bracts. Pseudospikelets with
2 flowers. Glume 1, with an obtuse apex. Lemma
ovate, the margins convolute, the apex obtuse. 86. Pseudobambusa T. Q. Nguyen
Lodicules absent. Stamens 6. Stigmas 1. Ovary
Pseudobambusa T. Q. Nguyen, Bot. Žurn. (Moscow &
with a rounded projection at the apex, pubescent
Leningrad) 76: 992 (1991).
at the apex. Pericarp free from the seed coat, soft.
One sp., O. buchwaldii K. Schum., Tanzania, Culms erect. Branches 1 to 4 per node, subequal.
Uganda, Zimbabwe, Malawi, Zambia. Culm leaves densely hairy, the sheaths with auri-
cles and dense fimbriae. Foliage leaves with
84. Oxytenanthera Munro blades with 10 to 12 secondary veins. Inflores-
cence a cluster of pseudospikelets, subtended by
Oxytenanthera Munro, Trans. Linn. Soc. London 26: 126 bracts. Pseudospikelets with 3 or 4 flowers.
(1868).
Glumes several, the apex of the distal one acute
and mucronate. Lemma apex acute and mucro-
Culms erect. Branches many per node, in an nate. Palea longer than the lemma. Lodicules 0 or
irregular line, one branch dominant. Culm leaves 3. Ovary pubescent.
without auricles, the blade erect on lower leaves, One sp., P. schizostachyoides (Kurz) T. Q.
becoming reflexed on more apical ones. Foliage Nguyen, southern Burma and Vietnam.
leaves with blades deciduous, the blades with
faint cross-veins, with fimbriae. Inflorescence a
capitate cluster of pseudospikelets at the ends of 87. Pseudoxytenanthera Soderstr. & R. P. Ellis
lateral branches. Pseudospikelets with 2 to 5 flow-
Pseudoxytenanthera Soderstr. & R. P. Ellis, Smithsonian
ers, the lower 1 to 3 sterile. Glumes 2, hispidulous. Contr. Bot. 72: 52 (1988).
Lemmas awned. Lodicules absent. Stamens with
filaments fused at the base. Ovary with a long Culms erect below, scandent above. Branches
hollow apical extension, glabrous. many per node, the branches more or less equal
One sp., O. abyssinica (A. Rich.) Munro, or one branch dominant. Culm leaves with
Africa. sheaths deciduous, leaving a persistent rim or
Molecular data (Yang et al. 2010) place this girdle, with auricles, the blade horizontal or
genus within Dendrocalamus, but without sup- reflexed. Foliage leaves with auricles, with fim-
port. The ovary extension is sketched by Holttum briae. Inflorescence unbranched, with clusters of
(1956); it is hollow, with the hollow portion not pseudospikelets at the nodes. Pseudospikelets
contiguous with the ovary locule.
Fig. 24. Bambusoideae-Bambuseae-Bambusinae. Oreo- H Spikelet cluster, lateral view. I Spikelet cluster partly
bambos buchwaldii. A Branch with foliage leaves. B dissected. J Two spikelets after removal of bracts. K, L
Foliage leaves. C Detail of abaxial surface of leaf blade Bracts. M Upper glume. N Lemma. O Palea. P Gynoecium
showing tessellate venation. D Flowering branches and androecium. Q Ovary. R Portions of anther. S Termi-
showing clusters of spikelets. E Spikelet cluster, viewed nal flower showing rachilla extension. (From Clayton
from the adaxial side. F Bract removed from spikelet 1970)
cluster. G Spikelet cluster, viewed from the abaxial side.
with 1 to 3 flowers, the distal ones reduced. 90. Vietnamosasa T. Q. Nguyen

Glumes 2 to several, the upper glume mucronate.
Lemma muticous or mucronate. Lodicules absent. Vietnamosasa T. Q. Nguyen, Bot. Žurn. (Moscow & Lenin-
Stamens with filaments fused. Stigmas sometimes grad) 75: 221 (1990); tax.: Haevermans et al. (2013); Xia
(1996).
1 or 2. Ovary with or without a swollen apex,
glabrous, or pubescent on the apex, or pubescent
Culms generally less than 2 m tall. Branches many,
throughout. Pericarp sometimes free from the
dendroid. Culm leaves with auricles, the blade
seed coat.
erect. Foliage leaves with the blade persistent or
Twelve spp., southern India, Sri Lanka.
deciduous at the ligule. Inflorescence branches
The number of species in Pseudoxyte-
branched. Spikelets with 5 to 9 flowers plus addi-
nanthera is based on Bamboo Phylogeny Group
tional reduced flowers distally. Glumes 2 to several.
(2012).
Lodicules 3. Ovary apex without an appendage.
Three spp., Vietnam.
88. Temburongia S. Dransf. & K. M. Wong Vietnamosasa is placed in Bambusinae by the
phylogenetic analyses of Sungkaew et al. (2009).
Temburongia S. Dransf. & K. M. Wong, Sandakania 7: 55 Although Watson and Dallwitz (1992 onward)
(1996).
describe the rhizomes as leptomorph, McClure
(1966) shows them clearly as pachymorph, con-
Culms scandent. Nodes with a narrow shelf-like
sistent with the placement of the genus here in
extension. Branches several, clumped, with one
Bambusinae.
branch dominant, from a single bud. Culm leaves
with small rounded auricles, without fimbriae,
BAMBUSINAE: HK CLADE sensu Goh et al. (2013).
the blade erect. Foliage leaves with auricles. Inflo-
HOLTTUMOCHLOA + KINABALUCHLOA
rescence with clusters of 1 to 3 conventional spi-
kelets, bracteate, without buds in the axils of the
bracts. Spikelets with 3 flowers, the proximal and 91. Holttumochloa K. M. Wong
distal ones sterile, the middle one fertile. Glumes
2. Lodicules 3. Apex of ovary with an umbo, Holttumochloa K. M. Wong, Kew Bull. 48: 518 (1993).
glabrous.
One sp., T. simplex S. Dransf. & K. M. Wong, Culms climbing or scrambling. Lateral branches
Brunei. many per node, more or less equal, unbranched,
Molecular data (Goh et al. 2013) place this from multiple buds. Culm leaves with auricles
species in Bambusinae but not closely related to and fimbriae, with blades reflexed. Foliage leaves
other sampled genera. with or without auricles. Pseudospikelets in a
linear inflorescence or clustered at the nodes,
each subtended by 1 to 3 sterile bracts, 1 or
89. Temochloa S. Dransf. 2 bracts with buds in their axils, and 1 to 2 glumes,
with 2 to 5 flowers, the distal ones reduced. Palea
Temochloa S. Dransf., Thai Forest Bull., Bot. 28: 179
(2000). with 10 veins. Lodicules 3. Ovary apex pubescent,
with a rounded outgrowth.
Culms erect below, scrambling above, ca. 2 m tall; Three spp., peninsular Malaysia.
internodes slender. Branches 2 to 4 from a single Holttumochloa and Kinabaluchloa are sisters
bud, subequal, with secondary branches. Culm in the molecular phylogeny of Goh et al. (2010).
leaves with deciduous sheaths. Foliage leaves
lacking auricles or fimbriae. Inflorescence 92. Kinabaluchloa K. M. Wong
unbranched, with 1 to 4 spikelets. Spikelets con-
ventional, with 1 fertile proximal flower and 1 Kinabaluchloa K. M. Wong, Kew Bull. 48: 523 (1993).
sterile distal flower. Glumes 2. Ovary glabrous.
One sp., T. liliana S. Dransf., Thailand. Culms leaning or scrambling, with internodes up
to 1.5 m long. Nodes ringed with spreading bris-
tles. Branches many, more or less equal, from a
single bud. Culm leaves with auricles. Foliage
leaves with or without auricles, the ligule mem- 94. Mullerochloa K. M. Wong
branous or a fringed membrane. Inflorescence a
cluster of pseudospikelets, subtended by a large Mullerochloa K. M. Wong, Blumea 50: 434–435, f. 1–9
bract. Individual pseudospikelets subtended by a (2005).
bract as long as the lemma, followed by 1 or
2 bracts with buds in their axils, and 1 glume; Culms with the base of the internodes swollen.
flowers 1 to 2. Palea with 10–16 veins, the apex Branches 1 per node, or 1 dominant and 2 second-
dentate. Lodicules 3. Ovary with or without a ary branches, from a single bud. Culm leaves with
rounded projection at the tip, sometimes sparsely sheaths with auricles inconspicuous, the blades
pubescent. spreading to reflexed. Pseudospikelets in clusters
Two spp., Malay peninsula and Borneo. at the nodes, bracteate, with 4 to 9 flowers. Palea
longer than the lemma, densely pubescent adaxi-
BAMBUSINAE: DMNS CLADE sensu Goh et al. (2013). ally. Lodicules 3. Stamens 4, filaments fused.
DINOCHLOA + NEOLOLEBA + MULLEROCHLOA + SPHAER- Ovary glabrous, the apex not thickened.
OBAMBOS
One sp., M. moreheadiana (F. M. Bailey) K.
M. Wong, North Queensland, Australia.
Culms climbing or scrambling. Culm leaf sheath with a Sphaerobambos also has paleas longer than
characteristic wrinkled zone at the base.
the lemmas, but unlike Mullerochloa, the palea
Chloroplast and nuclear DNA sequence data
keels are winged. The densely pubescent adaxial
support the close relationship of the following
side of the palea is also seen in Soejatmia.
four genera.
95. Neololeba Widjaja

93. Dinochloa B€
use
Neololeba Widjaja, Reinwardtia 11: 112 (1997).
Dinochloa B€
use, Pl. Jungh. 387 (1854), rev.: Dransfield
(1981).
Culms branching only above several long proxi-
mal internodes. Branches 2 to 6, very steeply
Culms spiraling around tree trunks. Branches 3 to ascending, with one dominant, the dominant
18 per node, from a single bud, more or less equal one branching again, the secondary branches
in size. Culm leaves with sheaths persistent or unbranched. Culm leaves with light brown hairs,
deciduous, with a dark girdle between the node with sheaths persistent or deciduous, auricles
and the base of the sheath, with or without auri- present, fimbriae present or absent, the blade
cles, the auricles bearing fimbriae, the blade con- erect. Foliage leaves with or without auricles.
stricted at the base or not, erect or deflexed. Inflorescence a cluster of pseudospikelets, most
Foliage leaves with or without auricles, with fim- sessile but one pedicellate. Pseudospikelets with 3
briae. Inflorescence elongate, up to 3 m long, the to 12 flowers, the distal flowers reduced in one
branches branched, each branch subtended by a species. Glumes 2 or 3, the apices long acuminate,
sheath and a prophyll, the pseudospikelets in mucronate. Lemma acuminate. Lodicules absent.
clusters separated by long internodes. Pseudospi- Ovary with a rounded projection at the apex,
kelets with 1 fertile flower. Glumes 2 or 3. Lodi- pubescent at the apex or throughout.
cules 0 or 3. Stigmas sometimes 2. Ovary with or Five spp., tropical Asia, Australia, Pacific.
without an appendage, glabrous. Pericarp free
from the seed coat, fleshy.
Thirty-one spp., mostly tropical Asia. 96. Sphaerobambos S. Dransf.
The fruit of Dinochloa is unusual in that the
Sphaerobambos S. Dransf., Kew Bull. 44: 428 (1989).
scutellum enlarges at the expense of the endo-
sperm, and fills almost the entire fruit (Rudall
and Dransfield 1989). This characteristic also Culms with branches several to many per node,
appears in Ochlandra, Melocalamus and Melo- dendroid, with one branch dominant. Culm leaf
canna. The base of each culm internode is swollen sheaths with fimbriate auricles, the blade erect,
and the base of the sheath is wrinkled. This sometimes becoming reflexed. Foliage leaves with
appears to function as a pulvinus, allowing the fimbriate auricles. Inflorescence with clusters of a
culm to twine around trees. few pseudospikelets at the nodes. Pseudospikelets
with 3 to 5 flowers. Glumes 2 to several. Palea

with narrow winged keels, the palea longer than
the lemma at maturity. Lodicules absent. Ovary
with or without a solid swollen structure at the
apex, glabrous.
Three spp., Borneo, the Philippines, Sulawesi.
See comments under Mullerochloa.
BAMBUSINAE: BDG COMPLEX sensu Goh et al. (2010,

2013). ALL REMAINING GENERA.
Nuclear and chloroplast phylogenies are
strongly incongruent for this group (Goh et al.
2013), suggesting either incomplete lineage sort-
ing or hybridization. This could justify combin-
ing all taxa into a single large genus.
97. Bambusa Schreb. Fig. 25
Bambusa Schreb., Gen. Pl. 1: 236 (1789), nom. conserv.;

phylog.: Goh et al. (2010).
Ischurochloa B€use, Pl. Jungh.: 389 (1854).
Leleba Rumph., Catal. Plantarum in Horto Botanico
Bogoriensi 20 (1866).
Tetragonocalamus Nakai, J. Jap. Bot. 9: 86, 88–89 (1933).
Lingnania McClure, Lingnan Univ. Sci. Bull. 9: 34 (1940).
Dendrocalamopsis Q.H. Dai & X.L. Tao, Acta Phytotax.
Sin. 20: 210 (1982).
Neosinocalamus Keng f., J. Bamboo Res. 2: 12 (1983).
Phuphanochloa Sungkaew & Teerawat., Kew Bull. 63: Fig. 25. Bambusoideae-Bambuseae-Bambusinae. Bam-
669–671, f. 1 (2008). busa chungii. A Portion of culm base and rhizome. B
Portion of culm, showing branching pattern. C Portion of
young culm, showing culm sheath. D Abaxial view of apex
Culms erect or scrambling. Branches many per of culm sheath. E Same, adaxial view. F Portion of inflores-
node, sometimes with 1 to 3 dominant. Culm cence. G Floret. H Lemma. I Palea. J Lodicules. K Stamens.
leaves with sheaths deciduous, with or without L Pistil. (From Wu et al. 2007, p. 23, with permission from
auricles, with marginal fimbriae, the blades the Missouri Botanical Garden Press, St. Louis, and Science
Press, Beijing; drawn by Huang Yingqin)
erect. Foliage leaves without auricles, with or
without fimbriae. Inflorescence with clusters of
pseudospikelets at the nodes, with a 2-keeled 98. Dendrocalamus Nees
prophyll. Pseudospikelets with 2 to many flowers,
with a sterile or imperfect terminal flower; proxi- Dendrocalamus Nees, Linnaea 9: 476 (1835); tax.: Staple-
ton (1994a).
mal 1 to several flowers sterile. Rachilla nodes Klemachloa R. Parker, Indian Forester 58: 7 (1932).
disarticulating. Lodicules 2 or 3. Ovary borne on Sinocalamus s.s., McClure, Lingnan Univ. Sci. Bull. 9: 66
a stalk, sometimes with an enlarged apical por- (1940).
tion, the apex pubescent. Neosinocalamus Keng f., J. Bamboo Res. 2: 12 (1983).
One hundred species, tropical and subtropi-
cal Asia. Culms arborescent, caespitose, apex of the stem
The inclusion of Dendrocalamopsis in Bam- generally pendulous. Branches several to many at
busa is supported by the molecular phylogeny of each node, none to 3 dominant. Culm leaves with
Yang et al. (2010). Although they could not rule sheaths deciduous, auricles absent or reduced,
out monophyly for Bambusa s.s., Goh et al. (2010) fimbriae present or absent, the blade reflexed or
find no support for the existing subgeneric clas- erect. Foliage leaves with or without short auri-
sification. cles, without fimbriae, sometimes lacking a pseu-
dopetiole. Inflorescence generally unbranched,
bearing clusters of pseudospikelets at the nodes. from a single bud. Culm leaves with sheaths bear-
Pseudospikelets with 1 to 2 fertile flowers, sessile. ing low rim-like auricles, with blades lanceolate.
Rachilla nodes not disarticulating. Lemma with a Inflorescence a cluster of pseudospikelets. Pseu-
long mucro. Lodicules 0 to 3. Ovary on a stalk, the dospikelets with 1 to 2 flowers. Glumes 3 to 5, the
apex thick, pubescent. 2n ¼ 70 + 2. distalmost 2 as large as the lemmas. Lodicules 3.
Forty-one species, Asia. Ovary with a rounded projection at the apex.
In the phylogenies of H.-Q.Yang et al. (2008) One sp., M. montana (Ridl.) K. M. Wong,
and J.-B. Yang et al. (2010), species of Dendroca- Malay Peninsula.
lamus form a clade within which members of
Gigantochloa and Oxytenanthera are embedded.
101. Soejatmia K. M. Wong
Unifying the three genera was suggested by
Soderstrom and Ellis (1987), although Yang Soejatmia K. M. Wong, Kew Bull. 48: 530 (1993).
et al. (2010) argue that this conclusion is still
premature. Yang et al. (2010) show that the type Culms climbing or scrambling. Branches 3 per
of Sinocalamus should be included in Dendroca- node. Culm leaves with sheaths with a wrinkled
lamus, but that other species assigned to Sinoca- zone at the base, with auricles and fimbriae, the
lamus belong to Bambusa. The chromosome blade erect. Foliage leaves with fimbriae, with
number report is based on Li et al. (2001). large, papery, ovate to rounded auricles. Inflores-
cence with clusters of pseudospikelets at the
99. Gigantochloa Kurz ex Munro nodes. Pseudospikelets with 4 to 6 flowers.
Glumes several. Palea apex with two hooked pro-
Gigantochloa Kurz ex Munro, Trans. Linn. Soc. London
26: 123 (1868).
jections. Lodicules 3. Apex of ovary thickened,
pubescent.
Culms erect, densely caespitose. Internodes often One sp., S. ridleyi (Gamble) Wong, Malay
with yellow stripes. Branches several per node, 1 Peninsula and Singapore.
dominant. Culm leaves with deciduous, densely Clayton et al. (2006 onward) synonymize Soe-
hairy sheaths, auricles absent or reduced, the jatmia with Kinabaluchloa, but Goh et al. (2013)
blade recurved or erect. Foliage leaves without show that the two genera are unrelated with Soe-
auricles, with or without fimbriae. Inflorescence jatmia falling here in the large BDG complex.
with clusters of pseudospikelets at the nodes.
Pseudospikelets with 2 to 5 flowers, the terminal 102. Thyrsostachys Gamble
one reduced to a lemma. Empty bracts proximal
to the glumes. Rachilla nodes not disarticulating. Thyrsostachys Gamble, Indian Forester 20: 1 (1894).
Lodicules absent. Stamens with filaments fused.
Ovary on a stalk, the apex thick and hairy, stigma Culms erect, caespitose. Branches several to many
1. 2n ¼ 76. per node, with 1 dominant. Culm leaves with
Thirty spp., India and southeast Asia. persistent sheaths, without fimbriae, with incon-
Molecular data (Yang et al. 2010) place this spicuous auricles, the blade erect or recurved.
genus in a clade with Dendrocalamus. It is possi- Foliage leaves without auricles or fimbriae. Inflo-
ble that the two genera should be merged, but the rescence elongate with branched branches. Pseu-
phylogeny is not sufficiently well resolved to be dospikelets in clusters of 1 to 3 fertile ones and 1
convinced of this. or 2 sterile ones, the cluster subtended by a pro-
phyll. Each pseudospikelet with a prophyll and
100. Maclurochloa K. M. Wong 2 to 4 flowers. Paleas of lower flowers deeply cleft.
Glumes 2 to 4. Lodicules 0 to 3. Ovary on a stalk,
Maclurochloa K. M. Wong, Kew Bull. 48: 528 (1993). glabrous. 2n ¼ 76.
Two spp., Burma, Thailand, China.
Culms climbing or scrambling, with a thick
band of silvery white hairs below each node.
Branches several per node, with 1 dominant,
V.5. SUBTRIBE MELOCANNINAE Benth. (1881) Culms arching. Branches several per node, with 1
branch dominant, from a single bud. Culm leaves
Schizostachyidinae Soderstr. & R. P. Ellis in Soderstr. with sheaths deciduous, leaving a persistent gir-
(1987).
dle or rim, the blades horizontal or reflexed.
Culms hollow. Foliage leaf blades generally with a Foliage leaves with sheaths with auricles, with
narrow waxless stripe along the abaxial margin. fimbriae, the blade deciduous. Inflorescence a
Flowers borne in pseudospikelets, in clusters at dense cluster of pseudospikelets, pseudospikelets
the nodes, subtended by bracts, the spikelets gen- with 1 to 3 flowers, the distal ones reduced.
erally with 2 glumes. Stamens 6. Style elongate, Glumes several. Lodicules 3. Stigmas 3.
hollow. Stigmas generally 3. Ovary glabrous. Mid- One sp., D. attenuata (Thwaites) Soderstr. &
rib with complex vasculature. Leaf cross section R. P. Ellis, Sri Lanka.
asymmetrical around the midrib, with a set of
prominent bulliform cells on one side only, creat- 105. Melocanna Trin.
ing a concave outline on one side of the midrib
and convex on the other. 2n ¼ 72. Melocanna Trin., Neue Entdeck. Pflanzenk. 2: 43 (1820);
Nine genera, 83 species. tax.: Stapleton (1994a); flowering: Ramanayake and
Melocanninae are sister to all other paleotro- Weerawardene (2003).
pical woody bamboos in the analyses of Sung-
kaew et al. (2009). Triplett et al. (2014) found a Culms erect, arborescent. Rhizomes with elon-
similar result albeit with a more limited sample of gated necks. Branches several to many per node,
taxa. The asymmetric leaf cross section is more or less equal. Culm leaves with persistent
described by Soderstrom and Ellis (1987) as an sheaths, with 1 to 2 strong undulations toward the
S-shaped midrib, a term that describes the shape apex, without auricles or fimbriae, the blade erect,
when viewed in cross section. at least proximally. Foliage leaves without auri-
cles, with fimbriae, with or without obvious cross
103. Cephalostachyum Munro veins. Inflorescence elongate, the branches
branched, bearing clusters of pseudospikelets.
Cephalostachyum Munro, Trans. Linn. Soc. London 26: Pseudospikelets with 1 fertile flower plus several
138 (1868); tax.: Stapleton (1994a). sterile flowers distally. Glumes 2 to 4, with buds
in their axils. Paleas lacking keels. Lodicules 2.
Culms erect or scandent. Branches many per Stigmas 2 to 4. Pericarp fleshy, with an apical
node, subequal. Culm leaf sheaths with 1 to
beak.
2 strong undulations near the apex, deciduous,
Two spp., eastern India to Burma.
with auricles and fimbriae, the blade reflexed.
Foliage leaves with or without auricles and fim-
briae, the blades without distinct cross veins. 106. Neohouzeaua A. Camus
Inflorescence capitate, or an unbranched axis,
with capitate clusters of pseudospikelets. Pseu- Neohouzeaua A. Camus, Bull. Mus. Natl. Hist. Nat. 28: 100
(1922).
dospikelets with 1 fertile flower plus 1 or 2 distal
reduced flowers. Glumes 2 or 3, awned. Lodicules
Culms erect or scandent. Branches 1 to several
3. Stigmas 2 or 3. Ovary on a stalk. Pericarp free
per node, more or less equal or one dominant.
from the seed coat.
Culm leaves with persistent or deciduous
Twelve spp., southeast Asia.
sheaths, with auricles and fimbriae, the blade
Two Madagascan species formerly placed in
reflexed. Foliage leaves with or without auricles,
Cephalostachyum belong in Hickeliinae and will
with short fimbriae. Inflorescence elongate,
need a new generic name (L.G. Clark, pers. comm.).
branched. Pseudospikelets with 1 flower. Glumes
3 or 4 with buds in their axils. Palea with 2 awns.
104. Davidsea Soderstr. & R. P. Ellis Lodicules 0 or 2. Stamens with filaments fused.
Stigmas 3. Pericarp free from seed coat.
Davidsea Soderstr. & R. P. Ellis, Smithsonian Contr. Bot. Seven spp., temperate to tropical Asia.
72: 59 (1988).
107. Ochlandra Thwaites
Ochlandra Thwaites, Enum. Pl. Zeyl. 376 (1864).
Culms erect. Branches several per node, in a line,

with one dominant. Culm leaves with sheaths
persistent or deciduous, with auricles and fim-
briae, the blade reflexed. Foliage leaves with or
without auricles, without fimbriae, the blades
persistent or deciduous, the venation with or
without cross veins. Inflorescence unbranched,
with clusters of pseudospikelets at the nodes.
Pseudospikelets more or less terete, with several
sterile flowers proximally, and 1 fertile flower.
Lodicules 1 or 3. Stamens numerous. Stigmas 3
to 9. Caryopsis with a fleshy pericarp.
Nine spp., southern India and Sri Lanka.
108. Pseudostachyum Munro
Pseudostachyum Munro, Trans. Linn. Soc. London 26:

141, pl. 4 (1868); tax.: Stapleton (1994a).
Culms erect, drooping at the apex. Rhizomes with

elongated necks. Branches many per node, sub-
equal. Culm leaves with the sheath deciduous,
with small auricles, with fimbriae, the blade
erect. Foliage leaf blades without auricles or fim-
briae. Inflorescence branches branched, each
branch subtended by a bract. Pseudospikelets Fig. 26. Bambusoideae-Bambuseae-Melocanninae. Schi-
pedicellate, with 1 flower. Glume 1, mucronate zostachyum dumetorum. A Portion of culm and culm
sheath, showing reflexed leaf blade. B Branch with foliage
to long awned. Lodicules 3 to 5. Stigmas 2. Fruit leaves. C Flowering branch. D Prophyll. E Bract. F Apical
globose; pericarp free from seed coat, brittle. rachis joint. G Lemma. H Apex of lemma. I Palea. J Apex of
One sp., P. polymorphum Munro, Bhutan, palea. K Stamen. L Pistil. M, N S. hainanense. M Abaxial
China, NE India, Myanmar, Vietnam. view of culm sheath. N Adaxial view of apical portion of
culm sheath. (From Wu et al. 2007, p. 55, with permission
In the protologue, Munro comments that from the Missouri Botanical Garden Press, St. Louis, and
“perfect flowers are very rarely found in this Science Press, Beijing)
species.”
tions near the apex, deciduous, leaving a narrow
109. Schizostachyum Nees Fig. 26 rim, auricles inconspicuous, with fimbriae, the
blade reflexed. Foliage leaves with or without
Schizostachyum Nees, Fl. Bras. Enum. Pl. 2: 535 (1829); auricles and fimbriae. Inflorescence an
phylog.: Yang et al. (2007). unbranched axis, each node with one or a few
Dendrochloa C. E. Parkinson, Indian Forester 59: 707 elongate pseudospikelets. Pseudospikelets with 1
(1933).
Leptocanna L. C. Chia & H. L. Fung, Acta Phytotax. Sin.
to several bracts with buds in their axils, with up
19: 212–213 (1981). to 4 flowers but only the terminal 1 or 2 fertile.
Glumes generally absent. Lodicules 0 to 3. Ovary
Culms erect to scandent, or drooping at the apex. on a stalk. Style 1, stigmas 3. Fruit fusiform, the
Branches many per node, subequal, from a single style base persistent.
bud. Culm leaf sheaths with 1 to 2 strong undula- Fifty spp., Asia, Pacific islands.
Although most species have culm leaves with conventional spikelets. Spikelets with one fertile
the blade reflexed, in S. chinense Rendel the culm flower. Glumes 4 to 6. Stamens 6. Style 1. Stigmas
sheath blade is erect. Dendrochloa is placed in 3.
synonymy here following the suggestion of Soder- Eight genera, 11 spp., Africa, Madagascar,
strom and Ellis (1987), although the Bamboo and Réunion.
Phylogeny Group (2012) recognize it as distinct.
112. Cathariostachys S. Dransf.
110. Stapletonia P. Singh, S. S. Dash & P. Kumari
Cathariostachys S. Dransf., Kew Bull. 53: 388–389 (1998).
Stapletonia P. Singh, S. S. Dash & P. Kumari, Nelumbo 51:
241 (2009). Culms erect, the tips arching; young shoots with
black hairs. Rhizome necks up to 4 m long.
Culms erect below, scandent above. Rhizomes Branches several to many at each node, with 1
with elongate necks. Branches (4) 6 per node, dominant, from a single bud. Culm leaf sheaths
subequal, from a single bud. Culm leaf sheaths persistent. Inflorescences dense, capitate, the
deciduous. Foliage leaves with auricles bearing branches branched, lateral branches arranged in
1 cm long fimbriae. Inflorescence a large head. tight fans. Spikelets with 5 glumes. Lodicules
Pseudospikelets with 1 flower, with a rachilla absent. Stigmas 3. Fruit stalked.
extension. Glumes 6 to 8, the apex aristate. Two spp., Madagascar.
Ovary on a stalk. Style 1, stigmas 2. Fruit fleshy, One species is eaten by endemic lemurs.
ca. 5 cm across.
One sp., S. arunachalensis (H.B. Naithani) P.
113. Decaryochloa A. Camus
Singh, S.S. Dash & P. Kumari, Arunachal Pradesh,
India. Decaryochloa A. Camus, Bull. Soc. Bot. France 93: 242
Stapletonia is segregated from Schizosta- (1947); tax.: Dransfield (1997).
chyum.
Culms scandent; internodes covered with black
111. Teinostachyum Munro hairs when young. Rhizome necks elongate.
Branches many in a clump. Culm leaf sheaths
Teinostachyum Munro, Trans. Linn. Soc. London 26: 142, auriculate, the auricles adaxially pubescent, with
t. 3 (1868); tax.: Stapleton (1994a). fimbriae. Foliage leaves with erect auricles, with
fimbriae. Inflorescence unbranched, surrounded
Culms erect below, scandent above, slender. by a spathe, with 1 to 4 spikelets, the spatheate
Branches several per node, subequal or with one units clustered. Glumes 3 or 4. Lodicules 3. Stig-
dominant. Culm leaves with the sheath margin mas 3. Ovary umbonate, pubescent. Fruit fleshy.
thickened, fimbriae present, the blade rolled. One sp., D. diadelpha A. Camus, Madagascar.
Foliage leaf sheaths with fimbriae. Inflorescence The fimbriae on the auricles of the foliage
unbranched, the pseudospikelets solitary or in leaves are 8–12 mm long, which according to the
clusters. Pseudospikelets with several flowers protologue “permettant de reconnaı̂tre cette
separated by long rachilla internodes. Glumes espèce en l’absence d’épillets”. In one specimen
2 to many. Lodicules 3. Stigmas 2 or 3. Fruit observed, the stamens are diadelphous, in two
baccate, with a beak. groups of three; in other specimens, however,
Two spp., Burma and India. the stamens are free.
114. Hickelia A. Camus Fig. 27

V.6. SUBTRIBE HICKELIINAE A. Camus (1924)
Hickelia A. Camus, Compt. Rend. Hebd. Séances Acad.
Nastinae Soderstr. & R. P. Ellis in Soderstr. (1987).
Sci. 179: 479 (1924); rev.: Dransfield (1994).
Pseudocoix A. Camus, Compt. Rend. Hebd. Séances Acad.
Rhizomes pachymorph. Foliage leaf blades with Sci. 179: 478 (1924).
adaxially projecting midribs. Flowers borne in
the blade erect initially, sometimes spreading

at maturity. Foliage leaves lacking auricles.
Inflorescence unbranched or the branches
branched, if the latter then each branch sub-
tended by a bract and prophyll. Glumes 3 to 5.
Lodicules 3. Ovary umbonate, the apex pubes-
cent, stigmas 3.
Four spp., Madagascar and eastern Africa.
115. Hitchcockella A. Camus
Hitchcockella A. Camus, Compt. Rend. Hebd. Séances

Acad. Sci. 181: 253 (1925).
Culms scandent. Rhizomes unknown. Branches 3

to several. Inflorescence bracteate, terminal on
lateral branches. Spikelets with 1 sterile flower
proximal to the fertile one. Glumes 2. Lodicules
3. Stigmas 2. Ovary glabrous, rostrate.
One sp., H. baronii A. Camus, Madagascar.
Camus describes Hitchcockella as similar to
Perrierbambus; Hitchcockella is distinguished
from the latter genus by its strongly laterally
compressed spikelets with keeled glumes and
lemmas (vs. nearly terete and non-keeled in Per-
rierbambus) and its long style that persists in the
fruit. The protologue provides little information
on the leaves.
116. Nastus Juss., s.s.
Nastus Juss., s.s., Gen Pl. 34 (1789).
Culms arborescent or scandent. Lateral branches

branched, 3 to many, subequal or one or two
dominant, generally arising at the node. Culm
sheath deciduous or not, auriculate. Foliage
leaves with sheaths with or without auricles,
with or without fimbriae. Inflorescence
unbranched or the branches branched, sur-
Fig. 27. Bambusoideae-Bambuseae-Hickeliinae. Hickelia rounded by the subtending leaf sheath. Spikelets
africana. A Inflorescence. B Branches. C Culm sheath. D with rachilla extension absent. Glumes 5 or 6.
Spikelet before anthesis. E Anthetic spikelet. F Lemma. G
Palea. H Apex of palea, viewed from inside. I Gynoecium. Lodicules 3. Ovary with or without an enlarged
J Lodicules. K Caryopsis. (From Dransfield 1994, drawn apex, the apex pubescent or not.
by the author) Twenty spp., Réunion, Madagascar.
The type species of Nastus, N. borbonicus J.F.
Culms erect, the upper part leaning. Rhizomes Gmel., is clearly placed here in Hickeliinae (L.G.
elongate or thickened. Branches many per node, Clark, pers. comm.), along with the Madagascan
one dominant, the others thinner, from a single species of the genus. The southeast Asian species,
bud. Culm leaves with sheaths lacking auricles, however, belong in Bambusinae and need to be
placed in a separate genus; available names are One sp., S. parvifolia (Munro) S. Dransf.,
Chloothamnus B€ use and Oreiostachys Gamble. Madagascar.
These are based on Southeast Asian material
and are not synonyms of Nastus s.s. 119. Valiha S. Dransf.
In some species, the nodal line is curved
downward. This has led some authors to report Valiha S. Dransf., Kew Bull. 53: 380 (1998).
the bud complement incorrectly as being below
the node, whereas it is the nodal line itself whose Culms erect, the tips arching. Rhizomes with
position has shifted. This character also appears necks elongate. Branches many per node, the
convergently in some species of Chusquea. central one dominant, from a single bud below
the node. Culm leaf sheaths with auricles. Foliage
117. Perrierbambus A. Camus leaves with sheaths lacking auricles. Inflorescence
branches branched, bracteate, the bracts lacking
Perrierbambus A. Camus, Bull. Soc. Bot. France 71: 697 axillary buds. Spikelets sometimes subterete.
(1924). Glumes 5 or 6. Lemma with 29 to 41 veins. Lodi-
cules 3. Stigmas 3. Fruit with pericarp free from
Culms erect. Branches numerous, thinner than seed coat.
the culm. Culm leaves without auricles or fim- Two spp., Madagascar.
briae. Foliage leaves with conspicuous transverse
veins. Inflorescences terminal on each lateral
branch, unbranched, with 1 to 3 spikelets hidden
by the distalmost leaves of the branch. Glumes 1 V.7. SUBTRIBE RACEMOBAMBOSINAE Stapleton
or 2, their apices acuminate. Lodicules 3. Stigmas (1994)
2 or 3, style branches 2 or 3, distinct to the base.
Ovary apex puberulent, extended to a narrow Culms generally scandent, over 3 m tall. Rhi-
beak. Fruit with pericarp free from seed coat. zomes pachymorph. Branches several to many,
Two spp., Madagascar. with one branch dominant, from a single bud.
The beak of the ovary may actually corre- Culm leaves with sheaths deciduous, with small
spond to a short style. The protologue notes rim-like fimbriate auricles, the blades erect or
“folia caduca”, but it is unclear whether this reflexed. Foliage leaf sheaths generally with auri-
refers to the culm leaves or foliage leaves. cles and fimbriae. Inflorescence branched or
unbranched (“racemose”). Spikelets conven-
tional, with 3 to 8 flowers plus one distal reduced
118. Sirochloa S. Dransf. flower. Glumes 2 or 3. Stamens 6. Stigmas 3.
Lodicules 3. Ovary apex generally pubescent.
Sirochloa S. Dransf., Kew Bull. 57: 965–966, f. 1 (2002).
One genus, 17 spp.
This subtribe was originally erected to
Culms clumped, erect below, scrambling above.
include Neomicrocalamus, Racemobambos, and
Branches several per node with one dominant,
Vietnamosasa, which were thought to be distin-
from a single bud. Culm leaves with sheaths
guished from members of Bambusinae by having
with a rugose basal portion, “breaking up during
conventional spikelets. The analyses of Yang et al.
branch development”, without auricles, with
(2008) place Neomicrocalamus plus two species
blades spreading. Foliage leaves with auricles
formerly in Racemobambos with Bonia in Bam-
prominent, with fimbriae. Inflorescence terminal
businae. Specifically, Racemobambos yunnanen-
on lateral branches, unbranched, bracteate. Spi-
sis is sister to Melocalamus, whereas R. prainii is
kelets with a rachilla extension. Glumes 4. Lodi-
sister to Neomicrocalamus, with which it has been
cules 3. Stigmas 3. Ovary glabrous, style short or
combined. Vietnamosasa also is now placed in
elongate. Fruit with pericarp free from seed coat,
Bambusinae. Once these species are removed,
brittle.
Racemobambos appears to be monophyletic.
Culms herbaceous, rarely reed-like, caespitose.

Culm leaves usually not differentiated from
foliage leaves. Outer ligule absent. Leaf sheaths
generally lacking auricles and fimbriae. Inflores-
cences generally axillary and terminal, sometimes
borne on a leafless culm separate from the vege-
tative culm. Plants monoecious, generally with
large pistillate spikelets and small staminate spi-
kelets. Spikelets often dorsiventrally compressed,
with one flower, without a rachilla extension. Pis-
tillate spikelets with the pedicel apex generally
wide and club-shaped; glumes 2, generally longer
than the flower; lodicules 3; stigmas 2 (rarely
three). Staminate spikelets generally lacking
glumes; lemma and palea membranous; lodicules
0 or 3; stamens 2 or 3.
Twenty-one genera, 122 species.
We know little about the phylogeny of this
group. Buergersiochloa has been supported as
sister to the rest by both morphological (Kellogg
and Watson 1993) and molecular (Bouchenak-
Khelladi et al. 2008; Oliveira et al. 2014; Zhang
and Clark 2000) data, and is the only genus that
does not occur in the New World. The remaining
genera share the synapomorphy of crenate and
cross-shaped silica bodies, and all but Pariana
and Eremitis have hardened lemmas and paleas
in the pistillate spikelets. The latter two genera
are clearly sisters, with strong morphological and
molecular synapomorphies. Sucrea and Raddia
Fig. 28. Bambusoideae-Bambuseae-Racemobambosinae. are sisters in molecular phylogenies, a relation-
Racemobambos hepburnii. A Flowering branches. B ship that is consistent with their distinctive glume
Glumes. C Floret and rachilla internode. D Lemma. E
Palea. F Ovary, style removed. (From Dransfield 1992)
morphology (Oliveira et al. 2014; Zhang and
Clark 2000). “Olyra” is apparently paraphyletic
with Lithachne and Arberella derived within it
(Oliveira et al. 2014), consistent with the sugges-
120. Racemobambos Holttum Fig. 28 tion of Judziewicz et al. (1999). As currently cir-
cumscribed, “Olyra” is defined as much by plant
Racemobambos Holttum, Gard. Bull. Singapore 15: 268
(1956); rev.: Dransfield (1992). size as anything else.
Characters as for the subtribe. The Bamboo Phylogeny Group (2012) note
that there is no morphological synapomorphy
Seventeen spp., southeast Asia. for this tribe, although monoecy may be synapo-
The dominant branch at a given node some- morphic within the context of the subfamily. A
times develops late, so that the minor branches major criterion for distinguishing genera of Olyr-
appear to form a cluster. eae is the distribution of the sexes in the inflores-
cence. This character may simply be highly labile
over evolutionary time, and may not be a good
VI. TRIBE OLYREAE Martinov (1820) indicator of relationship. However, testing this
requires more phylogenetic, developmental, and
Buergersiochloeae S.T. Blake (1946). genetic work.
Parianeae C.E. Hubb. (1934).
The treatment here is based heavily on that of

Judziewicz et al. (1999), which should be con-
sulted for more extensive descriptions of genera,
species, their geographic ranges, and their ecol-
ogy.
VI.1. SUBTRIBE BUERGERSIOCHLOINAE (S. T. Blake) L. G.

Clark & Judz. (2007)
Culms dimorphic, the vegetative stems leafy,

flowering stems with leaf blades strongly reduced
or entirely lacking. Inflorescence branches
branched, bearing pistillate spikelets distally, sta-
minate proximally. Pistillate spikelet with glumes
2, shorter than the flower; lemma leathery but not
hardened, pubescent, awned; staminodes 3; ovary
with style 1, stigmas 2. Staminate spikelet with
glumes lacking; lemma with a mucro or awn;
stamens 2 or 3, the filaments becoming fused
near anthesis. Lodicules 3.
121. Buergersiochloa Pilg. Fig. 29
Buergersiochloa Pilg., Bot. Jahrb. Syst. 52: 167 (1914).
Characters of the subtribe.

One sp., B. bambusoides Pilg., New Guinea.
Fig. 29. Bambusoideae-Olyreae-Buergersiochloinae. Buer-
The leaf epidermal structure and internal leaf gersiochloa bambusoides. A Habit. B Leaf. C Inflorescence.
anatomy are unusual and not obviously similar to D Staminate spikelet. E Lemma of staminate spikelet. F
other Bambusoideae (Soderstrom and Ellis 1987). Palea of staminate spikelet. G Androecium. H Pistillate
spikelets. J, K Glumes of pistillate spikelets. L Lemma. M
Remaining genera: Leaf epidermis with cross- Palea. N Pistil, lodicules and staminodes. O Base of pistil. P
shaped and crenate silica bodies, the former over Half-mature fruit. (From Pilger 1915, with permission from
E.Schweizerbart’sche Verlagsbuchhandlung OHG, www.
the vascular tissue (costal zone) and the latter over schweizerbart.de)
the mesophyll (intercostal zone).
All species are restricted to the New World,
except for “Olyra” latifolia, which also occurs in lets with flattened, hardened pedicels. Stamen
Africa where it is either native or naturalized. filaments fused.
122. Eremitis D€oll
VI.2. SUBTRIBE PARIANINAE Hack. in Engler & Prantl oll, Fl. Bras. 2, 2: 338 (1877); tax.: Ferreira et al.
Eremitis D€
(1887) (2013).
Plants caespitose, the clumps connected by rhi- Inflorescences on erect leafy culms, on trailing
zomes or stolons. Top of leaf sheath with thick- culms with reduced leaves, and on subterranean
ened auricles bearing fimbriae. Inflorescence culms with inflated spathe-like leaves. Pistillate
terminal only, unbranched. Pistillate spikelet ter- spikelets with glumes with 1 vein. Staminate spi-
minal, surrounded by whorls of staminate spike- kelets on long fused pedicels in distal whorl,
shorter pedicels in lower whorls; lodicules 0 or Leaves with the top of the sheath variously
similar to stamens; stamens 2. 2n 60, 67–94. decorated with translucent swellings (“lunar
Three spp., Atlantic Brazil. marks”). Inflorescence cylindrical, with a whorl
Two of the three species of Eremitis have of five staminate spikelets surrounding a
papillae on the adaxial surfaces of their leaves; single sessile pistillate spikelet and the associated
these papillae refract light and create blue irides- rachis internode. Pistillate spikelets with three or
cence, which may be an adaptation to shade (Fer- more lodicules. Staminate spikelets with pedicels
reira et al. 2013). often fused; stamens numerous, showy. 2n ¼ 22,
44, 38.
123. Pariana Aubl. Fig. 30 Thirty-five spp., Costa Rica and Trinidad,
south through tropical South America.
Pariana Aubl., Hist. Pl. Guiane 2: 876, pl. 337 (1775). Pariana is presumed to be insect pollinated
(Soderstrom and Calderón 1971).
VI.3. SUBTRIBE OLYRINAE Kromb. (1875)
Pistillate spikelet with lemma and palea hard-

ened. Staminate flowers with 3 stamens.
124. Agnesia Zuloaga & Judz.
Agnesia Zuloaga & Judz., Novon 3: 306 (1993).
Proximal leaves lacking blades, blades on leaves

only above the middle of the culm. Inflorescences
axillary, with one terminal pistillate spikelet and
several proximal staminate spikelets, the latter on
slender pedicels. Glumes of pistillate spikelet with
long stiff awns; pistillate flowers covered with
long hairs, awned.
One sp., A. lancifolia (Mez) Zuloaga & Judz.,
Peru, Colombia, Brazil.
125. Arberella Soderstr. & C.E. Calderón
Arberella Soderstr. & C.E. Calderón, Brittonia 31: 433

(1979).
Inflorescences with few flowers, borne at all nodes

or only at upper and lower ones, with one termi-
nal pistillate spikelet and several proximal stami-
nate spikelets. Pistillate spikelets with glumes
with many veins, with a prominent rachilla inter-
node below the flower; lemma smooth, bearing
hairs. 2n ¼ 22, 24.
Fig. 30. Bambusoideae-Olyreae-Parianinae. Pariana Seven spp., Central and South America.
radiciflora. A Whole plant. B Staminate (above) and pis- Arberella may be derived within “Olyra” (Oli-
tillate (below) spikelets. (From Davidse et al. 2004, with
permission from the Missouri Botanical Garden Press; veira et al. 2014).
drawn by Bruno Manara)
126. Cryptochloa Swallen One sp., F. boutelouoides G.A. Black, French

Guiana and adjacent Brazil.
Cryptochloa Swallen, Ann. Missouri Bot. Gard. 29: 317 The terminal inflorescence is showy, with the
(1942). spikelets arranged so that they look like a single
large spikelet rather than multiple small ones.
Inflorescences terminal and axillary, often partly
hidden in leaf sheaths, the terminal inflorescence
generally with only staminate spikelets, the axil- 129. Lithachne P. Beauv.
lary ones with spikelets of both sexes mixed.
Lithachne P. Beauv., Ess. Agrostogr. 135, 166, 168, t. 24, f.
Pistillate spikelets with glumes with 5 veins, the 2 (1812).
rachilla internode below the flower generally
thick and oil-containing, falling with the flower; Plants generally less than 30 cm tall. Leaf blade
lemma and palea becoming dark mottled at with the base asymmetrical. Inflorescences termi-
maturity. Staminate spikelets without glumes, nal and axillary, the terminal inflorescence gen-
with 3 or rarely with 2 stamens. 2n ¼ 22. erally with only staminate spikelets, the axillary
Eight spp., Central to South America. ones with distal pistillate spikelets and proximal
The thickened oil-bearing rachilla internode staminate ones. Pistillate spikelet with glumes
is absent in C. capillata (Trin.) Soderstr. and C. elongate, membranous, longer than the flower;
soderstromii Davidse. rachilla internode below the flower hardened;
lemma and palea white (becoming mottled), the
127. Ekmanochloa Hitchc. lemma gibbous. 2n ¼ 22.
Four spp., Central and South America, West
Ekmanochloa Hitchc., Man. Grasses W. Ind. 374 (1936). Indies.
The odd shape of the pistillate lemma in
Vegetative culms with multiple foliage leaves, Lithachne makes the flower look rather like a
flowering culms with few leaves, these lacking large tooth. Lithachne is monophyletic and possi-
blades. Inflorescence terminal, with 2 branches, bly derived within “Olyra” (Oliveira et al. 2014;
one bearing staminate spikelets and the other, Zhang and Clark 2000).
slightly longer, bearing pistillate spikelets. Pistil-
late spikelets elongate, narrow; lemma awned, the 130. “Olyra” L. Fig. 31
awn slender, much longer than the body of the
lemma. Staminate spikelets elliptical, sometimes Olyra L., Syst. Nat. (ed. 10), 2: 1253, 1261, 1379 (1759).
with 2 stamens. Fusoid cells lacking.
Two spp., eastern Cuba. Culms often up to 3 m tall or more. Inflorescences
Ekmanochloa occurs only on serpentine soils terminal or axillary from upper leaves, with pis-
(Zuloaga et al. 1993). tillate spikelets terminal in each inflorescence or
on each major inflorescence branch, staminate
128. Froesiochloa G.A. Black spikelets proximal. Pistillate spikelets on sturdy
pedicels. Staminate spikelets without glumes.
Froesiochloa G.A. Black, Bol. Tecn. Inst. Agron. N. 20: 29 2n ¼ 14, 22, 23, 40, 44.
(1950). Twenty-four spp., Central and South Amer-
ica, West Indies.
Leaf blades broad, with short petioles, deciduous. Judziewicz et al. (1999) suggest that Olyra
Inflorescence terminal, a cluster of spikelets, might be para- or polyphyletic, a hypothesis con-
the terminal spikelet pistillate, proximal ones firmed by the data of Oliveira et al. (2014). Arber-
staminate, spirally arranged. Pistillate spikelet ella and Lithachne are derived within “Olyra”,
with the rachilla internode below the flower gen- while the data on Cryptochloa are ambiguous.
erally thick and oil-containing; lodicules absent. Agnesia could also be derived from within
Staminate spikelets with stamens with filaments “Olyra” but data are not available to test this.
fused. Some species are visited and possibly pollinated
131. Parodiolyra Soderstr. & Zuloaga
Parodiolyra Soderstr. & Zuloaga, Smithsonian Contr. Bot.

69: 64 (1989).
Culms scrambling. Inflorescence lax, the

branches branched, distal branches with mixed
staminate and pistillate spikelets, or pistillate spi-
kelets only, the proximal branches with staminate
spikelets only. Pistillate spikelets becoming black
at maturity, falling with the glumes; rachis inter-
node between glumes swollen. 2n ¼ 36.
Five spp., Costa Rica through tropical South
America.
132. Raddiella Swallen
Raddiella Swallen, Bull. Torrey Bot. Club 75: 89 (1948).
Culms short, leaves <1.5 cm long in most spe-

cies. Inflorescences terminal and axillary, with
few spikelets, with staminate and pistillate spi-
kelets mixed in the inflorescence, or terminal
inflorescences wholly staminate, axillary wholly
pistillate. Pistillate flower glabrous, pale but
becoming dark at maturity in some species.
2n ¼ 20.
Fig. 31. Bambusoideae-Olyreae-Olyreae. Olyra micrantha. Eight spp., Panama, Trinidad, tropical South
A Inflorescence. B Leaf blade, with pseudopetiole and America.
ligule. C Base of the plant. D Portion of a branch of
inflorescence with staminate spikelet. E Staminate spikelet.
F Pistillate spikelet. G Lemma of staminate spikelet. H 133. Piresia Swallen
Palea of staminate spikelet, viewed from the adaxial side.
I Lower glume of pistillate spikelet. J Upper glume of pistil-
Piresia Swallen, Phytologia 11: 152 (1964); tax.: de Car-
late spikelet. K Lemma and palea of pistillate flower, left
abaxial side, right adaxial side. L Three caryopses in side valho et al. (2012).
view. (Reprinted from Soderstrom and Zuloaga 1989, cour-
tesy of Smithsonian Institution; drawn by G.B. Threlkeld) Culms dimorphic. Leaves of upright culms with
broad blades; leaves of decumbent culms with
blades reduced or lacking. Inflorescences termi-
nal and axillary, most commonly on decumbent
by insects (Soderstrom and Calderón 1971). culms, but sometimes on upright culms as well.
Although many species of “Olyra” range from Pistillate and staminate spikelets morphologically
0.3–3 m in height, some species are reported to distinct, or morphologically apparently bisexual
reach 5 or 6 m. but functionally unisexual, one distal pistillate
spikelet and several proximal staminate spikelets
PARODIOLYRA + RADIELLA near the apex of the inflorescence, sometimes
additional staminate and pistillate spikelets inter-
Leaf blades with apex asymmetrical, apiculate.
Parodiolyra is paraphyletic with Raddiella mixed proximally. Pistillate lemma densely
derived from within it (Oliveira et al. 2014). The pubescent. 2n ¼ 22.
two genera should perhaps be combined. Five spp., lowland tropics of South America.
134. Reitzia Swallen either sister to or derived from within Sucrea

(Oliveira et al. 2014).
Reitzia Swallen, Sellowia 7: 7 (1956).
Culms <30 cm tall. Inflorescence terminal, with MNIOCHLOA + PIRESIELLA + REHIA

several branches, each with a terminal pistillate Plants with corms.
spikelet and one or two proximal staminate spi-
kelets. Pistillate flower shorter than the glumes,
becoming darkly marbled at maturity. 137. Mniochloa Chase
One sp., R. smithii Swallen, southern Atlantic
Mniochloa Chase, Proc. Biol. Soc. Wash. 21: 185 (1908).
Brazil.
Culms dimorphic, vegetative culms leafy, flower-
SUCREA + RADDIA
ing culms with leaves lacking blades. Inflores-
Glumes of pistillate spikelets with thickened mar- cence with 2 branches, one bearing pistillate
gins extending into long twisted apices. spikelets, the other bearing staminate ones; spi-
kelets on thickened pedicels. Pistillate spikelet
Diaspores (anthecia) in these two genera are with glumes longer than the flower. Staminate
forcibly ejected from the glumes as the glumes spikelets much smaller than pistillate ones.
dry and curl (Sendulsky 1993). This mode of One sp., M. pulchella (Griseb.) Chase, eastern
dispersal, ballistochory, is otherwise unknown Cuba; possibly extinct.
in the grasses.
138. Piresiella Judz., Zuloaga & Morrone
135. Sucrea Soderstr.
Piresiella Judz., Zuloaga & Morrone, Ann. Missouri Bot.
Gard. 80: 856 (1993).
Sucrea Soderstr., Brittonia 33: 200 (1981).
Plants with obvious prop roots. Leaves one or a Culms short, often only a few centimeters tall,
few per culm, sheaths slightly inflated. Inflores- dimorphic, vegetative culms leafy, flowering
cence branches branched, with pistillate spikelets culms with leaves lacking blades, with slender
mostly on distal branches, staminate generally on stolons. Inflorescences with 2 branches, one bear-
proximal branches. 2n ¼ 22. ing pistillate spikelets, the other bearing stami-
Three spp., southeastern Brazil. nate ones. Pistillate spikelet with glumes longer
Sucrea may be paraphyletic with Raddia than the flower, the lemma apiculate; stigmas 3.
derived from within it (Oliveira et al. 2014). Staminate spikelets smaller than pistillate, lack-
ing glumes; stamens 2.
One sp., P. strephioides (Griseb.) Judz.,
136. Raddia Bertol. Zuloaga & Morrone, western Cuba.
Raddia Bertol., Opusc. Sci. 3: 410 (1819); tax.: Oliveira

et al. (2008). 139. Rehia Fijten
Leaves arranged along the culm with precise Rehia Fijten, Blumea 22: 416 (1975).
spacing, looking superficially similar to a large
compound leaf. Inflorescences terminal and axil- Culms short, caespitose. Inflorescence terminal,
lary, the terminal inflorescence with branches with several branches, each with a terminal pistil-
branched, bearing staminate spikelets, the axil- late spikelet and one or two proximal staminate
lary ones unbranched, with pistillate spikelets, or spikelets. Pistillate flowers dark marbled color at
the branches branched and bearing staminate maturity, glabrous except for a few hairs at the
spikelets. 2n ¼ 22. base, apex and margins of lemma. 2n ¼ 20.
Nine spp., French Guiana, Brazil. One sp., R. nervata Fijten, northeastern Brazil
Molecular data from all nine species of Rad- and adjacent French Guiana, Surinam, and
dia show that the genus is monophyletic and Guyana.
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VI. Subfamily Pooideae Benth. (1861)
Festucoideae Link (1827).
Annuals or perennials, caespitose, rhizomatous nomically and ecologically important of all tem-
or stoloniferous. Culms hollow. Ligule membra- perate grasses. These studies have substantially
nous, sometimes with a ciliolate margin but not revised our understanding of the history and
long ciliate. Leaf blades lacking pseudopetioles. classification of this group. In all molecular phy-
Leaf sheaths lacking auricles. Inflorescence logenies, Brachyelytrum is sister to the rest of the
unbranched, or of primary branches only, or the subfamily, with Nardeae (Nardus plus Lygeum)
branches branched. Spikelets with 1 to many the next diverging branch (Davis and Soreng
flowers, reduced flowers, if present, distal to the 2007; GPWG 2001; GPWG II 2012; Romaschenko
fertile ones. Rachilla generally breaking up et al. 2012; Schneider et al. 2009, and many
between the flowers, i.e., disarticulation above others). The relative positions of Meliceae, Phae-
the persistent glumes. Lemma awned or not. nospermateae, and Stipeae differ among analyses
Palea with 2 veins, often keeled. Lodicules 2, and likely reflect differences in taxon sampling
membranous, not vascularized. Stamens 3. Stig- and molecular markers. For example, the GPWG
mas 2, borne on style branches that are inserted II (2012), using three chloroplast loci, finds Sti-
separately into the ovary apex. Fruit with a linear peae, Phaenospermateae and Meliceae as succes-
hilum. Embryo small, with an epiblast, lacking an sive sister groups to the remainder of the pooids
internode between the coleoptilar and scutellar (Diarrheneae not included), whereas Schneider
vein, the embryonic leaf margins overlapping. et al. (2009, 2011) find the order to be Meliceae,
C3. Fusoid cells and arm cells absent. Leaf epider- Phaenospermateae, Stipeae, Diarrheneae and the
mal papillae absent. rest. Other authors find other arrangements,
One hundred seventy-seven genera, 3850 spp. which are often poorly resolved or weakly sup-
Pooideae are monophyletic in all molecular ported. For the moment then, the order of
phylogenies to date. The character “loss of stylar branching is unknown, which affects efforts to
fusion” appears to be synapomorphic (GPWG optimize characters on this part of the phylogeny.
2001), although anatomical interpretation of this After divergence of the tribes mentioned
character is unclear. It seems most likely that it above, most studies agree that Brachypodium is
represents a change in the zone of growth of the sister to the rest of the subfamily (e.g., GPWG II
developing carpels so that fusion occurs below 2012). After the divergence of Brachypodium, the
the separate lobes rather than above. In this remaining species are marked by a sharp increase
case, the style “branches” might properly be in genome size (Bennetzen and Kellogg 1997;
called stylodia, but this term is not used regularly Kellogg and Bennetzen 2004). Littledalea, Bromus
in the grass literature so I use the term style and Triticeae form a clade that is sister to the
branches here. Poeae s.l. (Schneider et al. 2009, 2011). The latter
Comprehensive phylogenetic studies of this group includes the former Aveneae as well as
subfamily have appeared only in the last few several smaller tribes, which are clearly non-
years, even though this group is the most eco- monophyletic and hence not recognized here.
200 VI. Subfamily Pooideae Benth. (1861)
Poeae s.l. has been supported as monophyletic in TRIBES AND SUBTRIBES OF POOIDEAE
all molecular analyses.
Hybridization is common in Pooideae, as VII. Tribe Brachyelytreae
noted years ago by Stebbins (1950), and in a phy- VIII. Tribe Nardeae
IX. Tribe Phaenospermateae
logenetic context by Kellogg and Watson (1993); X. Tribe Stipeae
hybridization is often associated with polyploidy. XI. Tribe Meliceae
Many taxa exhibit morphological characters XII. Tribe Diarrheneae
that combine characteristics of the putative paren- XIII. Tribe Brachypodieae
tal taxa, leading to many named hybrid genera XIV. Tribe Bromeae
XV. Tribe Triticeae
(e.g., xElyhordeum Mansf. ex Tsitsin & K. A. XVI. Tribe Poeae
Petrova, xPucciphippsia Tzvelev). Additional evi- Incertae sedis
dence for hybridization has come from cytogenet- XVI.1 Subtribe Torreyochloinae
ics and isozymes, but the most compelling data XVI.2. Subtribe Aveninae
have come from DNA sequences. Discrepancies XVI.3. Subtribe Phalaridinae
XVI.4. Subtribe Anthoxanthinae
between phylogenies produced by nuclear and XVI.5. Subtribe Agrostidinae
chloroplast genes are increasingly demonstrating XVI.6. Subtribe Scolochloinae
reticulation. In addition, single copy nuclear genes XVI.7. Subtribe Airinae
are clearly effective in resolving complex reticulate XVI.8. Subtribe Ammochloinae
phylogenies (see, for example, studies on Elymus; XVI.9. Subtribe Cynosurinae
XVI.10. Subtribe Loliinae
Fan et al. 2013; Hu et al. 2013; Mason-Gamer et al. XVI.11. Subtribe Dactylidinae
2002, 2010a, 2010b), and these will need to be XVI.12. Subtribe Parapholiinae
applied to problems throughout the subfamily. XVI.13. Subtribe Sesleriinae
Several large polyploid genera appear to be poly- XVI.14. Subtribe Coleanthinae
phyletic (e.g., “Calamagrostis”; Saarela et al. 2010); XVI.15. Subtribe Poinae
the morphological characters associated with such
genera are clearly homoplasious, creating difficul-
ties with diagnosis. KEY TO THE GENERA OF POOIDEAE
Generic limits are a problem throughout
1. Base of inflorescence surrounded by a trumpet-shaped
Pooideae, perhaps more so than in any other
bract originating from the basal inflorescence node;
subfamily of grasses except the bamboos. Some plants of the Mediterranean 302. Cornucopiae
large genera (e.g., Festuca, Poa) are paraphyletic 1. Inflorescence without a trumpet-shaped bract; plant
as traditionally circumscribed, with monotypic distribution various 2
genera with one or two unusual characters having 2 (1). Inflorescence composed of a single spikelet, barely
extending above the leaves 162. Aciachne
been segregated from the main genus. Mono-
2. Inflorescence with more than 1 spikelet 3
phyly can be restored by placing the segregates 3. Inflorescence unbranched, with spikelets sessile on the
into synonymy, although this sometimes means rachis, or with short pedicels Group I
that no obvious morphological character remains 3. Inflorescence branched 4
to identify the sensu lato genus. 4 (3). Spikelets without awns Group II
4. Spikelets with awns 5
All three phenomena – paraphyly, polyphyly,
5 (4). Awns straight Group III
and frequent intergeneric hybridization – 5. Awns curved, twisted, or geniculate Group IV
suggest that the subfamily taxonomy would be
clarified by recognizing fewer, more broadly
defined genera. While the classification pre- Group I – Unbranched inflorescences
sented here is a step in that direction, it certainly 1. Glumes and/or lemmas with awns 2
does not go far enough. As phylogenies are pro- 1. Spikelets entirely without awns 40
duced with more extensive species sampling, 2 (1). Spikelets on short pedicels 3
2. Spikelets sessile 12
including the many monospecific genera, and
3 (2). Flowers 1 per spikelet 4
as morphological studies become more detailed, 3. Flowers more than 1 per spikelet 5
the limits of the sensu lato genera are likely to 4 (3). Glumes large, enclosing the flower; awn from back
become clearer. of lemma; Afghanistan to western China 148. Duthiea
KEY TO THE GENERA OF POOIDEAE 201
4. First glume missing, second glume minute; awn an 23 (20). Plants annual 24
extension of the lemma; eastern Asia and eastern 23. Plants perennial 26
North America 142. Brachyelytrum 24 (23). Glumes with 1 vein 268. Loliolum
5 (3). Lemma with 5 to 9 awns 24. Glumes with >1 vein, these often prominent 25
194. Bromus (sect. Boissiera) 25 (24). Glumes oblong, elliptic, or ovate; awned or not,
5. Lemma with 1 awn 6 their apices truncate 196. Aegilops
6 (5). Distal portion of glumes hyaline; apex of upper 25. Glumes linear, extending to an attenuate apex
glume generally ragged; upper lemma margin hyaline; 199. Crithopsis
plants of wet sites 188. Pleuropogon 26 (23). Spikelets strongly divergent from the rachis, or
6. Glumes and lemmas membranous or firm, but not even somewhat reflexed 198. Australopyrum
hyaline; plants of mesic to dry sites 7 26. Spikelets appressed to rachis, not reflexed 27
7 (6). Lemma awned from sinus or back 8 27 (26). Glumes subulate to narrowly lanceolate, 1-veined
7. Lemma awn terminal 10 at mid-length 28
8 (7). Spikelet with 2 flowers; lemma with 11 rib-like veins 27. Glumes never subulate, variously lanceolate, rectan-
186. Lycochloa gular, ovate or obovate, generally with 3 or more veins
8. Spikelet with 4 or more flowers; lemma with few veins, at mid-length 29
these not rib-like 9 28 (27). Inflorescence erect; plants of cool temperate
9 (8). Ligule 7 mm long; lodicules 2; western China and regions but not New Zealand 207. Leymus
Himalayas 150. Pseudodanthonia 28. Inflorescence nodding or drooping; plants of New
9. Ligule <3 mm long; lodicules 3; Balkans and Caucasus Zealand 212. Stenostachys
147. Danthoniastrum 29 (27). Spikelets about twice the length of inflorescence
10 (7). Spikelets with 3 flowers, the lower 2 sterile; awn of internodes or more, the spikelets thus overlapping in
upper flower ca. 2 cm; plants of China and Japan the inflorescence 30
184. Brylkinia 29. Spikelets shorter than inflorescence internodes, the
10. Spikelets with 2 to many flowers, but never with the inflorescence thus appearing sparse 32
2 lower ones sterile; if 3 flowers, then awn of upper 30 (29). Leaves >0.5 mm wide when dry 201. Elymus p.p.
flower much less than 2 cm 11 30. Leaves 0.5 mm wide when dry 31
11 (10). Spikelets with 2 or 3 flowers; delicate annuals with 31 (30). Spikelets with 3 to 5 flowers 203. Festucopsis
leaves generally less than 1 mm wide 266. Festuca 31. Spikelets with 5 to 10 flowers 208. Peridictyon
11. Spikelets with more than 5 flowers; perennials or 32 (29). Rachis tough 210. Pseudoroegneria
sturdy annuals with leaves much wider than 1 mm 32. Rachis fragile 273. Agropyropsis
193. Brachypodium 33 (12). Plants annual 34
12 (2). Spikelets 1 per node 13 33. Plants perennial 37
12. Spikelets more than 1 per node, at least in the middle 34 (33). Spikelets embedded in hollows in the rachis
of the inflorescence 33 258. Scribneria
13 (12). Spikelets with glumes minute or absent 14 34. Rachis terete, spikelets not embedded 35
13. At least the first glume present, well developed 16 35 (34). Lemmas bearing long tuberculate hairs
14 (13). Plants annual 266. Festuca (former Psilurus) 205. Heteranthelium
14. Plants perennial, caespitose 15 35. Lemmas glabrous or scabrous, but not with long hairs
15 (14). Spikelets borne on one side of the inflorescence 36
axis; Europe 143. Nardus 36 (35). Awns many times longer than the lemmas,
15. Spikelets borne on opposite sides of the inflorescence extending at least twice the length of the inflorescence,
axis; New Zealand 212. Stenostachys tangled 213. Taeniatherum
16 (13). Lemma awned from the back or from a sinus 17 36. Awns up to twice as long as lemmas, extending no
16. Lemma awnless or with an apical awn 19 more than twice the length of the inflorescence and
17 (16). Plants annual, generally less than 30 cm tall 18 often much less, straight 206. Hordeum
17. Plants perennial, generally well over 30 cm tall 37 (33). Spikelets three per node, the central 1 sessile and
149. Metcalfia the lateral 2 pedicellate; spikelets with 1 flower
18 (17). Spikelets not sunken, borne on distinct pedicels 206. Hordeum
226. Trisetaria 37. All spikelets at the node sessile, each with 2 or more
18. Spikelets sunken in the inflorescence axis flowers, although the distal ones sometimes reduced
258. Scribneria and sterile 38
19 (16). Spikelet edgewise to rachis 267. Lolium 38 (37). Rachis disarticulating, rachilla tough; spikelets
19. Spikelet broadside to rachis 20 with 1 or 2 flowers 209. Psathyrostachys
20 (19). Lemmas and often glumes strongly keeled 21 38. Rachis tough, rachilla disarticulating; spikelets gener-
20. Neither lemmas nor glumes strongly keeled 23 ally with >2 flowers 39
21 (20). Margins of glumes and distal margins of lemmas 39 (38). Glumes with 1 vein; leaf blades stiff, often bluish
with long silky hairs 200. Dasypyrum with wax 207. Leymus
21. Margins of glumes and lemmas not long hairy 22 39. Glumes with 3 veins; leaf blades lax, green 201. Elymus
22 (21). Glumes subulate, lemmas lanceolate 211. Secale 40 (1) Spikelets 1 or 2 per inflorescence 41
22. Glumes lanceolate or oblong, lemmas lanceolate, 40. Spikelets more than 2 per inflorescence, the inflores-
ovate, or oblong 215. Triticum cences extending well above the leaves 42
41 (40). Inflorescences not or scarcely extending above 58 (55). Glumes stiff, obtuse to truncate 214. Thinopyrum
the leaves; leaves terete, their apices subulate, plants 58. Glumes and lemmas acute to acuminate, sometimes
cushion-forming; plants of the high Andes mucronate or awn-tipped, flexible 59
162. Aciachne 59 (58). Spikelets borne edgewise to the rachis, or if
41. Inflorescences borne well above the leaves; spikelet broadside to it, then the base corm-like and glumes
solitary, terminal, covered with dense golden or white membranous 267. Lolium
hairs the length of the spikelet; spikelet subtended by a 59. Spikelets borne broadside to the rachis, the base not
spathe; leaves flat; Mediterranean 144. Lygeum corm-like; glumes as firm as or firmer than the lemmas
42 (40). Spikelets partially embedded in hollows in the 60
rachis 43 60 (59). Spikelets scarcely overlapping along the inflores-
42. Rachis terete, spikelets not embedded 47 cence axis, the apex of one just reaching the base of the
43 (42). Spikelets on one side of the inflorescence axis, next 210. Pseudoroegneria
each with 4 to 6 flowers 60. Spikelets overlapping along the inflorescence axis, the
266. Festuca (former Narduroides) apex of one reaching about the mid-point of the next
43. Spikelets on opposite sides of the inflorescence axis, 201. Elymus
each with 1 or 2 flowers 44
44 (43). Culm internodes solid; lower glume absent on all
spikelets except the terminal one 276. Hainardia Group II – Inflorescence axis branched, spikelets
44. Culm internodes hollow; lower glume present on all without awns
spikelets 45
45 (44). Leaf sheath with falcate auricles 204. Henrardia 1. Glumes and lemmas broader than long, appearing
45. Leaf sheath lacking auricles 46 inflated 2
46 (45). Inflorescence axis not disarticulating; spikelets 1. Glumes and lemmas longer than broad, variously lan-
with 2 flowers 310. Pholiurus ceolate or oblong, but not appearing inflated 4
46. Inflorescence axis disarticulating; spikelets with 1 2 (1). Glumes longer than lemmas; flowers 1 per spikelet
flower 277. Parapholis 299. Beckmannia
47 (42). Plants annual 48 2. Glumes shorter than lemmas; flowers more than 1 per
47. Plants perennial 53 spikelet 3
48 (47). Inflorescence pectinate, the spikelets densely 3 (2). Lemma apex abruptly narrowed to an acute or
packed and stiffly divaricate 202. Eremopyrum acuminate tip 231. Chascolytrum
48. Inflorescence not pectinate, the spikelets overlapping 3. Lemma apex broad, obtuse or very broadly acute
only slightly if at all and not divaricate 49 230. Briza
49 (48). Lemma tuberculate 264. Castellia 4 (1). Plants with 1 flower per spikelet 5
49. Lemma smooth 50 4. Plants with more than 1 flower per spikelet 21
50 (49). Spikelets with 2 or more flowers; lemmas mem- 5 (4). Lemma coriaceous, noticeably harder than the
branous 51 glumes 6
50. Spikelets with 1 flower; lemmas translucent 52 5. Lemma similar in texture to the glumes, or hyaline 8
51 (50). Spikelets edgewise to the rachis, the lower glume 6 (5). Lemma glabrous, shining, dorsiventrally com-
lacking 267. Lolium pressed; most species north temperate 7
51. Spikelets broadside to the rachis, both glumes present 6. Lemma puberulent, at least near the apex, laterally
266. Festuca (former Micropyrum) compressed; plants of Argentina and Chile
52 (50). Leaf sheaths open 310. Poa 232. Relchela
52. Leaf sheaths closed 280. Mibora 7 (6). Inflorescence an open panicle; spikelets borne on
53 (47). Spikelets >1 per node, at least in the middle of the the distal half of the branches 306. Milium
inflorescence 54 7. Inflorescence a dense contracted panicle; branches
53. Spikelets 1 per node 55 spikelet-bearing to the base 227. Phalaris
54 (53). Plants rhizomatous; rachilla disarticulating, 8 (5). Inflorescence dense, cylindrical, much longer than
rachis tough broad 308. Phleum
207. Leymus (some Elymus may key here as well) 8. Inflorescence open, much branched, most spikelets
54. Plants caespitose; rachilla tough, rachis disarticulating pedicellate 9
209. Psathyrostachys 9 (8). Disarticulation below the glumes 10
55 (53). Spikelets densely packed in the inflorescence, the 9. Disarticulation above the glumes 12
internodes much shorter than the spikelets 56 10 (9). Spikelets laterally compressed 301. Cinna
55. Spikelets overlapping but largely erect, no more than 10. Spikelets globose 11
about twice the length of the internodes 58 11 (10). Leaf blades with pseudopetioles
56 (55). Glumes and lemmas firm, lanceolate 145. Phaenosperma
197. Agropyron 11. Leaf blades without pseudopetioles 238. Cyathopus
56. Glumes and lemmas membranous to hyaline, ovate 57 12 (9). Spikelets dorsiventrally compressed; plants annual
57 (56). Inflorescence with 2 bracts at the base 293. Zingeria
282. Sesleria 12. Spikelets laterally compressed; plants annual or peren-
57. Inflorescence without basal bracts 281. Oreochloa nial 13
13 (12). Glumes shorter than the flower 14 30. Lemma callus glabrous 41
13. Glumes longer than the flower 19 31 (30). Spikelets dorsiventrally compressed
14 (13). Palea with 1 vein 15 231. Chascolytrum
14. Palea with 2 veins or keels 16 31. Spikelets laterally compressed 32
15 (14). Plants erect, rhizomatous; Arctic 32 (31). Lemma apex truncate, ragged; plants of SE
314. Arctagrostis Australia 265. Dryopoa
15. Plants decumbent; New Zealand 312. Simplicia 32. Lemma apex entire or bidentate, but not truncate and
16 (14). Plants 0.5–1 m tall; leaf sheaths open ragged; plants of various localities 33
296. Aniselytron 33 (32). Leaf sheaths closed for more than ½ their length
16. Plants much less than 0.5 m tall; leaf sheaths often 34
closed >½ their length 17 33. Leaf sheaths open, or closed less than ½ their length
17 (16). Glumes less than 1/3 the length of the flower, the 35
lower glume sometimes absent; plants of the Arctic, 34 (33). Inflorescence branches spreading 286. Hyalopoa
Argentine Andes 288. Phippsia 34. Inflorescence branches upright 303. Dupontia
17. Glumes well developed; plants neither arctic nor 35 (33). Leaf blades with a prow-shaped apex, generally
Andean 18 with a translucent line on either side of the mid-vein;
18 (17). Plants caespitose, or if rhizomatous then fruit inflorescence branches generally smooth 310. Poa
lacking a beak 292. Colpodium 35. Leaf blades with apices acute or acuminate, not prow-
18. Plants rhizomatous; fruit with a small beak shaped, lacking a translucent line on either side of the
287. Paracolpodium mid-vein; inflorescence branches generally scabrous
19 (13). Palea minute or absent 233. Agrostis 36
19. Palea well developed, at least ½ the length of the 36 (35). Rachilla glabrous or scabrous 37
lemma 20 36. Rachilla pubescent 38
20 (19). Callus hairs up to 2/3 the length of the lemma; 37 (36). Leaf blades with prominent cross veins; plants of
Australia 243. “Lachnagrostis” Queensland and New South Wales 318. Sylvipoa
20. Callus hairs short, much less then 2/3 the length of the 37. Leaf blades lacking prominent cross veins; plants of
lemma; western hemisphere 245. Podagrostis various localities 226. Trisetaria
21 (4). First glume greater than ½ the length of the first 38 (36). Leaf blades less than 1 mm wide
lemma 22 300. Bellardiochloa
21. First glume ½ the length of the first lemma or less 49 38. Leaf blades more than 1 mm wide 39
22 (21). Plants annual 23 39. Plants with sturdy rhizomes 299. Arctopoa
22. Plants perennial 28 39. Plants generally caespitose, if rhizomatous then the
23 (22). Inflorescence branches disarticulating at their rhizomes short 40
bases, as well as at the bases of the pedicels and flowers 40. Apex of lemma minutely bifid, with an abaxial awn or
274. Cutandia mucro 226. Trisetaria
23. Inflorescence branches not disarticulating 24 40. Apex of lemma entire, unawned 222. Graphephorum
24 (23). Inflorescence with spikelets on short pedicels or 41 (30). Sheath margins fused; rachilla ending in a club-
short sturdy primary branches 275. Desmazeria like cluster of lemmas 187. Melica
24. Plants slender, the inflorescence much branched, with 41. Sheath margins fused less than ½ their length; rachilla
spikelets at the end of slender pedicels 25 terminating in a reduced flower, but not a cluster 42
25 (24). Glumes slightly shorter than the spikelet 26 42 (41). Callus broader than long, forming a smoothly
25. Glumes longer than the spikelet 27 rounded ridge at the base of the lemma 43
26 (25). Lemma pubescent, callus glabrous; flowers more 42. Callus about the same width as length, not obviously
than 2 in most spikelets 310. Poa smoothly rounded 45
26. Lemma glabrous, callus pubescent; flowers exactly 43 (42). Plants dioecious 263. Leucopoa
2 per spikelet 256. Molineriella 43. Plants bisexual 44
27 (25). Flowers separated by an internode 248. Antinoria 44 (43). Fruit free from the palea 266. Festuca s.s.
27. Flowers scarcely separated 229. Airopsis 44. Fruit adhering to the palea 267. Lolium s.l.
28 (22). Glumes notably dissimilar in shape, the first 45 (42). Lemma and glume margins ciliate 46
slender and linear, the second broadly oblanceolate 45. Lemma and glume margins not ciliate (although
to flabellate 225. Sphenopholis lemma often pubescent elsewhere) 47
28. Glumes similar in shape 29 46 (45). Rachilla internodes densely pubescent; plants of
29 (28). Lemma indurate, much firmer than the glumes; Australia 317. Saxipoa
puberulent; plants of Argentina and Chile 46. Rachilla internodes glabrous, scabrous, or puberulent;
232. Relchela plants of the Arctic 298. Arctopoa
29. Lemma similar in texture to the glumes; glabrous to 47 (45). Leaf blades with a translucent line on either side
pubescent; distribution of plants various 30 of the midvein; blade tip prow-like; lemma apex entire
30 (29). Lemmas with hairs on callus 31 310. Poa
47. Leaf blades lacking a translucent line; blade tip acute 1. Margins of leaf fused less than ½ their length, if at all 3
to acuminate but not curved like the prow of a boat; 2 (1). Leaf sheath forming a tube, the point of fusion of the
lemma apex often minutely bifid 48 margins not distinct 194. Bromus
48 (47). Basal leaves often with sheaths persisting and 2. Leaf sheath margins fused by a hyaline membrane
forming bulb-like structures; callus glabrous 189. Schizachne
284. Catabrosella 3 (1). Spikelets in pairs, with the outer and more obvious
48. Basal leaves not forming bulbs; callus pubescent member of the pair made up solely of sterile lemmas,
226. Trisetaria these well separated on the rachilla and stiffly divari-
49 (21). Lemma tuberculate 264. Castellia cate 261. Cynosurus
49. Lemma smooth, scabrous or pubescent, but not tuber- 3. Spikelets not paired, lacking sterile spikelets with divar-
culate 50 icate lemmas 4
50 (49). Leaf sheath margins fused more than ½ their 4 (3). Inflorescence capitate, more or less cylindrical 5
length 51 4. Inflorescence sparse or dense, if the latter then clearly
50. Leaf sheath margins free for at least ½ their length longer than broad 8
52 5 (4). Lemmas stiff, coriaceous to indurate, divided into 5
51 (50). Flowers 2 to 20 per spikelet; lemma with (5) 7 (11) to 7 stiff acuminate lobes 279. Echinaria
veins 185. Glyceria 5. Lemmas membranous to coriaceous, ending in a mucro
51. Flowers 1 to 3 per spikelet; lemma with 3 veins or single awn 6
283. Catabrosa 6 (5). Plants perennial; Australia, New Guinea, New Zeal-
52 (50). Second glume and lemmas much >1 cm long, the and 240. Echinopogon
glumes papery; leaf sheath with prominent auricles 6. Plants annual, often minute; Mediterranean and Middle
195. Littledalea East 7
52. Second glume and lemmas <1 cm long; leaf sheaths 7 (6). Spikelets with 4 to 14 flowers 260. Ammochloa
not auriculate 53 7. Spikelets with 1 flower 311. Rhizocephalus
53 (52). Inflorescence one-sided; base of style branches 8 (4). Spikelets with one flower 9
prominent and persistent in fruit 291. Sclerochloa 8. Spikelets with more than one flower 23
53. Inflorescence not one-sided; base of style branches not 9 (8). Tiny annuals; spikelets in clusters, appearing more
persistent 54 or less verticillate; glumes absent, lemmas translucent
54 (53). First glume minute, hyaline 272. Sphenopus 285. Coleanthus
54. First glume well developed 55 9. Plants perennial, or if annual, glumes present and
55 (54). Lemmas keeled, their apices generally acute, the spikelets not in clusters 10
veins easily visible, converging toward the apex 56 10 (9). Plants perennial, strongly rhizomatous; beaches in
55. Lemmas rounded on the back, their apices generally temperate regions 234. Ammophila
obtuse to truncate, the veins obscure, where visible 10. Plants annual or caespitose perennials; habitats vari-
then more or less parallel 59 ous, but not beaches 11
56 (55). Plants generally >1 m tall 265. Dryopoa 11 (10). Disarticulation below the glumes; glumes and
56. Plant generally <1 m tall 57 lemmas strongly compressed and keeled, the glumes
57 (56). Leaf blades narrow, thread-like or bristle-like often partially fused near their bases 295. Alopecurus
300. Bellardiochloa 11. Disarticulation above the glumes; glumes and lemmas
57. Leaf blades not thread-like or bristle-like 58 terete, if somewhat laterally compressed then not
58 (57). Lemma apex obtuse; apex of leaf blades not prow- strongly so, not keeled, glume bases not fused 12
like 289. Pseudosclerochloa 12 (11). Glumes awned; plants annual
58. Lemma apex acute; apex of leaf blades prow-like 233. Agrostis (former Chaetopogon)
310. Poa 12. Glumes lacking awns; plants annual or perennial 13
59 (55). Plants mat-forming; spikelets with 1 or 2 flowers; 13 (12). Callus hairs absent or less than 1/3 the length of
plants of New Zealand 312. Simplicia the lemma 14
59. Plants erect, not mat-forming; spikelets generally with 13. Callus hairs 1/3 the length of the lemma or more 21
more than 2 flowers; plants elsewhere 60 14 (13). Lemma with margins fused proximally 15
60 (59). Lemma veins generally 7 to 9, rarely 5 14. Lemma margins not fused 16
218. Torreyochloa 15 (14). Basal leaves with blades lacking or <2 cm long
60. Lemma veins generally 5 (3–7) 61 170. Patis
61 (60). Culms decumbent but not rhizomatous; plants of 15. Basal leaves with blades present, generally >2 cm
saline or alkaline habitats 290. Puccinellia 166. Piptatheropsis
61. Plant rhizomatous or stoloniferous; plants not of 16 (14). Glumes shorter than the flower 165. Ortachne
saline or alkaline habitats 292. Colpodium 16. Glumes as long as or longer than the flower 17
17 (16). Awn arising from the abaxial side of the lemma
245. Podagrostis
Group III – Inflorescence branched, spikelets 17. Awn apical or arising from the sinus between two
awned, the awn straight lemma teeth 18
18 (17). Spikelets cylindrical 19
1. Margins of leaf sheaths fused for most of their length 2 18. Spikelets laterally compressed 20
19 (18). Plants caespitose but <2 m tall, not forming 34. Callus hairs less than 2/3 the length of the flower 35
massive tussocks; plants of the New World 35 (34). Awns of adjacent flowers coiling around each
163. Anatherostipa other; callus elongate 190. Streblochaete
19. Plants to 2.5 m tall, forming massive tussocks; plants 35. Awns not coiling around each other; callus short 36
of Eurasia 154. “Neotrinia” 36 (35). Lemma awn terminal 37
20 (18). Plants over 1 m tall; lemma awn 4–9 mm long; 36. Lemma awn abaxial or from a sinus between two teeth
stamen 1 174. Anemanthele 40
20. Plants much less than 0.4 m tall; lemma awn less than 4 37 (36). Lemmas with three prominent green nerves; grain
mm long 294. Agrostopoa with a prominent indurate apical structure (“beak”)
21 (13). Callus hairs longer than the lemma 192. Diarrhena
237. “Calamagrostis” 37. Lemmas with five or more nerves, these not promi-
21. Callus hairs from 1/3 the length of the lemma to nearly nent; grain lacking a beak 38
as long, but not longer 22 38 (37). Plants of tropical Africa and Madagascar; leaf
22 (21). Lemma stiff, hardened, scabrous 239. Deyeuxia blades with obvious transverse veins
22. Lemma membranous, soft 243. Lachnagrostis 269. Pseudobromus
23 (8). Inflorescence a dense, spike-like panicle, with two 38. Plants of temperate regions worldwide, not tropical;
sterile bracts at the base 282. Sesleria leaf blades lacking obvious transverse veins 39
23. Inflorescence open or contracted, but not spike-like, 39 (38). Glumes and lemmas keeled; plants dioecious
lacking basal bracts 24 263. Leucopoa
24 (23). Spikelets borne in dense clusters at the ends of 39. Glumes and lemmas rounded on the back; plants
secondary branches 25 bisexual 266. Festuca
24. Spikelets not notably clustered, borne evenly along 40 (36). Plants with stout rhizomes; apex of leaf blade
branches 26 sharp pointed 316. Nicoraepoa
25 (24). Spikelet clusters one-sided; all spikelets in a 40. Plants generally caespitose, if rhizomatous then the
cluster fertile; plants perennial 270. Dactylis apex of the leaf blade not sharp pointed 41
25. Spikelet clusters not obviously one-sided; proximal 41 (40). Palea not tightly clasped by the lemma, clearly
spikelets in a cluster sterile; plants annual visible behind the lemma at maturity 225. Trisetaria
271. Lamarckia 41. Palea tightly clasped by the lemma, the flower thus
26 (24). Plants annual 27 opening only at anthesis, but otherwise closed 42
26. Plants perennial 31 42 (41). Spikelets with 2 or 3 flowers, 5–7 mm long; plants
27 (26). Lemma awn terminal 28 of North America 222. Graphephorum
27. Lemma awned from the sinus or from the abaxial side 42. Spikelets with 3 to 5 flowers, 7–14 mm long; plants of
29 eastern Australia 315. Hookerochloa
28 (27). Lemmas with 3 veins; inflorescence branches
disarticulating 274. Cutandia
28. Lemmas with 5 veins; inflorescence branches persis- Group IV – Inflorescence branched, spikelets
tent 266. Festuca awned, the awn curved, twisted or geniculate
29 (27). Lemma awned from the abaxial side; spikelets
with 2 flowers 257. Periballia 1. Plants sweetly scented with coumarin; spikelets with
29. Lemma awned from a short sinus; spikelets generally more than one flower, but only the distal one bisexual;
with more than 2 flowers 30 proximal flowers sterile or staminate
30 (29). Inflorescence with the proximal branches sterile 228. Anthoxanthum
307. Nephelochloa 1. Plants not scented; spikelets with one flower, or if more
30. Inflorescence with all branches spikelet-bearing than one, then proximal flowers bisexual 2
278. Vulpiella 2 (1). Disarticulation below the glumes; spikelets dis-
31 (26). Awn from the back of the lemma, with a crown of persed as a unit 3
hairs at the middle, the tip flattened and spathulate; 2. Disarticulation above the glumes; flowers dispersed
glumes hyaline, vase-shaped, noticeably longer than individually 4
the flowers 253. Corynephorus 3 (2). Tufted annuals; plants of the southern U.S. and
31. Awn terminal or from the back of the lemma, but Mexico 305. Limnodea
never with a crown of hairs or with a flattened tip; 3. Perennials with short rhizomes; plants of Siberia
glume texture various, but glumes not vase-shaped; 304. Limnas
glumes as long as or slightly shorter than the flowers 4 (2). Inflorescence dense, lanceolate; glumes linear and
32 covered with long hairs so that the entire inflorescence
32 (31) Culm internodes solid 155. Ampelodesmos appears furry 224. Lagurus
32. Culm internodes hollow 33 4. Inflorescence open or compact; glumes generally gla-
33 (32). Lemmas broader than long brous, if pubescent then the hairs not so dense as to
231. Chascolytrum (former Erianthecium) make the entire inflorescence appear furry 5
33. Lemmas longer than broad 34 5 (4). Spikelets with one fertile flower 6
34 (33). Callus hairs more than 2/3 the length of the flower 5. Spikelets with more than one fertile flower 53
243. Lachnagrostis 6 (5). Glumes shorter than the flower 7
6. Glumes as long as or longer than the flower 14 24 (20). Lemma awn subapical or from a sinus 25
7 (6). Rachilla extended beyond the distalmost flower, 24. Lemma awn terminal 41
ending blindly or in a rudiment 8 25 (24). Plants annual 297. Apera
7. Rachilla extension lacking 11 25. Plants perennial 26
8 (7). Lemmas deeply two-lobed, with a line of long hairs 26 (25). Vegetative prophylls extending out of the leaf
around the base of the lobes, awned from the sinus; sheaths 27
China and Central Asia 152. Stephanachne 26. Vegetative prophylls shorter than the leaf sheaths 28
8. Lemma apex entire or notched, lacking a line of long 27 (26). Inflorescence open, the branches spreading; awns
hairs; plants of Australia and the Pacific 9 twice geniculate 176. Celtica
9 (8). Lemma apex entire, acuminate, awned from the 27. Inflorescence narrow, the branches upright; awns once
apex; plants of Juan Fernández Island, possibly extinct geniculate 153. Macrochloa
217. Podophorus 28 (26). Callus glabrous 29
9. Lemma apex notched or ragged, not acuminate, awned 28. Callus pubescent 31
from the back; plants of Australia and New Guinea 10 29 (28). Plants rhizomatous; glume apices obtuse; plants
10 (9). Lemma apex notched, the awn abaxial, hooked; of Mongolia 157. Psammochloa
callus hairs short 235. Ancistragrostis 29. Plants caespitose; glume apices acute to acuminate;
10. Lemma apex ragged, the abaxial awn straight or genic- plants of various localities 30
ulate; callus hairs at least 1/3 the length of the flower 30 (29). Spikelets dorsiventrally compressed; awn decidu-
239. Deyeuxia ous 177. Oloptum
11 (7). Palea with two lobes 191. Triniochloa 30. Spikelets laterally compressed; awn persistent
11. Palea not lobed 12 294. Agrostopoa
12 (11). Lemma apex with a cylindrical beak, with retrorse 31 (28). Lemma glabrous or scabrous 32
spines; plants of the Himalayas 159. Orthoraphium 31. Lemma pubescent 33
12. Lemma not beaked; plants of Central and South 32 (31). Rachilla extension present; callus hairs at least 1/3
America 13 the length of the lemma 239. Deyeuxia
13 (12). Awn clearly divided into proximal and distal 32. Rachilla extension absent; callus hairs much less than
segments, once or twice geniculate; lemma much 1/3 the length of the lemma 174. Anemanthele
more firm than the glumes 182. “Jarava” 33 (31). Awn covered with hairs over its entire length, the
13. Awn not clearly divided into segments; lemma not hairs 1 mm long or more 160. Ptilagrostis
hardened 165. Ortachne 33. Awn glabrous, or pubescent only in the proximal part
14 (6). Lemma with at least three awns, a central one that 34
is apical or abaxial, and two or four slender lateral 34 (33). Lemma margins fused proximally
ones 15 166. Piptatheropsis
14. Lemma with only one awn 17 34. Lemma margins free 35
15 (14). Glumes ca. 4 cm long; major lemma awn from the 35 (33). Palea much shorter than the lemma, glabrous
sinus; plants of southeast Australia 146. Anisopogon 182. “Jarava”
15. Glumes less than 1 cm long; major lemma awn abaxial; 35. Palea about the same length as the lemma or longer,
plants of South America or the Mediterranean 16 often pubescent 36
16 (15). Palea absent or nearly so; plants of South America 36 (35). Ligules densely pubescent; proximal segment of
236. Bromidium the awn with hairs 4–9 mm long 167. Pappostipa
16. Palea present; plants Mediterranean, introduced else- 36. Ligules glabrous or sparsely pubescent; proximal seg-
where 247. Triplachne ment of awn with hairs short or lacking 37
17 (14). Lemma hyaline 18 37 (36). Lemma margins overlapping for most or all of
17. Lemma membranous to coriaceous 20 their length at maturity 38
18 (17). Glumes mucronate or awned 246. Polypogon 37. Lemma margins not overlapping 39
18. Glumes neither mucronate nor awned 19 38 (37). Plants annual 178. Stipellula
19 (18). Spikelets vase-shaped, the glumes inflated around 38. Plants perennial 173. Achnatherum s.s., 179.
the fruit and narrowed above, then flaring distally; “Timouria”, and 172. “Stillmania” (¼ “Ach-
palea present 241. Gastridium natherum” stillmanii)
19. Spikelets ovate to oblong or lanceolate, but not vase- 39 (37). Awns deciduous 171. Piptatherum
shaped; palea absent 233. Agrostis 39. Awns persistent 40
20 (17). Lemma awn abaxial, inserted at or below the mid- 40 (39). Lemma lobes <3 mm long; stigmas 2
point of the lemma 21 161. Trikeraia
20. Lemma awn terminal or from a sinus 24 40. Lemma lobes 3–6 mm long; stigmas often 3
21 (20). Lemma crustaceous at maturity 242. Hypseochloa 151. Sinochasea
21. Lemma not crustaceous, although sometimes hard- 41 (24). Palea deeply grooved, with lemma margins
ened 22 inrolled into the groove; palea apex extending beyond
22 (21). Callus hairs generally longer than the flower the lemma margins at the lemma apex
237. “Calamagrostis” 168. Piptochaetium
22. Callus hairs shorter than the flower 23 41. Palea and lemma margins flat, the latter overlapping
23 (22). Palea as long as the lemma 239. Deyeuxia or not; palea apex not extending beyond the lemma
23. Palea shorter than the lemma 243. Lachnagrostis apex 42
TRIBES AND GENERA OF POOIDEAE 207
42 (41). Flag leaf blades ca. 1 cm long or less lemma and glumes remaining on the plant at maturity
156. Oryzopsis 313. Ventenata
42. Flag leaf blades >1 cm 43 59. Curved or twisted abaxial awn on all lemmas; all
43 (42). Spikelets dorsiventrally compressed, the palea flowers falling off at maturity 60
about the same length as the lemma and fully visible 60 (59). Glumes about the same length as the lowest
171. Piptatherum lemma, but generally not extending beyond the flowers
43. Spikelets terete, the palea generally shorter than the 226. Trisetaria
lemma, partially or wholly surrounded by it 44 60. Glumes extending beyond the flowers, enclosing them
44 (43). Apex of lemma with a circular rim or “crown”, the 61
awn originating within the rim 45 61 (60). Spikelets generally much >1 cm long; ovary
44. Apex of lemma acute to acuminate, without a circular pubescent 231. Avena
rim 47 61. Spikelets generally <0.5 cm long; ovary glabrous
45 (44). Lemma margins overlapping at maturity, 250. Aira
completely concealing the palea 183. Nassella 62 (58). First glume as long as or longer than the first
45. Lemma margins only partially overlapping, the palea lemma 63
partially visible at maturity 46 62. First glume about ½ the length of the first lemma or
46 (45). Flowers large, 7.5–25 mm long 164. Hesperostipa less 71
46. Flowers generally less than 7 mm long 63 (62). Flowers more than two per spikelet 64
180. “Eriocoma” group 63. Flowers exactly two per spikelet 67
47 (44). Callus of flower obtuse, blunt 48 64 (63). Awn from a sinus between two lobes, the lobes
47. Callus of flower acute, often sharp-pointed 51 more than ¼ the length of the lemma 65
48 (47). Vegetative prophylls ending in two awns, 2–5 cm 64. Awn abaxial, the lemma apex entire or with very short
long 163. Anatherostipa lobes 66
48. Vegetative prophylls with two teeth, but not awned 49 65 (64). Ligule membranous, lacking cilia; plants of tropi-
49 (48). Spikelets laterally compressed; lemma chartac- cal Asia 150. Pseudodanthonia
eous, keeled 294. Agrostopoa 65. Ligule ciliate, ragged; plants primarily of temperate
49. Spikelets terete; lemma hardened 50 habitats, Eurasia 223. Helictotrichon
50 (49). Plants with knotty rhizomes; leaf blades with the 66 (64). Awn from about ½ way down the back of the
apices sharp–pointed 181. Amelichloa lemma; rachilla pubescent on one side only
50. Plants lacking knotty rhizomes; leaf blades with apices 252. Avenula
acuminate but not with sharp points 158. Stipa 66. Awn subapical; rachilla pubescent on all sides
51 (47). Spikelets generally >1 cm long, awns generally 226. Trisetaria
>10 cm long, plumose throughout 158. Stipa 67 (63). Lower flower staminate, with a prominent twisted
51. Spikelets <1 cm long, awns <10 cm long, if plumose awn; upper flower bisexual, with a fragile straight awn
then only on the basal segment 52 220. Arrhenatherum
52 (51). Plants not branching from the upper nodes 67. Both flowers with twisted awns, both bisexual 68
182. “Jarava” 68 (67). Panicle narrow, the branches upright; stamens
52. Plants branching from the upper nodes, the branches 2 226. Trisetaria (former Peyritschia)
stiff 175. Austrostipa 68. Panicle open, generally lax; stamens 3 69
53 (5). Disarticulation below the glumes, the spikelet 69 (68). Ligule acute 254. Deschampsia
falling entire 255. Holcus 69. Ligule obtuse to truncate 70
53. Disarticulation above the glumes, flowers falling indi- 70 (69). Ligule obtuse 251. Avenella
vidually 54 70. Ligule truncate with irregular teeth 259. Vahlodea
54 (53). Glumes aristate to long awned 55 71 (62). Leaves narrow, folded, and with a long sharp tip
54. Glumes not awned 56 266. Festuca (former Dielsiochloa)
55 (54). Lemma keeled; lodicules 2; plants of Juan Fer- 71. Leaves broad, flat, not aristate 72
nandez Island 216. Megalachne 72 (71). Callus glabrous 187. Melica
55. Lemma rounded on the back; lodicules 3; plants of 72. Callus pubescent 218. Amphibromus
Mexico 149. Metcalfia
56 (54). Lemma apex divided into four slender aristae
above the large abaxial awn 244. Pentapogon
56. Lemma apex often bilobed, but not with four aristae 57 TRIBES AND GENERA OF POOIDEAE
57 (56). Callus with hairs 2/3 the length of the lemma
243. Lachnagrostis
57. Callus glabrous or pubescent with hairs less than ½ VII. TRIBE BRACHYELYTREAE Ohwi (1941)
the length of the lemma 58 Perennials with short rhizomes. Leaf blades
58 (57). Plants annual 59
58. Plants perennial 62 broad, lanceolate. Inflorescence branches
59 (58). Second lemma with a twisted abaxial awn, other branched. Spikelets terete to dorsiventrally com-
lemmas with a terminal long slender straight awn; first pressed, with one flower, with a rachilla
Characters as for the tribe. 2n ¼ 22.

Three spp., eastern Asia, eastern North
America.
Brachyelytrum was previously placed in the
Bambusoideae on the basis of its embryo structure
(Campbell et al. 1986), but is clearly sister to all
other Pooideae. The presence of an epiblast, and
the lack of an internode between the vascular trace
to the coleoptile and that for the scutellum are
shared with all other Pooideae. The character over-
lapping embryonic leaf margins is plesiomorphic.
Campbell et al. (1986) show a clear demarcation
between the scutellum and the coleorhiza, but
others report that this is lacking (Reeder 1957);
although Campbell et al. (1986) suggest that this
character may be polymorphic, it seems equally
possible that the observation of Reeder (1957) was
in error. Campbell et al. (1986) report parallel-
sided subsidiary cells, a character that is otherwise
synapomorphic for the clade consisting of Brachy-
podieae, Bromeae, Triticeae, and Poeae.
Remainder of the subfamily:
Inflorescence phyllotaxis initially two-ranked, even
if subsequent branching makes them appear spiral
(Kellogg et al. 2013). Spikelets laterally compressed.
Embryo without a scutellar cleft, embryonic leaf
margins non-overlapping.
VIII. TRIBE NARDEAE W. D. J. Koch (1837)
Lygeeae J. Presl (1846).
Fig. 32. Pooideae-Brachyelytreae. Brachyelytrum erec- Perennials. Leaves slender, leathery. Rachilla
tum. A Flowering plant. B Adaxial leaf surface. C Flower. extension absent. Lower glume reduced to a rim,
(From Barkworth et al. 2007, drawn by L.A. Vorobik)
the upper glume absent or minute. Lodicules
absent; stigma one.
Here the two genera are combined in a single
extension. Lower glume minute or absent, upper tribe, following D€oring et al. (2007).
glume much shorter than the flower. Callus of
flower pubescent. Lemma apex entire, awned, 143. Nardus L. Fig. 33
the awn longer than the lemma. Fruit fusiform,
grooved, pubescent, beaked. Embryo with or Nardus L., Sp. Pl. 1: 53–54 (1753).
without a scutellar cleft. Microhairs lacking.
Plants caespitose. Inflorescence unbranched,
142. Brachyelytrum P. Beauv. Fig. 32 ending in a barren tip, the spikelets borne on
one side, in two rows. Spikelet dorsiventrally
Brachyelytrum P. Beauv., Ess. Agrostogr. 39, 155, pl. 9, f. compressed, with 1 flower. Upper glume minute.
2 (1812); rev.: Saarela et al. (2003). Lemma apex acute, awned. 2n ¼ 26.
One sp., N. stricta L., Europe.
Remaining tribes and genera:

Microhairs lacking.
Microhairs occur in a few Stipeae, where they
appear to be secondarily derived; otherwise all
remaining Pooideae lack them. If Stipeae diverged
before Phaenospermateae, as suggested by GPWG II
(2012), then loss of microhairs could have occurred
after the divergence of Stipeae.
IX. TRIBE PHAENOSPERMATEAE Renvoize & Clayton

(1985)
oser & J. Schneider (2011).

Duthieeae M. R€
Perennials. Ligule membranous. Lodicules 0,

2 or 3.
Eight genera, 14 spp.
Membership of this subtribe is gradually
becoming clearer as sampling has improved in
molecular phylogenetic studies. The GPWG
(2001) found a sister relationship between Aniso-
pogon and Phaenosperma, Davis and Soreng
(2007) confirmed the addition of Sinochasea and
Duthiea to the group, and D€oring et al. (2007)
found a clade including all four plus Danthonias-
trum. The thorough study of Schneider et al. (2011)
added the remaining genera listed below and dis-
cussed their placement, morphology and cytology.
Schneider et al. (2011) argue for separating Phae-
nospermateae and Duthieeae because of the very
different spikelet morphology of Duthieeae, and
because the combined tribe lacks an obvious
morphological synapomorphy. Like Soreng et al.
(2012 onward), I combine them here to avoid a
Fig. 33. Pooideae-Nardeae. Nardus stricta. A Habit. B monogeneric Phaenospermateae. In addition the
Inflorescence. C Portion of the inflorescence axis, the synapomorphies of Duthieae s.s. (i.e., without
spikelets fallen but the lower glumes retained. D Floret,
style exserted. E Anthetic floret, palea to the left, lemma at Phaenosperma) are not particularly distinctive
right. (From Cope and Gray 2009) within the context of Pooideae as a whole.
144. Lygeum Loefl. ex L. 145. Phaenosperma Munro ex Benth.

Lygeum Loefl. ex L., Gen. Pl. (ed. 5) 27, (522) (1754). Phaenosperma Munro ex Benth., J. Linn. Soc. Bot. 19: 59
(1881).
Plants with short rhizomes. Inflorescence with Euthryptochloa Cope, Kew Bull. 42(3): 707 (1987).
one spikelet, subtended by a leaf-like spathe.
Spikelet laterally compressed, with 2 to 3 flowers, Plants 1–3 m tall. Leaf blade pseudopetiolate and
their lemmas fused to form a cup or urn and resupinate, with primary veins extending at an
paleas fused to form a septum. Upper glume acute angle from the midrib. Inflorescence
absent. 2n ¼ 40. branches branched. Spikelets spherical to some-
One sp., L. spartum L., Mediterranean. what laterally or dorsiventrally compressed, with
1 flower; disarticulating below the glumes; 147. Danthoniastrum (J. Holub) J. Holub
rachilla extension lacking. Glumes shorter than
the flower, hyaline. Lemma apex entire, obtuse, Danthoniastrum (J. Holub) J. Holub, Folia Geobot. Phy-
awnless. Lodicules 3. Stigmas 2. Ovary glabrous. totax. 5: 435 (1970).
Style bases persistent in fruit as a small beak;
pericarp separable from seed. 2n ¼ 24. Inflorescence unbranched or the branches occa-
One sp., P. globosa Munro ex Benth., Assam sionally branched, generally with fewer than 5
to Japan. spikelets in the entire inflorescence. Spikelets
The resupinate leaf blades are similar to those with 4 to 9 flowers. Callus of flower with dense
of Pharus. Phaenosperma was previously placed hairs. Glumes shorter than or as long as the flow-
in Bambusoideae on the basis of embryo charac- ers, their apices attenuate, unawned. Lodicules 3.
ters, but it shares with the Pooideae the presence Stigmas 2 or 3. Ovary pubescent at the apex or all
of an epiblast and a short mesocotyl internode. over. Hilum linear. 2n ¼ 24.
The scutellar cleft and overlapping embryonic Two spp., Balkans and Caucasus.
leaf margins are plesiomorphic so not indicative The two species of Danthoniastrum were con-
of relationship. The lack of microhairs and fusoid sidered a subgenus of Helictotrichon by J. Holub
cells, and ample molecular data put Phaenos- (H. subg. Danthoniastrum Holub), whereas Clay-
perma in Pooideae. Watson and Dallwitz (1992 ton et al. (2006 onward) and Clayton and Renvoize
onward) report that Phaenosperma has ruminate (1986) placed them in synonymy under Metcalfia.
endosperm, a condition that is otherwise Neither placement is supported by the molecular
unknown in the grasses. phylogeny (Schneider et al. 2009). The leaf blades
of D. compactum disarticulate from the sheath,
Remaining genera of the tribe: and the anatomy of the disarticulation zone is
Disarticulation above the glumes. Rachilla extended described by R€oser and Heklau (2011).
beyond the distalmost flower. Lemma apex with two
elongate lobes or teeth, awned from the sinus, the
awn twisted. 148. Duthiea Hack. Fig. 34
Disarticulation above the glumes is likely to be
Duthiea Hack., Verh. K.K. Zool.-Bot. Ges. Wien 45: 200
plesiomorphic, but the pattern of gain and loss of (1896).
this character among the early-diverging tribes of
Poeae is uncertain. Inflorescence unbranched, with some pedicels
subtended by bracts. Spikelets sometimes nearly
terete, with 1 to 5 flowers. Glumes as long as or
146. Anisopogon R. Br. longer than the flowers, their apices acute. Callus
of flower pubescent. Lodicules 0, 2 or 3. Ovary
Anisopogon R. Br., Prodr. 176 (1810).
pubescent. Stigmas 2 or 3. Base of stylar branches
persistent in the fruit, forming a beak. 2n ¼ 14.
Leaf blades stiff and sharp-pointed. Ligule with a
Three spp., mountainous areas of Afghani-
ciliate margin. Inflorescence branches branched.
stan to western China.
Spikelets with one flower. Glumes longer than the
The bracts subtending the pedicels of the
flower, their apices obtuse. Callus of flower
spikelets are generally interpreted as being the
sharply pointed, pubescent. Lemma wrapped
vestiges of sterile spikelets. The persistent base
around and entirely concealing the palea, pubes-
of the stylar branches also appears in Phaenos-
cent, the lobes awn-like. Lodicules 3. Stigmas 2 or
perma and in Sclerochloa (Puccinellinae).
3. Ovary pubescent. Pericarp separable from the
seed.
Two spp., Southeastern Australia. 149. Metcalfia Conert
The single-flowered spikelet and sharp-
pointed callus led previous authors to place this Metcalfia Conert, Willdenowia 2: 417 (1960).
genus in Stipeae, but molecular data clearly place
it here in Phaenospermateae (Schneider et al. Inflorescence branches branched. Spikelets with 3
2009, 2011). or 4 flowers. Glumes as long as the flowers, their
Glumes as long as or longer than the flowers,

their apices attenuate. Callus of flower pubescent.
Lodicules 2. Stigmas 3. Ovary pubescent.
One sp., P. himalaica (Hook. f.) Bor & C. E.
Hubb., montane areas of western China and NW
Himalaya.
151. Sinochasea Keng
Sinochasea Keng, J. Wash. Acad. Sci. 48: 115, f. 1 (1958).
Plants with short rhizomes. Leaves with blades

stiff. Inflorescence branches branched. Spikelet
with one flower; rachilla extension minute or
absent on some spikelets. Glumes longer than the
flower, their apices acuminate. Callus of flower
pubescent. Lemma pubescent. Lodicules 2 or 3.
Stigmas 2 or 3. Ovary glabrous.
One sp., S. trigyna Keng, temperate Asia.
Sinochasea is the sister genus to Stepha-
nachne in the molecular study of Schneider
et al. (2011).
152. Stephanachne Keng
Stephanachne Keng, Contr. Biol. Lab. Chin. Assoc.

Advancem. Sci., Sect. Bot. 9: 134 (1934).
Inflorescence unbranched. Spikelets with one

flower. Glumes longer than the flower, their api-
ces acute to acuminate. Callus of flower pubes-
Fig. 34. Pooideae-Phaenospermateae. Duthiea oligosta- cent. Lemma with a tuft of long hairs near the
chya. A Habit. B Spikelet. C, D Glumes. E Lemma. F
Palea. G Pistil. (From Aitchison 1882)
apex, the hairs nearly as long as the slender lobes.
Lodicules 2 or 3. Stamens 1 or 3. Stigmas 2. Ovary
glabrous.
apices acute, short-awned. Callus of flower pubes- Three spp., Central Asia and western China.
cent. Lodicules 3. Stamens 2. Stigmas 2. Ovary
pubescent all over. Hilum linear. X. TRIBE STIPEAE Dumort. (1824)
One sp., M. mexicana (Scribn.) Conert, Perennials, generally caespitose but a few rhizo-
Mexico. matous, most with stiff slender leaves. Inflores-
See comments under Danthoniastrum. cence branches branched. Spikelets with one
flower, without a rachilla extension. Glumes gen-
150. Pseudodanthonia Bor & C. E. Hubb. erally longer than the flower. Lemma generally
wrapped around and often concealing the palea,
Pseudodanthonia Bor & C. E. Hubb., Kew Bull. 12: 425 pubescent, the apex entire or with two teeth;
(1958). awned from the apex, the awn straight or twisted,
often once or twice geniculate, scabrid to plu-
Leaf blades leathery. Inflorescence unbranched or mose. Lodicules 3. Anthers 3. Stigmas 2 (3, 4).
with branched branches. Spikelet with 4 to 8 flow- Embryo less than 1/3 the length of the fruit.
ers plus additional reduced flowers distally. Twenty-eight genera, 530 spp.
The one-flowered spikelet and lack of a 153. Macrochloa Kunth

rachilla extension may be synapomorphic for
this tribe, although both characters appear else- Macrochloa Kunth, Révis. Gramin. 1: 58 (1829); anat., tax.:
where in the Pooideae. Stipeae include Ampelo- Vázquez and Barkworth (2004).
Stipa subgen. Macrochloa (Kunth) Steud., Syn. Pl. Glum.
desmos, which has spikelets with several flowers, 1: 132 (1854).
which would then represent a reversal of the
tribal synapomorphy. While all genera have Plants up to 2 m tall; prophylls prominent, 1–2
pubescent lemmas, the trichomes may be in cm long, pubescent. Leaf blades hard, convolute,
lines or distributed evenly. disarticulating from the sheath; ligules velutinous.
Generic limits within this tribe are problem- Ligule a line of hairs. Glumes longer than the
atical and in a state of flux (Barkworth et al. 2008; flower, their apices attenuate. Callus of flower
Romaschenko et al. 2008, 2010, 2011, 2012). Tra- sharp. Lemma thick to hardened, pubescent, the
ditional taxonomy separated plants with short margins not overlapping, the apex with two teeth,
roundish spikelets from those with narrow elon- awned from the sinus, the awn twisted, once-
gate spikelets, but this distinction is wholly geniculate, the proximal portion plumose. Lodi-
artificial and does not indicate relationship cules 3. Hilum as long as the fruit. Midvein of leaf
(Romaschenko et al. 2011). The phylogenetic with a prominent adaxial rib, in cross section with
analyses of Romaschenko et al. (2010, 2011, a single vascular bundle at the top of the rib and
2012) place Macrochloa sister to all other mem- several pairs of bundles below it. 2n ¼ 24, 66, 72.
bers of the tribe, which is then divided into four Two spp., Mediterranean.
clades. Clade I includes an Old World subclade This genus is sister to all other Stipeae s.s.
(Stipa s.s., Ampelodesmos, Psammochloa, Ory- (Romaschenko et al. 2010, 2012). Vázquez and
zopsis s.s., Trikeraia, Orthoraphium, and Ptila- Barkworth (2004) draw attention to the long velu-
grostis, plus the New World Oryzopsis s.s.), and a tinous vegetative prophylls of this species, which
New World subclade (Aciachne, Anatherostipa, are distinctive in the context of the Stipeae. The
Hesperostipa, Lorenzochloa, Ortachne, Piptather- leaf blades of M. tenacissima (Loefl. ex L.) Kunth
opsis, Piptochaetium, and Pappostipa). Clade II disarticulate from the sheath. The anatomy of the
includes only Patis, and Clade III only Pip- disarticulation zone is described by R€oser and
tatherum s.s., now restricted to include only Heklau (2011).
Old World species, plus “Achnatherum” stillma-
nii. Clade IV, the “achnatheroid grasses”, Remainder of Stipeae:
includes Anemanthele, Austrostipa, Oloptum, Clade I, Eurasian subclade
Timouria, “Achnatherum” (including “Pip- Lemma lobes generally well developed. Long
tatherum” sect. Virescentia), and the Major cells of the lemma epidermis with sinuous walls.
American Clade (MAC), which includes the The sinuous walls on the long cells of the lemma
“Eriocoma group” of Achnatherum, plus “Jar- epidermis represent the plesiomorphic condition.
ava”, Amelichloa, and Nasella. Clade IV is distin- All genera are Eurasian except for the North Ameri-
guished by the epidermal morphology of the can Oryzopsis.
lemmas. In the ancestral lemma epidermal mor-
phology, the long cells have sinuous walls, a
condition that the authors described as “saw- 154. “Neotrinia”
like”. In the derived condition, which appears
in Clade IV, as well as in Pappostipa of Clade I, (“Neotrinia group” of “Achnatherum”)
the cells are straight-walled and isodiametric or
may be broader than long; this is the “maize- Plants robust, to 2.5 m tall, forming massive tus-
like” pattern because of its apparent similarity socks. Sheaths of basal leaves persistent, fibrous.
to the kernels on a maize ear. “Achnatherum” Glumes longer than the flower, their apices acute,
will ultimately be divided into several genera, as the lower with 1 vein, the upper with 3 veins.
indicated below. Callus of flower obtuse, pubescent. Lemma
Putative hybrid genera include Achnella densely pubescent, membranous, with two
Barkworth, a hybrid between “Achnatherum” lobes, the awn straight, deciduous. Palea extend-
and Nasella. ing slightly beyond the lemma apex. 2n ¼ 42, 48.
One sp., “Achnatherum” splendens (Trin.) awn bent, deciduous. Palea with 5 to 7 veins,
Nevski, Eurasia. similar to lemma, acute. Lodicules 3.
This species is unrelated to Achnatherum s.s. One sp., P. villosa (Trin.) Bor, Mongolia.
but is strongly supported as sister to Psammo- Psammochloa is clearly part of Stipeae, and
chloa villosa (Hamasha et al. 2012; Romaschenko sister to “Neotrinia” (Romaschenko et al. 2012;
et al. 2011, 2012). D€oring et al. 2007).
155. Ampelodesmos Link 158. Stipa L., s.s.
Ampelodesmos Link, Hort. Berol. 1: 136 (1827). Stipa L., Sp. Pl. 1: 78–79 (1753); tax.: Gonzalo et al. (2012,
2013); Vázquez Pardo and Gutiérrez Esteban (2011).
Plants robust, to 3.5 m tall, rhizomatous. Culm
solid. Leaf bases becoming hard and curved. Spi- Plants caespitose. Leaves often with blades con-
kelets with 2 to 6 flowers, with a rachilla exten- volute or involute. Glumes acuminate, much
sion, the rachilla pubescent. Glumes shorter than longer than the flower, with 3 to 7 veins. Callus
the flowers, their apices attenuate. Lemma apex of flower long, sharp, with the tip oblique or
with two teeth, awned or mucronate from the slightly curved to a sharp point, generally with
sinus. Awn twisted, not geniculate. Lodicules 3, long hairs. Lemma with the margins overlap-
vascularized. Ovary apex pubescent. 2n ¼ 48. ping, the apex lacking lobes, the awn once or
One sp., A. mauritanicus (Poir.) T. Durand & twice geniculate, the column twisted. Palea as
Schinz, Mediterranean. long as the lemma or slightly longer, the apex
The spikelets with multiple flowers have been somewhat wrinkled. Lodicules 3. Stigmas 2 or 3
used to exclude this species from Stipeae, but (4). 2n ¼ 44.
molecular data are unequivocal linking it with One hundred ten spp., Eurasia and North
Psammochloa and “Neotrinia” (Romaschenko Africa.
et al. 2012). The spikelet number thus appears Stipa s.l. has been dismembered over the last
to be a reversal. decade, but remains a large genus. Stipa s.s. now
includes sections Barbatae, Leiostipa, Pseudopti-
lagrostis, Regelia, Stipa, and Smirnowia. In its
156. Oryzopsis Michx. strict sense, the genus is monophyletic (Hamasha
et al. 2012; Romaschenko et al. 2012), but sub-
Oryzopsis Michx., Fl. Bor.-Amer. 1: 51 (1803).
divisions within the genus are not consistently
supported by available molecular data.
Inflorescence branches branched. Spikelets terete to
laterally compressed. Glumes shorter than or equal ORTHORAPHIUM + PTILAGROSTIS + TRIKERAIA
to the flower, not hardened, with 6 to 10 veins, their
apices mucronate. Callus of flower pubescent with a These three genera form a clade
ring of hairs 1 mm long. Lemma leathery, pubes- (Romaschenko et al. 2010, 2012). Hamasha et al.
cent, the margins overlapping and slightly involute, (2012) identified a similar group, although with
the apex lobed, awned from the sinus, the awn two species currently classified in Stipa plus two
straight, deciduous. 2n ¼ 46. from “Achnatherum”.
One sp., O. asperifolia Michx., North America.
159. Orthoraphium Nees
157. Psammochloa Hitchc.
Orthoraphium Nees, Proc. Linn. Soc. Lond. 1: 94 (1841).
Psammochloa Hitchc., J. Wash. Acad. Sci. 17: 140 (1927).
Plants caespitose. Glumes shorter than the flower,
Plant robust, rhizomatous. Leaf blades flat. their apices acute to obtuse. Callus of flower
Glumes about the same length as the flower, obtuse, pubescent. Lemma papery, margins over-
their apices obtuse. Callus of flower obtuse, gla- lapping only at the apex, the apex with a cylindri-
brous. Lemma not enclosing the palea, the apex cal beak, with stout retrorse spines, awned, the
with two short teeth, awned from the sinus, the awn straight, persistent. Lodicules 3.
One sp., O. roylei Nees, Himalayas above 2700

m altitude.
160. Ptilagrostis Griseb., s.s. Fig. 35
Ptilagrostis Griseb., Fl. Ross. 4: 447 (1852).
Plants caespitose. Leaf blades filiform, convolute.

Glumes longer than the flower, their apices
obtuse to acute. Callus of flower blunt, pubescent.
Lemma papery, the margins not overlapping, the
apex with two teeth, awned from the sinus, the
awn once- or twice-geniculate, covered with hairs
for its entire length. Palea pubescent. Lodicules 3.
Ovary glabrous. Stigmas 2. 2n ¼ 22.
Eight spp., mountains of Asia.
Ptilagrostis is probably monophyletic as long as
“P”. kingii is excluded (Romaschenko et al. 2010).
161. Trikeraia Bor
Trikeraia Bor, Kew Bull. 9: 555, f. s.n. (1954).
Plants robust, rhizomatous. Leaf blades rolled.

Glumes longer than the flower, their apices acu-
minate. Spikelets weakly dorsiventrally com-
pressed. Callus of flower obtuse, pubescent.
Lemma covering the edges of the palea, with two
short slender lobes flanking a weakly geniculate
awn, the awn persistent. Lodicules 2 or 3.
Three spp., Tibet and Himalayas.
Clade I, New World subclade
Indurate lemmas.
This clade has been identified by several
molecular phylogenetic studies, although support
varies (Cialdella et al. 2010; Romaschenko et al.
2010, 2012). Cialdella et al. (2010) suggest that the
presence of indurate lemmas may be synapo-
morphic for this clade, although the character
reverses in Anatherostipa and Ortachne.
162. Aciachne Benth.
Aciachne Benth., Hooker’s Icon. Pl. 14, t. 1362 (1881).
Fig. 35. Pooideae-Stipeae. Ptilagrostis pelliotii. A Habit.

B Ligule of basal leaf. C Ligule of culm leaf. D Spikelet. E
Glumes. F Flower. G Anther. (From Wu et al. 2007, p. 279,
with permission from the Missouri Botanical Garden
Press, St. Louis, and Science Press, Beijing)
Cushion plants. Leaf blades narrow and needle- Five spp., North America and Mexico.
like, with a sharp point. Inflorescence no taller Lemma epidermal morphology of Hespero-
than leaves, with only 1 or 2 spikelets. Glumes, stipa is similar to that of the fossil Berriochloa,
lemma and palea indurate. Flower cylindrical, from the late Miocene (Thomasson 1978, 1985).
somewhat gibbous. Lemma with a long point.
Three spp., Costa Rica to Argentina and 165. Ortachne Nees ex Steud.
Chile.
Romaschenko et al. (2010, 2012) find Ortachne Nees ex Steud., Syn. Pl. Glumac. 1: 121 (1854).
Aciachne to be monophyletic in their molecular
phylogenetic analysis, whereas Cialdella et al. Leaves filiform. Spikelets cylindrical. Glumes
(2010) suggest that it may be poly- or paraphy- shorter than the flower. Callus of flower blunt,
letic. pubescent. Lemma not hardened, awned, the
awn not clearly differentiated into a base and
163. Anatherostipa (Hack. ex Kuntze) Peñailillo tip. Lodicules 2 or 3.
Two spp., Costa Rica to montane South
Anatherostipa (Hack. ex Kuntze) Peñailillo, Gayana, Bot. America.
53: 277 (1996).
Stipa sect. Anatherostipa Hack. ex Kuntze, Revis. Gen. Pl.
3(3): 372 (1898). 166. Piptatheropsis Romasch., P. M. Peterson &
Stipa sect. Obtusae Parodi, Revista Argent. Agron. 17(3): Soreng
201 (1950), invalid.
Lorenzochloa Reeder & C. Reeder, Bol. Soc. Argent. Bot. Piptatheropsis Romasch., P. M. Peterson & Soreng, Taxon
11(4): 239 (1969). 60: 1712 (2011).
Leaf blades narrow, filiform, convolute. Glumes Spikelets dorsiventrally compressed or terete.
papery, as long as or longer than the flower. Glumes as long as or longer than the flower,
Callus of flower blunt. Lemma margins not with 1 to 3 veins. Callus of flower blunt, glabrous
inrolled, not covering the palea, the apex awned, or pubescent. Lemma leathery, pubescent, lobed
the awn straight or curved or reduced to a mucro. at the apex, with margins fused below, the awn
Lodicules 2 or 3. Hilum linear. persistent and straight or deciduous and twisted.
Nine spp., New World. Lodicules 3. Stigmas 2. Hilum linear. 2n ¼ 20,
Lorenzochloa is placed within Anatherostipa 22, 24.
by molecular data (Romaschenko et al. 2010, Five spp., North America.
2012). Plastid sequences hint that Anatherostipa Romaschenko et al. (2010) describe the fused
may be paraphyletic, with Aciachne derived from portion of the lemma margins as a “small fleshy
within it, but the phylogeny is not well supported wart”.
(Romaschenko et al. 2012). The two genera are
kept separate here pending additional data.
167. Pappostipa (Speg.) Romasch., P. M. Peterson &
Soreng
164. Hesperostipa (M. K. Elias) Barkworth
Pappostipa (Speg.) Romasch., P. M. Peterson & Soreng, J.
Hesperostipa (M. K. Elias) Barkworth, Phytologia 74: 15 Bot. Res. Inst. Texas 2: 181 (2008).
(1993).
Leaves with apices acute, sometimes sharp;
Glumes extending to a long slender tip. Flower ligules densely pubescent. Glumes longer than
cylindrical, narrow, unusually large, 7.5–25 mm the flower. Flower terete, fusiform, the callus
long, the callus sharp-pointed, with dense hairs. obtuse to truncate, pilose. Lemma with long
Lemma hard, the margins flat and slightly over- hairs at the apex, with two short lobes, lacking a
lapping, the apex without lobes, with a papillose, crown-like rim; awn once geniculate, plumose.
ciliate crown, awned, the awn twice geniculate. Palea about the same length as the lemma or
Palea pubescent. Fruit fusiform. 2n ¼ 38, 44, 46. shorter. Lodicules 3.
Thirty-one spp., South America, SW United Plants caespitose or short rhizomatous. Basal
States and northern Mexico. leaves with blades lacking or less than 2 cm long;
Members of this genus have lemma epidermal cauline leaves with blades >10 cm long and >5
cells that are straight-walled and nearly isodia- mm wide. Spikelets dorsiventrally compressed or
metric or broader than long (Romaschenko et al. terete. Glumes shorter than to much longer than
2008). This lemma epidermal pattern appears in the flower, with transverse veins. Callus of flower
parallel in Clade IV. Discrepancies between gene obtuse. Lemma with margins overlapping, fused
trees led Romaschenko et al. (2012) to suggest below, the awn persistent, straight. Lodicules 3.
that Pappostipa may be the product of ancient Stigmas 2. 2n ¼ 46, 48.
allopolyploidy. Three spp., eastern Asia and eastern North
America.
Patis was recircumscribed by Romaschenko
168. Piptochaetium J. Presl
et al. (2011) and now includes species placed by
Piptochaetium J. Presl, Reliq. Haenk. 1: 222 (1830), nom. previous authors in Oryzopsis or Piptatherum. It
conserv.; Cialdella et al. (2007, 2010), phylog., key. is not closely related to any other genus in the
tribe (Romaschenko et al. 2012). The short rhi-
Glumes longer than the flower. Callus of flower zomes may be uniquely derived here.
blunt or sharp, glabrous or pubescent. Lemma
Clade III
apex forming a firm crown-like rim, awned, the
awn deciduous or persistent, twice geniculate and
twisted. Palea grooved, extending beyond the 171. Piptatherum P. Beauv., s.s.
lemma; lemma margins involute and fitting into
the palea groove. Lodicules 2 or 3. Stigmas 2. 2n ¼ Piptatherum P. Beauv., s.s., Ess. Agrostogr. 17, 173 (1812).
28, 42.
Thirty-five spp., temperate North and South Plants caespitose or rhizomatous. Spikelets dorsi-
America. ventrally compressed. Glumes with 3 to 9 veins,
longer than the flower. Callus of flower blunt,
flattened, glabrous. Lemma leathery, the margins
169. “Ptilagrostis” kingii (Bol.) Barkworth not fused, the awn straight, deciduous. Palea visi-
ble proximally, not enclosed by the lemma at the
“Ptilagrostis” kingii (Bol.) Barkworth
base. Lodicules 3. Stigmas 2, exserted laterally.
Hilum linear. 2n ¼ 24.
Leaves with blades convolute, their apices stiff.
Thirty-two spp., Eurasian.
Glumes slightly longer than the flower, their api-
As traditionally circumscribed, Piptatherum
ces rounded to acute. Callus of flower blunt,
is polyphyletic (Romaschenko et al. 2010). The
pubescent. Lemma membranous, with margins
delimitation and description followed here are
not overlapping, the awn once or twice genicu-
based on Romaschenko et al. (2011), who provide
late, persistent, scabrous. Lodicules 3. Stigmas 2.
a detailed description of the taxonomic history of
2n ¼ 22.
the group and the reasons behind the problemat-
One sp., “P.” kingii (Bol.) Barkworth, Sierra
ical classifications. Piptatherum s.s. excludes the
Nevada, California.
former sections Miliacea and Virescentia, which
Although Romaschenko et al. (2011, 2012)
both belong in the achnatheroid grasses on the
place “P.” kingii sister to Piptatheropsis, support
basis of their basally fused lemma margins,
for this is weak. There is little morphological
lemma epidermal micromorphology, and DNA
similarity between the two.
sequences. Hamasha et al. (2012) also identify a
Clade II monophyletic Piptatherum clade.
170. Patis Ohwi 172. “Stillmania”
Patis Ohwi, Actta Phytotax. Geobot. 11: 181 (1942). “Achnatherum” stillmanii (Bol.) Barkworth
Plants 0.6–1.5 m tall, with short rhizomes. Glumes flower short, obtuse, pubescent. Lemma margins
longer than the flower, each with 1 vein. Callus of convolute, not covering the palea, the awn
flower blunt. Lemma pubescent, the margins straight, deciduous. Stamen 1. 2n ¼ 40–44.
overlapping for most or all of their length, the One sp., A. lessoniana (Steud.) Veldkamp,
apex with distinct slender lobes, awned, the awn New Zealand.
once or twice geniculate. Palea about as long as or
longer than the lemma, the keels extended to
175. Austrostipa S. W. L. Jacobs & J. Everett
about the length of the lemma lobes.
One sp., Achnatherum stillmanii (Bol.) Bark- Austrostipa S. W. L. Jacobs & J. Everett, Telopea 6: 582
worth, California. (1996); tax., micromorphology: Bustam (2010).
This species is placed sister to Piptatherum s.
s. by the analyses of Romaschenko et al. (2012). Plants often with knotty bases; culms branching
from the upper nodes. Glumes with apices acute
Clade IV “achnatheroid grasses” sensu
to acuminate, rarely muticous or mucronate. Cal-
Romaschenko et al. (2012)
lus of flower long, sharp-pointed, pubescent.
Lemma epidermis with cells generally isodia-
Lemma tough, dark at maturity, often with
metric or broader than long.
brownish pubescence, the margins generally con-
Clade IVA. Core achnatheroid subclade volute, rarely involute, the apex sometimes with
Glumes longer than the flower. two minute lobes, awned, the awn once or twice
geniculate. Lodicules 2 or 3.
Sixty-three spp., Australasia, several intro-
173. Achnatherum P. Beauv. s.s. duced elsewhere.
Achnatherum P. Beauv. s.s., Ess. Agrostogr. 19, 146, pl. 6,
f. 7 (1812). 176. Celtica F. M. Vázquez & Barkworth
Aristella Bertol., Fl. Ital. [Bertoloni] I. 690 (1833).
Piptatherum sect. Virescentia Roshev. ex Freitag, Notes Celtica F. M. Vázquez & Barkworth, Bot. J. Linn. Soc. 144:
Royal Bot. Gard., Edinburgh 33: 363 (1975). 491 (2004).
Plants caespitose. Spikelets fusiform. Glumes Plants to 2.5 m tall. Glumes with apices acuminate
with apices acute to acuminate. Callus of flower or caudate. Callus of flower sharp-pointed,
acute to obtuse, pubescent. Lemma with margins pubescent. Lemma leathery, the margins flat,
not overlapping, the apex without a ciliate crown- overlapping, the apex with two teeth, awned
like rim; awn persistent, not strongly twisted, from the sinus, the awn twisted and twice genicu-
straight, flexuous or once geniculate. 2n ¼ 28, 36. late. Palea veins extended into awns. Lodicules 3.
Twenty-one spp., Asia. Stigmas 2. Hilum as long as the fruit. 2n ¼ 96.
“Achnatherum” in the sense of Flora of China One sp., C. gigantea (Link) F. M. Vázquez &
(Wu and Phillips 2006) is polyphyletic Barkworth, Iberian Peninsula and northern
(Romaschenko et al. 2010). Achnatherum s.s. Africa, cultivated elsewhere.
includes only sections Achnatherum, Achnather- Vázquez and Barkworth (2004) note that Cel-
opsis, and Aristella (Romaschenko et al. 2011, tica has unusually long prophylls (14–15 cm) that
2012). Hamasha et al. (2012) suggest recognizing terminate in two equally long teeth. This charac-
sect. Aristella as a separate genus, but acknowl- ter is rarely reported in species descriptions of
edge that it may equally well be accommodated grasses, but may be diagnostic. The high chromo-
within Achnatherum s.s., as done here. some number of this species points to a possible
allopolyploid origin. This species is cultivated, as
174. Anemanthele Veldkamp Stipa gigantea Link.
Anemanthele Veldkamp, Acta Bot. Neerl. 34: 107 (1985).

177. Oloptum M. R€oser & H. R. Hamasha
Plants over 1 m tall. Spikelets laterally com- oser & H. R. Hamasha, Pl. Syst. Evol. (2012).
Oloptum M. R€
pressed. Glumes with apices acute. Callus of
Plants 1–1.5 m tall. Ligule membranous. Leaf This clade is strongly supported by chloro-
blades stiff, scaberulous. Spikelets dorsiventrally plast DNA sequences but not by nuclear ribo-
compressed. Glumes with apices acuminate. Cal- somal DNA (Cialdella et al. 2014; Romaschenko
lus of flower glabrous. Lemma coriaceous, dark et al. 2012). Relationships within the clade are
brown, shiny, with central vein prominent, the unclear as are the delimitation of groups within
basal margins fused, the awn flexuous, deciduous. it. One possibility would be to include all mem-
Anther apices minutely pubescent. 2n ¼ 24. bers of the clade in a single large genus.
One sp., Oloptum miliaceum (L.) R€oser & H.
R. Hamasha, worldwide.
The genus Oloptum was erected to accommo- 180. “Eriocoma group”
date the taxa of the Miliacea group of “Pip-
tatherum”, which is clearly unrelated to Spikelets fusiform or obovoid. Lemma with mar-
Piptatherum s.s. (Hamasha et al. 2012; gins only partially overlapping, the apex lacking
Romaschenko et al. 2011, 2012). lobes, with a ciliate crown, the awn persistent or
deciduous. Palea short.
Twenty-nine spp., New World.
178. Stipellula M. R€oser
The “Eriocoma group” includes species for-
Stipellula M. R€oser, Schlechtendalia 4: 91–93 (2012). merly placed in Achnatherum; these form a clade
Stipella (Tzvelev) M. R€oser & H. R. Hamasha, nom. illegit. separate from Achnatherum s.s. (Romaschenko
Stipa sect. Stipella Tzvelev, Novosti Sist. Vyssh. Rast. 11: et al. 2012).
15 (1974).
Annuals. Glumes with apices acuminate. Callus of 181. Amelichloa Arriaga & Barkworth
flower sharp-pointed. Lemma hard, constricted
below the apex, with lines of hairs, with margins Amelichloa Arriaga & Barkworth, Sida 22: 146 (2006).
overlapping, the apex awned; awn twice genicu-
late, twisted, the proximal portions pubescent, Plants with knotty rhizomes. Leaf blades stiff,
the distal portion glabrous. Palea much shorter inrolled, sharp-pointed. Ligule ciliate. Cleistoga-
than the lemma. Lodicules 2. 2n ¼ 36. mous spikelets often present in upper leaf
Two spp., Mediterranean. sheaths. Spikelets terete. Glumes longer than the
Stipellula is clearly unrelated to Stipa s.s. The flower, their apices acute to acuminate. Callus of
two species belong in the achnatheroid grasses on flower pubescent, blunt. Lemma pubescent,
the basis of lemma epidermal pattern and molec- awned, the awn twisted. Palea pubescent. Lodi-
ular data (Romaschenko et al. 2011). cules 3.
Five spp., South America, Mexico.
Clade IVB It is unclear whether Amelichloa is sister to
Nasella or embedded within it. The genus is ten-
179. Timouria Roshev. tatively recognized here, following Romaschenko
et al. (2012), although data presented by Cialdella
Timouria Roshev., Fl. Aziatsk. Ross. 2: 173, pl. 12 (1916). et al. (2014) argue for combining the two genera.
Leaf blades setaceous. Glumes with 3 veins. Callus

of flower short, obtuse. Lemma short (3–4 mm 182. “Jarava” Ruiz & Pav.
long), densely pubescent, the margins overlap-
ping, the apex lacking a ciliate crown, with apical “Jarava” Ruiz & Pav., Fl. Peruv. Prodr. 2 (1794).
lobes, the awn deciduous.
Five spp., Asia. Leaf blades convolute, their apices pointed.
See discussion in Romaschenko et al. (2011) Glumes shorter than to longer than the flower.
for species included here. Callus of flower with long white hairs. Lemma
with margins not overlapping at maturity, the
Clade IVC. Major American Clade apex minutely bilobed, bearing long hairs, the
Plants of the New World. awn terminal, deciduous or persistent, once or
twice geniculate. Palea much shorter than the length. Ovary glabrous. Stigmas and style
lemma, glabrous. Lodicules 2 or 3. branches 2, free to the base, the bases persistent.
Thirty spp., South America. Eight genera, 163 spp.
Monophyly of “Jarava” is uncertain; it may The sister taxon relationship between Brylk-
be paraphyletic with Nassella and Amelichloa inia caudata (formerly placed in its own tribe)
derived from within it (Romaschenko et al. and Meliceae has been shown by Schneider
2012). In addition, a group of at least seven spe- et al. (2009, 2011). The closed sheath margins
cies of “Achnatherum”, the “Pseudoeriocoma are synapomorphic. The lodicules of Brylkinia
group”, apparently fall within “Jarava” s.l. are thick and fused proximally, although they
are membranous and free distally. Previous
reports suggesting that they are entirely free
183. Nassella (Trin.) E. Desv.
are in error.
Nassella (Trin.) E. Desv., Fl. Chil. 6: 263 (1854); tax.:
Barkworth (1990); phylog.: Cialdella et al. (2014). 184. Brylkinia F. Schmidt
Spikelets obovoid or terete. Glumes longer than Brylkinia F. Schmidt, Mém. Acad. Imp. Sci. Saint Péters-
the flower, their apices awned or awnless. Callus bourg VII, 12: 199 (1868).
of flower blunt or sharp, glabrous or pubescent.
Lemma hardened, the margins convolute and Rhizomatous perennials. Inflorescence
completely wrapped around the palea and fruit, unbranched. Spikelets with two sterile flowers
the apex bearing a crown-like rim, the awn termi- proximally, and one bisexual flower distally, dis-
nal, deciduous or persistent, twice geniculate. articulating at the base of the pedicel. Lower
Palea reduced, without veins. Lodicules 2 or 3. glume acuminate, the upper narrowing to an
Anthers 1 or 3. 2n ¼ 28, 34, 36, 40, 42, 64, 70, attenuate apex. Lemma of sterile flower acumi-
82, 88. nate; lemma of bisexual flower with two tiny
One hundred seventeen spp., throughout the hyaline lobes, awned from the sinus, the awn
Western Hemisphere. straight, as long as or longer than the lemma.
Barkworth (1990) notes that Nassella has an Lodicules membranous and free distally, thick
unusually long epiblast on the embryo, a charac- and fused proximally. 2n ¼ 40.
ter that it shares with Hesperostipa, some species One sp., B. caudata (Munro) F. Schmidt,
of Achnatherum and two of Piptochaetium. China and Japan.
Romaschenko et al. (2012) observe that the
Remaining genera of the tribe:
lemma epidermis of Nassella is unique, with
Lodicules lacking a distal membranous portion. Spi-
short cells lacking silica bodies and “long” cells
kelets ending in a reduced flower or flower-like clus-
wider than long, forming a ladder-like pattern.
ter of lemmas.
Cialdella et al. (2014) show that members of
Nasella fall into two clades, one of which is largely
185. Glyceria R. Br. Fig. 36
restricted to high elevations.
Glyceria R. Br., Prodr.: 179 (1810), nom. conserv.; tax.,
XI. TRIBE MELICEAE Rchb. (1828) phylog., Whipple et al. (2007).
Brylkinieae Tateoka (1960). Perennials, rarely annual, caespitose, rhizoma-

tous. Inflorescence branches branched. Spikelets
Annuals or more often perennials. Sheath mar- with 2 to 20 flowers, laterally compressed or sub-
gins fused, often joined by a hyaline membrane. terete. Glumes shorter than the spikelet, their
Inflorescence branched or unbranched. Glumes apices obtuse to acute. Lemmas with 5 to 11
generally shorter than the adjacent lemmas. veins, these generally prominent and not converg-
Lemma membranous below, papery distally, ing at the apex. Stamens 2 or 3. 2n ¼ 20, 40, 60.
veins generally not converging at the apex. Lodi- Forty-eight spp., wet sites in temperate
cules thick, truncate, fused for part of all of their regions worldwide.
Lemma with 11 rib-like veins, the apex with two

teeth, with an abaxial awn.
One sp., L. avenacea Samuelsson, Syria.
187. Melica L.
Melica L., Sp. Pl. 1: 66–67 (1753); Hempel (2011), rev.
Perennials, caespitose or with short rhizomes,

sometimes with the lowest culm internode enlarged
and bulb-like. Inflorescence branches branched.
Spikelets laterally or dorsiventrally compressed,
with 1 to 8 flowers, disarticulating above or below
the glumes, the rachilla terminating in a club-like
cluster of lemmas. Glumes shorter or longer than
the spikelet, membranous to papery, the apices
translucent, obtuse to acuminate. Callus of flower
glabrous. Lemma muticous to awned. 2n ¼ 18, 36.
Ninety spp., worldwide, temperate areas.
Disarticulation below the glumes occurs only
in species with sharply bent pedicels.
188. Pleuropogon R. Br.
Pleuropogon R. Br., Chlor. Melvill.: 31–32, pl. D (1823).

Inflorescence unbranched. Spikelets laterally com-
pressed, with 5 to 20 flowers. Glumes shorter than
the flowers, hyaline, their apices ragged. Callus of
flower glabrous. Lemma apex ragged or toothed,
sometimes awned, the awn terminal or from a
sinus. Palea keels awned. 2n ¼ 16, 32, 36, 40, 42.
Six spp., wet areas of the Arctic, Pacific Coast
of North America south to California.
Fig. 36. Pooideae-Meliceae. Glyceria multiflora. A Culm 189. Schizachne Hack.

with inflorescence. B Spikelet. C Glumes. D Lemma.
E Palea and rachilla. F Fused lodicules. G Caryopsis, Schizachne Hack., Repert. Spec. Nov. Regni Veg. 7: 322
showing embryo. H Caryopsis, showing hilum. (From (1909).
Zuloaga et al. 2012)
Caespitose perennials. Inflorescence unbranched,

186. Lycochloa Samuelsson or with branched branches, sparse. Spikelets lat-
erally compressed, with 3 to 6 flowers. Glumes
Lycochloa Samuelsson, Ark. Bot. 25(8): 4 (1933). shorter than the spikelet, papery proximally, hya-
line distally, their apices acute. Callus of flower
Perennials. Inflorescence unbranched. Spikelets densely pubescent. Lemma apex bilobed, awned
laterally compressed, with 2 flowers. Glumes from the sinus, the awn straight. 2n ¼ 20.
shorter than the spikelet, papery, their apices One sp., S. purpurascens (Torr.) Swallen,
acuminate. Callus of flower densely pubescent. North America, north temperate Asia.
The leaf sheath margins are fused by a hyaline-

membrane that appears to disintegrate eventually.
190. Streblochaete Hochst. ex Pilg.
Streblochaete Hochst. ex Pilg., Bot. Jahrb. Syst. 37, Beibl.

85: 61 (1906).
Perennials. Inflorescence branches branched.

Spikelets laterally compressed, with 2 to 6 flow-
ers. Glumes shorter than the spikelet, their apices
acute. Callus of flower sharp-pointed, pubescent.
Lemma with 7 veins, with two apical teeth and an
abaxial awn; awns of adjacent flowers coiling
around each other. Lodicules not fused. 2n ¼ 20.
One sp., S. longiaristata (A. Rich.) Pilg., trop-
ical Africa, Réunion, southeast Asia.
Streblochaete is strongly supported as being
part of Meliceae, but its precise position within
the tribe is not certain (Schneider et al. 2011). The
lack of fusion in the locidules is probably a rever-
sal, but might prove to be plesiomorphic.
191. Triniochloa Hitchc.
Triniochloa Hitchc., Contr. U.S. Natl. Herb. 17: 303 (1913).
Perennials, caespitose or rhizomatous. Inflores- Fig. 37. Pooideae-Diarrheneae. Diarrhena obovata. A

cence branches branched. Spikelets with one Plant. B Spikelet. C Flower. D Caryopsis. (From Bark-
flower. Callus of flower pubescent. Glumes worth et al. 2007, drawn by H. Paźdirková)
shorter than or as long as the spikelet, their apices
acute to acuminate. Lemma with two slender
lobes, awned; the awn abaxial or from the sinus. One genus, five spp.
Palea with two slender lobes. 2n ¼ 32. The embryo of Diarrhena has been described
Six spp., Mexico, Central and South America. as lacking (Reeder 1957) or having (Macfarlane
Triniochloa is sister to Glyceria (Schneider and Watson 1980) a scutellar cleft, and embry-
et al. 2011), indicating that the one-flowered onic leaf margins meeting (Reeder 1957) or over-
spikelet is derived and synapomorphic here. lapping (Macfarlane and Watson 1980). The
genus may be polymorphic, or some of the obser-
XII. TRIBE DIARRHENEAE C.S. Campb. (1985) vations may be erroneous.
Perennials, rhizomatous or loosely clumped;

culm nodes concealed by leaf sheaths. Leaf blades 192. Diarrhena P. Beauv. Fig. 37
glossy, narrowing toward the ligule, the midvein Diarrhena P. Beauv., Ess. Agrostogr.: 142, 160, 162 (1812),
usually excentric. Inflorescence branches branched. nom. conserv.; rev.: Brandenburg et al. (1991b).
Spikelets laterally compressed, with 2 to 7 flowers, Neomolinia Honda, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 3:
the distal ones reduced and sterile. Glumes shorter 110 (1930).
than the flowers, their apices acute. Callus of flower
glabrous, rounded. Lemma with three prominent Characters as for the tribe.
veins, the apex acute, sometimes mucronate. Five spp., eastern Asia, eastern North America.
Stamens generally 2. Fruit apex with an indurate The East Asian species form a monophyletic
beak, pericarp free from seed. 2n ¼ 38, 60. group (Schneider et al. 2009) and are sometimes
segregated as Neomolinia, but the clade is clearly

monophyletic and there seems little reason to
divide this tiny genus into two.
Remaining genera of the subfamily:
Parallel-sided subsidiary cells.
XIII. TRIBE BRACHYPODIEAE Harz (1880)

Inflorescence unbranched. Spikelets laterally com-
pressed, with (3) 5 to 24 flowers, the distal flowers
reduced. Glumes shorter than the flowers, their
apices obtuse to acute or acuminate. Callus of flower
glabrous. Lemma veins not converging toward the
apex, the apex obtuse to acute, awned or not. Ovary
apex pubescent. 2n ¼ 10, 14, 16, 18, 20, 28, 30.
One genus, 16 spp.
193. Brachypodium P. Beauv. Fig. 38
Brachypodium P. Beauv., Ess. Agrostogr.: 100, 15, pl. 19, f.

35 (1812); phylog.; Catalán and Olmstead (2000), Catalán
et al. (2012).
Characters as for the tribe.

Sixteen spp., temperate regions of Eurasia,
Mexico, Central and South America.
The annual species Brachypodium distachyon
has become a valuable model system for genomic
and genetic research. Its tiny stature and small
genome make it a more tractable study system
than others in Triticeae and Poeae (Mur et al.
2011). Plants previously classified as B. distach-
yon have 10, 20 or 30 pairs of chromosomes;
while these had been treated as chromosomal
races, they now appear to constitute three sepa-
rate species (Catalán et al. 2012).
Remaining genera of the subfamily:
Lemma with veins converging distally. Chromosome
base number 7; chromosomes large. Genomes large.
See data and discussion of genomic synapo-
morphies in Bennetzen and Kellogg (1997) and Kel-
logg and Bennetzen (2004).
BROMEAE + TRITICEAE
Apex of ovary pubescent.
Fig. 38. Pooideae-Brachypodieae. Brachypodium dis-

tachyon. A Plant. B Inflorescence. C Adaxial leaf blade
surface. D Leaf sheath, blade, and ligule. E Culm node. F Stamen. (From Barkworth et al. 2007, drawn by C. Roché
Spikelet. G Flower, abaxial side. H Flower, adaxial side. I and A. Miller)
XIV. TRIBE BROMEAE Dumort. (1824)

Annuals or perennials, caespitose, sometimes rhi-
zomatous. Inflorescence unbranched, or with
branches branched. Spikelets generally laterally
compressed (sometimes terete), with 2 or more
flowers, the distal ones progressively reduced;
rachilla extension present. Glumes shorter than
the flowers. Callus of flower glabrous. Lemma
apex entire or with two teeth or lobes, awned
from the sinus or abaxially just below the apex,
the awn generally straight.
Two genera, one hundred sixty-two species.
Inclusion of Littledalea here renders Bromeae
paraphyletic (Davis and Soreng 2007; Schneider
et al. 2009). Ultimately either Littledalea should
be placed in its own tribe, or Bromeae as recog-
nized here should be incorporated in Triticeae as
recommended by Schneider et al. (2009). The for-
mer solution would be better for nomenclatural
stability, as well as making distinctive tribes. Bro-
meae as described here have no synapomorphies.
194. Bromus L. Fig. 39
Bromus L., Sp. Pl. 1: 76–78 (1753); phylog.: Saarela et al.

(2007).
Boissiera Hochst. ex Steud., Nomencl. Bot. (ed. 2) 1: 213
(1840).
Annuals or perennials, caespitose or rhizomatous. Fig. 39. Pooideae-Bromeae. Bromus berteroanus. A

Plant. B Ligule region. C Spikelet. D Anthecium. (From
Leaf sheath margins fused. Inflorescence branches Zuloaga et al. 2012, drawn by J. Coliera)
branched or unbranched. Spikelets with 3 to 30
flowers. Glume apices acute, sometimes mucro- flowers. Glumes papery, shorter than the flowers,
nate. Lemma entire to bidentate, awnless to more their apices obtuse to acute. Lemmas papery, over
often awned, the awn from a sinus or abaxial just 1 cm long, the apex obtuse or truncate, ragged,
below the apex. Style branches originating on the awnless or mucronate.
abaxial side of the ovary, the ovary extending Four spp., Central Asia.
adaxial to the style branches and ending in two Photographs of the ovary of L. alaica (Korsh.)
lobes. 2n ¼ 14, 28, 42, 56, 70, 84. Petrov ex Nevski in Schneider et al. (2009) show
One hundred sixty-one spp., temperate clearly that it lacks the ovary appendage charac-
regions worldwide. teristic of Bromus.
Boissiera is clearly derived within Bromus
(GPWG II 2012) and is thus placed in synonymy XV. TRIBE TRITICEAE Dumort. (1824)
here.
Annuals or perennials, caespitose or rhizoma-
195. Littledalea Hemsl. Fig. 40 tous. Leaf sheaths open, often with conspicuous
extensions (auricles) from the apex of the sheath
Littledalea Hemsl., Hooker’s Icon. Pl. 25: t. 2472 (1896). and surrounding the stem. Inflorescence
unbranched, with spikelets crosswise to the
Perennials, caespitose or rhizomatous. Leaf rachis. Spikelets 1 to 5 per node, generally later-
sheath with prominent auricles. Inflorescence ally compressed. Lodicules ciliate. Embryo about
with branched branches. Spikelets with 2 to 11 1/3 the length of the fruit; starch grains simple.
has been influential it is rarely followed blindly.

Generic limits here largely follow those laid out by
Barkworth (2007c), except that I take a broader
view of Elymus, which here includes most poly-
ploids with an St genome from Pseudoroegneria
irrespective of which other genomes are present.
The central problem is created by placing Hor-
deum, Triticum, and Secale in separate genera;
were they not such important crops, their names
could easily be changed and the entire tribe could
be placed in a single genus as repeatedly suggested
by Stebbins (Stebbins 1956; Stebbins and Snyder
1956; Stebbins, Jr. and Walters 1949). However,
maintaining nomenclatural stability for the major
crops forces inherent ambiguity in the classifica-
tion of the rest of the tribe. The evolutionary
history of the group is clearly highly reticulate
and any attempt to force a reticulum into a taxo-
nomic hierarchy becomes arbitrary (Kellogg et al.
1996). The website at http://herbarium.usu.edu/
Triticeae/ provides useful species descriptions
and other information relevant to the tribe.
Named intergeneric hybrids include xTritico-
secale Wittm. ex A. Camus, xPseudelymus Bark-
worth & D. R. Dewey, xElyhordeum Mansf. ex
Tsitsin & K. A. Petrova, xElyleymus B. R. Baum,
xPascoleymus (B. Boivin) Barkworth, and xLey-
deum Barkworth. These are sterile and morpho-
logically intermediate between their parental
genera, and are not described here.
196. Aegilops L.
Fig. 40. Pooideae-Bromeae. Littledalea tibetica. A Habit. Aegilops L., Sp. Pl. 2: 1050 (1753); tax.: Kimber and Feld-
B Ligule. C Spikelet. D Lemma. E Palea with pistil inside. F man (1987), van Slageren (1994).
Stamen. G Lodicules and pistil. (Hooker’s Icon. Pl. 25: t. Ambylopyrum (Jaub. & Spach) Eig, Agric. Rec. P.Z.E. Inst.
2472 (1896); from Hemsley 1896) Agric. Nat. Hist. (Tel Aviv) 2: 199 (1929).
Annuals. Leaves with falcate auricles. Inflores-

Twenty genera, 374 spp. cence axis disarticulating at the nodes or not, or
The challenges of generic delimitation in this shed as a unit. Spikelets one per node, laterally
tribe are summarized by Barkworth (2007c). compressed to terete, often partially embedded in
Intergeneric hybridization and polyploidy are the rachis, with 2 to 7 flowers. Glumes often with
common. Extensive cytogenetic studies have prominent veins, their apices truncate, toothed or
assessed genomic groups within which chromo- awned, or awned only on the distal spikelets.
somes pair freely and between which chromo- Lemma apex toothed, awnless or with 1 to 3
some pairing is reduced or absent (Dewey 1984; awns. 2n ¼ 14, 28, 42, 48.
L€ove 1984). L€ove (1984) and Barkworth and Twenty-eight spp., Europe, Mideast, North
Dewey (1985) have suggested that each distinct America.
genome or distinct combination of genomes Aegilops includes plants with genomes desig-
should be the basis of a genus. While this approach nated B, C, D, S, T, U, M, and N.
197. Agropyron Gaertn. above the nodes. Spikelets two per node, dorsi-
ventrally compressed, with 1 fertile flower and 1
Agropyron Gaertn., Novi Comment. Acad. Sci. Imp. Pet- reduced flower distally. Glumes longer than the
rop. 14: 539 (1770). flowers, linear, their apices attenuate. Lemma
apex attenuate, awned, the awns several times
Perennials, caespitose or rhizomatous. Leaf sheaths the length of the lemma. 2n ¼ 14.
with auricles. Inflorescence axis not disarticulating. One sp., C. delileana (Schult.) Roshev.,
Spikelets one per node, generally more than 3 times Morocco to Afghanistan, but most common in
as long as the internodes, laterally compressed, the eastern Mediterranean.
with 3 to 16 flowers. Glumes shorter than the flow- Crithopsis includes plants with the K genome.
ers, asymmetrically keeled, their apices acute, Attempts to cross C. delileana with other Triti-
awned or not. Lemma asymmetrically keeled, the ceae have been unsuccessful (Frederiksen 1993).
apex acute to awned. 2n ¼ 14, 28, 42.
Fifteen spp., Eurasia, introduced in North
200. Dasypyrum (Coss. & Durieu) T. Durand
America.
Agropyron includes plants with the genome Dasypyrum (Coss. & Durieu) T. Durand, Index Gen.
designated P. Phan.: 504 (1888); tax.: Frederiksen (1991a).
The name Agropyron was at one time applied Haynaldia Schur, Verh. K.K. Zool.-Bot. Ges. Wien 16: 37
to any perennial member of the Triticeae with one (1866).
spikelet per node, and thus included species now
placed in Pseudoroegneria, Thinopyrum, Elymus, Annuals or perennials, caespitose or rhizomatous.
Eremopyrum, and Pascopyrum. This use of the Inflorescence with one spikelet per node, disarticu-
name still appears in some treatments (e.g., Clay- lating at the nodes. Spikelets laterally compressed,
ton et al. 2006 onward), but such a circumscription with 2 flowers plus 1 or 2 reduced flowers distally.
renders Agropyron polyphyletic. As described Glumes shorter than the flowers, 2-keeled, apices
here, Agropyron includes only perennial species tapering into long awns, margins with long silky
with asymmetrically keeled glumes and lemmas hairs. Lemma apex acuminate, awned, the awn
(the crested wheatgrasses); the estimated number much longer than the lemma. 2n ¼ 14.
of species follows Chen and Zhu (2006). Two spp., Mediterranean.
Dasypyrum includes plants with the V genome.
198. Australopyrum (Tzvelev) Á. L€ove
201. Elymus L.
Australopyrum (Tzvelev) Á. L€ove, Feddes Rep. 95: 442
(1984). Elymus L., Sp. Pl. 1: 83 (1753); phylog.: Fan et al. (2013);
Hu et al. (2013); Mason-Gamer et al. (2002, 2010a, 2010b);
Petersen and Seberg (2008); Sun et al. (2008).
Perennials, caespitose. Inflorescence with one Hystrix Moench., Methodus: 294 (1794).
spikelet per node, not disarticulating, the spike- Elytrigia Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: 190
lets broadly spreading or reflexed, much longer (1810).
than the internodes. Spikelets disarticulating Sitanion Raf., J. Phys. Chim. Hist. Nat. Arts 89: 103 (1819).
Roegneria K. Koch, Linnaea 21: 413 (1848).
below the glumes, with 3 to 9 flowers. Glumes Anthosachne Steud., Syn. Pl. Glumac. 1: 237 (1855).
subulate. Lemma with short awns. 2n ¼ 14. Campeiostachys Drobow, Fl. Uzbekistan. 1: 300, 540 (1941).
Five spp., Australia, New Guinea, New Zealand. Pascopyrum Á. L€ ove, Taxon 29: 547 (1980).
Australopyrum includes plants with the W Kengyilia C. Yen & J.L. Yang, Canad. J. Bot. 68: 1897 (1990).
genome. Douglasdeweya C. Yen, J.L. Yang & B. R. Baum, Canad. J.
Bot. 83: 416 (2005).
Connorochloa Barkworth, S.W.L. Jacobs & H. Q. Zhang,
199. Crithopsis Jaub. & Spach Breed. Sci. 59: 685–686 (2009).
Crithopsis Jaub. & Spach, Ill. Pl. Orient. 4: 30 (1851); tax.: Perennials, caespitose, rhizomatous or stolonifer-
Frederiksen (1993). ous. Auricles often present. Inflorescence with 1
to 3 (5) spikelets per node, generally not disarti-
Annuals, less than 30 cm tall. Leaf sheath with culating at the nodes. Spikelets with 1 to 12 flow-
falcate auricles. Inflorescence disarticulating just ers, the distal ones often reduced. Glumes linear
or awl-like, awned or not, sometimes absent or One sp., Festucopsis serpentini (C.E. Hubb.)
reduced. Lemma apex acute, entire or with two Melderis, Albania.
tiny teeth, awned from the apex or sinus, or Festucopsis and Peridictyon are sister taxa in
awnless. 2n ¼ 28, 42, 56. molecular phylogenies (Petersen and Seberg
One hundred fifty spp., north temperate 2008), but are distinguished by the presence or
regions worldwide. absence of awns, the disarticulation of the rachis,
Elymus is defined by its genomic constitution and the number of flowers in each spikelet.
as including only polyploids; these all include at
least one diploid ancestor related to Pseudoroeg- 204. Henrardia C. E. Hubb.
neria (the St genome). Some taxa also include a
genome from Hordeum sect. Critesion (the H Henrardia C. E. Hubb., Blumea, Suppl. 3: 15 (1946); tax.:
genome), whereas others include the P genome Frederiksen (1993).
from Agropyron, and/or the Y genome, which has
not yet been found in any diploid. Species with the Annuals, less than 35 cm tall. Leaf sheath with
genomic constitution StY have sometimes been falcate auricles. Inflorescence disarticulating
segregated as Roegneria but are retained here in below the nodes. Spikelets one per node, dorsi-
Elymus. Elymus smithii is an allooctoploid with the ventrally compressed, embedded in the inflores-
St, H, Ns, and Xm genomes (Dewey 1975); it has cence axis, with 1 or 2 flowers. Glumes placed side
been segregated into its own genus (Pascopyrum) by side, their margins overlapping, slightly longer
on the basis of its novel genomic constitution, but than the flowers, their apices obtuse to acute.
is scarcely distinct morphologically. No compre- Lemma apex acute. 2n ¼ 14.
hensive taxonomic treatment exists for this genus. Two spp., southwest and central Asia.
Henrardia includes diploid plants with the O
202. Eremopyrum (Ledeb.) Jaub. & Spach genome. H. persica will not cross with Aegilops
species, Elymus tsukushiense, Eremopyrum bonae-
Eremopyrum (Ledeb.) Jaub. & Spach, Ann. Sci. Nat., Bot. partis, Heteranthelium, or Triticum boeoticum, but
III, 14: 360 (1851); tax.: Frederiksen (1991b). forms sterile hybrids with Eremopyrum orientale
(Sakamoto 1972). A few seeds were formed in a
Annuals. Leaf sheath with inconspicuous auricles. cross with Taeniatherum (Frederiksen 1993).
Inflorescence with one spikelet per node, disarti-
culating at the nodes or not, internodes flat, much 205. Heteranthelium Hochst. in Kotschy
shorter than the spikelets, the spikelets broadly
spreading. Spikelets laterally compressed, with Heteranthelium Hochst. in Kotschy (ed. Hohenacker), Pl.
2 to 5 flowers, sometimes with additional reduced Aleppo exsiccatae 130a (1843); tax.: Frederiksen (1993).
flowers distally. Glumes shorter than the flowers,
their apices acute to acuminate, muticous or with Annuals. Leaf sheath and blade densely pubescent,
a short awn. Lemma apex muticous to awned. Palea the sheath with falcate auricles. Inflorescence with
keels sometimes extended into teeth. 2n ¼ 14, 28. one spikelet per node, fertile and sterile spikelets
Four spp., North America, central Asia. interspersed, disarticulating at some nodes, so that
Eremopyrum includes plants with the F and the dispersal unit includes both fertile and sterile
Xe genomes. spikelets; distal inflorescence nodes with spikelets
reduced to awns. Spikelets laterally compressed,
203. Festucopsis Melderis with 1 or 2 fertile flowers and a rachilla ending in a
cluster of awns. Glumes longer than the flowers,
Festucopsis Melderis, Bot. J. Linn. Soc. 76: 317 (1978). slender, awl-shaped, densely pubescent. Lemma
with long tuberculate hairs, the apex with a long
Caespitose perennials. Leaf sheaths lacking auri- awn. 2n ¼ 14.
cles, leaf blades filiform. Inflorescence with one One sp., H. piliferum (Banks & Sol.) Hochst.,
spikelet per node, not disarticulating. Spikelets Turkey to Pakistan.
with 3 to 5 flowers. Glumes with acute apices, Heteranthelium includes plants with the Q
unawned. Proximal lemma of spikelet with a genome. H. piliferum will produce sterile hybrids
short awn, distal lemma lacking an awn. 2n ¼ 14. with Eremopyrum bonaepartis (both diploid and
eral two generally sterile and pedicellate. Spikelets

dorsiventrally compressed, with 1 flower. Glumes
shorter or more often longer than the spikelet,
generally subulate to linear, their apices long atten-
uate or extending into an awn. Rachilla extension
generally present on central spikelet. Lemma apex
attenuate, generally with a long apical awn, the awn
straight or curved. 2n ¼ 14, 28, 42.
Forty-three spp., temperate and subtropical
regions of Eurasia, North and South America,
Africa, introduced elsewhere.
Hordeum vulgare, barley, was domesticated
in the Fertile Crescent about 8500 years ago (Har-
lan and Zohary 1966), with a possible second
center of domestication in the Zagros Mountains
in present day Iran (Morrell and Clegg 2007).
Other species of the genus are weedy. Hordeum
includes plants with genomes designated H, I, X,
and Y. Twenty species are diploid and the
remainder are polyploid, so the history of the
genus is reticulate (Blattner 2006, 2009). Species
with the H genome have hybridized in the past
with Pseudorogeneria (St genome; see below) to
form allopolyploids classified here as Elymus
(Mason-Gamer et al. 2010a, 2010b).
207. Leymus Hochst.
Leymus Hochst., Flora 31: 118 (1848); phylog.: Petersen

and Seberg (2008).
Hordelymus (Jess.) Harz, Landw. Samenk. 2: 1147 (1885).
Eremium Seberg & Linde-Laursen, Syst. Bot. 21: 10, f. 4
(1996).
Fig. 41. Pooideae-Hordeeae. Hordeum brevisubulatum
subsp. turkestanicum. A Habit. B Spikelet. C Middle
flower. D Lateral flower. (From Wu et al. 2007, with Perennials, rhizomatous or sometimes caespitose.
permission from the Missouri Botanical Garden Press, Leaf blades stiff, leaf sheath with auricles. Inflo-
St. Louis, and Science Press, Beijing) rescence with 1 to 8 spikelets per node, occasion-
ally branched, not disarticulating. Spikelets sessile
tetraploid) and Hordeum depressum (a tetra- or on pedicels up to 2 mm long, laterally com-
ploid) but not with E. triticeum (diploid), Hen- pressed, with 2 to 12 flowers, the distal flower(s)
rardia persica (diploid) or Elymus tsukushiense reduced. Glumes with apices acute, acuminate,
(hexaploid) (Frederiksen 1993; Sakamoto 1974). awl-like, or tapering to a short awn. Lemma
apex acute, awned or awnless, the awn generally
206. Hordeum L. Fig. 41 short but sometimes as long as the lemma,
straight. 2n ¼ 28, 42, 56.
Hordeum L., Sp. Pl. 1: 84–85 (1753); tax.: Blattner (2009). Fifty-four spp., temperate regions of the
Critesion Raf., J. Phys. Chim. 89: 103 (1819). northern hemisphere.
Leymus includes polyploid plants with the Ns
Annuals or perennials, caespitose, rarely rhizo- genome, derived from Psathyrostachys. A second
matous. Leaf sheath with or without auricles. genome may be present but the data are ambiguous
Inflorescence axis generally disarticulating at the (B€odvarsdottir and Anamthawat-Jónsson 2003;
nodes; spikelets in groups of three at each node, Wang and Jensen 1994). Hordelymus is included
with the central spikelet fertile and sessile, the lat- here (rather than in Elymus) because of the presence
of an N genome and apparent lack of an St genome Pseudoroegneria includes plants with the St
(Petersen and Seberg 2008; Zhang and Sun 2010). genome. It is clearly one progenitor of most of
the polyploids in Elymus s.l. (Mason-Gamer et al.
208. Peridictyon Seberg, Fred. & Baden 2002, 2010a, 2010b) and Thinopyrum (Liu and
Wang 1993; Mahelka et al. 2011). Not surprisingly,
Peridictyon Seberg, Fred. & Baden, Willdenowia 21: 96 because it has so many polyploid derivatives, the
(1991). exact delimitation of Pseudoroegneria and its des-
cendants has been controversial, with some spe-
Caespitose perennials. Leaf sheaths with auricles, cies placed in Elytrigia and others in Elymus.
leaf blades filiform. Inflorescence with one spike-
let per node, disarticulating at the nodes. Spike- 211. Secale L.
lets with 5 to 10 flowers. Glumes shorter than the
flowers, their apices acute, with short awns. Secale L., Sp. Pl. 1: 84 (1753); tax.: Frederiksen and Peter-
Lemma apex awned. 2n ¼ 14. sen (1998).
One sp., Peridictyon sanctum (Janka) Seberg,
Fred. & Baden, Northern Greece and southwest- Annuals or perennials. Leaf sheath with falcate
ern Bulgaria. auricles. Inflorescence with one spikelet per node,
See comments under Festucopsis. disarticulating at the nodes in wild species, not
disarticulating in cultivated ones. Spikelets later-
209. Psathyrostachys Nevski ally compressed, with 2 flowers. Glumes subulate,
keeled, shorter than the flowers, their apices
Psathyrostachys Nevski, Trudy Bot. Inst. Akad. Nauk S.S.S. extending into a terminal awn. Lemma keeled,
R., Ser. 1, Fl. Sist. Vyssh. Rast. 2: 57 (1934); tax.: Baden (1991). acuminate, awned. 2n ¼ 14.
Eight spp., Mediterranean and western Asia,
Perennials, caespitose, rhizomatous, or stolonif- introduced and often cultivated elsewhere.
erous. Leaf sheath with or without auricles. Inflo- Secale includes plants with the R genome.
rescence axis disarticulating at the nodes.
Spikelets borne in groups of two or three on the
212. Stenostachys Turcz.
rachis, each with 1 or 2 (3) flowers. Glumes as
long as the flowers or longer, both abaxial to the Stenostachys Turcz., Bull. Soc. Imp. Naturalistes Moscou
rest of the spikelet, subulate. Lemma apex entire 35: 330 (1862); phylog.: Petersen et al. (2011).
or with minute lobes, awned from the apex or
from the sinus, the awns straight. 2n ¼ 14, 28. Stoloniferous perennials. Leaves with auricles
Nine spp., northern Asia and the Caucasus, minute. Inflorescence drooping or nodding, not
south to Turkey and Afghanistan. disarticulating at the nodes. Spikelets one per
This genus includes diploids and tetraploids node. Glumes shorter than the flower or absent,
bearing the N genome. awl-shaped or lanceolate, their apices attenuate.
Lemma with a short awn. 2n ¼ 28.
210. Pseudoroegneria (Nevski) Á. L€ove Four spp., New Zealand.
Stenostachys includes polyploid plants with
Pseudoroegneria (Nevski) Á. L€ove, Taxon 29: 168 (1980). the H and W genomes.
Perennials, caespitose or rhizomatous. Leaf sheaths 213. Taeniatherum Nevski

with auricles. Inflorescence with one spikelet per
node, the rachis not disarticulating, with spikelets Taeniatherum Nevski, Trudy Sredne-Aziatsk. Gosud.
appressed to the axis, about the same length as the Univ., Ser. 8b, Bot. 17: 38 (1934); tax.: Frederiksen (1986).
internodes or only slightly longer. Spikelets laterally
compressed, with 4 to 9 flowers. Glumes shorter than Annuals. Leaf sheath with falcate auricles. Inflores-
the flowers, their apices acute to obtuse or truncate. cence not disarticulating at the nodes. Spikelets
Lemma apex acute, awned or unawned, the awn 2 (rarely 3) per node, dorsiventrally compressed,
straight and erect or bent and divergent. 2n ¼ 14, 28. with 1 flower plus 1 additional reduced flower distally.
Fifteen spp., Eurasia, Asia, North America. Glumes placed side by side, longer than the flowers,
awl-like, their bases fused. Lemma apex acuminate, Desf. is tetraploid, and is the major wheat used
awned, the awn many times longer than the lemma; for pasta, semolina and bulghur.
awns becoming entangled at maturity. 2n ¼ 14.
One sp., T. caput-medusae (L.) Nevski, Medi- XVI. TRIBE POEAE R. Br. (1814), s.l
terranean and mid-East to Pakistan; introduced
Annuals or perennials. Leaf sheaths generally
elsewhere.
lacking auricles. Inflorescence branches branched
The entangled awns are responsible for the Latin
in most species. Spikelets laterally compressed,
and common names of this grass, Medusa-head.
generally disarticulating above the glumes. Hilum
Taeniatherum includes plants with the Ta genome.
punctate; endosperm with lipid.
One hundred-six genera, 2578 spp.
214. Thinopyrum Á. L€ove This tribe includes most of the species in the
subfamily, following the circumscription of Soreng
Thinopyrum Á. L€ove, Taxon 29: 351 (1980). et al. (2007, 2012 and onward). Synapomorphies of
Lophopyrum Á. L€ove, Taxon 29: 351 (1980). the tribe include a punctate hilum and lipid in the
endosperm (Soreng et al. 2007), although these
Perennials, caespitose or rhizomatous. Auricles characters reverse in some taxa. Abaxial awn attach-
present or absent. Inflorescence generally with ment, absence of cilia on the lodicules, lack of ovary
one spikelet per node, not disarticulating at the pubescence, and presence of compound starch
nodes. Spikelets appressed to ascending, 1 to 3 grains may also be derived here, although they are
times as long as the internodes, with 3 to 12 flow- homoplasious and thus their position on the phy-
ers. Glumes thick, stiff, keeled, the apices truncate logeny depends on the method of optimization.
to acute, sharp-pointed but awnless. Lemma thick, The classification of Soreng et al. (2007), con-
stiff, the apex truncate to acute, mucronate or with tinually updated by Soreng et al. (2012 and
short awns. 2n ¼ 14, 42, 43, 56, 69, 70. onward), is used as the basis of the classification
Ten spp., Mediterranean to western Asia, presented here, although some differences are
introduced elsewhere. noted below. Soreng et al. (2012 and onward)
Thinopyrum includes plants with the E genome; divide Poeae into 19 subtribes, of which 15 are
the polyploids also incorporate the St genome from recognized here. Within tribe Poeae s.l., Gillespie
Pseudoroegneria, the P genome from Agropyron, et al. (2008) used both nuclear (ITS) and plastid
and maybe the L genome of Festucopsis. (trnT-trnL-trnF) sequences to assess relation-
ships within and among genera in subtribe Poi-
215. Triticum L. nae, extending work done by the same group of
investigators (Soreng et al. 2007) proposing a
Triticum L., Sp. Pl. 1: 85–87 (1753). subtribe phylogeny for tribe Poeae. Sequences of
the plastid genome fall into two clades, which are
Annuals. Leaf sheaths with auricles. Inflorescence noted here; however, these clades may not reflect
with one spikelet per node, disarticulating at the the evolution of the nuclear genome.
nodes, the spikelets generally falling with the Molecular phylogenetic studies consistently
internode below but sometimes with the adjacent identify a clade within Poeae that has been called
internode, or the entire inflorescence shed as a PPAM, an acronym for Poinae, Puccinelliinae, Alo-
unit. Spikelets with 2 to 9 flowers, the distal ones pecurinae and Miliinae (all tribes sensu Soreng et al.
often sterile. Glumes keeled, the apex with a tooth 2007); ITS data also place Cinninae in this group.
or awn. Lemma keeled, the proximal ones awned, Relationships among the five former subtribes vary
the distal ones awned or not. 2n ¼ 14, 28, 42. depending on the molecular marker used, but they
Eighteen spp., western and central Asia, culti- seem to form a coherent group (albeit without an
vated and escaped in temperate regions worldwide. obvious morphological synapomorphy or even a
Triticum includes diploid plants with the A diagnostic combination of characters), with Pucci-
genome and polyploids with the A, B, D, and G nelliinae (now recognized as Coleanthinae) sister to
genomes; the B and D genomes are derived from the rest. Accordingly, here I place the entire PAM
species of Aegilops. T. aestivum L. (bread wheat) clade plus Cinninae in subtribe Poinae.
is a hexaploid species that is one of the most Named hybrid genera include xAgropogon P.
widely cultivated grasses in the world. T. durum Fourn., xArctodupontia Tzvelev, xDupoa J. Cay. &
Darbysh., and xPucciphippsia Tzvelev. These are

not described here.
INCERTAE SEDIS
216. Megalachne Steud.
Megalachne Steud., Syn. Pl. Glumac. 1: 237 (1854).
Perennials. Spikelets with 3 to 6 flowers plus

additional reduced flowers distally. Glumes
about as long as flowers, their apices acuminate
to attenuate, with aristae or long awns. Callus of
flower pubescent. Lemma keeled, the apex acumi-
nate, awned from the apex, the awn long and
curved. Ovary apex pubescent.
Two spp., Juan Fernández Island.
217. Podophorus Phil.
Podophorus Phil., Bot. Zeitung (Berlin) 14: 648 (1856).
Perennials <40 cm tall. Spikelets with 1 flower

plus 1 reduced flower distally. Glumes shorter
than or as long as the flowers, their apices acumi-
nate but unawned. Callus of flower pubescent.
Lemma rounded, the apex acuminate, awned,
the awn bent. Ovary apex pubescent.
One sp., P. bromoides Phil., Juan Fernández
Island.
Only a handful of specimens exist of this
species, which has not been collected since the
late 19th century (Baeza et al. 2002). It is pre-
sumed to be extinct.
GROUP WITH AVENEAE-TYPE PLASTID SEQUENCES
This clade is retrieved in all molecular studies Fig. 42. Pooideae-Poeae-Torreyochloinae. Amphibromus
scabrivalvis. A Plant. B Spikelet. C Lemma. D Palea. E
of the plastid (e.g., Davis and Soreng 2007; Gillespie Palea, lodicules and anthers. F Lodicules, callus, and
et al. 2008; Soreng and Davis 2000; Soreng et al. base of the palea. (From Zuloaga et al. 2012)
2007), but appears to have no morphological syna-
pomorphies. The group is supported poorly or not Two genera, 16 species.
at all by analyses of nuclear gene sequences (Gille- This subtribe is placed sister to the rest of the
spie et al. 2008; Quintanar et al. 2007; Saarela et al. Avena-plastid-type clade in chloroplast molecu-
2010), with the position of Sesleriinae being partic- lar phylogenies (Davis and Soreng 2007; Soreng
ularly different between the two sorts of analyses. and Davis 2000).
218. Amphibromus Nees Fig. 42

XVI.1. SUBTRIBE TORREYOCHLOINAE Soreng & J. I. Davis
(2003) Amphibromus Nees, London J. Bot. 2: 420 (1843); tax.:
Jacobs and Lapinpuro (1986).
Aquatic or semi-aquatic perennials. Glumes shorter
than the flowers, unawned. Lemma awned or Perennials, caespitose, rhizomatous or stolonifer-
unawned. Hilum 1/3 to 1/2 the length of the fruit. ous, often with cleistogamous inflorescences
enclosed by the lower leaf sheaths. Spikelets with or bidentate, with or without a small awn. Ovary
3 to 10 flowers; rachilla extension present or pubescent. Hilum long-linear. 2n ¼ 14, 28, 42.
absent. Glumes with apices obtuse to acute. Cal- Eight spp., Europe, mid-East.
lus of flower pubescent. Lemma awned from the Clayton and Renvoize (1986) argue that
abaxial side, the awn twisted and geniculate, Arrhenatherum is simply a segregate from Helic-
spreading or recurved, lemma apex with 2 to 4 totrichon and division between the two might be
teeth or lobes, the lobes obtuse or with aristae. unwarranted. D€oring et al. (2007) present data
Ovary glabrous. 2n ¼ 42. indicating that Arrhenatherum could be linked
Twelve spp., Australia, New Zealand, South with Pseudarrhenatherum, and the two could per-
America. haps be included in Helictotrichon. Saarela et al.
Amphibromus is morphologically similar to (2010) resolve Arrhenatherum as monophyletic,
Helictotrichon, but with a glabrous ovary; the two not forming a clade with Helictotrichon. Arrhe-
genera are not related (Saarela et al. 2010). natherum is thus recognized here.
219. Torreyochloa G. L. Church 221. Avena L.
Torreyochloa G. L. Church, Amer. J. Bot. 36: 163 (1949); Avena L., Sp. Pl. 1: 79–81 (1753); tax.: Baum (1977).
tax.: Church (1952).
Annuals. Spikelets with 1 to 6 flowers plus a rachilla
Perennials, rhizomatous. Spikelets with 2 to 8 flow- extension. Glumes generally enclosing the flowers,
ers; rachilla extension present. Glumes with apices but sometimes shorter, their apices obtuse to acu-
acute. Callus of flower glabrous. Rachilla glabrous. minate. Callus of flower glabrous or pubescent.
Lemma with 7 to 9 prominent veins, the veins Lemma apex with two teeth or slender setae,
parallel and not converging toward the apex, awned, the awn curved or twisted, generally genicu-
lemma apex truncate to acute, sometimes with a late. Ovary pubescent. Hilum linear. 2n ¼ 14, 28, 42.
ragged edge. Ovary pubescent. 2n ¼ 14. Twenty-four spp., Europe, Mediterranean
Four spp., North America, Northeast Asia. areas, northern Africa.
Oats, Avena sativa L., are an important grain,
particularly in temperate regions of the world.
XVI.2. SUBTRIBE AVENINAE J. Presl (1830) The species of Avena form a complex series of
polyploids; although there is one phylogeny of
Annuals or perennials. Lemma awnless, mucronate, these (Peng et al. 2010), many questions remain.
or with an abaxial awn, the awn geniculate. Hilum
short or linear, the latter state presumably a reversal. 222. Graphephorum Desv.
Seven genera, 324 species.
A detailed phylogeny of this subtribe is pre- Graphephorum Desv., Nouv. Bull. Sci. Soc. Philom. Paris
sented by Saarela et al. (2010). 2: 189 (1810); tax.: Finot et al. (2005).
Caespitose perennials. Spikelets with 2 or 3 flow-

220. Arrhenatherum P. Beauv.
ers; rachilla with dense hairs. Glumes shorter
Arrhenatherum P. Beauv., Ess. Agrostogr. 55: 152–153 (1812). than or as long as the spikelet, their apices acute
Pseudarrhenatherum Rouy, Bull. Soc. Bot. France 68:401 to acuminate. Lemma apex obtuse to acute, muti-
(1921). cous or with an awn just below the apex.
Two spp., North and Central America.
Perennials, caespitose or rhizomatous, the base of Plastid data place Sphenopholis and Graphe-
the plant sometimes forming corms. Spikelets with phorum as sisters and the two as sister to Trise-
2 flowers, the proximal one staminate, the distal tum s.l., whereas ITS data put Graphephorum
one pistillate or bisexual. Glumes as long as the within Trisetum s.l. and Sphenopholis sister to
flowers, their apices acute to acuminate. Callus of the Graphephorum/Trisetum clade (Quintanar
flower pubescent. Lemma of proximal flower with et al. 2007, 2010). In the absence of a more con-
the apex acute, awned, the awn twisted, geniculate; sistent phylogenetic signal, I keep Sphenopholis
lemma of distal flower with the apex acute, entire and Graphephorum separate here.
223. Helictotrichon Besser
Helictotrichon Besser, Mant. 3: 526, in obs. (1827); tax.:

Lange (1995).
Caespitose perennials. Ligule ciliate, ragged. Spi-

kelets with 2 to 8 flowers plus additional reduced
flowers distally; rachilla pubescent. Glumes as
long as or longer than the flowers, their apices
acute to acuminate, sometimes mucronate or
awned. Callus of flower acute, with straight
hairs. Lemma apex with two teeth, awned, the
awn twisted and geniculate. Ovary pubescent.
Hilum long-linear. 2n ¼ 14, 28, 42.
One hundred two spp., temperate regions of
Eurasia.
Based on molecular data, Helictotrichon in its
traditional circumscription is paraphyletic, with
subg. Pratavenastrum Holub more closely related
to Deschampsia than to subg. Helictotrichon or
subg. Tricholemma M. R€oser (Grebenstein et al.
1998). The three subgenera are all strongly sup-
ported as monophyletic. Therefore, Helictotri-
chon is restricted to the latter two sections, and
subg. Pratavenastrum is placed in Avenula; see
comments under the latter genus.
224. Lagurus L.
Fig. 43. Pooideae-Poeae-Aveninae. Trisetum barbinode
Lagurus L., Sp. Pl. 1: 81 (1753). Trin. A Plant. B Spikelet. C Lowermost flower. (From
Zuloaga et al. 2012)
Tufted annuals. Inflorescence dense, ovoid. Spi- rachilla extension. Glumes shorter than or as long
kelets with 1 flower and a pubescent rachilla as the flowers; lower glume slender, the apex
extension. Glumes longer than the flower, covered acute, upper glume broader above than below,
with wooly hairs, their apices acuminate, awned, broadly oblanceolate to flabellate, appearing
the awns covered with hairs. Lemma pubescent, wedge shaped in lateral view, the apex obtuse to
the apex with two teeth or slender setae, awned, acute. Callus of flower glabrous. Lemma apex
the awn twisted. 2n ¼ 14. obtuse to acute, with a small mucro or a twisted
One sp., L. ovatus L., Mediterranean but awn. Fruit with liquid endosperm. 2n ¼ 14.
widely cultivated. Eight spp., North America and Mexico.
The wooly hairs on the glumes make the See comments under Graphephorum. Spheno-
entire inflorescence look furry, giving rise to the pholis is distinguished from other members of
common name “rabbit’s foot”. this subtribe by having spikelets that disarticulate
below the glumes.
225. Sphenopholis Scribn.
226. Trisetaria Forssk. Fig. 43
Sphenopholis Scribn., Rhodora 8: 142 (1906), tax.: Erdman
(1965); Finot et al. (2004).
Trisetaria Forssk., Fl. Aegypt.-Arab. 60: 27 (1775); tax.:
Finot et al. (2004, 2005).
Perennials, rarely annual. Spikelets with 2 or 3 Trisetum Pers., Syn. Pl. 1: 97 (1805), nom. conserv.
flowers, disarticulating below the glumes, with a Koeleria Pers., Syn. Pl. 1: 97 (1805).
Gaudinia P. Beauv., Ess. Agrostogr. 95, 164 (1812). several species of Calamagrostis (subtribe Agros-
Rostraria Trin., Fund. Agrost. 149 (1820). tidinae). In the future these clades may be recog-
Avellinia Parl., Pl. Nov. 59 (1842).
Peyritschia E. Fourn., Mexic. Pl. 2: 109 (1886).
nized as sister genera or subgenera. No molecular
xTrisetokoeleria Tzvelev, Novosti Sist. Vyssh. Rast. 7: 73 data are available for Leptophyllochloa; it is
(1970 [1971]). placed here based on morphology. Species for-
Leptophyllochloa C. E. Calderón, Fl. Patagónica 3: 69 (1978). merly placed in Gaudinia are distinctive in that
Parafestuca E.B. Alexeev, Bjull. Moskovsk. Obač. Isp. the inflorescence is unbranched and the rachis
Prir., Otd. Biol. 90: 107–108 (1985).
disarticulates just above the spikelet.
Perennials (most species) or annuals, caespitose,
rhizomatous or stoloniferous. Inflorescences
unbranched (in some annuals) or the branches XVI.3. SUBTRIBE PHALARIDINAE Fr. (1835)
branched. Spikelets with 2 to 7 (11) flowers;
rachilla extension present, glabrous or pubescent, Annuals or perennials, caespitose or rhizoma-
sometimes ending in a reduced flower. Glumes tous, in some perennials the lowest internode
shorter than to longer than the flowers, their enlarged and corm-like. Inflorescence dense, cap-
apices obtuse to attenuate, generally mucronate itate or spike-like. Spikelets bisexual, or in clus-
but rarely awned. Callus of flower generally ters with the proximal ones staminate and the
pubescent. Lemma apex entire or with two teeth, terminal pistillate or bisexual; with 1 to 3 (4)
awnless or with a mucro or awn from the abaxial flowers; disarticulation above the glumes, or
side. Ovary glabrous or pubescent. Fruit some- beneath the distalmost spikelet of the cluster, or
times with liquid endosperm. 2n ¼ 14, 28, 42. the clusters shed as a unit. Glumes longer than the
One hundred seventy-nine spp., temperate flowers, keeled, their apices truncate to attenuate.
regions worldwide. Proximal two flowers sterile, reduced to awl-
Neither Trisetum nor Koeleria is monophy- shaped or flap-like lemmas. Lemma and palea of
letic in any molecular phylogeny to date, but distal bisexual flower polished and hard, the
together they comprise a clade as long as several lemma apex generally acute to acuminate or
smaller genera are included as well (Grebenstein beaked. Hilum long-linear.
et al. 1998; Quintanar et al. 2007, 2010; Saarela One genus, 22 species.
et al. 2010; Soreng et al. 2007). Species of Ros-
traria, Gaudinia, Avellinia, Peyritschia, and Par- 227. Phalaris L. Fig. 44
afestuca are intermixed with those of Trisetum
and Koeleria (Quintanar et al. 2007; Saarela Phalaris L., Sp. Pl. 1: 54–55 (1753); tax., Baldini (1995);
et al. 2010) so are placed in synonymy here. phylog.: Voshell et al. (2011).
Among these smaller genera is Trisetaria, a
name that has priority over Trisetum. The annual Characters as for the subtribe. 2n ¼ 12, 14, 28,
species of Trisetaria are clearly embedded in Tri- 29, 42.
setum, as noted by Quintanar et al. (2010). Twenty-two spp., north temperate and tem-
Although Trisetum is a nomen conservandum, it perate regions of South America.
is not conserved against Trisetaria. The nomen- Chromosome numbers reported for P. arun-
clatural solutions are either to conserve Trisetum dinacea include the aneuploid counts 27, 29, 31,
against Trisetaria, or to transfer all Trisetum spe- and 35 (Barkworth 2007b).
cies to the older genus; a few such transfers have
already been done (Paunero Ruiz 1950).
As delimited here, Trisetaria corresponds to XVI.4. SUBTRIBE ANTHOXANTHINAE A. Gray (1856)
the subtribe Koeleriinae (Quintanar et al. 2010),
except that Graphephorum and Sphenopholis are Annuals or perennials, caespitose or rhizoma-
kept separate pending additional data. Saarela tous. Inflorescence sometimes unbranched, or
et al. (2010) identify two well-supported clades the branches branched. Spikelets with 3 flowers,
within Trisetum s.l., an Old World group and a the 2 proximal ones staminate or sterile, if sterile
New World one; the New World clade includes then consisting solely of a lemma; rachilla
extension absent. Glumes as long as or longer

than the flowers, their apices acute to acuminate,
mucronate. Callus glabrous or hairy. Lemma of
sterile flowers longer than that of the upper one,
with long hairs, the apex bilobed, unawned or
awned from the abaxial side; lemma of bisexual
flower hardened, shiny, with margins enclosing
the palea, the apex notched, awnless. Lodicules
absent or 2. Stamens 2 or 3.
One genus, 52 species.
228. Anthoxanthum L. Fig. 45
Anthoxanthum L., Sp. Pl. 1: 28 (1753); tax.: Pereira et al.

(2007); phylog.: Pimentel et al. (2013).
Hierochloe R. Br., Prodr.: 208 (1810), nom. conserv.
Characters as for the subtribe. 2n ¼ 10, 20, 28,

42, 56.
Fifty-two spp., temperate and arctic regions
throughout the world, widely introduced for hay.
Species in this genus produce coumarin (pre-
sumed to be a synapomorphy), which makes
them sweetly scented and accounts for the pleas-
ant odor of some hay. Species formerly in Hiero-
chloe were and are used by Native Americans to
make baskets and decorations. The genus Hiero-
chloe has been separated from Anthoxanthum on
the basis of the proximal flowers of the spikelet,
which are staminate in the former and sterile in
the latter. However, Schouten and Veldkamp
(1985) suggest that the two intergrade in parts
of their range, and the genera are treated as a
single genus by Allred and Barkworth (2007).
Merging the two is also supported by molecular
data (Quintanar et al. 2010; Saarela et al. 2010).
While most reported chromosome numbers are
euploid, those for A. monticola are cited as 56, 58,
63, 66, 72 (Allred and Barkworth 2007).
XVI.5. SUBTRIBE AGROSTIDINAE Fr. (1835)
Subtribe Brizinae Tzvelev, Bot. Žurn. (Moscow & Lenin-

grad) 53: 310 (1968).
Fig. 44. Pooideae-Poeae-Phalaridinae. Phalaris canarien-

sis. A Habit. B Sterile flowers and a fertile one. C Spikelet.
(From Barkworth et al. 2007, drawn by L.A. Vorobik and
H. Paźdirková)
Lemma awnless or with an abaxial awn, the awn

often geniculate. Palea generally hyaline, the mar-
gins often wider than the space between the veins.
Twenty-one genera, 681 species.
Saarela et al. (2010) show that Brizinae and
Agrostidinae are intermixed and argue against
recognizing both. The two are combined here.
229. Airopsis Desv.
Airopsis Desv., J. Bot. (Desvaux) 1: 200 (1809).
Annuals. Spikelets nearly spherical, with 2 flowers.

Glumes longer than the flowers, their apices
obtuse. Lemma orbicular, the apex obtuse, with
three broad lobes. 2n ¼ 8.
One sp., A. tenella Coss. & Durand, Mediter-
ranean.
Briza s.s. and Airopsis form a weakly sup-
ported clade in the analyses of Saarela et al. (2010).
230. Briza L.
Briza L., Sp. Pl. 1: 70–71 (1753); phylog.: Essi et al. (2008).
Annuals or perennials, caespitose. Spikelets oval

to elliptic in outline, sometimes as broad as long,
with 3 to 12 (15) flowers that diverge strongly
from the rachilla; rachilla extension absent.
Glumes shorter than the flowers, their apices
obtuse. Callus of flower glabrous. Lemma
inflated, the base often cordate, the margins
translucent, the apex unawned. 2n ¼ 10, 14, 28.
Five spp., Mediterranean, introduced and
sometimes cultivated elsewhere.
231. Chascolytrum Desv.
Chascolytrum Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2:

190 (1810); tax.: Essi et al. (2011).
Calotheca Desv., Nouv. Bull Sci. Soc. Philom. Paris 2: 190
(1810).
Rhombolytrum Link, Hort. Berol. 2: 296 (1833).
Poidium Nees, Intr. Nat. Syst. Bot.: 450 (1836).
Erianthecium Parodi, Notas Mus. La Plata, Bot. 8: 75 (1943).
Fig. 45. Pooideae-Poeae-Anthoxanthinae. Anthox-

anthum odoratum. A Plant. B Ligule region. C Inflores-
cence. D Spikelet. E Sterile flowers. F Caryopsis. (From
Barkworth et al. 2007, drawn by L.A. Vorobik and H.
Paźdirková)
Microbriza Parodi ex Nicora & Rúgolo, Darwiniana 23: intergrading nature of morphological variation in
292 (1981). this group.
Gymnachne Parodi, Notas Mus. La Plata, Bot. 3: 29 (1938).
233. Agrostis L.
Perennials, with culms sometimes forming
corms. Inflorescence branched, condensed. Spi- Agrostis L., Sp. Pl. 1: 61–63 (1753).
kelets with 2 to 4 (to 10) flowers, ovate or orbicu- Chaetopogon Janch., Eur. Gatt. Farn. Bl.-Pfl. (ed. 2): 33
lar; flowers barely separated on the rachilla. (1913).
Glumes shorter than the flowers, often keeled,
their apices acute to obtuse. Callus hairy or not. Perennials, rarely annuals, caespitose, rhizoma-
Lemmas broader than long, contracted distally to tous or stoloniferous. Spikelets generally less than
an acute or acuminate apex, or with 2 teeth or 7 mm long, without a rachilla extension. Glumes
lobes, mucronate or with a short awn from the generally longer than the flower, with one vein,
sinus. Rachilla disarticulating. 2n ¼ 28. their apices acute to acuminate, muticous or
Twenty-eight spp., South America. rarely awned. Callus of flower glabrous or with
Clayton et al. (2006 onward) and Clayton and straight hairs less than ½ the length of the lemma.
Renvoize (1986) consider Chascolytrum to be Lemma generally hyaline or thinly membranous,
synonymous with Briza, but molecular data with 3 or 5 veins, the apex truncate or toothed,
(Davis and Soreng 2007; Essi et al. 2008) indicate awned from the apex or more often abaxially, the
that the two genera are distinct. Essi et al. (2008) awn straight or geniculate. Palea generally minute
show convincingly that the two species formerly or lacking. Fruit with endosperm hard, soft or
placed in Microbriza (¼ B. poaemorpha and B. liquid. 2n ¼ 14, 28, 35, 42, 56.
brachychaete) are not sisters in either chloroplast Two hundred sixty-four spp., worldwide in
or nuclear gene trees. Erianthecium and Rhombo- temperate regions.
lytrum (represented by Gymnachne koelerioides) Agrostis intergrades morphologically with
also fall in this clade. Erianthecium was Calamagrostis, but Saarela et al. (2010) find that
segregated from Chascolytrum because it forms most species of Agrostis form a reasonably well-
bulbs at the base of the culm, and the lemmas are supported clade, distinct from Calamagrostis.
awned. Chaetopogon falls within Agrostis in the trees of
Saarela et al. (2010) and the morphology is fully
compatible with including the former in the latter
232. Relchela Steud. genus.
Relchela Steud., Syn. Pl. Glumac. 1: 101 (1855).
234. Ammophila Host
Stoloniferous perennials. Spikelets with 1 or
Ammophila Host, Icon. Descr. Gram. Austriac. 4: 24
2 flowers; rachilla extension present or absent. (1809); tax.: Delisle-Oldham et al. (2008).
Glumes as long as or longer than the flowers,
their apices acute. Lemma hard, the margins Perennials with strong rhizomes. Leaf blades
inrolled and largely covering the palea, the apex strongly inrolled, with sharp points. Inflorescence
obtuse. Ovary apically pubescent. dense, narrow. Spikelet with a rachilla extension.
One sp., R. panicoides Steud., Argentina and Glumes as long as or longer than the flowers, their
Chile. apices acute to acuminate, unawned. Callus of
Remaining genera of the subtribe: flower with long hairs. Lemma apex entire or
Spikelets with 1 flower. with two teeth, mucronate, the mucro attached
Named hybrids include xAgrocalamagrostis Asch. & abaxially. Hilum about 2/3 as long as the fruit.
Graebn., xAgropogon P. Fourn., and xAmmocala- 2n ¼ 14, 28, 56.
magrostis P. Fourn. These have not been verified Three spp., ocean dunes of Europe, North
by molecular data, but their existence shows the Africa, North America. Introduced elsewhere.
Hybridization with Calamagrostis has been flower pubescent, the hairs generally longer than
documented by Reznicek and Judziewicz (1996). the flower. Lemma generally more firm than
The hybrids are morphologically intermediate glumes, generally with 5 veins, the apex generally
between the two parents and apparently largely with two lobes or irregular teeth, awned from the
sterile. apex or from the abaxial side, the awn straight or
bent.
235. Ancistragrostis S.T. Blake Ninety-eight spp., worldwide in temperate
regions.
Ancistragrostis S.T. Blake, Blumea, Suppl. 3: 56 (1946). This genus is polyphyletic in the analyses of
Saarela et al. (2010), who indicate that Deyeuxia
Tiny perennials, <10 cm tall. Leaves filiform. should be excluded from “Calamagrostis”, a con-
Inflorescence branches branched. Spikelets with clusion followed here. Some New World species
a rachilla extension, the rachilla extension with currently classified as “Calamagrostis” appear to
long hairs. Glumes shorter than the spikelet, trans- belong instead to Trisetaria, q.v. (as Trisetum s.l.,
lucent, their apices acute. Lemma apex notched, Saarela et al. 2010). The number of species belong-
awned from the abaxial side; the awn hooked. ing to Calamagrostis s.s. is uncertain; the number
One sp., A. uncinioides S.T. Blake, New here is based on the number in GrassBase (Clayton
Guinea and Australia. et al. 2006 onward) excluding Deyeuxia, but Lu
(2006) suggests that there are only 20 species.
236. Bromidium Nees & Meyen
I have found no indication that any species of
“Calamagrostis” is diploid; all appear to be tetra-
Bromidium Nees & Meyen, Gramineae 22–23 (1841); tax.: ploid or higher-order polyploid. Polyploidy and
Rúgolo de Agrasar (1982). apomixis are well documented, suggesting that
“Calamagrostis” may be a genus entirely of allo-
Annuals or perennials, caespitose or rhizoma- polyploid origin, analogous to Elymus, so it is
tous. Inflorescence dense. Spikelets without a not surprising that the evolutionary history and
rachilla extension. Glumes longer than the flower, taxonomy are complex.
the apices unawned. Callus of flower with hairs
less than 1/3 the length of the lemma, evenly 238. Cyathopus Stapf
distributed or in two tufts. Lemma apex with
four teeth or awns, plus a central awn from the Cyathopus Stapf, Hooker’s Icon. Pl. 24: t. 2395 (1895).
abaxial side, the awn twisted, geniculate.
Five spp., South America. Tufted perennials. Inflorescence branches
Bromidium is sometimes included in Agrostis numerous at most nodes. Spikelets without a
but is kept separate here because of the distinc- rachilla extension, disarticulating below the
tive long awns on the lemma. No molecular data glumes. Glumes slightly longer than the flower,
are available. their apices extended into long tails. Lemma apex
acute, awnless.
237. “Calamagrostis” Adans. One sp., C. sikkimensis Stapf, Sikkim, Bhutan,
China.
“Calamagrostis” Adans., Fam. Pl. 2: 31, 530 (1763). Clayton et al. (2006 onward) indicate that the
primary inflorescence branches are whorled, but
Perennials, caespitose or rhizomatous. Spikelets this is unlikely. In other supposedly “whorled”
occasionally with 2 flowers; rachilla extension pooids, the primary branch complexes are on
present, glabrous. Glumes longer than the flower, either side of the inflorescence suggesting that
generally with one vein, their apices acute to the branches initiate in a distichous phyllotaxis
attenuate, muticous or rarely awned. Callus of as in all other pooids.
239. Deyeuxia Clarion ex P. Beauv. Fig. 46
Deyeuxia Clarion ex P. Beauv., Ess. Agrostogr. 43 (1812);

tax.: Phillips and Chen (2003).
Dichelachne Endl., Prodr. Fl. Ins. Norf.: 20 (1833).
Perennials, caespitose or rhizomatous, rarely

annual. Spikelets usually with a slender rachilla
extension. Glumes somewhat shorter or longer
than the flower, their apices acute to attenuate.
Callus of flower pubescent, the hairs from 1/3 the
length of the flower to about the same length.
Lemma apex ragged, with minute teeth, awned
from the abaxial side, the awn straight or genicu-
late. Stamens sometimes reduced to 2 or 1. 2n ¼
28, 42, 56, 70.
Two hundred-seven spp., temperate regions
worldwide, tropical mountains.
This genus appears to be entirely polyploid;
no diploids are listed either in the Flora of China
treatment (Lu et al. 2006) or in the Index to Plant
Chromosome Numbers. Species formerly recog-
nized as Dichelachne are reported to have 2n ¼ 70
(Beuzenberg and Hair 1983; Murray et al. 2005).
240. Echinopogon P. Beauv.
Echinopogon P. Beauv., Ess. Agrostogr.: 42, 148, 161

(1812).
Perennials, caespitose or rhizomatous. Spikelets

with a rachilla extension. Glumes as long as or
longer than the flower, their apices obtuse to
attenuate. Callus of flower pubescent. Lemma
apex entire or with two teeth or lobes, awned
from the apex or the abaxial side, the awn stiff,
the lobes also awned in some species. 2n ¼ 42. Fig. 46. Pooideae-Poeae-Agrostidinae. Deyeuxia zang-
Seven spp., New Guinea, Australia, and New xiensis. A Habit. B Ligule. C Spikelet. D Glumes. E Flower.
Zealand. F Anthers. (From Wu et al. 2007, with permission from
the Missouri Botanical Garden Press, St. Louis, and Sci-
ence Press, Beijing)
241. Gastridium P. Beauv.
ulate; awned and unawned lemmas in the same
Gastridium P. Beauv., Ess. Agrostogr.: 21, 164 (1812). inflorescence. 2n ¼ 14.
Two spp., Europe, North Africa, mid-East.
Annuals. Spikelets vase-shaped, with or without a Saarela et al. (2010) place Gastridium sister to
rachilla extension. Glumes inflated around the Triplachne.
fruit and narrowed above, then flaring distally,
hardened and enlarged proximally, membranous 242. Hypseochloa C. E. Hubb.
distally, their apices acuminate, unawned. Callus
of flower glabrous. Lemma apex with small teeth, Hypseochloa C. E. Hubb., Bull. Misc. Inform. Kew 1936:
muticous or with an abaxial awn, the awn genic- 300, f. 1 (1936).
Annuals. Spikelets with a rachilla extension. 245. Podagrostis (Griseb.) Scribn. & Merr.
Glumes longer than the flower, their apices acu-
minate. Callus of flower pubescent. Lemma Podagrostis (Griseb.) Scribn. & Merr., Contr. U.S. Natl.
becoming indurate and crustaceous at maturity Herb. 13: 58 (1910).
with involute margins, the apex with two teeth,
with an abaxial awn, the awn twisted and Perennials, caespitose or rhizomatous. Spikelets
geniculate. with or without a rachilla extension. Glumes lon-
Two spp., Mount Cameroon and Tanzania. ger than the flower, their apices acute to acumi-
Hubbard segregated Hypseochloa from Agros- nate. Callus of flower glabrous or with short hairs.
tis and Aira on the basis of its 5-veined glumes. Lemma with the apex truncate to acute, awned or
unawned, the awn abaxial. 2n ¼ 14.
Six spp., New World.
243. “Lachnagrostis” Trin. Podagrostis is distinguished from Agrostis
by the relatively long palea, and from “Calama-
“Lachnagrostis” Trin., Fund. Agrost.: 128, pl. 10 (1820); grostis” and “Lachnagrostis” by the short callus
tax.: Jacobs (2001).
hairs.
tous. Inflorescence often shed and dispersed as a 246. Polypogon Desf.
whole. Spikelets sometimes with 2 flowers; with a
rachilla extension. Glumes longer than the flower, Polypogon Desf., Fl. Atlant. 1: 66 (1798).
their apices acuminate, unawned. Callus of flower
pubescent, the hairs up to about 2/3 the length of Annuals or perennials. Spikelets without a
the lemma. Lemma with minute teeth at the apex, rachilla extension, disarticulating midway up the
unawned or with an abaxial awn, the awn straight pedicel. Glumes longer than the flower, their api-
or geniculate. 2n ¼ 56. ces entire or emarginate or bilobed, mucronate or
Twenty spp., Australasia. awned from the sinus or apex, the awn flexuous.
“Lachnagrostis” is polyphyletic in the ana- Lemma apex entire to lobed, muticous to awned,
lyses of Saarela et al. (2010) with some species the awn generally just below the apex or from the
related to Polypogon and others to Agrostis. sinus. 2n ¼ 14, 28, 35, 42, 56,
Jacobs (2001) distinguishes “Lachnagrostis” Twenty-six spp., worldwide in warm temper-
from Agrostis by its much better developed ate regions and on tropical mountains.
palea. He also compares the lemma epidermal Some species currently classified in “Lachna-
pattern of “Lachnagrostis” to that in Deyeuxia grostis” should perhaps be placed in or near Poly-
and Agrostis. pogon (Saarela et al. 2010).
244. Pentapogon R. Br. 247. Triplachne Link
Pentapogon R. Br., Prodr.: 173 (1810); tax.: MacFarlane Triplachne Link, Hort. Berol. 2: 241 (1833).
and Hearn (2007).
Annuals. Glumes longer than the flower, with one
Perennials. Spikelets without a rachilla extension. vein, their apices acute. Callus of flower pubes-
Glumes as long as the flowers, with 1 vein, their cent. Lemma with lateral awns on either side of a
apices acuminate, awned. Callus of flower pubes- central, abaxial awn. 2n ¼ 14.
cent. Lemma with margins convolute, covering One sp., T. nitens (Guss.) Link, Mediterra-
the palea, with four apical aristae and a fifth nean.
abaxial. Saarela et al. (2010) find Triplachne to be
One sp., P. quadrifidus (Labill.) Baill., SE strongly supported as sister to Gastridium.
Australia.
GROUP WITH POEAE-TYPE PLASTID SEQUENCES
This clade is retrieved in all molecular studies of

the plastid (e.g., Davis and Soreng 2007; Gillespie
et al. 2008; Soreng and Davis 2000; Soreng et al.
2007), but appears to have no morphological
synapomorphies.
Incertae sedis in Poeae-type plastid clade:
248. Antinoria Parl.
Antinoria Parl., Fl. Palerm. 1: 92 (1845).
Annuals or perennials. Spikelets with 2 flowers,

without a rachilla extension. Glumes longer than
the flower, their apices obtuse. Lemma apex trun-
cate, unawned, with three teeth. 2n ¼ 18.
Two spp., Mediterranean.
Antinoria is similar to Airopsis, but the flow-
ers are separated by a short internode. This genus
is sister to Loliinae in the analyses of Quintanar
et al. (2007) and to the Parapholiinae/Cynosuri-
nae clade in Inda et al. (2008); both placements
are based solely on an ITS sequence for A. agros-
tidea (DC) Parl.
XVI.6. SUBTRIBE SCOLOCHLOINAE Tzvelev (1987)
Perennials up to 2 m tall, with fleshy rhizomes.

Spikelets with 3 or 4 flowers; rachilla extension
present. Glumes shorter than or as long as the
flowers, their apices acute to acuminate. Callus of
flower pubescent, with stiff hairs. Lemma apex with
3 lobes or teeth, unawned. Ovary apex pubescent.
249. Scolochloa Link Fig. 47
Scolochloa Link, Hort. Berol. 1: 136 (1827).
Characters as for the subtribe. 2n ¼ 28.

One sp., S. festucacea (Willd.) Link, north
temperate regions.
Scolochloa looks superficially like Festuca.
XVI.7. SUBTRIBE AIRINAE Fr. (1835)
Annuals or perennials. Spikelets generally with

Fig. 47. Pooideae-Poeae-Scolochloinae. Scolochloa festu-
cacea. A Habit. B Spikelet. C Flower. (From Barkworth 2 flowers. Lemma generally with an abaxial awn,
et al. 2007, drawn by L.A. Vorobik) the awn geniculate.
Ten genera, 97 spp.
250. Aira L. Avenula is similar to Helictotrichon, but the

latter has ligules that are truncate to rounded, leaf
Aira L., Sp. Pl. 1: 63–66 (1753). blades that are ribbed, rachillas that are pubes-
cent on all sides, a long hilum, and solid endo-
Delicate tufted annuals. Spikelets generally <5 sperm. Avenula pubescens falls in conflicting
mm long, lacking a rachilla extension. Glumes positions in phylogenetic trees, and may be an
longer than the flowers, their apices obtuse to intergeneric hybrid (Gillespie et al. 2008; Soreng
acute, unawned. Callus of flower pubescent. and Davis 2000). It is probably more closely
Lemma apex bilobed. 2n ¼ 14, 28. related to Poinae than to Avenula s.s., but is
Eight spp., Europe, Mediterranean, Iran and retained here pending a better placement. The
weedy elsewhere in the world. closed sheaths are unusual for this subtribe.
Aira includes plants that are distinctively del-
icate, with capillary inflorescence branches and
pedicels. 253. Corynephorus P. Beauv.
Corynephorus P. Beauv., Ess. Agrostogr.: 90, 159 (1812),

251. Avenella (Bluff & Fingerh.) Drejer nom. conserv.
Avenella (Bluff & Fingerh.) Drejer, Fl. Excurs. Hafn.: 32 Annuals or perennials. Spikelets with a rachilla
(1837); tax.: Chiapella and Zuloaga (2010).
extension. Glumes hyaline, vase-shaped, longer
than the flowers. Callus of flower pubescent.
Caespitose perennials, with or without rhizomes.
Lemma apex acute, awned from the abaxial side
Spikelets with a pubescent rachilla extension.
just above the base, the awn articulated in the
Glumes as long as the flowers, their apices acute.
middle, twisted below, club-shaped above, with
Callus of flower pubescent. Lemma apex truncate
hairs at the connection between the two portions.
or toothed, the awn hair-like. 2n ¼ 28.
2n ¼ 14.
One sp., A. flexuosa (L.) Drejer, widespread in
Five spp., coastal dunes of Europe, Mediter-
temperate and cold regions worldwide, including
ranean, mid-East.
Antarctic islands.
The molecular analyses of Hoffmann et al.
Clayton and Renvoize (1986) synonymize this
(2013) place Corynephorus in Poinae, but on a
genus with Deschampsia, whereas Chiapella
very long branch. Because this placement contra-
(2007) presents molecular phylogenetic data sup-
dicts other information on the genus, it is
porting the distinction of the two and a close
retained here pending additional data.
relationship instead with Aira.
252. Avenula (Dumort.) Dumort. 254. Deschampsia P. Beauv. Fig. 48
Avenula (Dumort.) Dumort., Bull. Soc. Roy. Bot. Belgique Deschampsia P. Beauv., Ess. Agrostogr.: 91–92, pl. 18, f. 3
7: 68 (1868); cytogenetics: Winterfeld (2006); tax.: Lange (1812); tax.: Chiapella and Zuloaga (2010).
(1995); phylog.: Winterfeld et al. (2012).
Helictotrichon subg. Pratavenastrum (Vierh.) Holub, Phi- Annuals or perennials, caespitose. Spikelets gen-
lipp Maxm. Opiz Bedeut. Pfl.-tax.: 125 (1958). erally with a pubescent rachilla extension.
Glumes generally longer than the spikelet, their
Perennials, caespitose or stoloniferous. Sheaths apices acute to acuminate, muticous or mucro-
open or closed. Spikelets with 2 to 7 flowers; nate. Callus of flower pubescent. Lemma apex
rachilla glabrous when adjacent to the palea, truncate or toothed, the abaxial awn hair-like,
pubescent elsewhere. Glumes as long as or longer straight or geniculate and twisted. 2n ¼ 26, 52.
than the flowers. Callus of flower pubescent. Forty spp., temperate areas worldwide.
Lemma apex obtuse, with two teeth, awned, the Barkworth (2007a) cites various aneuploid
awn from about ½ way down the abaxial side. chromosome counts for several species but
Endosperm liquid. 2n ¼ 14, 28. notes that the voucher specimens have not been
Thirty spp., Eurasia, North Africa, North checked. These counts are not listed here.
America.
256. Molineriella Rouy
Molineriella Rouy, Fl. France 14: 102 (1913).
Annuals. Spikelets without a rachilla extension.

Glumes shorter than or about the same length as
the flowers, their apices obtuse to acute. Callus of
flower pubescent. Lemma apex obtuse, unawned.
2n ¼ 8.
One sp., M. minuta (L.) Rouy, Mediterranean,
Australasia.
257. Periballia Trin.
Periballia Trin., Fund. Agrost.: 133 (1820).
Annuals. Spikelets without a rachilla extension.

Glumes shorter than or about the same length as
the flowers, their apices acute. Callus of flower
pubescent. Lemma awned from the abaxial side,
the awn straight, or the apex acute, mucronate. 2n
¼ 8, 14, 18.
Two spp., Mediterranean.
258. Scribneria Hack.
Scribneria Hack., Bot. Gaz. 11: 105 (1886).
Fig. 48. Pooideae-Poeae-Airinae. Deschampsia antarc- Tufted annuals. Leaf blades filiform. Inflorescence
tica. A Plant. B Spikelet, lateral view. C Flowers. D Base unbranched, the spikelets distichous and sunken
of the flower, viewed from the side of the palea and in the rachis, generally one per node, but some-
rachilla. E Palea, internal face, and lodicules. F Caryopsis,
scutellar view. G Same, hilar view. (From Zuloaga et al.
times up to 4; rachis not disarticulating. Spikelets
2012) with 1 flower; rachilla extension present. Glumes
longer than the flower, coriaceous, unawned. Cal-
255. Holcus L. lus of flower pubescent. Lemma apex with two
teeth, awned from the sinus. Stamen 1. 2n ¼ 26.
Holcus L., Sp. Pl. 2: 1047 (1753), nom. conserv. One sp., S. bolanderi (Thurb.) Hack., western
U.S.
Perennials, rarely annual, caespitose or rhizoma- Scribneria is placed here on the basis of the
tous. Spikelets disarticulating below the glumes, data of Schneider et al. (2012).
with the proximal flower(s) bisexual and the dis-
tal one staminate. Glumes as long as or longer 259. Vahlodea Fr.
than the flowers, keeled, their apices obtuse to
acuminate, unawned. Callus of flower glabrous Vahlodea Fr., Bot. Not. 1842: 141, 178 (1842); tax.: Chia-
or pubescent. Lemma firm, shiny, the apex acute pella and Zuloaga (2010).
or bidentate, the proximal lemma in the spikelet
unawned, the distal one with an abaxial awn. 2n ¼ Caespitose perennials, the leaves mostly cauline.
14, 28 (35, 42, 49). Spikelets with a rachilla extension. Glumes as
Eight spp., Europe, north Africa, mid-East. long as or longer than the flowers, their apices
See comments under Vahlodea. acute to acuminate. Callus of flower pubescent,
the hairs about ½ the length of the lemma.

Lemma apex with minute irregular teeth, the
awn hair-like, twisted, geniculate. Hilum linear.
2n ¼ 14.
One sp., V. atropurpurea (Wahlenb.) Fr. ex
Hartm., temperate regions throughout the world.
Clayton and Renvoize (1986) synonymize
Vahlodea with Deschampsia, whereas molecular
phylogenetic data (Chiapella 2007; D€oring et al.
2007; Schneider et al. 2009) support the distinc-
tion of the two, and place Vahlodea sister to
Holcus with strong support. In Vahlodea the
leaves are mostly cauline, whereas they are largely
basal in Deschampsia. The rachilla extension is
less than 0.5 mm long in Vahlodea and is gla-
brous, whereas it is generally more than 0.5 mm
long in Deschampsia and is pubescent.
XVI.8. SUBTRIBE AMMOCHLOINAE Tzvelev (1976)
Annuals, often tiny. Inflorescence capitate, sur-

rounded by leaf sheaths in some species. Spikelets
with 4 to 14 flowers plus additional reduced flow-
ers distally or a barren rachilla extension. Glumes
shorter than the flowers, sometimes each with an
asymmetrical wing on the keel, their apices
obtuse to acute, muticous to mucronate. Lemma
with broad margins, the apex entire or with two
teeth, mucronate or awned, the awn from the
apex or sinus. Lodicules absent. Base of style
branches persistent in fruit, forming a beak.
260. Ammochloa Boiss. Fig. 49
Ammochloa Boiss., Diagn. Pl. Orient. ser. 1. 2(13): 51

(1854).
Characters as for the subtribe.

Three spp., Mediterranean and mid-East.
In A. pungens the proximal 1 or 2 spikelets
are sterile.
Fig. 49. Pooideae-Poeae-Ammochloinae. Ammochloa. A

XVI.9. SUBTRIBE CYNOSURINAE Fr. (1835) A. palaestina. Habit. (From Boulos 2005). B–F Ammo-
chloa involucrata. B Capitulum of spikelets subtended
by an involucre. C Spikelet with two flowers. D Lower
Annuals or perennials, caespitose or rhizoma- glume viewed from the abaxial side. E Upper glume
tous. Inflorescence unbranched or the branches viewed from the abaxial side. F Caryopsis with stylopo-
branched, with paired spikelets, the proximal dium and stigmas. (From Murbeck 1899)
261. Cynosurus L. Fig. 50
Cynosurus L., Sp. Pl. 1: 72–73 (1753).
Characters as for the subtribe.

Ten spp., Europe and Mediterranean.
XVI.10. SUBTRIBE LOLIINAE Dumort. (1829)
Annuals or perennials. Leaf sheaths with or with-

out auricles. Spikelets with more than one flower.
Glumes generally shorter than the adjacent lemma.
Eight genera, 543 spp.
Loliinae are dominated by the large genus
Festuca, the circumscription of which varies
extensively in the literature. Molecular phyloge-
nies show that Festuca as traditionally defined is
paraphyletic, with many smaller genera derived
from within it (Catalán et al. 2004; Inda et al.
2008; Torrecilla and Catalán 2002; Torrecilla
et al. 2004). Catalán et al. (2004) identified a
large clade that they called the “the fine-leaved
fescues” and another “the broad-leaved fescues”;
these clades include most, but not all species
conventionally recognized as Festuca. The classi-
fication adopted here is influenced by the phylog-
eny, recognizing the “fine-leaved fescues” as
Festuca s.s. This clade includes Vulpia, which
has long been known to be related to Festuca, as
well as several other smaller genera. The “broad-
leaved fescues” include Lolium, also well known
as a close relative of species formerly classified as
Festuca; this clade thus becomes Lolium s.l. The
remaining species of Festuca fall into one of two
Fig. 50. Pooideae-Poeae-Cynosurinae. Cynosurus echina-
tus. A Habit. B Inflorescence. C Pair of spikelets, fertile poorly supported grades, here listed as the “inter-
spikelet with two spread glumes (right), and sterile spikelet mediate grade” and the “Leucopoa grade.” I have
with multiple glumes (left). D Lemma, at two different mag- not attempted to dismember these into subgroups.
nifications. (From Barkworth et al. 2007, drawn by S. Long) There are several reasonably well-supported sub-
clades within each of the grades, but all require
spikelet of each pair made up of a cluster of sterile additional taxon sampling and sequence data. A
lemmas, generally abaxial to and obscuring the more radical treatment than that adopted here
fertile distal spikelet. Proximal sterile spikelet would be to place the entire subtribe in a single
with 6 to 18 flowers, the glumes and lemmas genus, although even with this solution the genus
narrow, sometimes awned. Distal fertile spikelet would lack any obvious diagnostic characters.
with 1 to 5 flowers; rachilla extension present,
glabrous. Glumes shorter than the flowers, their 262. Incertae sedis – “intermediate grade” of Lolii-
apices acute to attenuate, sometimes awned. Cal- nae
lus of flower glabrous. Lemma apex entire or with
two teeth, unawned or awned from the apex or Festuca sect. Subulatae Tzvelev, Ukrain. Bot. Rev. 56(9):
sinus. 2n ¼ 14. 1253 (1971).
Festuca sect. Subuliflorae (E.B. Alexeev) Darbysh., Phyto- branches borne along a central axis. Spikelets
logia 82(2): 77 (1997). broadside to the rachis, with 6 to 12 flowers
Festuca sect. Amphigenes (Janka) Tzvelev, Bot. Žurn.
(Kiev) 56(9): 1253 (1971).
plus additional reduced flowers distally.
Glumes with apices obtuse to acute. Lemma
Catalán et al. (2004) and Inda et al. (2008) identify tuberculate, the apex obtuse to acute, unawned.
a grade that they called “intermediate”, which is a 2n ¼ 14.
set of poorly supported lineages sister to Festuca One sp., C. tuberculosa (Moris) Bor, Canary
s.s. (the fine-leaved fescues). Their analysis Is., Mediterranean, Sudan, Somalia, Pakistan.
includes representatives of the three sections Catalán et al. (2004) and Inda et al. (2008)
listed above, plus Castellia. I have not attempted place Castellia in a position between the broad-
a diagnostic description of members of this grade. leaved and fine-leaved fescues.
265. Dryopoa Vickery

263. “Leucopoa grade”
Dryopoa Vickery, Contr. New South Wales Natl. Herb. 3:
“Leucopoa grade” 195 (1963).
Leucopoa Griseb., Fl. Ross. 4: 383 (1852).
Drymochloa Holub., Folia Geobot. Phytotax. 19: 96 (1984).
Festuca subg. Drymanthele V.I. Krecz. & Bobrov, Fl. URSS Perennials. Spikelets with 3 to 7 flowers plus
2: 532 (1934). additional reduced flowers distally. Glumes with
Festuca sect. Montanae Hack., Monogr. Festuc. Eur. 195 apices acute. Lemmas with prominent raised
(1882). veins, the apex truncate, ragged, unawned.
Festuca subg. Obtusae E.B. Alexeev, Novosti Sist. Vyssh.
Rast. 17: 45 (1980).
Ovary apex pubescent.
Festuca sect. Scariosae Hack., Monogr. Festuc. Eur.: 193 One sp., D. dives (F. Muell.) Vickery, SE Aus-
(1882). tralia.
Festuca sect. Pseudoscariosa Krivot., Bot. Mater. Gerb. While chloroplast data place Dryopoa near or
Bot. Inst. Komarov.Akad. Nauk. SSSR 20: 61 (1960). in Loliinae (Gillespie et al. 2008), data from the
nuclear internal transcribed spacer (ITS) and
Perennials, generally caespitose, rhizomatous or external transcribed spacer (ETS) of the ribo-
not; plants dioecious or bisexual. Leaves rela- somal RNA place it sister to Agrostidinae (Birch
tively broad and flat, with sclerenchyma girders et al. 2014).
adaxial and abaxial to the veins. Auricles absent.
Spikelets with 2 to 10 flowers. Glumes shorter
than the flowers, their apices acute to acuminate, 266. Festuca L. Fig. 51
unawned. Callus of flower generally wider than
Festuca L., Sp. Pl. 1: 73–76 (1753).
long, glabrous. Lemma apex acute to attenuate, Wangenheimia Moench, Methodus: 200 (1794).
mucronate to awned. Apex of ovary densely Vulpia C. C. Gmel., Fl. Bad. 1: 8 (1805); rev.: Cotton and
pubescent, rarely glabrous. 2n ¼ 28, 42, 56. Stace (1967); Lonard and Gould (1974).
Thirty-seven spp., temperate regions. Micropyrum (Gaudin) Link, Linnaea 17: 397 (1844).
Lolium s.l. is derived from within a largely Ctenopsis De Not., Index Sem. Hort. Genuensis: 26 (1847).
Narduroides Rouy, Fl. France 14: 301 (1913).
unresolved grade including Leucopoa and several Hellerochloa Rauschert, Taxon 31: 561 (1982).
sections formerly assigned to Festuca (Catalán Dielsiochloa Pilg., Bot. Jahrb. Syst. 73: 99 (1943).
et al. 2004; Inda et al. 2008). I am calling this
simply “the Leucopoa grade”, pending further Annuals (some very tiny) or perennials, caespi-
study. Festuca subg. Montanae presumably falls tose or rhizomatous. Leaves generally slender.
here, based on its leaf anatomy, although its spe- Inflorescence unbranched or with branched
cies have not been placed phylogenetically. The branches. Spikelets with 1 to 11 (17) flowers, or
description offered here is only approximate. pseudoviviparous in a few species. Glumes
shorter than the flowers, their apices acuminate,
264. Castellia Tineo awned or unawned. Callus of flowers often wider
Castellia Tineo, Pl. Rar. Sicil. 2: 17 (1846). than long, pubescent or glabrous. Lemma apex
acute to acuminate, entire or sometimes minutely
Annuals. Leaf sheaths with falcate auricles. bidentate, mucronate to awned. Apex of ovary
Inflorescence unbranched, or of unbranched
several annual species each of which has some

distinctive morphological characteristic that led
it to be placed in its own genus at one time. These
genera are clearly all derived from within Festuca
and the distinctive characters are phylogeneti-
cally uninformative autapomorphies. While the
group recognized here as Festuca is clearly mono-
phyletic, it lacks any obvious morphological
synapomorphies. Darbyshire and Pavlick (2007)
describe patterns of leaf cross-sectional anatomy
that may be diagnostic for Festuca as circum-
scribed here, although comparable data are not
available for the segregate genera, which need to
be checked.
267. Lolium L.
Lolium L., Sp. Pl. 1: 83 (1753); tax.: Charmet et al. (1996);

Darbyshire (1993).
Schedonorus P. Beauv., Ess. Agrostogr.: 99, 162, 177
(1812).
Festuca subg. Schedonorus (P. Beauv.) Peterm., Deutschl.
Fl.: 643 (1849).
Micropyropsis Romero Zarco & Cabezudo, Lagascalia 11:
95 (1983).
Festuca sect. Subbulbosae Nyman ex Hack., Bot. Cen-
tralbl. 8: 413 (1881), p.p.
Annuals or perennials, caespitose, sometimes rhi-

zomatous. Leaf sheath with or without falcate
auricles. Inflorescence unbranched, or with
branched branches; if unbranched then spikelets
edgewise to the rachis. Spikelets with 2 to 22
flowers. Glumes generally shorter than the flow-
ers, their apices acute, unawned; in unbranched
Fig. 51. Pooideae-Poeae-Loliinae. Festuca ovina L. A
Habit. B Cross section of leaf blade. C Spikelet. D Flower. inflorescences the lower glume lacking except on
(From Wu et al. 2007, with permission from the Missouri the terminal spikelet, upper glume on side away
Botanical Garden Press, St. Louis, and Science Press, from rachis. Callus of flower glabrous or with
Beijing) sparse hairs. Lemma apex acute, awned or
unawned, the awn straight. Apex of ovary gla-
glabrous or pubescent. Stamens 1 or 3. 2n ¼ 14, brous. Hilum linear. 2n ¼ 14, 28, 42, 56, 63, 70.
28, 42, 44, 49, 50, 56, 63, 70. Twenty-six spp., temperate regions of Eura-
Ca. 430 spp., temperate regions of North sia; widely introduced as a lawn and forage grass.
America and Eurasia. Inclusion of Schedonorus in Lolium is sup-
Festuca as circumscribed here corresponds to ported by morphological and molecular data
the “fine-leaved fescues” (Catalán et al. 2004; Inda (Catalán et al. 2004); as noted by Darbyshire
et al. 2008; Torrecilla and Catalán 2002; Torrecilla (2007), the two “could appropriately be treated
et al. 2004), including taxa formerly classified as as congeneric subgenera . . .”. The characters
Festuca subgenus Festuca sects. Festuca, Aulaxy- shared by Lolium and Schedonorus and not with
per Dumort., Eskia Willk., and Dimorphae Joch. Festuca s.s. include the falcate leaf auricles, flat,
M€ull. & Catalán. In addition the group contains wide leaf blades, ovaries that are glabrous at the
apex, stylar attachment abaxial, below the apex of

the ovary, and fruit that adheres firmly to the
palea. Hybridization between species of Lolium
s.s. and S. arundinaceus (¼ Festuca arundinacea)
is common, further supporting the close relation-
ship between the two.
268. Loliolum V. I. Krecz. & Brobov.
Loliolum V.I. Krecz. & Bobrov, Fl. URSS 2: 544, 766

(1934).
Annuals. Inflorescence unbranched, the spikelets

all borne on one side. Spikelets wedge-shaped,
with 3 to 7 flowers plus additional reduced
flowers distally. Glumes longer than the flowers,
their apices attenuate, mucronate. Lemma apex
notched to truncate to acute, muticous or with a
short awn.
One sp., L. subulatum (Banks & Sol.) Eig.,
mid-East to central Asia.
No molecular data are available for L. subu-
latum, and it may be misplaced here. It has also
been placed in Festuca and Nardurus, the latter
now a synonym of Festuca.
269. Pseudobromus K. Schum.
Pseudobromus K. Schum., Pflanzenw. Ost-Afrikas C: 108

(1895).
Perennials. Leaf blades with transverse veins. Spi-

kelets with 1 to 4 flowers, the rachilla terminating
in a rudiment. Glumes shorter than the flowers, Fig. 52. Pooideae-Poeae-Dactylidinae. Dactylis glomer-
their apices acuminate. Callus of flower puberu- ata. A Plant. B Ligule area. C Open panicle. D Closed
lous. Lemma awned, the awn from the apex or panicle. E Spikelet. F Flower. (From Barkworth et al.
2007, drawn by L.A. Vorobik and K. Klitz)
just below. Palea longer than the lemma.
Six spp., tropical Africa and Madagascar.
No molecular data are available for Pseudo- 270. Dactylis L. Fig. 52
bromus; it is placed here based on morphology.
Dactylis L., Sp. Pl. 1: 71 (1753); tax.: Stewart and Ellison
(2011).
XVI.11. Subtribe DACTYLIDINAE Stapf (1898)
Perennials, caespitose or short rhizomatous.
Leaf sheaths closed for part of their length. Spike- Spikelet clusters borne on one side of the inflores-
lets borne in dense clusters. cence branches, the branches naked proximally.
Dactylidinae are strongly supported as mono- Spikelets with 2 to 6 flowers; rachilla extension
phyletic by molecular data (Catalán et al. 2004; lacking. Glumes shorter than the flowers, their
Inda et al. 2008). apices acute, mucronate or awn-tipped. Callus of
flower glabrous. Lemma apex acuminate, with a than ½ the length of the flowers, hyaline, their
short awn. Hilum round. 2n ¼ 14, 21, 27–31, 42. apices obtuse or the upper one acute. Lemma
Five spp., temperate Eurasia, widely intro- apex truncate or obtuse, muticous to mucronate.
duced elsewhere as a forage grass. 2n ¼ 12, 14, 28.
Two spp., Mediterranean and mid-East.
Sphenopus is sister to the rest of the subtribe
271. Lamarckia Moench
in molecular phylogenies (Inda et al. 2008; Soreng
Lamarckia Moench, Methodus: 201 (1794). et al. 2007).
Remaining genera of the subtribe:
Annuals. Inflorescence with all branches borne Inflorescence generally unbranched, often disarti-
on one side, secondary inflorescence branches culating at the nodes.
ending in clusters of spikelets, bent and disarticu-
lating below the clusters. Proximal spikelets 273. Agropyropsis (Trab.) A. Camus
within each cluster sterile, with 5 to 10 flowers,
distal spikelet fertile, with 2 flowers, the proximal Agropyropsis (Trab.) A. Camus, Bull. Soc. Bot. France 82:
flower bisexual, on an elongate rachilla internode, 11 (1935).
the distal flower reduced to an awned rudiment.
Glumes as long as or longer than the flower, their Perennials. Inflorescence unbranched, disarticu-
apices acuminate, mucronate to awned. Callus of lating at the nodes. Spikelets distichous, with 3 to
flower glabrous. Lemma apex with two minute 6 flowers plus additional reduced flowers distally.
teeth, awned from the abaxial side. 2n ¼ 14. Glumes borne side by side on the abaxial side of
One sp., L. aurea (L.) Moench, Mediterranean the spikelet, shorter than the flowers, their apices
and Middle East. obtuse. Callus of flower glabrous. Lemma apex
obtuse, unawned. Ovary with a conspicuous api-
cal appendage.
One sp., A. lolium (Balansa ex Coss. & Dur-
XVI.12. SUBTRIBE PARAPHOLIINAE Caro (1982)
ieu) A. Camus, Algeria.
Schneider et al. (2012) argue that Agropyrop-
Annuals or perennials. Inflorescences branched
sis should be placed in Loliinae, based on mor-
or unbranched, often disarticulating at the nodes.
phology. No molecular data are available, so the
Spikelets with one to many flowers. Glumes
genus is retained here for the time being.
shorter or longer than the flowers.
Many members of Parapholiinae were for- 274. Cutandia Willk.
merly placed in the tribe Hainardieae (Clayton
and Renvoize 1986), a group that has since been Cutandia Willk., Bot. Zeitung (Berlin) 18: 130 (1860).
shown to be polyphyletic (Schneider et al. 2012).
Narduroides is now included in Festuca s.s., Pho- Annuals. Inflorescence branches branched, disar-
liurus is in Poinae, whereas Scribneria is in Air- ticulating at the base of the branches, pedicels or
inae, unrelated to the others. The remainder form flowers. Spikelets with 2 to 9 flowers. Glumes
Parapholiinae. The subtribe includes the named shorter than the flowers, their apices acute to
hybrid xHainardiopholis Castrov. rounded, awned or not. Callus of flower glabrous.
Lemma keeled, with 3 conspicuous veins, the
apex rounded to bifid to acuminate and entire,
272. Sphenopus Trin. awned or awnless. 2n ¼ 14.
Six spp., Mediterranean and western Asia,
Sphenopus Trin., Fund. Agrost.: 135 (1820).
one species introduced to North America.
Suggestions for placing this genus near Vul-
Annuals. Leaf blades filiform. Inflorescence
pia (Catalán et al. 2004; Torrecilla et al. 2004)
branches branched. Spikelets with 2 to 6 flowers
were erroneous (Inda et al. 2008). It is clearly
plus additional reduced flowers distally; pedicels
placed here, sister to Desmazeria.
notably wider toward their apices. Glumes less
275. Desmazeria Dumort.
Desmazeria Dumort., Comment. Bot.: 26 (1822).

Catapodium Link, Hort. Berol. 1: 44 (1827).
Scleropoa Griseb., Spic. Fl. Rumel. 2: 431 (1846).
Annuals. Inflorescences branched or unbran-

ched, one-sided, not disarticulating at the nodes.
Spikelets with 4 to 25 flowers, the distal ones
reduced. Glumes shorter than the flowers, their
apices obtuse to acute. Callus of flower glabrous.
Lemma glabrous or pubescent with capitate hairs,
the apex obtuse to acute, with or without a mucro.
2n ¼ 14.
Eight spp., Mediterranean.
276. Hainardia Greuter Fig. 53
Hainardia Greuter, Boisseria 13: 178 (1967).
Annuals. Culm internodes solid. Inflorescence

unbranched, disarticulating at the nodes. Spike-
lets embedded in the inflorescence internodes,
with 1 or 2 flowers; rachilla extension present.
Lower glume absent except on the terminal spike-
let, upper glume longer than the flowers, leathery,
ribbed, the apex acute. Lemma membranous, the
apex acuminate, unawned. Stamens 1 or 3. Fruit
with an apical appendage. 2n ¼ 26.
One sp., H. cylindrica (Willd.) Greuter,
coastal areas of the Mediterranean.
Hainardia is strongly supported as sister to
Parapholis (Catalán et al. 2004; Schneider et al.
2012). The solid culm internodes for this species
are unusual in the Pooideae. The name Monerma
is sometimes misapplied to this genus; the latter
is a superfluous name for the chloridoid genus
Lepturus.
277. Parapholis C. E. Hubb.
Parapholis C. E. Hubb., Blumea, Suppl. 3: 14 (1946); tax.:

Runemark (1962).
Annuals, tufted. Inflorescence unbranched, dis-

articulating at the nodes. Spikelets embedded in
the inflorescence internodes, with one flower;
Fig. 53. Pooideae-Poeae-Parapholinae. Hainardia cylin- rachilla extension present or absent. Glumes
drica. A Plant. B Spikelet, with rachis internode (right) borne side by side on the abaxial side of the
and upper glume (left) enclosing the flower (tip visible). C spikelet, generally longer than the flower, coria-
Flower concealed by upper glume. (From Barkworth et al.
2007, drawn by L.A. Vorobik and H. Paźdirková) ceous, their apices acute to acuminate. Lemma
translucent, the apex obtuse to acute. 2n ¼ 14, apex as narrow lobes or awns. Palea with two
18, 38. awns. 2n ¼ 14, 18.
Six spp., coastal areas of Europe, Mediterra- One sp., E. capitata (L.) Desf., Mediterranean
nean, and the mid-East. and mid-East.
278. Vulpiella (Batt. & Trab.) Burollet 280. Mibora Adans.
Vulpiella (Batt. & Trab.) Burollet, Ann. Serv. Bot. Tunisie Mibora Adans., Fam. Pl. 495 (1763).
4: 68 (1927).
Tiny annuals, less than 15 cm tall. Inflorescence
Annuals. Spikelets with 5 to 18 flowers plus addi- unbranched, the spikelets imbricate, borne on
tional reduced flowers distally. Glumes shorter short pedicels on one side of the rachis. Spikelet
than the flowers, their apices acute, mucronate. with one flower; rachilla extension absent.
Lemma apex with two teeth, awned from the sinus. Glumes longer than the flower, their apices trun-
One sp., V. stipoides (L.) Maire, Western cate or obtuse. Callus of flower glabrous. Lemma
Mediterranean. translucent, pubescent, the apex truncate and
ragged. Palea pubescent. Lodicules absent. Style
1, stigmas 2. 2n ¼ 14.
XVI.13. SUBTRIBE SESLERIINAE Parl. (1845) Two spp., Western Europe and North Africa.
Miborinae Asch. & Graebn., Syn. Mitteleur. Fl. 2: 118

(1899). 281. Oreochloa Link
Annuals or perennials, most less than 40 cm tall. Oreochloa Link, Hort. Berol. 1: 44 (1827).
Leaf sheaths with fused margins. Inflorescence
dense, one-sided or capitate. Perennials, less than 40 cm tall. Inflorescence
Four genera, 33 spp. unbranched, one-sided, capitate. Spikelets with
This subtribe is placed within Aveninae in 3 to 7 flowers plus additional reduced flowers
ITS analyses, whereas it clearly has a plastid sim- apically. Glumes hyaline, shorter than the flowers,
ilar to that of Poinae (Quintanar et al. 2007; Sor- their apices acute. Callus of flower glabrous.
eng et al. 2007; Gillespie et al. 2008). Although the Lemmas hyaline, their apices acute, mucronate.
discrepancy between the ITS and plastid data Four spp., southern Europe.
suggests a hybrid origin for the tribe, the low
chromosome numbers indicate that polyploidiza-
282. Sesleria Scop. Fig. 54
tion may not be involved. Both Quintanar et al.
(2007) and Saarela et al. (2010) show that Mibora Sesleria Scop., Fl. Carniol.: 189 (1760); cytol.,: Lysak and
is derived within Sesleriinae. A few species of Dolezel (1998).
Sesleria reach 60 or 70 cm tall, but most members
of this subtribe are tiny. Perennials, caespitose, rhizomatous or stolonifer-
ous. Inflorescence spike-like or capitate, often
279. Echinaria Desf. with two bracts or scales at the base. Spikelets
with 2 to 5 flowers; rachilla extension present.
Echinaria Desf., Fl. Atlant. 2: 385 (1799). Glumes shorter than the flowers, their apices
acute to acuminate, muticous or awned. Callus
Annuals, less than 25 cm tall. Inflorescence capi- of flower glabrous, broadly rounded. Lemma
tate, prickly. Spikelets with 2 to 4 flowers plus apex entire or with 3 to 5 teeth, the teeth generally
additional reduced flowers distally. Glumes awned. Apex of ovary pubescent. 2n ¼ 28, 56.
shorter than the flowers, their apices obtuse, Twenty-six spp., Europe.
lower glume with two keels, upper glume with a The bracts at the base of the inflorescence are
short mucro. Callus of flower pubescent. Lemma generally interpreted as sterile spikelets (Clayton
leathery, with 5 to 7 veins extending beyond the and Renvoize 1986).
283. Catabrosa P. Beauv.
Catabrosa P. Beauv., Ess. Agrostogr.: 97 (1812); tax.: Sor-

eng and Fish (2011).
Perennials, sometimes stoloniferous. Leaves with

sheath margins fused most of their length. Spike-
lets with 1 to 3 flowers; rachilla extension present.
truncate to obtuse. Callus of flower glabrous.
Lemma with 3 prominent veins, the apex rounded
to truncate. 2n ¼ 20, 30.
Three spp., north temperate regions and Chile.
284. Catabrosella (Tzvelev) Tzvelev
Catabrosella (Tzvelev) Tzvelev, Bot. Žurn. (Moscow &

Leningrad) 50: 1320 (1965).
Caespitose perennials, basal leaves often with

sheaths persisting and forming bulb-like struc-
tures. Leaves with sheath margins fused 1/6 to 1/
4 of their length. Spikelets with 2 to 4 flowers.
Glumes much shorter than the flowers, the apices
obtuse to subacute. Callus of flower glabrous or
pubescent. Lemma apex obtuse. 2n ¼ 10.
Seven spp., Europe to the Himalayas.
285. Coleanthus Seidl Fig. 55
Coleanthus Seidl, Syst. Veg. 2: 11, 276 (1817).
Fig. 54. Pooideae-Poeae-Seslerinae. Sesleria autumnalis.

A Habit. B Inflorescence. C Spikelet. D Lemma. (From Tiny tufted annuals, mostly less than 7 cm tall.
Barkworth et al. 2007, drawn by A. Miller) Leaves with sheath margins fused most of their
length. Inflorescence dense, with spikelets in cap-
itate clusters, surrounded by the upper sheath.
XVI.14. SUBTRIBE COLEANTHINAE Rouy (1913) Spikelets with one flower, without rachilla exten-
sion. Glumes absent. Callus of flower glabrous.
Subtribe Puccinelliinae Soreng & J. I. Davis, Contr. U.S. Lemma translucent, the apex acute, awned. Lodi-
Natl. Herb. 48: 721 (2003).
cules absent. Stamens 2. 2n ¼ 14.
Annuals or perennials, tufted or rhizomatous. One sp., C. subtilis (Tratt.) Seidl, Central Eur-
Leaf sheaths often partially to completely closed. ope, northern Asia, and North America.
Spikelets with one to several flowers; awns absent.
Glumes often with obtuse apices, the veins prom- 286. Hyalopoa (Tzvelev) Tzvelev
inent or obscure, approximately parallel and not
reaching the apex. Lemma apex obtuse to acute, Hyalopoa (Tzvelev) Tzvelev, Bot. Žurn. (Moscow &
often scarious, veins approximately parallel and Leningrad) 50: 1320 (1965).
not reaching the apex.
Twelve genera, 156 spp. Rhizomatous perennials. Leaves with sheath mar-
Named hybrid: x Pucciphippsia Tzvelev gins fused 1/3 to 3/4 of their length. Spikelets with
2 to 4 flowers. Glumes shorter than flowers. Callus

of the flower pubescent. Lemma weakly keeled,
the apex obtuse to acute, sometimes with a sharp
point. 2n ¼ 28, 42.
Four spp., Central Asia.
Although Clayton and Renvoize (1986) place
this in the synonymy of Colpodium, Kim et al.
(2009) and Hoffmann et al. (2013) suggest it is
more closely related to Catabrosa, Paracolpo-
dium, and Catabrosella.
287. Paracolpodium (Tzvelev) Tzvelev
Paracolpodium (Tzvelev) Tzvelev, Bot. Žurn. (Moscow &

Leningrad) 50: 1320 (1965).
Rhizomatous perennials. Leaves with sheath mar-

gins fused ¼ to almost their entire length. Spike-
lets with one flower, rarely with a reduced flower
distally. Glumes almost as long as the flower.
Callus of flower glabrous or pubescent. Lemma
apex acute, the veins not reaching the apex. Fruit
with a small beak. 2n ¼ 28, 42.
Four spp., Caucasus to Himalayas.
Kim et al. (2009) show that Paracolpodium is
phylogenetically distinct from Colpodium, and
may be sister to Catabrosella.
288. Phippsia (Trin.) R. Br.
Phippsia (Trin.) R. Br., Chlor. Melvill.: 27 (1823); tax.:

Aares et al. (2000).
Tiny perennials, caespitose or mat-forming. Fig. 55. Pooideae-Poeae-Coleanthinae. Coleanthus subti-

lis. A Habit. B Single flowering shoot. C Spikelet with
Sheaths of culm leaves closed at least ½ their protruding caryopsis. D Lemma. E Palea. F Stamens. G
length. Spikelets with one flower; rachilla exten- Pistil. (From Wu et al. 2007, with permission from the
sion absent. Glumes less than 1/3 the length of the Missouri Botanical Garden Press, St. Louis, and Science
flower, without veins, their apices obtuse to acute; Press, Beijing)
lower glume sometimes absent. Callus of flower
glabrous. Lemma translucent to membranous, 289. Pseudosclerochloa Tzvelev
the apex truncate to acute. Stamens 1 or 2. Fruit
extending beyond the flower at maturity. Pseudosclerochloa Tzvelev, Bot. Žurn. (Moscow & Lenin-
Three spp., Arctic, Argentine Andes. 2n ¼ 28. grad) 89: 840 (2004).
Phippsia will hybridize with Puccinellia
(Steen et al. 2004). The species of Phippsia appear Annuals. Leaves with sheaths open. Spikelets with
to be allopolyploids but the parents are unknown 2 to 7 flowers, the distalmost ones reduced.
(Aares et al. 2000). Glumes shorter than the spikelet, their apices
obtuse to acute. Callus of flower pubescent. of flower glabrous. Lemma hardened, with 7 to
Lemma distally keeled, the apex obtuse. 9 prominent veins, the apex obtuse. Base of
Two spp., western Europe, China. style branches persistent in fruit, forming a beak.
These species have been segregated from Puc- 2n ¼ 14.
cinellia on the basis of their keeled lemmas. Two spp., southern Europe and mid-East,
weedy elsewhere.
Beaked fruits also appear in Duthiea, but
290. Puccinellia Parl.
have been acquired independently. S. dura (L.)
Puccinellia Parl., Fl. Ital. 1: 366 (1848), nom. conserv.; tax.: P. Beauv. is widely introduced and weedy.
Consaul et al. (2008a, 2008b).
COLPODIUM + ZINGERIA
Annuals or perennials, caespitose, occasionally Chromosome base number x ¼ 2.
stoloniferous. Leaves with sheaths closed 1/4 to Colpodium and Zingeria are sisters in molec-
1/3 of their length. Spikelets with 2 to 10 flowers. ular phylogenetic studies (Gillespie et al. 2008;
Glumes shorter than the spikelet, their apices Kim et al. 2009; Schneider et al. 2009). These
notched or truncate to acute or acuminate. Callus two genera were once placed in Miliinae (Soreng
of flower glabrous or pubescent. Lemma generally et al. 2007), but more recent data clearly place
with 5 veins, the lateral veins obscure, lemma them here in Coleanthinae (Hoffmann et al. 2013;
apex entire or ragged, notched or truncate to Schneider et al. 2009, 2012; Soreng et al. 2012 and
acute, unawned. 2n ¼ 14, 21, 28, 42, 56, 77. onward).
One hundred eleven spp., temperate regions
in salty soil. 292. Colpodium Trin.
Puccinella has lemmas rounded on the back,
which distinguished it from most species of Poa Colpodium Trin., Fund. Agrost.: 119, pl. 7 (1820); tax.:
in which the lemmas are generally keeled. It also Soreng and Fish (2011).
has obtuse lemma apices and rounded hila (the
latter synapomorphic for Poeae) that distinguish Perennials, <30 cm tall, caespitose, rhizomatous
if from Festuca, which has generally acute or or stoloniferous. Spikelets with 1 to 4 flowers,
acuminate lemma apices and elongate hila. Puc- with or without additional reduced flowers dis-
cinella has a 3-veined upper glume, which distin- tally. Glumes shorter or longer than the flowers,
guishes it from Glyceria in which the upper glume their apices obtuse to acute. Callus of flower gla-
has 1 vein. Morphological similarity and docu- brous or with cobweb-like hairs. Lemma glabrous
mented hybridization between Puccinellia and or pubescent, the apex truncate to acute, awnless
Phippsia have led to the suggestion that the two (mucronate in one species). 2n ¼ 4, 8, 20.
be combined (Steen et al. 2004); L€ove and L€ove Thirteen spp., central Asia to eastern Siberia,
(1976) provide some but not all of the necessary south to mountainous areas of Africa.
combinations. Puccinellia may be sister to Sclero-
chloa (Hoffmann et al. 2013).
293. Zingeria P. A. Smirn.
291. Sclerochloa P. Beauv. Zingeria P. A. Smirn., Bjull. Moskovsk. Obač. Isp. Prir.,
Otd. Biol. 51(2): 67 (1946).
Sclerochloa P. Beauv., Ess. Agrostogr.: 97, 177 (1812).
Delicate annuals. Spikelet dorsiventrally com-
Annuals, tufted. Leaves with sheaths closed ¼ to pressed, with one flower, without a rachilla exten-
½ of their length. Inflorescence unbranched and sion. Glumes shorter or longer than the spikelets,
one-sided, sometimes the branches branched. their apices obtuse to acute, upper glume apex
Spikelets with 2 to 7 flowers, proximal rachilla with a sharp point. Lemma apex obtuse, awnless.
internode noticeably thicker than the distal ones. 2n ¼ 4, 8, 12.
Glumes shorter than the flowers, with hyaline Five spp., the Caucasus, Anatolia, northern
margins, their apices obtuse to emarginate. Callus Iraq and Iran.
The hexaploid species Z. kochii (Mez) Tzvel. kelets with one flower; rachilla extension lacking;
is an allopolyploid with one genome donated by disarticulation below the glumes. Glumes as long
Colpodium versicolor (Kotseruba et al. 2010). as or longer than the flower, strongly compressed
and keeled, often partially fused near their bases,
their apices truncate to acute, muticous to awned.
Callus of flower glabrous. Lemma keeled, the mar-
XVI.15. SUBTRIBE POINAE Dumort. (1829)
gins often partially fused, the apex truncate to
Miliinae Dumort. (1829). acute, awned from the abaxial side, the awn
Alopecurinae Dumort. (1829). straight, curved or geniculate. Lodicules absent.
Cinninae Caruel (1892). Stamens 2 or 3. Style 1. 2n ¼ 14, 26, 28, 30, 42,
98, 100, 105, 112, 117, 119, 120.
Annuals or perennials. Leaf sheaths of upper Fifty-two spp., north temperate regions and
culm leaves often closed up to ¼ of their length. South America.
Flower callus generally pubescent. Lemma gener-
ally keeled, the apex with or without an awn, the
296. Aniselytron Merr.
awn generally terminal. Palea generally membra-
nous, green. Ovary glabrous. Aniselytron Merr., Philipp. J. Sci. 5: 328 (1910); anatomy:
Twenty-six genera, 636 spp. Ma et al. (2005).
This subtribe corresponds to the PAM clade
of Gillespie et al. (2008). The entire clade is well Perennials, the culms solitary or tufted, 0.5–1 m
supported (Gillespie et al. 2008; Hoffmann et al. tall. Leaf sheaths open. Spikelets with one flower,
2013), but recognition of the subtribes Alope- with a rachilla extension. Glumes shorter than the
curinae, Cinninae, and Miliinae renders Poinae spikelet, the margins hyaline, lower glume apex
paraphyletic so they are placed in synonymy truncate to acute, upper glume apex acute to
here. Named hybrids include xDupoa J. Cay. & acuminate. Lemma leathery, the apex entire
Darbysh. or with two teeth, acute to acuminate, awnless.
2n ¼ 42.
294. Agrostopoa Davidse, Soreng & P. M. Peterson Two spp., Asia.
Aniselytron – or at least A. treutleri (Kuntze)
Agrostopoa Davidse, Soreng & P. M. Peterson, Novon 19: Soják – is likely to be of hybrid origin. One parent
33 (2009). comes from the Poinae (minus Poa) clade,
whereas the other is related to Poa sect. Sylvestres
Annuals or perennials, caespitose. Spikelets with (Gillespie et al. 2008, 2010).
one flower, without a rachilla extension. Glumes
about the same length as the flower, their apices
297. Apera Adans.
acute to acuminate. Callus of flower glabrous.
Lemma apex entire or with two teeth, mucronate Apera Adans., Fam. Pl. 2: 495 (1763).
or awned from the sinus, the awn straight or
twisted. Annuals. Spikelet with one flower, with a rachilla
Three spp., Colombia. extension. Glumes slightly shorter or longer than
Davidse et al. (2008) observe that members of the flowers, their apices acute to acuminate. Cal-
this genus are somewhat similar to Old World lus of flower glabrous or with sparse hairs.
species of Colpodium. Lemma apex acute, awned, the awn originating
just below the lemma apex, generally several
295. Alopecurus L. times as long as the lemma. 2n ¼ 14, 28.
Five spp., Europe through the Caucasus to
Alopecurus L., Sp. Pl. 1: 60–61 (1753). Afghanistan.
Apera may be closely related to Nephelochloa
Annuals or perennials, caespitose or rhizomatous. and Bellardiochloa, as suggested by DNA
Inflorescence densely contracted, cylindrical. Spi- sequence data (Hoffmann et al. 2013).
298. Arctopoa (Griseb.) Prob. Four spp., southern Europe and Middle East.
In molecular analyses, Bellardiochloa is sister
Arctopoa (Griseb.) Prob., Novosti Sist. Vyssh. Rast. 11: 49 to Nephelochloa (Schneider et al. 2012), or to
(1974). Nephelochloa plus Apera (Hoffmann et al. 2013).
Rhizomatous perennials. Spikelets with 2 to 6 (8)
flowers, the rachilla pubescent. Glumes acute to 301. Cinna L.
acuminate. Callus of flower glabrous or with a
ring of short hairs. Lemma keeled, the apex Cinna L., Sp. Pl. 1: 5 (1753); tax.: Brandenburg et al.
(1991a).
acute. 2n ¼ 28, 29, 42, 62.
Five spp., North America and Asia.
Perennials, caespitose or rhizomatous. Spikelets
Some authors recognize this as Poa subgenus
with one flower, the rachilla internode notably
Arctopoa, but it is most likely to be intergeneric
elongate below the flower; rachilla extension pres-
hybrid. The plastid is similar to Poa sect. Sylves-
ent; disarticulation below the glumes. Glumes as
tres, but the nuclear gene (ITS) sequences are
long as the flowers or longer, their apices acute.
similar to Cinna (Gillespie et al. 2008, 2010). This
Callus of flower glabrous. Lemma apex acute,
is the opposite of what is found for Aniselytron.
muticous to awned, the awn subapical. Stamens
The sterile hybrid between A. eminens and Dupon-
1 or 2. Endosperm liquid. Fruit often with a beak.
tia fisheri is known as xDupoa J. Cay. & Darbysh.
2n ¼ 28.
Four spp., north temperate regions, Mexico,
299. Beckmannia Host Central America and northern South America.
Beckmannia Host, Icon. Descr. Gram. Austriac.: 3: 5
(1805). 302. Cornucopiae L.
Tufted annuals or rhizomatous perennials. Pri- Cornucopiae L., Sp. Pl. 1: 54 (1753).
mary inflorescence branches unbranched, borne
on an elongated central axis, the spikelets borne Annuals, with branched culms. Upper leaf sheaths
on one side, imbricate. Spikelets orbicular or cir- surrounding 1 or more head-like inflorescences;
cular and slightly flattened, with 1 or 2 flowers; stem below the inflorescence expanded into a
rachilla extension absent; disarticulation below broad cup. Spikelets with one flower, without a
the glumes. Glumes gibbous, shorter than or as rachilla extension. Glumes as long as or longer
long as the flowers, their apices obtuse to acute. than the flowers, keeled, partially fused along
Callus of flower glabrous. Lemma apex muticous. their margins, the apices truncate to acute, muti-
2n ¼ 14. cous to mucronate. Lemma keeled, the margins
Two spp., north temperate regions. fused, the apex truncate, awned from the abaxial
Beckmannia is sister to Pholiurus in the study side, the awn straight or twisted. Palea minute or
of Hoffmann et al. (2013), but the two genera are lacking. Lodicules absent.
quite different morphologically. Two spp., eastern Mediterranean region.
The branched culms are unusual among
pooid grasses.
300. Bellardiochloa Chiov.
Bellardiochloa Chiov., Stud. Veg. Piemonte: 60 (1929). 303. Dupontia R. Br.
Densely tufted perennials. Leaf blades 1 mm wide; Dupontia R. Br., Chlor. Melvill.: 32–33 (1823).
bristle-like. Spikelets with 2 to 8 flowers, the Arctophila (Rupr.) Rupr. ex Andersson, Pl. Scand. Gram.
x: 48 (1852).
rachilla pubescent. Glumes shorter than or about
as long as the flowers, their apices acuminate.
Rhizomatous perennials. Leaf sheaths closed for
Callus of flower pubescent. Lemma with the apex
more than ½ their length. Spikelets with 2 to 7
entire or with two minute teeth, acute, unawned or
flowers, with a rachilla extension or the rachilla
with a short mucro from the sinus. 2n ¼ 14, 28.
ending in a vestigial flower. Glumes shorter than keep them separate. The difference in length of
or as long as the flower, their apices acute to the rachilla internode below the lemma (long in
obtuse. Callus of flower glabrous or pubescent. Cinna, short in Limnodea), and the beaked fruit
Lemma keeled, apex obtuse to acute or acumi- in Cinna differentiate the two genera. No molec-
nate, awnless or with a short awn tip. 2n ¼ 42, 44, ular data are available for Limnodea.
63, 66, 84, 88.
Two spp. and a named hybrid, Arctic. 306. Milium L.
Specimens of the putative hybrid between
Arctophila fulva and Dupontia fisheri (xArctodu- Milium L., Sp. Pl. 1: 61 (1753).
pontia scleroclada (Rupr.) Tzvelev) are misiden-
tified material of Arctophila fulva, except for the Annuals or perennials, caespitose or rhizoma-
type, whose identity is uncertain (Brysting et al. tous. Spikelets dorsiventrally compressed, with
2003). one flower, without a rachilla extension. Glumes
Dupontia fisheri and Arctophila fulva are as long as or longer than the flowers, their apices
strongly supported as sisters in the molecular obtuse to acute. Callus of flower glabrous. Lemma
phylogenies of Gillespie et al. (2010), and ITS and palea hardened, smooth, shiny, the lemma
sequences from the two are intermixed in the apex obtuse. 2n ¼ 14, 18, 28.
work of Hoffmann et al. (2013). Morphologically Five spp., north temperate Eurasia and east-
they differ principally in the relative length of the ern North America.
glumes. This close relationship, their morpholog- The flowers of Milium are superficially simi-
ical similarity and their ability to hybridize sup- lar to those in the Panicoideae, but there is only
ports their combination. one flower per spikelet.
304. Limnas Trin. 307. Nephelochloa Boiss.
Limnas Trin., Fund. Agrost. 116, t. 6 (1820). Nephelochloa Boiss., Diagn. Pl. Or. 1: 72 (1844).
Perennials with short rhizomes. Inflorescence Annuals, less than 25 cm tall. Inflorescence
densely contracted. Spikelets with one flower, branches appearing to be in whorls, the proximal
without a rachilla extension, disarticulating one or two sets of branches generally sterile. Spi-
below the glumes. Glumes as long as the flower, kelets with 3 to 6 flowers, rachilla internode with
their apices acute. Lemma apex acute, awned a ring of hairs. Glumes shorter than the flowers,
from the abaxial side, the awn twisted. Stamens 2. their apices acute. Callus of flower pubescent.
Three spp., Siberia. Lemma apex with two lobes, awned from the
sinus. Stamens 2.
305. Limnodea L.H. Dewey One sp., N. orientalis Boiss., Turkey.
Mill (1985) describes the lodicules of this
Limnodea L.H. Dewey, Contr. U.S. Natl. Herb. 2: 518 species as being fused and thickened at the base,
(1894). an unusual character for the tribe. Nephelochloa
is sister to Bellardiochloa in molecular analyses
Tufted annuals. Internodes solid. Spikelets with (Schneider et al. 2012).
one flower, with a rachilla extension, disarticulat-
ing below the glumes. Glumes as long as the
308. Phleum L.
flower, coriaceous, their apices acute. Callus of
flower glabrous. Lemma apex entire or with two Phleum L., Sp. Pl. 1: 59–60 (1753).
teeth, awned from the abaxial side, the awn Pseudophleum Dogan, Notes Roy. Bot. Gard. Edinburgh
twisted and geniculate. 2n ¼ 14. 40: 75 (1982).
One sp., L. arkansana (Nutt.) L. H. Dewey,
southern U.S. and Mexico. Annuals or perennials, caespitose or rhizoma-
Limnodea was included in Cinna by Tucker tous. Inflorescence dense, cylindrical, spike-like.
(1996), whereas Brandenburg and Thieret (2000) Spikelets with one flower; rachilla extension
309. Pholiurus Host ex Trin.
Pholiurus Host ex Trin., Fund. Agrost.: 131 (1820).
Annuals or perennials. Inflorescence unbran-

ched, the rachis not disarticulating. Spikelets on
opposite sides of the rachis, with 2 flowers;
rachilla extension absent. Glumes borne side by
side on the abaxial side of the spikelet, longer
than the flower, coriaceous, their apices obtuse.
Lemma apex acute, unawned. 2n ¼ 14.
One sp., P. pannonicus (Host) Trin., eastern
Europe and central Asia.
Pholiurus has been placed here, rather than in
Parapholiinae, based on the work of Schneider
et al. (2012) and Hoffmann et al. (2013).
310. Poa L. Fig. 56
Poa L., Sp. Pl. 1: 67–70 (1753); tax.: Soreng et al. (2010);
phylog.: Gillespie et al. (2008), Birch et al. (2014).
Anthochloa Nees & Meyen, Reise Erde: 2: 14 (1834).
Dissanthelium Trin., Linnaea 10: 305 (1836).
Aphanelytrum (Hack.) Hack., Oesterr. Bot. Z. 52: 12 (1902).
Libyella Pamp., Bull. Soc. Bot. Ital. 1925: 151 (1925).
Eremopoa Roshev., Fl. URSS 2: 429, 756 (1934).
Lindbergella Bor, Svensk. Bot. Tidskr. 63: 368 (1969).
Austrofestuca (Tzvelev) E.B. Alexeev, Bjull. Moskovsk.
Obač. Isp. Prir., Otd. Biol. 81: 55 (1976).
Parodiochloa C. E. Hubb., Bull. Brit. Mus. (Nat. Hist.),
Bot. 8: 395 (1981).
Tovarochloa T. D. Macfarl. & But, Brittonia 34: 478 (1982).
Neuropoa Clayton, Kew Bull. 40: 728 (1985).
Fig. 56. Pooideae-Poeae-Poinae. Poa lilloi. A Habit, with Tzvelevia E. B. Alexeev, Bjull. Moskovsk. Obač. Isp. Prir.,
perfect flowers. B Habit, with pistillate flowers. C, D Lig- Otd. Biol. 90: 103 (1985).
ule. E Spikelet. F Pistillate flower with large stamens. G Oreopoa H. Scholz & Parolly, Willdenowia 34: 146, f. 1
Basal perfect flower. H Perfect flower with 3 stamens and (2004).
lodicules. I Detail of the lodicules. (From Zuloaga et al.
2012) Perennials or annuals, caespitose, rhizomatous,
or stoloniferous; plants bisexual, monoecious,
present or absent. Glumes shorter or more com-
dioecious, or gynodioecious. Spikelets with (1)
monly longer than the flower, strongly com-
2 to 6 (13) flowers; rachilla extension present or
pressed, keeled, keels extending into stiff awns,
absent. Glumes generally shorter than the flowers,
glume apices truncate. Callus of flower glabrous.
rarely longer, their apices obtuse to acuminate,
Lemma white, the apex acute, muticous or with a
sometimes muticous, rarely mucronate. Callus of
short awn. 2n ¼ 14, 28, 42.
flower glabrous, or with long cobweb-like hairs, or
Sixteen spp., north temperate regions and
with a tuft of hairs. Lemma with a strong keel in
South America.
many species, rounded in others, rarely flabellate,
P. pratense L. is widely cultivated as a pasture
the apex obtuse to acuminate, generally awnless.
grass. Pseudophleum was segregated from
2n ¼ 14, 28, 42, 56, 70, as well as virtually all
Phleum because the glumes are shorter than the
aneuploid numbers up to and including 147.
flower, but there is no other reason to separate it.
500 spp., worldwide in cool temperate regions. remains to be done, particularly on the evolution
In its current circumscription, Poa is the larg- of the many polyploid complexes. The genus is
est genus of grasses. Five subgenera are recog- the result of rapid and recent diversification, with
nized by Soreng et al. (2010): Ochlopoa, Poa, much of the speciation occurring in the last 2 mil-
Pseudopoa, Stenopoa, and Sylvestres. Of these, lion years (Hoffmann et al. 2013).
subgenus Poa is by far the largest and is divided
into two supersections, Homalopoa (called HAM-
311. Rhizocephalus Boiss.
BADD by Gillespie et al. 2007) and Poa (called
POM by Gillespie et al. 2007). Rhizocephalus Boiss., Diagn. Pl. Orient. 1: 68 (1844).
The work of Gillespie et al. (2007, 2008, 2009,
2010) and Soreng et al. (2010, 2011) has gone a Tiny annuals, less than 2 cm tall. Inflorescence
long way to resolving phylogenetic issues within capitate, surrounded by a leaf sheath. Spikelets
Poa and its many segregate genera. The synonymy with one flower, without a rachilla extension.
presented here largely follows that of Gillespie Glumes shorter than the flower, with fused mar-
et al. (2008). Libyella and Lindbergella are placed gins, their apices muticous. Lemma with 5 veins,
in synonymy based on morphology (R. Soreng pubescent, with club-shaped hairs, the apex atten-
and L. Gillespie, pers. comm.) and molecular uate, with a mucro. Lodicules absent. Stamens 2.
data (Hoffmann et al. 2013), although the DNA Caryopsis with an apical beak. 2n ¼ 14.
sequences are quite short and other loci may need One sp., R. orientalis Boiss., eastern Mediter-
to be explored. Anthochloa was segregated from ranean region and Iran.
Poa on the basis of its flabellate lemma, and
Aphanelytrum because of its awl-shaped lemma.
Oreopoa was segregated based on its convolute 312. Simplicia Kirk
leaf blades and somewhat rounded lemmas, but
these characters also appear in some species of Simplicia Kirk, Trans. Proc. New Zealand Inst. 29: 497
(1897); tax.: Smissen et al. (2011).
Poa, and the genus was placed in synonymy with
Poa by Soreng et al. (2012 and onward); the
Decumbent, mat-forming perennials. Spikelets
molecular data on which the placement is based
with 1 or 2 flowers. Glumes shorter than the
are unpublished. Austrofestuca was segregated
flowers, the distal one less than ½ the length of
based on having five or more veins on the
the flower; glume apices acute. Lemma apex
lemma, but it clearly belongs in Poa (Jacobs
acute, with or without a mucro. Palea with one
et al. 2008). In addition, Gillespie et al. (2008)
vein. Stamens 2 or 3. 2n ¼ 28.
and Refulio-Rodriguez et al. (2012) provide con-
Two spp., New Zealand.
vincing evidence for placing Dissanthelium and
Simplicia is morphologically similar to “Cala-
Tovarochloa in Poa. The former genus had been
magrostis”, but has a 3-veined lemma, and is
segregated (and even placed in a different tribe)
similar to Poa but has a 1-keeled palea.
because its glumes are longer than the flowers,
while the latter was recognized on the basis of its
tiny stature and capitate inflorescence. However, 313. Ventenata Koeler
both are clearly derived from ancestors that must
have looked like conventional Poa. It is not clear Ventenata Koeler, Descr. Gram.: 272 (1802), nom. conserv.
why Eremopoa or Neuropoa were segregated. Gaudiniopsis (Boiss.) Eig., Repert. Spec. Nov. Regni Veg.
26: 74 (1929), nom. invalid.
Libyella was segregated based on its unbranched
inflorescence and lack of lodicules.
Tufted annuals. Spikelets with 2 to 10 flowers,
Species of Poa are often polyploid, and hybri-
with reduced flowers distally or with a rachilla
dization and apomixis are common (e.g., Huff
extension; the most proximal flower remaining on
2010; Rodionov et al. 2010; Soreng et al. 2010).
the plant after the other flowers have been shed,
The morphological characters that separate the
eventually falling with the glumes. Glumes
species are subtle and generally quantitative.
shorter than or as long as the flowers, their apices
Despite the enormous strides made in disentan-
acute to acuminate. Callus of flower pubescent.
gling the phylogeny of this genus, much work
Proximal lemmas unawned or with a straight cate, awned, the awn subapical or from the sinus.
terminal awn; distal lemmas with a twisted abax- Palea tightly clasped by the lemma.
ial awn; in some species, additional awns devel- Two spp., Australia, New Zealand.
oped on the lemma lobes. 2n ¼ 14. Festucella was synonymized with Hookero-
Thirteen spp., Eurasia. chloa by Jacobs et al. (2008).
Gillespie et al. (2008) place Gaudiniopsis in
Poinae, but keep it distinct from Ventenata, cit-
316. Nicoraepoa Soreng & L. J. Gillespie
ing data from D€oring et al. (2007). However, the
latter authors do not mention Gaudiniopsis, and Nicoraepoa Soreng & L. J. Gillespie, Ann. Missouri Bot.
no molecular data are available for it. The five Gard. 94: 842 (2007).
species of Gaudiniopsis are synonymized here
with Ventenata following (Clayton and Renvoize Dioecious, gynodioecious or hermaphrodite per-
(1986), pending further information. ennials, with stout rhizomes. Ligule pubescent
abaxially. Leaf blade with sharp-pointed apex;
HSAQN clade
adaxial surface with prominent ridges. Spikelets
The following four genera form a strongly
with 2 to 5 flowers. Callus of flower glabrous or
supported clade, called HSAQN, in the study of
with short hairs. Lemmas acute, mucronate or
Gillespie et al. (2009). The clade is sister to Cinna,
with a short awn from a shallow sinus.
and HSAQN plus Cinna is sister to Aniselytron.
Six spp., Argentina, Chile, Islas Malvinas.
The HSAQN species share no obvious morpho-
logical characters, although they are generally
similar to Poa from which they have all been 317. Saxipoa Soreng, L. J. Gillespie & S. W. L. Jacobs
segregated. All except for Arctagrostis are native
to the Southern Hemisphere. Saxipoa Soreng, L. J. Gillespie & S. W. L. Jacobs, Austral.
Syst. Bot. 22: 406–407, f. 1C, 3 (2009).
314. Arctagrostis Griseb. Caespitose perennials. Spikelets with 2 to 4 flow-

ers. Glumes shorter than the flowers, margins
Arctagrostis Griseb., Fl. Ross. 4: 434 (1852).
scarious, their apices acute to obtuse. Callus of
flower with a few straight hairs. Rachilla densely
Rhizomatous perennials. Spikelets with one pubescent. Lemma margin ciliate, the apex acute,
flower; rachilla extension present or the rachilla sometimes with a tiny subapical mucro.
ending in a vestigial flower. Glumes shorter than One sp., S. saxicola (R. Br.) Soreng, L. J.
the flower, their apices acute to obtuse. Callus of Gillespie & S. W. L. Jacobs, Australia.
flower glabrous. Lemma translucent distally, Saxipoa is similar to Hookerochloa in having
keeled, the apex obtuse to acute, awnless. Palea leaf sheaths open almost their entire length, and
with one vein, folded. Hilum less than 1/3 the the rachilla densely pubescent; the two also share
length of the fruit. 2n ¼ 28, 42, 56. minor characters of the lodicules and lemma
Two spp., Arctic. margins. Their habitat is quite different however,
and the two are not sisters in the analyses of
315. Hookerochloa E. B. Alexeev Gillespie et al. (2009).
Hookerochloa E. B. Alexeev, Bjull. Moskovsk. Obač. Isp.

Prir., Otd. Biol. 90: 106 (1985). 318. Sylvipoa Soreng, L. J. Gillespie & S. W. L.
Festucella E. B. Alexeev, Byull. Moskovsk. Obshch. Isp. Jacobs
Prir. Otd. Biol. 90: 104 (1985).
Sylvipoa Soreng, L. J. Gillespie & S. W. L. Jacobs, Austral.
Caespitose perennials. Spikelets with 3 to 5 flow- Syst. Bot. 22: 404, f. 1D (2009).
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acute, margins ciliolate. Rachilla internodes Caespitose perennials. Leaves with sheaths open
densely pubescent. Callus of flower pubescent. to the base, blades with obvious cross veins. Spike-
Lemma apex entire or with 2 teeth, acute or trun- lets with 2 to 4 (7) flowers. Glumes shorter than
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PACMAD Clade
Embryonic mesocotyl internode elongate. Plants flap at the base, with three awns, these separate or
often C4. fused at the base. Palea less than half the length of
the lemma. Style branches 2. Hilum linear.
The PACMAD clade appeared as a strongly
Embryo with epiblast absent, scutellar cleft pres-
supported group in some of the earliest molecular
ent or absent, embryonic leaf margins meeting.
phylogenies of the grasses, and remains a well-
Three genera, 365 spp.
documented clade (Davis and Soreng 1993;
Placement of the three genera now in Aristi-
GPWG 2001; GPWG II 2012). Perhaps surpris-
doideae has been a long-standing problem in
ingly for a clade with such firm molecular sup-
grass taxonomy. They are superficially similar to
port, there are no obvious morphological
Stipeae, in that the spikelets are often fusiform in
synapomorphies, with the possible exception of
shape, and the callus is often sharp-pointed.
the elongate mesocotyl internode in the embryo.
However, the presence of microhairs on the epi-
All 24+ origins of C4 photosynthesis occur in this
dermis, the embryo characters, and the C4 photo-
clade. While the photosynthetic pathway is not
synthesis in Aristida and Stipagrostis exclude
universal, it is derived wherever it occurs. Thus,
Aristidoideae from Pooideae. Once DNA
the propensity to evolve C4 may indeed be a
sequences became available for Aristida and Sti-
synapomorphy. While the subfamilies within the
pagrostis, it was clear that Aristidoideae formed a
clade are all clearly monophyletic, relationships
separate lineage within the PACMAD clade
among them have been unclear and highly sensi-
(Barker et al. 1995; Clark et al. 1995; GPWG
tive to sampling. Addition of a large amount of
2001), and the subfamily now appears as sister
data and a broad sample of species has shown
to all other subfamilies in the clade (GPWG II
that Aristidoideae are sister to all other members
2012). The subfamily itself is clearly monophy-
of the clade (GPWG II 2012) (Fig. 9).
letic, and each of the three included genera is also
monophyletic (Cerros-Tlatilpa et al. 2011).
C4 photosynthesis has been derived twice
VII. SUBFAMILY ARISTIDOIDEAE Caro (1982) within Aristidoideae, once in Aristida and once
in Stipagrostis. Stipagrostis is sister to the C3
Annuals or perennials, herbaceous. Ligule a genus Sartidia, and the C3 species Aristida long-
fringe of hairs, or a fringed membrane. Leaf ifolia is sister to all remaining species of Aristida
blades generally narrow. Inflorescence branches (Cerros-Tlatilpa and Columbus 2009; GPWG II
branched. Spikelets cylindrical or laterally com- 2012). As in all C4 species, Stipagrostis and Aris-
pressed, with one flower and no rachilla exten- tida have co-opted an existing isoform of PEP-
sion. Glumes generally longer than and enclosing carboxylase for carbon fixation. However, they
the flower. Disarticulation above the glumes. Cal- have co-opted different paralogues, consistent
lus of flower short or long, blunt or sharp- with the observation of independent origins of
pointed, pubescent. Lemma terete, leathery, the C4 (Christin and Besnard 2009). Leaf anatomy
margins enfolding the palea, with a germination and gene expression differ between the two as
268 PACMAD Clade
well. All Aristidoideae, whether C3 or C4, have two

sheaths around the vascular bundles (Cerros-Tla-
tilpa and Columbus 2009; Hattersley and Watson
1992). The inner sheath of plants in the Sartidia/
Stipagrostis clade and in A. longifolia consists of
thick-walled cells lacking chloroplasts (a mes-
tome sheath), whereas the outer sheath is paren-
chymatous and contains chloroplasts (Cerros-
Tlatilpa and Columbus 2009). However, in C4
Aristida both sheaths are parenchymatous. Dec-
arboxylating enzymes appear only in the inner
sheath (Sinha and Kellogg 1996; Voznesenskaya
et al. 2005b), pointing to that sheath as the site of
carbon reduction. Rubisco is expressed in both
sheaths, suggesting that it may capture carbon
leaked out of the inner sheath.
KEY TO THE GENERA OF ARISTIDOIDEAE
1. Awns plumose 321. Stipagrostis

1. Awns glabrous 2
2 (1). Glumes with 3 veins or more; leaves with C3 anat-
omy 320. Sartidia
2. Glumes with 1 vein; leaves of most plants with unusual
C4 leaf anatomy 319. Aristida
GENERA OF ARISTIDOIDEAE
319. Aristida L.
Aristida L., Sp. Pl.: 82 (1753); phylog.: Cerros-Tlatilpa

et al. (2011).
Fig. 57. Aristidoideae. Stipagrostis hirtigluma subsp.
patula. A Habit. B Ligule. C Glumes. D Flower. (From
Annuals or perennials, mostly caespitose. Leaf Launert 1971)
apex often sharp-pointed. Ligule a fringe of
hairs, or a fringed membrane. Glumes generally
with one vein, their apices often mucronate or Perennials. Ligule a fringe of hairs. Glumes with 3
awned. Lemma awns often fused to form a twisted or 5 veins, their apices acute to long attenuate or
column. C4, NADP-ME subtype, except for A. with an awn tip. C3. 2n ¼ 22.
longifolia. 2n ¼ 22, 24, 36, 44, 48, 66. Five spp., Zambia, Angola, Congo, northeast-
Three hundred four spp., warm areas ern South Africa, Madagascar.
throughout the world. S. dewinteri occurs on serpentine soils in
Cerros-Tlatilpa et al. (2011) find no evidence South Africa (Balkwill et al. 2011).
to support previously recognized subgenera in
Aristida. Instead they identify several large clades 321. Stipagrostis Nees Fig. 57
that primarily reflect geography.
Stipagrostis Nees, Linnaea 7: 290 (1832); phylog.: Cerros-
Tlatilpa et al. (2011).
320. Sartidia de Winter
Sartidia de Winter, Kirkia 3: 137 (1963); tax.: Balkwill Annuals or perennials, caespitose, rhizomatous
et al. (2011); phylog.: Cerros-Tlatilpa et al. (2011). or stoloniferous, sometimes becoming somewhat
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primary branches themselves branched to form Panicoideae (Clayton and Renvoize 1986), but are
higher-order branches; branch complexes some- clearly unrelated to the panicoids and are placed
times present in the axils of upper culm leaves. in Micrairoideae (Sánchez-Ken et al. 2007).
Spikelets laterally or dorsiventrally compressed, Chromosome numbers are generally consis-
disarticulating above or below the glumes. tent within major clades and are synapomorphic
Glumes two, membranous, hyaline or indurate, for some. Gynerieae, Chasmanthieae, Centothe-
with or without awns. Lemmas and paleas mem- ceae, and Tristachyideae have chromosome base
branous, hyaline or indurate, with or without numbers of 11, 12, 12, and 10–12, respectively.
awns. Paleas with 0 to 2 keels. Lodicules 2, fleshy. Andropogoneae and Paspaleae share a base chro-
Stamens 3, stigmas 2. Caryopsis with hilum punc- mosome number of 10, while Paniceae have a
tate. Embryo with a long mesocotyl, with a scu- base chromosome number of 9 (Giussani et al.
tellar cleft and with the embryonic leaf margins 2001). Both x ¼ 9 and x ¼ 10 are probably
overlapping; epiblast present, but lost in Panicoi- derived and synapomorphic.
deae s.s. Stomatal subsidiary cells triangular or C4 photosynthesis has originated 22 to 24
domed (a plesiomorphy for the family); micro- times within Panicoideae s.l. (Christin et al.
hair apical cells generally much longer than wide 2007; Edwards and Smith 2010; Giussani et al.
(“the panicoid type”), but this character variable. 2001; GPWG II 2012; Vicentini et al. 2008).
Midribs simple or complex. Ancestrally C3. Gynerieae, Chasmanthieae and Centotheceae are
Two hundred-twelve genera, 3316 species. entirely C3, whereas Andropogoneae are wholly
In its traditional, strict sense, Panicoideae C4, and Paniceae, Paspaleae, and Tristachyideae
include the tribes Andropogoneae, Paspaleae, each contain a mix of C3 and C4 lineages. All
and Paniceae; this clade is referred to here as known C4 subtypes are represented, but most C4
Panicoideae s.s. Recent molecular phylogenetic species are NADP-ME, with a single bundle
analyses, however, have led to the current broad sheath surrounding the veins. The NAD-ME and
circumscription of the subfamily, called here PCK subtypes are synapomorphic for Panicinae
Panicoideae s.l., which also includes the tribes and Melinidinae, respectively, both monophyletic
Gynerieae, Chasmanthieae, Centotheceae, and subtribes of Paniceae.
Tristachyideae (Morrone et al. 2012; Sánchez- Spikelet structure has been noted since the
Ken and Clark 2010) (Fig. 58). Both Panicoideae time of Robert Brown (1810, 1814) to be a defin-
s.s. and s.l. are consistently monophyletic in all ing characteristic of Panicoideae s.s. Spikelets in
272 VIII. Subfamily Panicoideae Link (1827)
Reinheimer et al. 2005; Zanotti et al. 2010). Devel-

opmentally, paired spikelets in all Panicoideae
form from a single primordium that appears to
divide later in development, giving rise to one
terminal spikelet and one lateral (Bommert et al.
2005; Kellogg 2000; Zanotti et al. 2010).
TRIBES AND SUBTRIBES OF PANICOIDEAE
XVII. Tribe Steyermarkochloeae

XVIII. Tribe Tristachyideae
XIX. Tribe Chasmanthieae
XX. Tribe Gynerieae
XXI. Tribe Centotheceae
Panicoideae s.s.
Panicoideae s.s. incertae sedis
XXII. Tribe Andropogoneae
XXII.1. Subtribe Arundinellinae
Andropogoneae s.s.
Andropogoneae s.s. incertae sedis
XXII.2. Subtribe Tripsacinae
XXII.3. Subtribe Rottboelliinae
XXII.4. Subtribe Ischaeminae
Fig. 58. Relationships of the subtribes and tribes of Pani- XXII.5. Subtribe Saccharinae
coideae. Tree topology based on figures in Morrone et al. XXII.6. Subtribe Germainiinae
(2012). XXII.7. Subtribe Andropogoninae
XXIII. Tribe Paspaleae
Panicoideae s.s. are two-flowered, with the upper XXIII.1. Subtribe Arthropogoninae
XXIII.2. Subtribe Otachyriinae
flower generally hermaphrodite and the lower XXIII.3. Subtribe Paspalinae
one staminate or sterile. Developmental studies XXIV. Tribe Paniceae
show that the upper flower develops ahead of the Incertae sedis
lower one, and exhibits a characteristic pattern of XXIV.1. Subtribe Anthephorinae
expression of genes that control floral organ iden- XXIV.2. Subtribe Boivinellinae
XXIV.3. Subtribe Neurachninae
tity (Doust and Kellogg 2002; Malcomber and XXIV.4. Subtribe Cenchrinae
Kellogg 2004; Reinheimer et al. 2006). Staminate XXIV.5. Subtribe Melinidinae
flowers in all species studied initiate both a pistil XXIV.6. Subtribe Panicinae
and stamens (Le Roux and Kellogg 1999; Reinhei-
mer et al. 2005; Zaitchik et al. 2000). The gynoe-
cium develops a clear nucellus, surrounded by an
KEY TO THE GENERA OF PANICOIDEAE
emerging ridge of tissue that would normally
form the ovary wall, but then the pistil ceases 1. Leaf blades absent from inflorescence-bearing culms or
development. Cessation of growth correlates from all culms 2
with loss of nuclei and cytoplasm in subepider- 1. Leaf blades present 4
mal cells, apparently a pattern of controlled cell 2 (1). Plants annual; leaf blades absent from upper
death. In Andropogoneae, the dead cells are in leaves, present on lower ones; axillary branch com-
plexes present; upper glume gibbous; plants of
the center of the gynoecium, whereas in the one India 363. Pogonachne
species of Panicum studied, cell death appeared 2. Plants perennial; blades absent from entire culms; axil-
near the base of the ovary. lary branch complexes absent; upper glume not gib-
Pairing of spikelets, with one sessile or on a bous; plants not occurring in India 3
short pedicel and the other on a longer pedicel, is 3 (2). Spikelets laterally compressed, with 4 to 9 flowers,
bisexual; plants of southwest Australia 339. Spartochloa
usually cited as a character useful for recognizing 3. Spikelets dorsiventrally compressed to terete, with 2 or
Andropogoneae. However, this character is much 3 flowers, most unisexual; plants of northern South
more widespread in the subfamily (Kellogg 2000; America 322. Steyermarkochloa
KEY TO THE GENERA OF PANICOIDEAE 273
4 (1). Upper surface of leaves with sinuous lamellae; 18. Plants perennial 20
plants aquatic, central Africa 342. Hydrothauma 19 (18). Base of leaf blades clasping the stem;
4. Upper surface of leaves flat or ribbed, but not with spikelets with 8 to 20 flowers; lemma without reflexed
sinuous lamellae; habitat various 5 bristles 340. Megastachya
5 (4). Inflorescence a dense spherical or hemispherical 19. Base of leaf blades narrow, not clasping; spikelets
head 6 with 1 to 4 flowers; lemma with reflexed bristles
5. Inflorescence variously elongate, open or contracted, 341. Centotheca
but not a dense head 12 20 (18). Plants monoecious, to 3 m tall, forming an ele-
6 (5). Spikelets with both flowers bisexual; plants of vated tussock, the base of the plant covered by rem-
India 323. Chandrasekharania nants of leaf sheaths and aerial roots; plants of central
6. Spikelets with only one flower bisexual; plants of Aus- Brazil 330. Arundoclaytonia
tralia or, if in India, then only on beaches 7 20. Plants with flowers bisexual, generally less than 3 cm tall,
7 (6). Plants annual 471. Thedachloa not forming elevated tussocks, not occurring in Brazil 21
7. Plants perennial 8 21 (20). All flowers fertile but the distal ones reduced;
8 (7). Lower glume absent, if present then thin, less than palea straight; stamens 3 338. Cyperochloa
½ the length of the spikelet, the apex obtuse to acute; 21. Proximal 1 to 2 flowers sterile; palea gibbous, arched;
leaf blades disarticulating 497. Cleistochloa stamens 1 331. Chasmanthium
8. Lower glume stiff, as long as the spikelet, the apex 22 (12). All or most spikelets associated with one or more
acuminate; leaf blades not disarticulating 9 bristle-like sterile branches 23
9 (8). Plants with unisexual flowers, dioecious; forming 22. Inflorescences without bristle-like sterile branches; if
clumps connected by long rhizomes; glumes membra- bristle-like branches present, then associated with
nous 10 only some of the spikelets, usually just the terminal
9. Plants with bisexual flowers; caespitose or stolonifer- ones on the lateral branches 32
ous; glumes coriaceous to indurate 11 23 (22). Lateral branches disarticulating below any bris-
10 (9). Pistillate spikelets subtended by papery bracts, tles and falling off as a unit 24
these acute to acuminate but not with long stiff tips; 23. Lateral branches not falling off as a unit, spikelets
staminate spikelets 1 per inflorescence branch; plants disarticulating below the glumes but above any asso-
of interior Australia 522. Zygochloa ciated bristles 27
10. Pistillate spikelets subtended by bracts with long stiff 24 (23). Leaves pseudopetiolate, the blades cordate or
tips; staminate spikelets several per inflorescence sagittate 516. Streptolophus
branch; plants of coastal dunes 521. Spinifex 24. Leaves lacking pseudopetioles, the blades narrow,
11 (9). Lower glume with a transverse line of hairs linear 25
above a hyaline patch; glume apices truncate to 25 (24). Spikelets associated with one bristle or a fan of
obtuse 499. Thyridolepis bristles, if the latter then the bristles not enclosing the
11. Lower glume lacking a transverse line of hairs spikelets; plants of Australia 508. Paractaenum
and lacking a hyaline patch; glume apices acumi- 25. Spikelets subtended and surrounded by a cluster of
nate 498. Neurachne bristles, these often forming a cup or involucre, but
12 (5). Spikelets with one to many flowers, if two flowered not a flattened fan; plants widespread 26
then the reduced flower distal to the bisexual one 13 26 (25). Spikelets bisexual; inflorescence branches contracted
12. Spikelets with exactly two flowers, the distal one fertile creating a bottle-brush appearance 501. Cenchrus
and generally bisexual, the proximal one generally 26. Spikelets unisexual, the plants dioecious;
staminate or reduced to a sterile lemma 22 inflorescence branches elongate, with 2 to 5 spike-
13 (12). Leaf blades with a pseudopetiole 14 lets 509. Pseudochaetochloa
13. Base of leaf blades not pseudopetiolate 17 27 (23). Spikelets and bristles strictly paired, the two
14 (13). Sterile lemmas awned, the awns strongly borne on a common pedicel 28
recurved 333. Lophatherum 27. Spikelets and bristles not in strict pairs, spikelets
14. Sterile lemmas unawned, or if awned then the awns either associated with multiple bristles, or some spike-
straight 15 lets lacking bristles entirely 30
15 (14). Flowers unisexual, the proximal flower in the 28 (27). Lateral branches of inflorescence elongate, with
spikelet pistillate with a gibbous lemma 335. Zeugites numerous spikelet/bristle pairs; plants of Central
15. At least some of the flowers in the spikelet bisexual, America, cultivated elsewhere 505. Ixophorus
none gibbous 16 28. Inflorescence slender, spike-like, lateral branches very
16 (15). Lemmas awned; spikelets with paleas free from short if developed at all; plants of the West Indies,
the rachilla 332. Chevalierella Pacific, South America or the Old World 29
16. Lemmas without awns; spikelets with palea keels fused 29 (28). Apex of first lemma rounded; plants of
to the rachilla 334. Orthoclada Australia 502. Chamaeraphis
17 (13). Spikelets with only one flower, sterile flowers 29. Apex of first lemma acuminate; plants of Old and New
lacking 347. Garnotia World tropics 507. Paratheria
17. Spikelets with more than one flower, although some 30 (27). First glume and lemma saccate, the sacs collaps-
flowers reduced 18 ing upon drying to form two abaxial protuberances;
18 (17). Plants annual 19 plants of Mexico 512. Setariopsis
30. First glume and lemma not saccate, not collapsing on 8 (4). Spikelets partially embedded in hollows in the broad
drying; worldwide distribution 31 flat inflorescence axis 514. Stenotaphrum
31 (30). Spikelets with the upper flower bisexual, the lower 8. Spikelets not embedded in the inflorescence axis 9
flower staminate or sterile 511. Setaria 9 (8). Both glumes present 523. Chaetium
31. Spikelets with both flowers bisexual 503. Dissochondrus 9. Lower glume absent or minute 10
32 (22). Primary branches of inflorescence broad, leaf- 10 (9). Lemma with margins flat, overlapping and partially
like, often drooping to surround the spikelets; covering the palea; lower lemma awned 513. Snowdenia
spikelets in clusters on the abaxial side of the 10. Lemma with margins inrolled, covering only the mar-
branches 478. Chlorocalymma gins of the palea; lower lemma unawned 11
32. Primary branches of inflorescence slender, if flattened 11 (10). Lower lemma adaxial 453. Axonopus
not drooping to surround the spikelets, spikelets not 11. Lower lemma abaxial 462. Paspalum
clustered abaxially 33 12 (1). Spikelets not obviously paired 13
33 (32). Inflorescence unbranched above the uppermost 12. Spikelets in sessile-pedicellate pairs, although the
spathe or leaf Group I (includes plants with paired pedicellate spikelet sometimes represented only by
spikelets borne on otherwise unbranched axes) the pedicel 16
33. Inflorescence with two or more branches above the 13 (12). Spikelets all pedicellate, strongly laterally com-
uppermost spathe or leaf 34 pressed 389. Dimeria
34 (33). Inflorescence axis and/or branches disarticulating 13. Spikelets all sessile, dorsiventrally compressed 14
at the nodes; spikelets generally in pairs, less fre- 14 (13). Lower glume smooth, not pitted or sculp-
quently single or in sets of 3 or more; dispersal unit tured 369. Oxyrhachis
including one or more spikelets and their associated 14. Lower glume sculptured or pitted 15
internodes Group II 15 (14). Spikelets numerous on each inflorescence
34. Inflorescence axis tough; spikelets borne singly or in branch 386. Ophiuros
pairs; dispersal units rarely including the inflorescence 15. Spikelets few (often <3) with a long naked peduncle
axis 35 below 387. Thaumastochloa
35 (34). Primary branches of inflorescences not them- 16 (12). Lower glume of sessile spikelet hard, shiny,
selves branched Group III and variously sculptured, with pits, hooks, warts, or
35. Primary branches of inflorescence branched, although ridges 17
branches sometimes very short Group IV 16. Lower glume of sessile spikelet smooth and shiny or
green and leaf-life, with or without prominent veins or
grooves, but not sculptured 22
Group I – Inflorescence unbranched 17 (16). Pedicel free from rachis internode 18
17. Pedicel fused partially to wholly to rachis internode 19
1. Glumes membranous, less firm than or about as firm 18 (17). Spikelets unawned 383. Rottboellia
as the lemmas, one or both often shorter than the 18. Spikelets with the pedicellate spikelet awned; awns
spikelet 2 generally also on the upper glume of the sessile spike-
1. Glumes coriaceous to indurate, much firmer than the let 370. Rhytachne
lemmas, generally longer than the spikelet 12 19 (17). Lower glume of sessile spikelet awned
2 (1). Inflorescence axes surrounded by spathe-like 376. Glyphochloa
bracts 3 19. Glumes unawned 20
2. Inflorescence axes not subtended by spathe-like 20 (19). Sessile spikelets two per node, on either
bracts 4 side of a slender flattened pedicel bearing a tiny
3 (2). Inflorescence axes in clusters at nodes, each axis flap at its summit representing the pedicellate
ending in a sharp point; all spikelets bisexual; plants of spikelet 381. Mnesithea
dry inland sites 519. Xerochloa 20. Sessile spikelets one per node; pedicellate spikelet well
3. Inflorescence axes not clustered, ending in a spikelet; developed with obvious glumes 21
proximal spikelets bisexual, distal ones staminate; 21 (20). Lower glume of sessile spikelet suborbicular 377.
plants of coastal dunes 528. Thuarea Hackelochloa
4 (2). Spikelets laterally compressed 5 21. Lower glume of sessile spikelet broadly lanceolate-
4. Spikelets dorsiventrally compressed 8 ovate 385. Heteropholis
5 (4). Lemmas awned 326. Jansenella 22 (16). Lemmas lacking awns; lower flower of sessile
5. Lemmas lacking awns 6 spikelet often with lemma, palea, and stamens 23
6 (5). Base of leaf blades cordate; glumes with tuberculate- 22. Lemmas of one or both spikelets awned; lower flower
based hairs 457. Echinolaena of sessile spikelet often reduced 34
6. Base of leaf blades narrow; glumes glabrous, or if 23 (22). Distal spikelet pairs of the inflorescence stami-
pubescent then the hairs not tuberculate-based 7 nate, proximal ones pistillate 351. Chionachne (note: if
7 (6). Culms generally over 25 cm tall; plants caespitose, Coix is interpreted as having an unbranched inflores-
rarely stoloniferous; plants of Central and South cence it will key here as well, but can be easily distin-
America 446. Mesosetum guished by the hard bead-like sheaths surrounding the
7. Culms less than 25 cm tall; plants stoloniferous, pistillate spikelets)
restricted to New South Wales 500. Alexfloydia
23. Spikelet pairs similar in sex expression throughout the 37 (36). Awn covered with short hairs 423. Themeda
inflorescence 24 37. Awn glabrous 421. Iseilema
24 (23). Lower glume of pedicellate spikelet extended into 38 (36). Pedicellate spikelet scarcely developed, forming
a long slender, tail-like awn, becoming recurved after a tiny point at the apex of the pedicel, or lacking
anthesis 371. Urelytrum entirely 39
24. Lower glume of pedicellate spikelet not awned, or the 38. Pedicellate spikelet developed, staminate or sterile 43
awn <5 mm long 25 39 (38). Inflorescence partially or almost wholly enclosed
25 (24). Sessile spikelets two per node 26 by the subtending leaf sheath or spathe; upper glume
25. Sessile spikelets one per node 28 with a slender awn 40
26 (25). Glumes acuminate 361. Lasiurus 39. Inflorescence not completely enclosed by leaves;
26. Glumes obtuse 27 upper glume of sessile spikelet awned or not 41
27 (26). Lower glume of sessile spikelet with two 40 (39). Plants annual; inflorescence reduced to a
keels, these expanding to narrow wings near the set of three spikelets; plants of northern
apex 383. Rottboellia Australia 353. Clausospicula
27. Lower glume of sessile spikelet lacking keels 381. 40. Plants perennial; inflorescence with more than three
Mnesithea spikelets; plants of tropical Asia 359. Kerriochloa
28 (25). Lower glume with a constriction about half way up, 41 (39). Inflorescence internodes as broad as the spike-
with an obtuse apex; plants of India 380. Manisuris let 390. Ischaemum
28. Lower glume without a constriction; plants of Old 41. Inflorescence internodes slender 42
World tropics 29 42 (41). Base of sessile spikelet sunken into a crater-
29 (28). Upper glume of sessile spikelet laterally com- like node; inflorescence axis with internodes cla-
pressed 350. Chasmopodium vate 433. Schizachyrium
29. Upper glume of sessile spikelet dorsiventrally com- 42. Base of sessile spikelet somewhat flattened,
pressed 30 connected apparently laterally to a filiform inflores-
30 (29). Lower glume of sessile spikelet more or less cence axis 413. Anadelphia
oblong, with a long extended apex 384. Hemarthria 43 (38). Lemma of pedicellate spikelet awned, that of
30. Lower glume with an acute to obtuse apex, not sessile spikelet unawned 412. Trachypogon
extended 31 43. Lemma of pedicellate spikelet unawned, that of sessile
31 (30). Upper part of plant with axillary branch spikelet awned 44
complexes, producing complex compound inflores- 44 (43). Proximal spikelet pairs without awns, both
cences 383. Rottboellia sessile and pedicellate spikelets of these pairs
31. Upper part of plant without axillary branch com- staminate 45
plexes, inflorescence terminal on the culm 32 44. Proximal spikelet pairs with awns, the sessile spikelets
32 (31). Pedicellate spikelets rudimentary, or reduced of these pairs bisexual, the pedicellate ones staminate
only to an awn, or lacking, leaving a barren pedicel; or sterile 47
nodes of inflorescence transverse 375. Eremochloa 45 (44). Callus rounded 422. Pseudodichanthium
32. Sessile and pedicellate spikelets morphologically simi- 45. Callus sharp, barbed 46
lar, the pedicellate well developed, the sessile with non- 46 (45). Pedicellate spikelets smaller than the sessile ones;
winged glumes; nodes of inflorescence oblique 33 lower glume of sessile spikelet not deeply grooved, the
33 (32). Upper part of inflorescence internode not sur- spikelet terete 420. Heteropogon
rounding the base of sessile spikelet above; lower 46. Pedicellate spikelets larger than sessile ones; lower
glume of sessile spikelet with a distinctly bifid glume of sessile spikelet deeply grooved, the groove
tip 355. Elionurus translucent 410. Agenium
33. Upper part of inflorescence internode forming a crater 47 (44). Upper glume of sessile spikelet with a slender
or cup, surrounding the base of sessile spikelet above; awn 48
lower glume of sessile spikelet entire 379. Loxodera 47. Upper glume of sessile spikelet lacking an awn 52
34 (22). Spikelets <5 mm long; glume apices obtuse to 48 (47). Inflorescence completely enclosed by a reddish
truncate 35 brown spathe 415. Monocymbium
34. Spikelets generally over 5 mm long; glume apices 48. Inflorescence exserted, not completely enclosed by a
generally acute to acuminate, rarely truncate 36 spathe 49
35 (34). Upper glume awned, sessile spikelets one per 49 (48). Pedicellate spikelets bisexual, similar to the ses-
node 405. Pogonatherum sile ones; lower glume of the pedicellate spikelet
35. Upper glume awnless, sessile spikelets two per about the same width and color as that of the sessile
node 364. Polytrias spikelet 362. Microstegium
36 (34). Base of spikelet-bearing branch with four 49. Pedicellate spikelets staminate or sterile, clearly dif-
unawned spikelets, sessile or short pedicellate, ferent from the sessile ones; lower glume of the pedi-
arranged in a whorl, surrounding the single awned cellate spikelet often a different size or color from that
spikelet, another two unawned spikelets terminating of the sessile spikelet 50
the branch 37 50 (49). Lower glume of pedicellate spikelet firm to
36. Awned and unawned spikelets not arranged as hard and shiny, the margins enclosing the upper
above 38 glume 413. Anadelphia
50. Lower glume of pedicellate spikelet broad, flat, firm to 5. Pistillate spikelets proximal to the staminate, in the
papery but not hard and shiny, not enclosing the same inflorescence 6
upper glume 51 6 (5). Pistillate spikelet surrounded by a hard round bead-
51 (50). Lower glume of pedicellate spikelet well devel- like sheath 354. Coix
oped, membranous, with prominent green veins con- 6. Pistillate spikelet with hardened glumes, but not sur-
verging toward the apex 365. Sehima rounded by a bead-like sheath 7
51. Lower glume of pedicellate spikelet papery, purplish, 7 (6). Pedicellate spikelets developed in all parts of the inflo-
without prominent veins 430. Diectomis rescence; glume of pistillate spikelet flattened above;
52 (47). Rachis internodes and pedicels broadly club- plants of India and Southeast Asia 351. Chionachne
shaped; sessile spikelet inserted into a crater-like 7. Pedicellate spikelets not developing in the lower
node; glumes of pedicellate spikelets with prominent (pistillate) portion of the inflorescence; glume of
wings 53 pistillate spikelet rounded; plants of the New
52. Rachis internodes and pedicels about the same diam- World 373. Tripsacum
eter throughout their length; sessile spikelet not 8 (4). Inflorescence branches themselves branched, the
inserted into a crater-like node; glumes without proximal part of the branches bare 9
wings 54 8. Inflorescence branches themselves unbranched; spike-
53 (52). Inflorescence exserted, not enclosed in a spathe; lets borne the full length of the branch 12
lower glume of sessile spikelet two-keeled with strong 9 (8). Plants annual: spikelets solitary 537. Arthragrostis
central groove; plants of Africa 388. Andropterum 9. Plant perennial; spikelets paired 10
53. Inflorescence subtended and partially enclosed by 10 (9). Ultimate inflorescence branches with more than 10
a broad spathe; lower glume of sessile spikelet spikelet pairs 402. Lasiorhachis
neither two-keeled nor grooved; plants of tropical 10. Ultimate inflorescence branches with fewer than 7
Asia 348. Apluda spikelet pairs 11
54 (52). Lower glume with a prominent circular pit; spike- 11 (10). Callus of spikelets with tiny hairs; sessile spikelet
lets strongly dorsiventrally compressed; plants of truly sessile 396. Sorghum
Africa 419. Eremopogon 11. Callus of spikelets with hairs about half the
54. Lower glume not pitted; spikelets more or less round length of the spikelet; “sessile” spikelet on a short
in cross section 55 stalk 399. Eriochrysis
55 (54). Plants perennial; plants of Africa and South 12 (8). Spikelets borne singly 517. Uranthoecium
America 358. Hypogynium 12. Spikelets borne in pairs, one sessile and one pedicel-
55. Plants annual; plants of India and Southeast Asia 56 late 13
56 (55). Plants trailing, <0.8 m tall, rooting from the nodes; 13 (12). Spikelets <5 mm long 14
pedicellate spikelet staminate 429. Pseudanthistiria 13. Spikelets 10 mm long or more 15
56. Plants upright, 0.5–2 m tall, with prop roots; pedicel- 14 (13). Inflorescence with two (to six) branches; sessile
late spikelet rudimentary 368. Triplopogon spikelets overlapping; sessile and pedicellate spikelets
morphologically different, the sessile with lower
glumes truncate 407. Germainia
Group II – Inflorescence branched; inflorescence 14. Inflorescences with many branches; “sessile” spikelets
or branch axes disarticulating at the nodes actually short pedicellate, not overlapping on the
branches; sessile and pedicellate spikelets morpholog-
1. Spikelets without awns 2 ically similar, with glumes acute to acuminate
1. Spikelets bearing awns from lemmas, glumes or both 17 403. Miscanthus
2 (1). Primary inflorescence branches broad and leaf-like; 15 (13). Lower glume of sessile spikelet with a long-
spikelets in clusters of 2 to 6 on the abaxial side of the flattened tail, several times longer than the spikelet
branches 482. Trachys 372. Vossia
2. Primary inflorescence branches narrow, not leaf 15. Lower glume of sessile spikelet often long acuminate,
like; spikelets single or paired but not in abaxial but the apical portion no longer than the spikelet 16
clusters 3 16 (15). Lower glume of sessile spikelet not
3 (2). Glumes with prominent silicified hooks on winged 382. Phacelurus
the margin, and often over the back of the lemma as 16. Lower glume of sessile spikelet with two winged
well 378. Jardinea keels 383. Rottboellia (former Thyrsia)
3. Glumes smooth, without prominent hooks 4 17 (1). Primary inflorescence branches unbranched, digi-
4 (3). Plants strictly monoecious, the pistillate spikelets tate, two to many on inflorescence axis, generally
often distinctly different from the staminate 5 spikelet-bearing to near the base 18
4. Plants andromonoecious, with the upper flower of each 17. Primary inflorescence branches themselves branched,
sessile spikelet bisexual, the lower staminate or with multiple orders of branching; spikelets often
reduced; sex expression of the pedicellate spikelet restricted to the distal half of the branch 44
various 8 18 (17). Glumes sculptured; inflorescence internodes
5 (4). Staminate and pistillate spikelets in separate inflor- club-shaped 367. Thelepogon
escences, the staminate inflorescence branched, the 18. Glumes smooth or with hooked prickles, but not
pistillate unbranched 374. Zea sculptured; inflorescence internodes slender 19
19 (18). Plants stoloniferous, culms weak, rooting from 30. Lower glume of sessile spikelet with a deep groove;
the nodes; leaves broadly lanceolate, <6 cm long, but callus of sessile spikelet obliquely attached to the rachis,
>1 cm wide, often clasping the stem 20 and extending beyond it 428. Parahyparrhenia
19. Plants caespitose or rhizomatous, culms robust; 31 (29). Inflorescence branches more than five; inflores-
leaves linear, generally <1 cm wide, if lanceolate then cence axis often elongate 32
much longer than 6 cm, generally not clasping the 31. Inflorescence branches generally 2 or 3, up to 5,
stem 22 digitate or nearly so 34
20 (19). Lemma awn originating from below the midpoint 32 (31). Pedicellate spikelet similar to sessile one in size
of the abaxial side 349. Arthraxon and sex expression; both spikelets of the pair with
20. Lemma awn originating from a short notch near the callus hairs as long as the spikelet 404. Saccharum
apex 21 32. Pedicellate spikelet differing from the sessile one,
21 (20). Rachis internodes slender; stamens generally 2; staminate or more often sterile, generally lacking an
glumes unawned 393. Leptatherum awn; spikelets generally with callus hairs shorter than
21. Rachis internodes clavate; stamens 2 or 3; upper the spikelet 33
glume often awned 362. Microstegium 33 (32). Pedicels and rachis with prominent marginal veins
22 (19). “Sessile” spikelets actually on a short pedicel 23 or keels and hyaline central region 425. Bothriochloa
22. Sessile spikelets truly sessile, without any subtending 33. Pedicels and rachis round in cross section, without
pedicel 24 prominent veins or keels 394. Pseudosorghum
23 (22). Inflorescence contracted, superficially spike-like, 34 (31). Sessile and pedicellate spikelets similar in mor-
covered in long silky white hairs 401. Imperata phology, sex expression and awn development 35
23. Inflorescence of spreading digitate branches, glabrous or 34. Sessile spikelets and pedicellate spikelets different;
pubescent but not with long silky hairs 390. Ischaemum one of the pair bisexual and awned, the other stami-
24 (22). Ultimate inflorescence branches exactly two, nate or sterile and unawned 38
closely subtended and often surrounded by a spathe- 35 (34). Glumes stiff, truncate, with three teeth; inflores-
like leaf 25 cence branches two, overlapping 408. Lophopogon
24. Ultimate inflorescence branches two to many, if 35. Glumes tapering, if truncate, then not stiff; inflores-
exactly two then spathe-like leaf, if present, not sur- cence branches generally more than two 36
rounding the branches 29 36 (35). Upper glume with a slender awn; lower glume
25 (24). Upper glume deeply three lobed, the central lobe with hairs in tufts 357. Eulaliopsis
awned 409. Bhidea 36. Upper glume unawned or muticous; hairs on glumes
25. Upper glume apex entire, if bifid only shallowly so, not tufted 37
generally without an awn 26 37 (36). Callus rounded; spikelets compressed dorsiven-
26 (25). Lower glume of sessile spikelet with a deep trally 400. Eulalia
groove; apex of inflorescence (between the branches) 37. Callus sharp-pointed; spikelets round in cross sec-
extended into a flattened appendage at least as long as tion 411. Homozeugos
spikelets 417. Hyperthelia 38 (34). Inflorescence branches paired, one spikelet-bear-
26. Lower glume without a groove, inflorescence branch ing to near the base, the other with a long naked
without a flattened extension 27 portion (over 1.5 cm); lowermost spikelets on each
27 (26). Base of spikelet pair inserted into a crater-like branch staminate, in a whorl of four 416. Exotheca
extension of the internode below; ultimate inflorescence 38. Inflorescence branches paired or up to 5, both spike-
branches generally reflexed at maturity; proximal, non- let-bearing to near the base; lowermost spikelets not in
spikelet-bearing portions of branches generally more or a whorl of four 39
less equal in length 418. Cymbopogon 39 (38). Lower glume of sessile spikelet with two raised
27. Base of spikelet pair not inserted into a crater-like margins (keels), with a hollow or groove between
extension of the internode below; ultimate inflores- them 40
cence branches usually not reflexed; proximal, non- 39. Lower glume of sessile spikelet rounded on the back,
spikelet-bearing portion of branches generally not grooved or hollowed 42
unequal in length 28 40 (39). Callus of sessile spikelet rounded, not extended
28 (27). Lower glume of sessile spikelet without a line of below the base of the glumes 432. Andropogon
hairs; glumes of pedicellate spikelets slender, tapering; 40. Callus of sessile spikelet elongate, ending in a sharp
fertile lemma with two teeth, the awn originating between point 41
them; plants of Africa and Asia 431. Hyparrhenia 41 (40). Pedicels the same color as the rachis; sessile
28. Lower glume of sessile spikelet with a line of hairs; spikelet inserted into a deep crater-like node; plants
glumes of pedicellate spikelets broad, acute to obtuse; of Africa 427. Diheteropogon
fertile lemma apex entire, not two-toothed, awned 41. Pedicels deeply pigmented; sessile spikelet not
from the apex; plants of Australia 398. Spathia inserted into a crater-like node; plants of South Amer-
29 (24). Lower glume of sessile spikelet with an awn 30 ica 410. Agenium
29. Lower glume of sessile spikelet unawned 31 42 (39). Lower glume with a truncate apex, winged in
30 (29). Lower glume of sessile spikelet without a groove; some species; pedicellate spikelet reduced to a pedicel,
callus of sessile spikelet not protruding beyond the not obvious, tucked behind and partially fused to the
rachis axis 414. Elymandra lower glume 406. Apocopis
42. Lower glume with a rounded to acute apex, without 3 (1). Primary branches of the inflorescence ending in a
wings; pedicellate spikelet clearly present 43 sterile point or flexuous bristle 4
43 (42). Inflorescence nodding; plants annual 356. Euclasta 3. Primary branches of the inflorescence terminated by a
43. Inflorescence upright; plants generally perennial 426. spikelet 8
Dichanthium 4 (3). Primary branches of the inflorescence very short,
44 (17). Spikelets in groups of three with one sessile and partially embedded in the broad flat inflorescence
two pedicellate, laterally compressed 352. Chrysopo- axis 514. Stenotaphrum
gon 4. Primary branches of the inflorescence not embedded in
44. Spikelets solitary or paired, if in groups of three then the inflorescence axis 5
these only at the ends of the branches that also bear 5 (4). Plants monoecious; spikelets distantly spaced along
pairs of spikelets; spikelets generally dorsiventrally the axis, the lower ones pistillate and smaller than the
compressed 45 upper, staminate ones; plants of wet sites in northern
45 (44). Pedicels and rachis internodes with a translucent Australia 504. Hygrochloa
median line 424. Capillipedium 5. Plants with bisexual flowers; habitat various 6
45. Pedicels and rachis internodes without a translucent 6 (5). Spikelets on slender pedicels often longer than the
median line 46 spikelet itself; underdeveloped spikelets common;
46 (45). Pedicels ending blindly, without a spikelet 47 lower lemma deeply grooved 468. Holcolemma
46. Pedicels terminated with a spikelet, although this 6. Spikelets on short pedicels, nearly sessile; lower lemma
sometimes rudimentary 49 without a groove 7
47 (46). Upper glume keeled abaxially; plants of Southeast 7 (6). Upper glume mucronate to awned 510. Pseudoraphis
Asia 391. Asthenochloa 7. Upper glume not mucronate or awned 511. Setaria
47. Upper glume not keeled; plants of Africa, SE Asia, and 8 (3). Lower glume minute (ca. 0.1 the length of the
the Americas 48 spikelet) or absent 9
48 (47). Sessile spikelet rounded in cross section; apex 8. Lower glume well developed 17
with two teeth or keels 397. Sarga 9 (8). Spikelets with a globose swelling at the base 525.
48. Sessile spikelet dorsiventrally compressed; apex taper- Eriochloa
ing, not toothed 395. Sorghastrum 9. Spikelets lacking a globose swelling at the base 10
49 (46). Spikelets unpaired 392. Cleistachne 10 (9). Upper lemma membranous to cartilaginous; mar-
49. Spikelets in sessile-pedicellate pairs 50 gins broad, overlapping, lying flat on the palea 11
50 (49). Sessile and pedicellate spikelets similar in shape, 10. Upper lemma indurate; margins narrow, not overlap-
size and sex expression 51 ping, inrolled on the edges of the palea 13
50. Sessile and pedicellate spikelets differing in size, shape 11 (10). Spikelets lacking awns 479. Digitaria
and sex expression; in some species the two superfi- 11. Spikelets awned on the lower lemma 12
cially similar, but the pedicellate spikelet staminate 12 (11). Spikelets in pairs 515. Stereochlaena
and lacking an awn 53 12. Spikelets solitary 513. Snowdenia
51 (50). Inflorescence not hairy; glumes with prominent 13 (10). Inflorescence branches capillary; callus of spikelet
veins 366. Spodiopogon with dense white hairs; plants of Madagascar 530.
51. Inflorescence and spikelets covered with long dense Yvesia
hairs; glumes without prominent veins 52 13. Inflorescence branches not capillary; callus of spikelet
52 (51). Inflorescence hairs reddish-brown or golden- glabrous or with short hairs; plants of various loca-
brown; primary branches short, dense 399. Eriochrysis tions 14
52. Inflorescence hairs silvery; primary branches spread- 14 (13). All spikelets with the lower lemma facing the
ing, diffuse 404. Saccharum branch axis (adaxial) 453. Axonopus
53 (50). Nodes with stiff spreading white hairs; awns more 14. All spikelets with the lower lemma facing away from
than 2 cm long; callus of sessile spikelet sharp-pointed the branch axis (abaxial), or spikelet orientation vari-
in many species 397. Sarga able 15
53. Nodes glabrous or pubescent, but without stiff white 15 (14). Spikelets hemispherical or orbicular in outline,
hairs; awns no more than 1.5 cm long; callus of sessile often flat on one side and convex on the other; primary
spikelet blunt 52 inflorescence branches often flattened or winged 462.
54 (53). Sessile spikelet well over 5 mm long 396. Sorghum Paspalum
54. Sessile spikelet 5 mm long or less 360. Lakshmia 15. Spikelets ovate to lanceolate, generally convex on both
sides; primary inflorescence branches terete or angu-
lar, not flattened or winged 16
Group III – Inflorescence branches unbranched, 16 (15). Upper glume and lower lemma pubescent 452.
not disarticulating Acostia
16. Upper glume and lower lemma glabrous to puberu-
1. Spikelets surrounded by a set of stiff flattened, glume- lous 454. Anthaenantiopsis s.l.
like bracts 2 17 (8). Spikelets bearing hooked hairs on one or both
1. Spikelets without a cluster of flattened bracts 3 glumes 496. Pseudechinolaena
2 (1). Upper lemma awned 477. Chaetopoa 17. Spikelets lacking hooked hairs 18
2. Upper lemma unawned 476. Anthephora 18 (17). One or both glumes clearly awned 19
18. Glumes unawned, although the apex attenuate in some 34. Upper glume and lower lemma with hairs not in lines
taxa 25 or glabrous; culms generally not woody 35
19 (18). Both glumes with long awns; spikelets paired 20 35 (34). Upper palea winged near the apex, the wings
19. Only one glume with an awn; spikelets solitary or visible at maturity 459. Hopia
paired 21 35. Upper palea not winged 36
20 (19). Plant erect, caespitose; upper lemma awned 36 (35). Apex of upper lemma with a tiny mucro
523. Chaetium 533. Moorochloa
20. Plants decumbent, trailing; upper lemma awnless 36. Apex of upper lemma acute to obtuse, not mucronate 37
492. Oplismenus 37 (36). Leaf blades rigid, sharp-pointed
21 (19). Awn from lower glume only, upper glume and 445. Keratochlaena
both lemmas with at most a short mucro; plants of 37. Leaf blades membranous, lacking shart points 38
Madagascar and Africa 22 38 (37). Upper lemma and palea indurate 39
21. Awn from upper glume and lower lemma, lower glume 38. Upper lemma and palea cartilaginous to coriaceous 41
awnless 23 39 (38). Spikelet with thickened rachilla; upper lemma
22 (21). Upper lemma and palea cartilaginous, not hya- and palea covered by bicellular microhairs and simple
line; spikelets paired, similar in sex expression papillae 464. Streptostachys
495. Poecilostachys 39. Spikelet with slender rachilla; upper lemma and palea
22. Upper lemma and palea hyaline; spikelets paired with without extensive bicellular microhairs; papillae sim-
the sessile one staminate or sterile 486. Cyphochlaena ple or compound 40
23 (21). Awn of lower lemma coiled 465. Acritochaete 40 (39). Spikelets borne on the abaxial side of the branch,
23. Awn of lower lemma straight 24 solitary or paired 529. Urochloa
24 (23). Upper palea reflexed; ligule often absent 40. Spikelets borne on both sides of the branch, soli-
467. Echinochloa tary 535. Panicum s.s. (only a few Panicum species
24. Upper palea erect; ligule present, a fringed mem- will key here; most will fall in Group IV)
brane; 438. Oplismenopsis 41 (38). Lemma margins flat, largely concealing the
25 (18). Upper lemma with an awn extending beyond the palea 42
apex of the glumes 26 41. Lemma margins involute, the palea generally
25. Upper lemma awnless; if awned then the awn minute, exposed 44
less than 1.2 mm long, not extending beyond the apex 42 (41). Lower glume as long as the spikelet
of the glumes 30 480. Megaloprotachne
26 (25). Spikelets strongly laterally compressed 42. Lower glume shorter than the spikelet 43
389. Dimeria 43 (42). Ligule a fringed membrane 444. Triscenia
26. Spikelets dorsiventrally compressed or scarcely com- 43. Ligule membranous 481. Tarigidia
pressed 27 44 (41). Proximal flower of spikelet staminate, with a
27 (25). Spikelets paired, the members of the pair notably palea; plants of Central and South America
different in morphology and sex expression 28 454. Anthaenantiopsis s.l.
27. Spikelets all alike in morphology and sex expression 29 44. Proximal flower of spikelet sterile, lacking a palea;
28 (27). Pedicellate spikelet bisexual, the sessile one sta- plants of Africa and Australia 488. Entolasia
minate or sterile 412. Trachypogon
28. Pedicellate spikelet sterile or rarely staminate, notably
smaller than the bisexual sessile spikelet 396. Sorghum Group IV – Primary branches of inflorescence
29 (27). Spikelets borne in clusters 484. Alloteropsis branched, although branches sometimes very
29. Spikelets borne in pairs, one nearly sessile and the short
other clearly pedicellate 403. Miscanthus
30 (25). Spikelets laterally compressed 457. Echinolaena 1. Plants tall (up to 10 m), reed-like; leaf blades disarticu-
30. Spikelets dorsiventrally compressed 31 lating 2
31 (30). Upper lemma with a prominent beak; ligule 1. Plants less than 2 m tall, usually much less than 1 m; leaf
generally absent 467. Echinochloa blades generally persistent 3
31. Upper lemma not beaked; ligule present 32 2 (1). Plants with flowers unisexual, dioecious; spikelets
32 (31). Rachilla internode elongate between the lower disarticulating above the glumes; upper glume cau-
and upper flowers 33 date, recurved 336. Gynerium
32. Rachilla internode between the lower and upper flow- 2. Plants with flowers bisexual; spikelets disarticulating
ers contracted 34 below the glumes; upper glume apex erect
33 (32). Stoloniferous perennials; upper glume and lower 337. Thysanolaena
lemma without net-like venation; plants of Central and 3 (1). Lemmas awned 4
South America 461. Ocellochloa 3. Lemmas lacking awns, although awns sometimes pres-
33. Annuals; upper glume and lower lemma with promi- ent on the glumes 13
nent net-like venation; plants of eastern Africa 4 (3). Lemma awned from the apex, apex entire 5
524. Eccoptocarpha 4. Lemma awned from the sinus of a two lobed lemma 7
34 (32). Upper glume and lower lemma with a transverse 5 (4). Glumes lacking awns, although the upper glume
line of hairs; culms often woody 531. Leucophrys apex often caudate 346. Arundinella
5. Glumes with awns about 10 times longer than the glume 24 (23). Spikelets laterally compressed 25
itself 6 24. Spikelets dorsiventrally compressed 27
6 (5). Callus of flower glabrous 436. Canastra 25 (24). Spikelet obovate, asymmetrical, the upper lemma
6. Callus of flower pubescent 435. Arthropogon gibbous, enlarged abaxially 487. Cyrtococcum
7 (4). Spikelets in groups of three, either all pedicellate 25. Spikelet lanceolate to ovate, symmetrical, the lemma
or all sessile; glumes firm, often with pustule-based not gibbous 26
hairs 329. Tristachya 26 (25). Leaf blades broadly ovate to elliptical, narrowed
7. Spikelets solitary or paired, if the latter then on pedicels toward the base, with a fringe of hairs on the abaxial
of unequal lengths; glumes various 8 side of the collar (contraligule); spikelet callus pubes-
8 (7). Callus of spikelet with two teeth, or a single sharp cent 491. Microcalamus
point 327. Loudetia 26. Leaf blades linear or lanceolate, not narrowed to the
8. Callus of spikelet blunt 9 base; no hairs on the abaxial side of the collar; spikelet
9 (8). Lower glume absent or a minute hair-covered callus glabrous 483. Acroceras
flap 532. Melinis 27 (24). Upper lemma and palea leathery; plants of Aus-
9. Lower glume present, although sometimes only about tralia 469. Homopholis
1/3 the length of the lower lemma 10 27. Upper lemma and palea indurate; widespread 28
10 (9) Palea absent; plants of Cuba 345. Reynaudia 28 (27). Spikelets borne all around the inflorescence
10. Palea present; plants of Africa and western Asia 11 branches; hairs on apex of upper lemma
11 (10). Keels of upper palea lacking wings flattened 527. Rupichloa
328. Trichopteryx 28. Spikelets borne on one side of the inflorescence
11. Keels of upper palea winged 12 branches; hairs on apex of upper lemma cylindri-
12 (11). Pedicels of spikelets fused to the rachis; lower cal 529. Urochloa
palea becoming thick and hard 325. Gilgiochloa 29 (23). Lower glume minute or absent 30
12. Pedicels of spikelets free from the rachis; lower palea 29. Lower glume present 33
membranous 324. Danthoniopsis 30 (29). Spikelets glabrous; plants with rhizomes ending
13 (3). Inflorescence branches verticillate, 10–60 per in cleistogamous spikelets 485. Amphicarpum
node 440. Stephostachys 30. Spikelets sparsely or densely pubescent; plants not
13. Inflorescence not markedly verticillate 14 producing cleistogamous spikelets 31
14 (13). Leaves stiff, coriaceous, without a clear distinction 31 (30). Upper lemma cartilaginous 448. Anthaenantia
between sheath and blade; ligule often lacking 15 31. Upper lemma indurate 32
14. Leaf blades stiff or lax, but clearly distinct from the 32 (31). Inflorescence open, the branches spreading;
sheath; ligule present 17 plants of Central America 455. Aakia
15 (14). Plants rhizomatous; apex of leaf blades obtuse; 32. Inflorescence narrow, the branches upright; plants of
rachilla extension present 490. Mayariochloa South America 454. Anthaenantiopsis
15. Plants caespitose; apex of leaf blades sharp-pointed; 33 (29). Spikelets attached obliquely to the pedicel 34
rachilla extension absent 16 33. Spikelets attached upright on the pedicel 35
16 (15). Panicle open, spreading 434. Apochloa 35 (33). Spikelets glabrous 443. Coleataenia
16. Panicle dense, contracted 463. Renvoizea 35. Spikelets with long hairs 447. Tatianyx
17 (14). Culms woody, leaning or scrambling; upper 35 (33). Inflorescence narrow, dense, highly branched but
lemma apex blunt, bearing short white hairs, with a the branches short, producing a bottle-brush appear-
concave depression 489. Lasiacis ance 449. Hymenachne
17. Culms herbaceous, erect; upper lemma apex various, 35. Inflorescence open; if narrow then the branching
but not both concave and with short white hairs 18 sparse, not brush-like 36
18 (17). Lower palea enlarged at maturity, hardened, 36 (35). Leaves with base of blades rounded to cordate,
winged and enclosing upper flower 450. Otachyrium clasping the stem and/or pseudopetiolate 37
18. Lower palea neither enlarged, hardened, nor winged 19 36. Leaves with base of blades narrow, neither cordate nor
19 (18). One or both glumes gibbous, sac-like 20 pseudopetiolate 46
19. Glumes not gibbous 22 37 (36). Spikelets minute, <2 mm long; plants of
20 (19). Lower glume absent 441. Cyphonanthus Africa 470. Hylebates
20. Lower glume present 21 37. Spikelets over 2 mm long; plants of the New World 38
21 (20). Leaf blades with a sharp tip; spikelets clearly 38 (37). Upper lemma transversely rugose
falcate in side view; lower lemma grooved, translucent 473. “Panicum” sects. Monticolae, Verrucosa,
472. Thyridachne Ovalifoliae
21. Leaf blades without a sharp tip; spikelets not falcate; 38. Upper lemma smooth, generally shiny 39
lower lemma not grooved 474. Sacciolepis 39 (38). Margins of upper lemma lying flat on the
22 (19). Glumes extended into a long acuminate tail, palea 437. Homolepis
otherwise unawned 344. Oryzidium 39. Margins of upper lemma involute on the palea 40
22. Glumes awnless, not extended to a long acuminate 40 (49). Upper lemma with a tiny mucro
tail 23 494. Parodiophyllochloa
23 (22). Upper lemma with a thickened crest at the tip 24 40. Upper lemma not mucronate 41
23. Upper lemma without a thickened crest 29
TRIBES AND GENERA OF PANICOIDEAE 281
41 (40). Spikelets borne abaxially on the inflorescence 56 (55). Upper lemma rugose, the apex acute to mucro-
branches, often crowded on short pedicels 42 nate 529. Urochloa
41. Spikelets borne on both sides of the inflorescence 56. Upper lemma smooth to slightly rough, but not
branches, generally spread out on long pedicels 44 strongly rugose, the apex obtuse 57
42 (41). Upper lemma and palea membranous; plants of 57 (56). Plants aquatic, the culms spongy, sheaths with
Central and South America, the Caribbean and Aus- aerenchyma 536. Louisiella
tralia 449. Hymenachne 57. Plants terrestrial, the culms solid or hollow but not
42. Upper lemma and palea indurate; plants of southeast- spongy; sheaths lacking aerenchyma 58
ern North America 43 58 (57). Ligule a fringed membrane 535. Panicum s.s.
43 (42). Glumes approximately equal in length 58. Ligule an unfringed membrane 456. Osvaldoa
439. Phanopyrum 59 (55). Inflorescence branches deciduous; upper
43. Lower glume less than 1/2 the length of the upper flower on an extended rachilla internode; plants of
one 451. Rugoloa Australia 537. Arthragrostis
44 (41). Basal leaves generally forming a rosette, morpho- 59. Inflorescence branches persistent; rachilla internode
logically distinct from cauline leaves; spikelets below the upper flower not extended; plants of the New
obovate 466. Dichanthelium World 454. Anthaenantiopsis
44. Basal leaves not rosette forming, not morphologically
distinct from the cauline leaves; spikelets generally
globose or elliptic 45
45 (44). Leaf blades lacking a pseudopetiole, persis- TRIBES AND GENERA OF PANICOIDEAE
tent 475. Trichanthecium
45. Leaf blades with a pseudopetiole, disarticulating from
the sheath 458. Gerritea XVII. TRIBE STEYERMARKOCHLOEAE Davidse & R.P.
46 (36). Both glumes less than 1/2 the length of the Ellis (1984)
spikelet 493. Ottochloa
46. One or both glumes as long as the spikelet 47 Caespitose perennials; monoecious. Vegetative
47 (46). Glumes with tubercle-based hairs, lower lemma and flowering culms morphologically distinct,
deeply sulcate 518. Whiteochloa the former with only one leaf, the latter with
47. Glumes lacking tubercle-based hairs, lower lemma multiple leaves that lack blades. Leaf sheaths
rounded 48 hard and longer than culm internodes. Ligule
48 (47). Abaxial ligule (contraligule) present, fringed;
plants of Madagascar 343. Lecomtella absent. Inflorescence branches branched, short
48. Abaxial ligule absent; plants of various regions 49 and appressed to the main axis of the inflores-
49 (48). Plants with bulbous corms at the base, or cence; staminate spikelets below, pistillate above.
with stout thick rhizomes with short internodes Spikelets dorsiventrally compressed to terete.
520. Zuloagaea Staminate spikelets with two fertile flowers and
49. Plants without corms 50
50 (49). Glumes covered with prominent bumps or one sterile one distally. Pistillate spikelets with
warts 473. “Panicum” sect. Verrucosa one fertile flower, with sterile flowers proximal
50. Glumes without warts 51 and distal. Glumes shorter than the spikelet.
51 (50). Spikelets laterally compressed 52 Lemma apex truncate or obtuse. C3.
51. Spikelets dorsiventrally compressed 54 One genus, one species.
52 (51). Spikelets paired; callus of upper flower
expanded into two wings or scars on the base of the Steyermarkochloeae are treated here as mono-
lemma 460. Ichnanthus generic, following the work of Sánchez-Ken and
52. Spikelets solitary; callus of upper flower not Clark (2010) and Morrone et al. (2012). Both
expanded 53 authors exclude Arundoclaytonia, based on phylo-
53 (52). Lower glume present; lemma of lower flower genetic data. Steyermarkochloa is clearly a member
awnless 534. Tricholaena
53. Lower glume tiny or absent; lemma of lower flower of Panicoideae, although its precise placement
awned 532. Melinis remains unclear; it is sister to Tristachyideae in
54 (51). Rachilla thickened above the upper glume and the combined morphological/ molecular analysis
below the lower flower 442. Oncorachis of Morrone et al. (2012), but support is lacking.
54. Rachilla not thickened above the upper glume and
below the lower flower 55
55 (54). Upper lemma indurate, generally shiny, more 322. Steyermarkochloa Davidse & R. P. Ellis Fig. 59
firm than the glumes 56
55. Upper lemma coriaceous to membranous, about as Steyermarkochloa Davidse & R. P. Ellis, Ann. Missouri
firm as the glumes or less firm 59 Bot. Gard. 71: 995 (1984).
with tufts of hair, with two teeth or lobes and

awned from the sinus. Ancestrally C3.
Most of the genera included here are mem-
bers of the former Arundinelleae. However, that
tribe is clearly an artificial assemblage represent-
ing at least two distinct lineages, Arundinella
versus everything else (Li and Phipps 1973).
Arundinella itself is sister to Andropogoneae
and is therefore included in the latter tribe. Chan-
drasekharania, Jansenella, Gilgiochloa, and Tri-
chopteryx are placed in Tristachyideae
provisionally, based on the similarity of their
lemmas to those of other genera included here.
However, Chandrasekharania and Jansenella
share some characters, including photosynthetic
pathway, with Micrairoideae and may ultimately
be better placed there.
323. Chandrasekharania V. J. Nair, Ramachandran,

Sreekumar
Chandrasekharania V. J. Nair, Ramachandran, Sreeku-

mar, Proc. Indian Acad. Sci. 91: 79 (1982).
Annuals. Ligule membranous. Inflorescence cap-

Fig. 59. Panicoideae-Steyermarkochloeae. Steyermarko- itate. Spikelets with two bisexual flowers. Glumes
chloa angustifolia. A Flowering plants. B Spikelet. C Flower. awned. Lemma with a short awn originating
(From Davidse et al. 2004, with permission from the Mis- between two teeth. C3.
souri Botanical Garden Press; drawn by Bruno Manara) One sp., C. keralensis V.J. Nair, V.S. Ramach.
& Sreek., India.
Characters of the tribe. No molecular data are available for this
One sp., S. angustifolia (Spreng.) Judz., genus. It is possible that it is misplaced here,
northern South America. given its C3 leaf anatomy and spikelets with a
pair of hermaphrodite flowers. Sánchez-Ken and
Clark (2010) suggest that Chandrasekharania
XVIII. TRIBE TRISTACHYIDEAE Sánchez-Ken & L.G. should be retained in Arundinelleae but without
Clark (2010) strong evidence. The genus could be a better fit
Annuals or perennials with erect culms; caespi- for Micrairoideae.
tose, stoloniferous or short rhizomatous. Leaves
without pseudopetioles, the ligule a line of hairs 324. Danthoniopsis Stapf
or membranous. Inflorescence unbranched to
highly branched, with the spikelets in groups of Danthoniopsis Stapf, Hooker’s Icon. Pl. 31: pl. 3075
two or three, the spikelet cluster falling as a unit. (1916).
Spikelets laterally compressed or not, with two
flowers, disarticulating above the glumes. Glumes Perennials or annuals. Ligule a fringe of hairs.
2, membranous, often a different color from the Leaf blades with or without pseudopetioles. Spi-
lemmas. Lemma of lower flower similar to the kelets laterally compressed, in pairs or groups of
upper glume. Upper flower bisexual, the lemma three, purplish. Callus of flower blunt, pubescent.
membranous to coriaceous, not indurate, often Upper lemma glabrous or with 2 to 8 tufts of
hairs, with two lobes, the margins clasping the pairs. Callus of the spikelet with two teeth or with
palea. Upper palea with winged keels, expanded a single oblique sharp point. Upper lemma gla-
above into a club-shaped swelling. C4, with a brous to pubescent, with two teeth. Upper palea
single bundle sheath. 2n ¼ 24. keels thick. Stamens 2. C4, with one bundle sheath;
Sixteen spp., Africa, east to Pakistan. isolated bundle sheath cells sometimes in place of
minor veins.
Twenty-five spp., Africa, Madagascar and
325. Gilgiochloa Pilg.
South America.
Gilgiochloa Pilg., Bot. Jahrb. Syst. 51: 415 (1914).
328. Trichopteryx Nees
Annuals. Leaf margins cartilaginous. Ligule a
fringe of hairs. Inflorescence narrow and spike- Trichopteryx Nees, Nat. Syst. Bot.: 449 (1836).
like, the pedicels largely fused to the rachis. Spike-
lets laterally compressed, disarticulating above Trailing annuals or perennials. Ligule a fringe
the glumes. Glumes awned. Callus of flower of hairs. Inflorescence branches branched,
pubescent. Lower palea becoming thick and spreading or upright. Spikelets laterally com-
hard. Upper lemma with tufts of hair at the mar- pressed or not compressed, disarticulating
gins and at the base of the awn, with two long above the glumes. Callus of spikelet blunt, pubes-
narrow lobes; the awn flattened and geniculate. cent. Upper lemma with tufts of hair at the mar-
Upper palea keels winged. C4, with a single bundle gins, with two long narrow lobes. C4, with one
sheath. bundle sheath; isolated bundle sheath cells some-
One sp., G. indurata Pilg., central Africa. times in place of minor veins.
Five spp., Africa and Madagascar.
326. Jansenella Bor
329. Tristachya Nees Fig. 60
Jansenella Bor, Kew Bull. 10: 96 (1955).
Tristachya Nees, Fl. Bras. Enum. Pl. 2: 458 (1829).
Mat-forming annuals. Ligule membranous. Spi- Loudetiopsis Conert, Bot. Jahrb. Syst. 77: 277 (1957).
kelets in pairs, but the inflorescence otherwise Dilophotriche (C. E. Hubb.) Jacq.-Fél., J. Agric. Trop. Bot.
Appl. 7: 407 (1960).
unbranched. Spikelets laterally compressed, dis- Zonotriche (C.E. Hubb.) J.B. Phipps, Kirkia 4: 113 (1964).
articulating above the glumes. Lower lemma with
an awn, upper lemma with two tufts of hair and Perennials, or a few species annual, caespitose or
two long narrow lobes. Upper palea papillose rhizomatous. Ligule a fringe of hairs. Spikelets in
pubescent, with winged keels. C3. 2n ¼ 20. groups of three. Callus of spikelet blunt to sharp-
One sp., J. griffithiana (M€
ull. Stuttg.) Bor, India. pointed. Lower lemma with 3 to 9 veins. Stamens
T€urpe (1970) provides a detailed description 2 or 3. C4, with one bundle sheath. 2n ¼ 20,
of the leaf anatomy of this species in comparison 24, 40.
to Arundinella and Danthoniopsis. Sánchez-Ken Forty spp., Central and southern Africa, Cen-
and Clark (2010) suggest that Jansenella should tral and South America, Madagascar.
be retained in Arundinelleae but without strong In contrast to Andropogoneae, in which spi-
evidence. The genus may be a better fit for kelets are in sessile and pedicellate pairs or tri-
Micrairoideae. plets, in Tristachya all three spikelets of a group
are pedicellate or (in some African species) all
327. Loudetia Hochst. ex Steud. sessile. In some species, the common branch
supporting the cluster of three is sharply bent at
Loudetia Hochst. ex Steud., Syn. Pl. Glumac. 1: 238 (1854). maturity, with a point of disarticulation in the
middle of the bend. The African species have
Perennials. Ligule a fringe of hairs. Inflorescence pustule-based macrohairs on the glumes, the pus-
branches branched or unbranched, the branches tule itself black and contrasting with the light-
spreading or upright; spikelets borne singly or in colored hairs and glume surface. These
pressed in most species, often strongly so.

Stamens 1, 2 or 3. C3.
Six genera, 25 species.
Chasmanthieae plus Zeugiteae form a clade
(GPWG II 2012; Sánchez-Ken and Clark 2010),
with reasonably good support in the GPWG II
analysis; accordingly, the two are combined
here. One species of Zeugites is annual, presum-
ably a derived condition.
330. Arundoclaytonia Davidse & R. P. Ellis
Arundoclaytonia Davidse & R. P. Ellis, Ann. Missouri Bot.

Gard. 74: 479 (1987).
Caespitose perennials, to 3 m tall, with an unusual

elevated tussock, the base of the plant covered by
remnants of leaf sheaths and aerial roots; monoe-
cious. Leaf insertion reported to be spiral. Ligule
a fringed membrane. Staminate and pistillate spi-
kelets in separate capitate clusters, each cluster
surrounded by bracts. Staminate spikelets with 3
to 9 flowers, glumes generally with cross veins;
lemmas mucronate. Pistillate spikelets with one
fertile flower, the sterile flowers both proximal
and distal; fertile lemma pubescent, with 9 to 11
veins and cross veins. Palea of the pistillate flower
longer than the lemma, curved, thickened and
smooth below, convolute. C3.
Fig. 60. Panicoideae-Tristachyideae. Tristachya decora. One sp., A. dissimilis Davidse & R. P. Ellis,
A Habit. B Rootstock. C Triad of spikelets. D Upper central Brazil.
lemma, viewed from the adaxial side. E Upper lemma, A partial sequence of ndhF places Arundoclay-
viewed from the abaxial side. (From Clayton et al. 1974,
drawn by P. Halliday) tonia near Chasmanthium (Sánchez-Ken and
Clark 2007); when the same sequence is combined
with morphological data, the species falls within
Chasmanthium (Morrone et al. 2012), a position
distinctive characters of the inflorescence have that needs to be confirmed. Arundoclaytonia was
led to recognition of several small segregate included with Steyermarkochloa in the tribe
genera, but the group as a whole appears to be Steyermarkochloeae, but the two genera do not
easily accommodated in a single genus. appear to be closely related (Morrone et al. 2012).
XIX. TRIBE CHASMANTHIEAE W. V. Br. & B. N. 331. Chasmanthium Link

Smith ex Sánchez-Ken & L. G. Clark (2010)
Chasmanthium Link, Hort. Berol. 1: 159 (1827).
Bromuniola Stapf & C.E. Hubb., Bull. Misc. Inform. Kew
Zeugiteae Sánchez-Ken & L. G. Clark (2010). 1926: 366 (1926).
Gouldochloa Valdés-Reyna, Morden & S.L. Hatch, Syst.
Perennials, caespitose, rhizomatous, or stolonif- Bot. 11: 112 (1986).
erous. Leaf blades broad, with or without pseu-
dopetioles. Inflorescence branches branched. Perennials. Culms hollow or solid. Ligule a
Spikelets with one to many flowers, laterally com- fringed membrane. Primary branches of the
inflorescence branched or unbranched. Spikelets

with 2 to 10 flowers, the proximal 1 or 2 sterile.
Paleas gibbous, the keels winged. Stamens 1 (3).
2n ¼ 24, 48.
Seven spp., eastern USA, Mexico, tropical
Africa.
Molecular data clearly unite Chasmanthium
with Bromuniola and Gouldochloa (Sánchez-Ken
and Clark 2007, 2010).
Leaves with a pseudopetiole. Spikelets disarti-
culating below the glumes.
332. Chevalierella A. Camus
Chevalierella A. Camus, Rev. Bot. Appl. Agric. Trop. 13:

421 (1933).
Perennials, 1–2 m tall. Culms hollow. Leaf blades

broad. Ligule a ciliolate membrane. Primary
branches of the inflorescence unbranched. Spike-
lets with one fertile flower plus a rudimentary one
distally. Glumes awned, the rachilla internode
between them elongated. Lemmas awned, palea
keels winged. Lodicules fused. Stamens 2.
One sp., C. dewildemannii (Vanderyst) Van
der Veken ex Compère, Zaire.
Mesophyll with distinct palisade and spongy
mesophyll layers.
Fig. 61. Panicoideae-Chasmanthieae. Orthoclada afri-
cana. A Rhizome. B Leafy culm. C Part of underside of
333. Lophatherum Brongn. leaf blade showing tessellate venation. D Pseudopetiole. E
Ligule. F Inflorescence. G Spikelet. H Lower glume. I
Lophatherum Brongn., Voy. Monde 2: 49 (1829) [1831]. Upper glume. J Lemma. K Palea and sterile lemmas. L
Lodicules. M Flower. (From Clayton 1970a)
Perennials with tuberous roots. Culms solid or
hollow. Ligule a ciliolate membrane. Primary
branches of the inflorescence unbranched, with Perennials. Culms hollow. Leaf blades narrow.
the spikelets borne on one side. Spikelets with one Ligule membranous. Primary inflorescence
fertile flower plus up to 9 sterile lemmas. Lemmas branches branched or unbranched. Spikelets
awned, those of sterile lemmas forming a set of with 1 to 4 flowers, the rachilla ending in a rudi-
recurved hooks. Stamens 2 or 3. 2n ¼ 48. mentary flower. Rachilla internode partially fused
Two spp., tropical Asia. to keels of palea. Lemmas acuminate but not
awned. Stamens 2 or 3. 2n ¼ 24.
Two spp., Tropical America and Africa.
334. Orthoclada P. Beauv. Fig. 61 Orthoclada is sister to Zeugites s.l. plus
Lophatherum in molecular phylogenies (Sán-
Orthoclada P. Beauv., Ess. Agrostogr. 69 (1812). chez-Ken and Clark 2007, 2010). It is
distinguished by its bisexual flowers and the

unusual fusion of its rachilla internode to the
palea keels.
335. Zeugites P. Browne
Zeugites P. Browne, Civ. Nat. Hist. Jamaica 341 (1756).

Calderonella Soderstr. & H.F. Decker, Ann. Missouri Bot.
Gard. 60: 427 (1973).
Pohlidium Davidse, Soderstr. & R.P. Ellis, Syst. Bot. 11:
131 (1986).
Annuals or perennials. Leaf base narrow, pseudo-

petiolate or not. Ligule membranous or a ciliolate
membrane. Inflorescence with branched branches.
Spikelets with 2 to 15 unisexual flowers, with both
sexes combined in a single spikelet. Lowermost
flower pistillate, distal flowers staminate. Pistillate
flower often with the lemma gibbous, sometimes
strongly so. Stamens 3. 2n ¼ 24, 48.
Twelve spp., mid-elevations in Central and
South America.
Zeugites, Calderonella, and Pohlidium clearly
form a monophyletic group in the data analyses
of Sánchez-Ken and Clark (2007, 2010) and
GPWG II (2012), and share the synapomorphy
of unisexual spikelets; they are accordingly
united here. Fig. 62. Panicoideae-Gynerieae. Gynerium sagittatum. A
Plant. B Portion of pistillate inflorescence. C Pistillate
spikelet, lateral view. D Flower of the pistillate spikelet.
XX. TRIBE GYNERIEAE Sánchez-Ken & L.G. Clark E Palea and rachilla, viewed from the adaxial side. F
(2001) Staminodes. (From Zuloaga et al. 2012)
Plants rhizomatous, perennial, reed-like, up to 10

m tall, dioecious. Culms solid. Leaf blades disar- Characters of the tribe. 2n ¼ 44.
ticulating from the sheaths. Ligule a line of hairs One sp., G. sagittatum (Aubl.) P. Beauv., Cen-
or a fringed membrane; abaxial ligule present or tral and South America, West Indies.
absent. Inflorescences highly branched. Spikelets
disarticulating above the glumes. Pistillate spike- XXI. TRIBE CENTOTHECEAE Ridl. (1907)
let with 2 flowers, laterally compressed, the upper
glume much longer than the lemmas, extended
into a long attenuate tip, recurved; lemma plu- Thysanolaeneae C. E. Hubb. (1934).
Cyperochloeae L. Watson & Dallwitz ex Sánchez-Ken &
mose, with attenuate tip; staminodes 2. Staminate L. G. Clark (2010).
spikelet with 2 to 4 flowers, disarticulating below
the most distal of these, glabrous or sparsely Annuals or perennials. Spikelets with few to many
pubescent, the glumes and lemmas not extended; flowers, laterally compressed, reduced flowers
stamens 2. C3. either proximal or distal to the bisexual ones. C3.
Five genera, eight species.
336. Gynerium Willd. ex P. Beauv. Fig. 62 This is a small tribe of only a few species of
Africa, southern Asia, Australia and the Pacific.
Gynerium Willd. ex P. Beauv., Ess. Agrostogr. 138 (1812). Although the gross morphology of the plants is
not at all similar, the group is strongly supported have a broad apical cell and may be similar to
by molecular data (GPWG II 2012; Morrone et al. those of Centotheca and Megastachya.
2012; Sánchez-Ken and Clark 2010). While the
group can be divided into three tribes, this
339. Spartochloa C. E. Hubb.
seems unnecessary given the small number of
genera and species. Spartochloa C. E. Hubb., Kew Bull. 7: 308 (1952).
337. Thysanolaena Nees Caespitose. Leaf blades lacking, the culms and
sheaths photosynthetic. Primary branches of
Thysanolaena Nees, Edinburgh New Philos. J. 18: 180 inflorescence branched. Glumes, lemmas and
(1835). paleas glabrous. Fruit trigonous, pitted.
One sp., S. scirpoidea (Steud.) C.E. Hubb., SW
Caespitose perennials, 2–4 m tall. Leaf blades Australia.
disarticulating, with cross veins. Ligule membra-
nous. Inflorescence branches branched, the MEGASTACHYA + CENTOTHECA
branches lax and spreading. Spikelets with two Mushroom-button microhairs on leaves.
flowers, disarticulating below the glumes.
Glumes about the same length, shorter than the 340. Megastachya P. Beauv.
spikelet. Lower flower reduced to a sterile lemma,
upper flower bisexual, the lemma lanceolate. C3, Megastachya P. Beauv., Ess. Agrostogr. 74 (1812).
mesophyll cells with invaginated cell walls.
2n ¼ 24. Annuals. Base of leaf blades clasping the stem.
One sp., T. latifolia (Roxb. ex Hornem.) Primary inflorescence branches branched or not.
Honda, tropical Africa, Asia, and the Pacific. Spikelets with 8 to 20 flowers, these all similar but
Although the spikelet morphology of this spe- the distal ones somewhat smaller. Lemmas
cies is superficially similar to that of Panicoideae mucronate.
s.s., the rachilla extends as a short stub beyond Two spp., forests of tropical Africa.
the upper flower, a character rarely seen in Pani-
coideae.
341. Centotheca Desv. Fig. 63
CYPEROCHLOA + SPARTOCHLOA
Perennials with wiry culms. Ligule a fringe of Centotheca Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2:
hairs. Spikelets with 4 to 9 flowers, the distal ones 189 (1810), nom. conserv.
smaller. Glumes approximately the same length,
shorter than the lowest lemma. Annuals. Leaf blades narrow, not clasping the
stem. Primary inflorescence branches branched
or not. Spikelets with 1 to 4 flowers, these all
338. Cyperochloa Lazarides & L. Watson similar but the distal ones somewhat smaller.
Lemma with reflexed bristles. Stamens 2.
Cyperochloa Lazarides & L. Watson, Brunonia 9: 216 Four spp., West Africa, Asia, Queensland,
(1986).
Pacific islands.
Plants rhizomatous. Leaf blades filiform, blades
lacking on leaves immediately subtending the
inflorescence. Inflorescence a cluster of 2 to 5 PANICOIDEAE S.S. CLADE
spikelets, these sessile or nearly so. Glumes
pubescent. Lemmas and paleas pilose. Flowers two per spikelet, the upper one hermaph-
One sp., C. hirsuta Lazarides & L. Watson, rodite, staminate, or pistillate, and the lower
SW Australia. either staminate or reduced to an empty lemma.
The microhairs shown in the figure accom- This clade is the group that is called Panicoi-
panying the original description of the genus deae in most of the grass literature. It is one of the
Aquatic annuals. Leaf blades floating, the adaxial

surface with sinuous flattened projections or
lamellae. Ligule membranous, auricles promi-
nent. Inflorescence with unbranched branches,
these appressed to the central axis. Spikelets ped-
icellate. Lower glume hyaline, without veins,
upper glume gibbous. Upper lemma and palea
leathery. C3.
One sp., H. manicatum C. E. Hubb., Zambia
and Zaire. Aquatic.
Molecular data are unavailable for Hydro-
thauma. In the analyses of Morrone et al.
(2012), it was resolved as sister to one or more
of the tribes of Panicoideae s.s.
343. Lecomtella A. Camus
Lecomtella A. Camus, Compt. Rend. Hebd. Séances Acad.

Sci. 181: 567 (1925).
Rhizomatous perennials, monoecious. Ligule a

fringe of hairs. Abaxial ligule a line of hairs.
Inflorescence branches with 1 to 4 staminate spi-
kelets proximally, and a single bisexual spikelet
distally. Bisexual spikelets with the lower flower
staminate, upper flower pistillate, lower and
upper glumes separated by a distinct internode,
rachilla internode below upper flower winged;
upper flower with lemma pubescent, tuberculate,
the apex ending in three appendages. C3. Meso-
phyll with non-radiate chlorenchyma.
One sp., L. madagascariensis A. Camus,
Madagascar.
Fig. 63. Panicoideae-Centotheceae. Centotheca lappacea Lecomtella was provisionally placed sister to
(L.) Desvaux. A Habit. B Spikelet. C Lower glume. D Gerritea in Paspalinae by Morrone et al. (2012)
Upper glume. E Unarmed lemma. F Palea. G Palea. H based on morphological data, but molecular data
Pistil. I Armed lemma. (From Bor 1960, drawn by S.
Ross-Craig) (Besnard et al. 2013) place it in an unresolved
position at the base of Panicoideae. The tribal
name Lecomtelleae Pilg. ex Potztal is available
most distinctive groups of grasses and has been
for the genus. However, the tribe is not recognized
recognized for two centuries, since the work of
here pending more definitive phylogenetic data.
Robert Brown (1810, 1814). The spikelet mor-
phology is diagnostic and synapomorphic.
344. Oryzidium C.E. Hubb. & Schweick.
Oryzidium C.E. Hubb. & Schweick., Bull. Misc. Inform.

INCERTAE SEDIS Kew 1936: 326 (1936).
342. Hydrothauma Hubbard Perennial, aquatic and floating. Ligule a fringe of

hairs. Inflorescence branches branched. Lower
Hydrothauma Hubbard, Hooker’s Icon. Pl. 35: t3458
glume a tiny scale, upper glume longer than the
(1947).
spikelet, awned. Rachilla internode elongated 90 genera, 1274 spp.

between flowers. Lower flower staminate, the Andropogoneae are monophyletic in all
upper pistillate. C4, with two bundle sheaths and molecular phylogenetic studies (Bomblies and
even outline. Doebley 2005; Estep et al. 2014; GPWG II 2012;
One sp., O. barnardii C.E. Hubb. & Schweick., Hodkinson et al. 2002; Kellogg 2000; Lukens and
Zambia and SW Africa. Doebley 2001; Mathews et al. 2002; Skendzic et al.
Leaf anatomy suggests that this species may 2007; Spangler et al. 1999; Teerawatananon et al.
be either in Melinidinae or Panicinae, whereas the 2011), with the genus Arundinella (formerly the
long internode between the flowers is reminiscent type genus of the tribe Arundinelleae) sister to
of Lecomtella. The phylogenetic study of Morrone remaining members of the tribe. Molecular data
et al. (2012) places it either within Melinidinae indicate that the tribe is the result of a rapid
(Paniceae), or sister to Reynaudia (Paspaleae) radiation, but the forces driving this radiation
based on morphology; molecular data are are unknown. Allopolyploidy is remarkably com-
unavailable. mon and can be documented for at least a third of
the species in the tribe (Estep et al. 2014).
Members of Andropogoneae have paired spi-
345. Reynaudia Kunth
kelets, one sessile and one pedicellate, although
Reynaudia Kunth, Révis. Gramin. 1: 72 (1829). this characteristic is shared with many other
Panicoideae (Kellogg 2000; Zanotti et al. 2010).
Perennials. Ligule a fringed membrane. Leaf Throughout the tribe, the pedicellate spikelet is
blades filiform. Inflorescence branches branched, often staminate, but in a few genera it is hermaph-
erect. Upper and lower glumes and the lower rodite and in still others pistillate. Anatomically,
lemma as long as the spikelet, bifid, each with a epidermal papillae appear to have been ancestrally
single awn. Upper lemma smooth, acute. Upper absent, and costal short cells ancestrally in long
palea reduced or lacking. Stamens 2. C4, with one rows (Watson and Dallwitz 1992 onward). In addi-
bundle sheath. tion, in most species the rachis (inflorescence axis)
One sp., R. filiformis (Spreng. ex Schult.) breaks up at maturity and the glumes are hard-
Kunth, West Indies. ened, particularly in comparison with the lemma.
Morrone et al. (2012) place Reynaudia sister The disarticulating main axis is lost independently
to Paspaleae but with weak to moderate support. in several genera, whereas in other genera the
The spikelet morphology is quite different from lateral branches disarticulate as well. There are
that of other Paspaleae, and the chromosome many changes in glume texture, from membra-
number is unknown. nous to coriaceous to extremely hard and back
(Preston et al. 2012). Most species have lemmas
that bear an awn that is usually twisted and hygro-
XXII. TRIBE ANDROPOGONEAE Dumort. scopic.
(1824) All Andropogoneae use the C4 photosynthetic
pathway with NADP-ME as a decarboxylating
Arundinelleae Stapf (1898).
Garnotieae Tateoka (1957). enzyme. Vascular bundles of the leaf have a single
sheath (Hattersley and Watson 1975). This pho-
Ligule membranous. Inflorescence complex and tosynthetic pathway and the accompanying leaf
difficult to describe using conventional terms (see anatomy are synapomorphies for the tribe
below). Spikelets paired, with one sessile or (GPWG II 2012).
nearly so, the other pedicellate; branches often The overall form of the inflorescence in
ending with three spikelets, one sessile and two Andropogoneae varies extensively even within
pedicellate. Glumes firm, sometimes indurate and clades, and this may be why so few of the Clayton
lignified. Lemma and palea hyaline. Rachis disar- and Renvoize (1986) subtribes, which rely heavily
ticulating so that the spikelet pair becomes the on inflorescence form, have proven to be mono-
dispersal unit. All taxa C4, NADP-ME subtype; phyletic (Kellogg 2000; Mathews et al. 2002;
veins with a single bundle sheath. Skendzic et al. 2007; Spangler et al. 1999).
Reduction of the pedicellate spikelet has occurred Many of the possible combinations of these
apparently more than once and does not by itself four characters are observed, making key con-
indicate relationship of genera. The tribe has struction difficult. The rapid radiation documen-
been heavily split such that many genera have ted by molecular phylogenies suggests that the
only one or two species. Watson and Dallwitz morphological diversity might be controlled by a
(1992 onward) recognize 111 genera in the tribe, handful of genes that were easily modified and/or
of which 32 are monotypic. Clayton and Renvoize polymorphic at one time in evolutionary history.
(1986) combine a number of these, and recognize For discussion on the valid name of the tribe
85 genera, of which 21 still include only a single Andropogoneae, see Welker et al. (2014).
species, and another eight include two species.
The complex form of “inflorescences” in
Andropogoneae reflects variation in several XXII.1. SUBTRIBE ARUNDINELLINAE Honda (1930)
characters:
1) Proliferation of branches at upper culm nodes. Ligule membranous, or a fringed membrane. Inflo-
Whereas most monocots, and indeed most rescence branches branched, neither the rachis nor
grasses, have a single axillary branch and adaxial the branches disarticulating at maturity. Spikelets
prophyll at a node, at the upper culm nodes of generally paired, the “sessile” one actually short
many Andropogoneae, the axillary branches pedicellate. Disarticulation above the glumes.
themselves branch, forming a prophyll and Leaves with minor veins reduced to single isolated
another branch but without a corresponding sub- bundle sheath cells (“distinctive cells”).
tending leaf. Each branch of this complex can Two genera, 87 species.
then terminate in an inflorescence, or can reiter- This subtribe is monophyletic in the analyses
ate the branch complex, creating compound of GPWG II (2012) and strongly supported as
structures. The leaves subtending the entire com- sister to all other Andropogoneae. While Teera-
plex are variously called leaves or spathes in the watananon et al. (2011) also find Arundinella to
literature, and may be developed or suppressed. be sister to all other Andropogoneae, they place
The entire upper culm, including all branch com- Garnotia sister to Eremochloa. Garnotia and
plexes, is often called an inflorescence in the Arundinella share enough morphological and
literature, but here the term inflorescence is anatomical characters that placing them together
reserved for whatever structure develops after seems plausible, although this needs to be
the uppermost leaf or spathe. checked with more data.
2) The ultimate inflorescences may produce
branches (often called “racemes” or “rames” in 346. Arundinella Raddi Fig. 64
the literature), which initiate in a spiral phyllo-
taxis. They may develop continuously with the Arundinella Raddi, Agrostogr. Bras. 36 (1823).
apical meristem, or branch elongation may be
delayed relative to the development of the pri- Callus pubescent. Glumes the same texture as or
mary axis. Each branch then produces spikelet thinner than the lemmas, the apex acute to muti-
pair meristems in a two-ranked phyllotaxis. cous, the upper glume apex often caudate. Upper
3) Both the sessile and pedicellate spikelets of a lemma apex generally awned. 2n ¼ 14, 20, 28,
pair may develop, or one member of the pair may 36, 56.
be reduced or suppressed entirely. Often the Fifty-seven spp., most Asian, others tropics
reduction occurs in the pedicellate spikelet, but and subtropics, north to Japan.
sometimes the sessile spikelet is the one that is
reduced. The two spikelets may be identical in
morphology and sex expression, but more often 347. Garnotia Brongn.
they are different.
4) Both flowers of a spikelet may develop or one Garnotia Brongn., Voy. Monde 2: 132–133, pl. 21 (1830).
may be suppressed.
ANDROPOGONEAE incertae sedis
348. Apluda L.
Apluda L., Sp. Pl. 1: 82 (1753).
Decumbent or scrambling perennials. Ligule a

fringed membrane. Axillary branch complexes
present. Inflorescences unbranched, each with a
spathe-like subtending leaf; internodes and pedi-
cels thickened, clavate. Spikelets borne in triads,
with one sessile and two pedicellate, pedicels flat
and broad. One pedicellate spikelet staminate, the
other vestigial, represented by a solitary glume.
Sessile spikelet with bulbous callus; lower glume
winged on keel. Upper glume gibbous. Upper
lemma muticous or awned from a sinus, the
awn twisted or straight. 2n ¼ 20, 40.
One sp., A. mutica L., tropical Asia.
349. Arthraxon P. Beauv.
Arthraxon P. Beauv., Ess. Agrostogr. 111 , pl. 11, f. 6

(1812).
Annuals or perennials, decumbent. Leaves

broadly lanceolate with cordate base. Ligule
membranous, or a fringed membrane or a fringe
of hairs. Inflorescence unbranched or the
Fig. 64. Panicoideae-Andropogoneae-Arundinellinae. A–C branches digitate. Pedicellate spikelets staminate,
Arundinella nepalensis. A Habit. B Spikelet pair. C Abaxial
and adaxial views of upper flower. D–F A. parviflora. D sterile, reduced to a pedicel or lacking. Sessile
Inflorescence. E Spikelet pair. F Abaxial and adaxial views spikelets with the callus glabrous or pubescent,
of upper flower. (From Wu et al. 2007, with permission the lower glume smooth or rugose or with
from the Missouri Botanical Garden Press, St. Louis, and
Science Press, Beijing; drawn by Zhang Jiade)
prickles, variously toothed on the margins, both
flowers generally lacking paleas. Upper lemma
apex entire or lobed, awned, the awn abaxial,
Spikelets occasionally solitary, in pairs or in attached well below the midpoint of the lemma.
groups of three. Spikelets laterally compressed, Stamens 2 or 3. 2n ¼ 18, 20, 36, 40.
with a single flower, the proximal sterile flower Twenty-seven spp., Old World tropics,
absent; callus glabrous or pubescent. Glumes thin- mostly India, some species weedy in other warm
ner than the lemmas, the apex muticous or awned. regions of the world.
Upper lemma apex muticous or awned. 2n ¼ 20.
Thirty spp., Africa, Asia, Australia, Pacific. 350. Chasmopodium Stapf
Chasmopodium Stapf, Fl. Trop. Afr. 9: 76 (1917).

ANDROPOGONEAE s.s. Robynsiochloa Jacq.-Fél., J. Agric. Trop. 7: 406 (1960).
Possible synapomorphies for this clade include a Annuals, 2–4 m tall. Ligule a fringe of hairs or
disarticulating rachis and a twisted hygroscopic membranous. Inflorescence branches unbranched.
awn on the lemma (Estep et al. 2014). Internodes and pedicels thickened, flattened.
Sessile spikelet callus pubescent, with a central plus Polytoca. Chionachne falls in an unresolved
peg. Pedicellate spikelets sterile. Lower glume of position among the awnless Andropogoneae in
sessile spikelet with narrow wings, upper glume molecular phylogenies (Estep et al. 2014; GPWG
laterally compressed, winged on keel. Lemma with- II 2012; Teerawatananon et al. 2011).
out an awn. Embryo 4/5 as long as caryopsis.
2n ¼ 16, 20.
352. Chrysopogon Trin.
Three spp., tropical Africa.
Clayton and Renvoize (1986) synonymize Chrysopogon Trin., Fund. Agrost. 187 (1820), nom. con-
Robynsiochloa with Rottboellia, but that seems serv.; rev.: Veldkamp (1999).
to be just one of several possible placements. Vetiveria Bory, Bull. Sci. Soc. Philom. Paris 1822: 43
Clayton (1970b) indicates that Robynsiochloa (1822).
and Chasmopodium are “barely distinct” in a
numerical taxonomic analysis, and later provides Annuals or perennials, caespitose, rhizomatous,
the necessary combination in Chasmopodium or stoloniferous. Ligule membranous, or a
(Clayton 1973). The winged upper glume is dis- fringed membrane, or a fringe of hairs. Inflores-
tinctive and possibly indicates monophyly. The cence branches branched, the spikelets in pairs or
internode and pedicel are fused in one species. triads, if the latter then one spikelet sessile and
two pedicellate. Spikelets laterally compressed, the
callus truncate to acute or pungent, the attach-
351. Chionachne R. Br. ment oblique or inserted into the cup-like inter-
node apex. Pedicellate spikelets staminate, sterile,
Chionachne R. Br., Pl. Jav. Rar. 15 (1838); rev., Jannink
and Veldkamp (2002). or reduced to glumes, the glumes muticous to
Sclerachne R. Br., Pl. Jav. Rar. 15 (1838). awned. Sessile spikelets with glumes membra-
Polytoca R. Br., Pl. Jav. Rar. 20 (1838). nous to leathery but not indurate, the upper one
Cyathorhachis Nees ex Steud., Syn. Pl. Glumac. 1: 403 mucronate to awned. Upper lemma awned from
(1854). the apex or from a sinus, the awn straight or
Trilobachne M. Schenck ex Henrard, Meded. Rijks-Herb.
67: 4 (1931). twisted. 2n ¼ 20, 40.
Forty-eight spp., tropical and subtropical
Annuals or perennials, monoecious. Ligule mem- throughout the world.
branous or a fringed membrane. Inflorescence Vetiveria is merged with Chrysopogon here
unbranched, with pistillate spikelets below, stami- following the revision by Veldkamp (1999), a
nate above, or with several primary branches result confirmed by molecular data (Adams et al.
bearing staminate spikelets only. Pistillate spike- 1998; Estep et al. 2014). Chrysopogon zizanioides
lets unpaired or with staminate pedicellate spike- (also known as vetiver) is economically important
lets; lower glumes hard, shining, surrounding the as a source of oil and also for erosion control (Joy,
rachis, enlarged to look reminiscent of the spathe undated; Panel on Vetiver et al. 1993).
in Coix. Staminate spikelets paired, two-flowered,
pedicellate spikelets reduced to an empty glume. 353. Clausospicula Lazarides
Lower glume of the pistillate spikelet lobed,
obtuse or notched at the apex, with two obvious Clausospicula Lazarides, Austral. Syst. Bot. 4: 399 (1991).
constrictions in one species. Lemmas without
awns. Caryopsis flattened, concave-convex. Annuals. Leaf blades with margins cartilaginous,
2n ¼ 20, 40. the ligule a fringed membrane. Inflorescence
Twelve spp., India, SE Asia, Australia, and unbranched, with one sessile and two pedicellate
Polynesia. spikelets, the set of three surrounded by the trum-
Four small genera are included here in Chio- pet-shaped tip of the peduncle. Pedicellate spikelets
nachne, based on their shared inflorescence char- well developed or rudimentary. Sessile spikelets
acters and overlapping geographic distribution. with a pubescent callus, sharp-pointed, the lower
Teerawatananon et al. (2011) and Estep et al. glume rugose, muticous or awned, upper glume
(2014) confirm the monophyly of Chionachne with a long awn. Both flowers of a spikelet lacking
paleas. Lemma apex awned, the awn geniculate, Annuals or perennials, caespitose. Ligule membra-
pubescent. nous, or a fringed membrane, or a fringe of hairs.
One sp., C. extensa Lazarides, N. Australia. Inflorescences unbranched or branched, the rachis
Clausospicula is somewhat similar to the south- flattened, white-hairy; apex of internodes not hol-
east Asian species Kerriochloa siamensis. Clayton lowed or rimmed, base of internodes oblique,
et al. (2006 onward) provide a slightly different hairy. Pedicellate spikelets staminate, the lower
interpretation of the inflorescence morphology, glume muticous or short-awned. Sessile spikelet
and suggest that the structure here described as bisexual, the lower glume with a bifid tip, with two
the tip of the peduncle is in fact a bract subtending keels with hairs in tufts or lines and an oil streak on
the branch of a highly branched inflorescence. the inner side of the keels; lower palea absent;
Another interpretation would suggest that the ped- lemmas hyaline, unawned. 2n ¼ 10, 20.
uncle tip is homologous to the flattened structure at Fifteen spp., Tropical Africa, America,
the base of the branches in Hyperthelia. Australia.
Elionurus falls in an unresolved position
among the awnless Andropogoneae in the molec-
354. Coix L.
ular phylogenies of GPWG II (2012), Teerawata-
Coix L., Sp. Pl. 2: 972 (1753). nanon et al. (2011), and Estep et al. (2014). The
chloroplast phylogeny of GPWG II (2012) places
Annuals or perennials, caespitose or rhizoma- it sister to Pseudopogonatherum contortum, a
tous, monoecious. Ligule membranous. Inflores- position that is hard to explain morphologically.
cence a complex branched structure with
staminate spikelets borne distally on the primary 356. Euclasta Franch.
axis; lateral branch demarcated by a prophyll,
forming in the axil of a highly modified leaf (“utri- Euclasta Franch., Bull. Soc. Hist. Nat. Autun 8: 335 (1895).
cle”); lateral branch bearing a triad of spikelets;
two pedicellate ones develop thickened pedicels Sprawling annuals. Ligule a fringed membrane.
and initiate spikelet parts but do not develop Inflorescence unbranched or with digitate
further. The sessile spikelet becomes pistillate. branches, nodding. Proximal 1 to 3 pairs of spike-
Glumes membranous, not indurate. Lemmas sim- lets entirely staminate. Distal pedicellate spikelets
ilar in texture to the glumes, without awns. staminate or sterile. Distal sessile spikelets bisexual
Embryo as long as the caryopsis. 2n ¼ 10, 20, 40. with a pubescent callus. Flowers lacking paleas.
Four spp., tropical Asia. Lemma apex awned, the awn geniculate. 2n ¼ 40.
The “utricle” is clearly a modified leaf sheath Two spp., tropical regions of Africa, south
with fused margins and only a rudimentary blade. Asia, and the Americas.
Sheaths elsewhere in the inflorescence have fused Watson and Dallwitz (1992 onward) report
margins as well. Many authors (e.g., Hitchcock that the shoots are aromatic.
1950; Watson and Dallwitz 1992 onward) have
suggested a relationship between this genus and
357. Eulaliopsis Honda
Zea, but molecular data do not support this idea.
Coix falls in an unresolved position among the Eulaliopsis Honda, Bot. Mag. (Tokyo) 38: 56 (1924).
awnless Andropogoneae in molecular phylogenies
(Estep et al. 2014; GPWG II 2012; Teerawatananon Perennials. Ligule a fringed membrane or a fringe
et al. 2011). The chloroplast phylogeny of GPWG II of hairs. Inflorescence branches unbranched, dig-
(2012) places it sister to Ischaemum afrum. itate. All spikelets bisexual, callus pubescent.
Lower glumes of both spikelets with many prom-
355. Elionurus Humb. & Bonpl. ex Willd. inent raised veins. Sessile spikelet with lower
glume with tufts of long hairs about midway up
Elionurus Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 941 abaxial side; upper glume thinner than lower,
(1806), nom. et orth. cons. with a slender awn. Lemma apex awned from a
sinus, the awn geniculate. Lodicules ciliate. Leaf 360. Lakshmia Veldk.
cross section with minor vascular bundles
crowded, almost superposed. Lakshmia Veldk., Rheedea 18: 81 (2013).
Two spp., India and SE Asia (Philippines),
China, Afghanistan. Caespitose perennials. Ligule membranous. Inflo-
rescence branches branched, the ultimate
branches with 3 to 7 spikelet pairs. Sessile spikelet
358. Hypogynium Nees callus short, blunt, with white hairs. Glumes char-
taceous, their apices acuminate. Upper lemma
Hypogynium Nees, Fl. Bras. Enum. Pl.: 364 (1829). with the apex bifid, awned from the sinus, the
awn geniculate. Pedicellate spikelets similar to
Caespitose perennials, monoecious. Ligule mem- the sessile ones in size and shape, but lacking
branous. Axillary branch complexes present. awns, staminate or sterile.
Inflorescence unbranched, the base of the dis- One species, L. venusta (Thwaites) Veldk.,
persal unit truncate. Sessile and pedicellate spike- India and Sri Lanka.
lets morphologically similar, the sessile ones Lakshmia venusta has been variously placed
pistillate, the pedicellate ones staminate. Glumes in Andropogon, Capillipedium, Bothriochloa,
and lemmas unawned. Pistillate flowers with 3 Vetiveria, and most recently Hemisorghum.
staminodes. 2n ¼ 30. Molecular data would be particularly useful to
Two spp., tropical Africa and America. place this species.
Some authors (e.g., Clayton and Renvoize
1986; Renvoize 1998) consider Hypogynium to
be a synonym of Andropogon, whereas others (e. 361. Lasiurus Boiss.
g., Rosengurtt et al. 1970; Watson and Dallwitz
Lasiurus Boiss., Diagn. Pl. Orient. 4: 145 (1859).
1992 onward) consider it to be separate on the
basis of its unisexual, unawned spikelets, and
Perennials with short rhizomes. Ligule a fringe of
unbranched inflorescence axes. No molecular
hairs. Inflorescence unbranched or of digitate
data are available to help with its placement.
primary branches, the internodes clavate, with
The chromosome number suggests an allopoly-
silky hairs. Spikelets in groups of three, two ses-
ploid origin.
sile and one pedicellate. Sessile spikelet callus
pubescent, with a central peg. Lower glume of
359. Kerriochloa C.E. Hubb.
sessile spikelet two keeled, flattened on the back
with a short apical extension; upper glume thin-
Kerriochloa C. E. Hubb., Hooker’s Icon. Pl. 35: pl. 3494 ner and boat-shaped. Lemma apex acute, without
(1951). an awn. 2n ¼ 18, 56.
One sp., L. scindicus Henrard, E. Africa to NE
Decumbent perennials, less than 25 cm tall. Lig- India.
ule membranous. Axillary branch complexes Faruqi et al. (1987) note that the cytotypes of
present. Inflorescence enclosed by the subtending this species correlate with geography; plants with
leaf sheath, unbranched, the rachis flattened; 2n ¼ 18 are found in India and Pakistan, whereas
internodes wedge-shaped. Sessile spikelet later- those with 2n ¼ 56 are in North Africa. No
ally compressed, the callus obtuse, pubescent, molecular data are available.
inserted into the cupulate apex of the internode
below; lower glume villous, upper glume awned. 362. Microstegium Nees
Lemma of upper flower awned from a shallow
sinus, the awn geniculate. Pedicellate spikelet Microstegium Nees, Nat. Syst. Bot. 447 (1836).
reduced to a tiny vestige with one glume. Ischnochloa Hook. f., Hooker’s Icon. Pl. 25: t. 2466 (1896).
One sp., K. siamensis C. E. Hubb., Thailand
and Indo-China. Annuals or perennials, caespitose. Leaf blades
Molecular data place K. siamensis sister to broadly linear to lanceolate, sometimes with a
Microstegium vimineum (Estep et al. 2014). short pseudopetiole. Ligule membranous, or a
fringed membrane or a fringe of hairs. Inflores- late, all bisexual. Spikelets lacking proximal
cence unbranched, or of digitate or subdigitate flower. Glumes pubescent. Upper lemma awned
primary branches. Rachis internodes enlarged, from a shallow sinus, the awn geniculate. Upper
clavate. Pedicellate spikelets and sessile spikelets palea tiny or absent. Lodicules absent.
similar, bisexual, the callus pubescent. Glumes One sp., P. indica (Houtt.) Veldkamp, South-
generally more firm than the lemmas, the upper east Asia.
glume often with a long awn, lower glume con- The spikelet arrangement in Polytrias is similar
cave between keels. Upper lemma awned from the to that in Mnesithea, Ratzeburgia, and
apex or from a sinus, the awn straight or genicu- some Coelorachis, but the plant is much more deli-
late. 2n ¼ 20, 40. cate, the glumes are truncate, the rachis is slender,
Twenty-seven spp., Asia, Africa, some species and the lemma bears an awn. Molecular data (Estep
weedy elsewhere. et al. 2014; Teerawatananon et al. 2011) place this in
Chen et al. (2009) separate Microstegium an unresolved position in the tribe.
from Leptatherum, based on morphological and
molecular data. The monotypic Ischnochloa (¼
365. Sehima Forssk.
Microstegium falconeri) is placed here, consistent
with Clayton et al. (2006 onward); the plant illu- Sehima Forssk., Fl. Aegypt.-Arab. 178 (1775).
strated in the protologue has a strongly flattened
rachis similar to that in Microstegium. The posi- Annuals or perennials, caespitose. Ligule a line of
tion of the lemma awn is similar to that in hairs or fringed membrane. Inflorescence
Arthraxon, but the lemma appears to be deeply unbranched and curved, the spikelets embedded
bilobed and the pedicellate spikelet is bisexual in the rachis; internodes and pedicels thickened,
and awned. Molecular data place M. vimineum clavate. Sessile spikelet with the lower glume with
sister to Kerriochloa siamensis but with little sup- two keels, the apex mucronate to awned; the
port (Estep et al. 2014). upper glume membranous, green, awned, with
prominent veins converging toward the apex;
363. Pogonachne Bor upper lemma awned from a sinus, the awn genic-
ulate. Pedicellate spikelet staminate or sterile,
Pogonachne Bor, Kew Bull. 4: 176 (1949). strongly flattened, the callus fitting into a concave
depression in the pedicel; glume with a terminal
Annuals to 1 m tall, with prop roots forming from bristle. 2n ¼ 34, 40.
the lower nodes. Upper leaves without blades. Five spp., Old World tropics.
Ligule membranous. Axillary branch complexes Andropogon sect. Notosolen looks similar to
present. Inflorescence unbranched, the rachis flat- Sehima but has more than one inflorescence
tened. Spikelets apparently unpaired, the sessile branch. Molecular data (Estep et al. 2014) put
spikelet reduced to small scales or absent. Pedicel- Sehima in an unresolved position in the tribe.
late spikelet laterally compressed; callus pubes-
cent, truncate. Lower glume hairy, upper glume
gibbous, with a tuft of hair. Upper lemma awned 366. Spodiopogon Trin.
from a sinus, the awn geniculate.
Spodiopogon Trin., Fund. Agrost. 192, pl. 17 (1820).
One sp., P. racemosa Bor, India. Eccoilopus Steud., Syn. Pl. Glumac. 1: 123 (1854).
364. Polytrias Hack. Annuals or perennials, caespitose or rhizoma-

tous. Leaf blades sagittate or not, pseudopetiolate
Polytrias Hack., Nat. Pflanzenfam. 2(2): 24 (1887). or not, rarely deciduous. Ligule membranous or a
fringed membrane. Inflorescence branches
Stoloniferous perennials. Ligule a fringed mem- branched, the spikelets set into a cup-like depres-
brane. Inflorescence unbranched. Spikelets in sion at the distal end of the rachis joint. All
threes, two apparently sessile but actually on a spikelets bisexual, callus pubescent. Lower
short flattened pedicel, the third clearly pedicel- glume scabrous and pubescent; veins all equally
prominent (no clear keel). Upper glume muticous parental genomes of the paleotetraploids Zea and
or mucronate. Lower lemma mucronate or Tripsacum (McKain and Kellogg, unpubl. data).
awned; upper lemma awned from a sinus, the The lack of awns may be synapomorphic,
awn geniculate. Lodicules ciliate. 2n ¼ 40, 42. although awns are also lost in Rottboelliinae.
Eighteen spp., SE Asia, temperate Asia, Mid-
dle East.
369. Oxyrhachis Pilg.
367. Thelepogon Roth Oxyrhachis Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 11:
655 (1932).
Thelepogon Roth, Syst. Veg. 2: 46 (1817).
Caespitose perennials. Leaf blades filiform, rigid,
Annuals, often with prop roots. Leaf blades the ligule a fringed membrane. Inflorescence single,
broad, with the base cordate. Ligule membranous unbranched, terminal. Pedicellate spikelets absent,
or a fringed membrane. Inflorescence branched, the pedicel sometimes visible but fused to the inflo-
the primary branches borne on a short rachis, the rescence axis. Sessile spikelets sunken in internode,
proximal portion of each branch without spike- producing a rat-tail inflorescence. Spikelet callus
lets. Sessile spikelet with the callus pubescent, the blunt, without a cylindrical peg, attached obliquely.
lower glume strongly sculptured with hooked pro- Lower flower of sessile spikelet sterile.
jections; upper lemma awned from a sinus, the One sp., O. gracillima (Baker) C.E. Hubb.,
awn geniculate. Pedicelled spikelet absent, the Africa and Madagascar.
pedicel ending blindly.
Two spp., tropical Africa, Indonesia, Australia. 370. Rhytachne Desv. ex Ham.
Molecular data place this genus sister to
Chrysopogon (Estep et al. 2014). Rhytachne Desv. ex Ham., Prodr. Pl. Ind. Occid. xiv, 11
(1825).
368. Triplopogon Bor Annuals or perennials, caespitose. Ligule mem-

branous or a fringed membrane. Inflorescence
Triplopogon Bor, Kew Bull. 9: 52 (1954).
unbranched, the spikelets embedded in hollows
in the rachis. Spikelets paired, the sessile spikelet
Plant annual, 0.5–2 m tall, with prop roots. Leaf
squashed between the internode and the pedicel.
blades narrow at the base, more or less pseudo-
Pedicellate spikelets bisexual or reduced, some-
petiolate. Ligule membranous. Axillary branch
times represented only by an awn, or by an empty
complexes present. Inflorescence unbranched,
pedicel, or lacking entirely. Sessile spikelet callus
the rachis flattened, the internodes long hairy.
with central peg. Upper glume awned in some
Sessile spikelet with callus pubescent; lower
species. Costal short-cells of the leaf epidermis
glume with a median groove, with tufts of hair
paired. 2n ¼ 20.
on the margins; upper glume mucronate, with
Twelve spp., tropical Africa and America.
one tuft of hair; upper lemma awned from a
sinus, the awn geniculate. Pedicellate spikelet
rudimentary. 371. Urelytrum Hack.
One sp., T. ramossimus (Hack.) Bor, India.
Urelytrum Hack., Nat. Pflanzenfam. 2: 22, 25 (1887).
XXII.2. SUBTRIBE TRIPSACINAE Dumort. (1829) Perennials, rarely annual, caespitose. Leaves
auriculate. Ligule membranous or a fringed
Inflorescence unbranched. Upper lemma hyaline, membrane. Inflorescence unbranched, or with
lacking awns. digitate branches, the internodes clavate with a
Five genera, 32 species. broad scarious rim, the proximal end with a peg-
Oxyrhachis, Rhytachne, Urelytrum and Vossia like knob after disarticulation. Pedicellate spike-
are well supported as being sister to one of the let staminate or sterile, the lower glume with a
long curved awn. Sessile spikelet with a pubescent

callus inserted into the distal end of the internode
below. Lower flower of sessile spikelet staminate,
with palea. 2n ¼ 20.
Seven spp., tropical Africa.
372. Vossia Wall. & Griff.
Vossia Wall. & Griff., J. Asiat. Soc. Bengal 5: 572 (1836).
Rhizomatous perennials, in wet sites or aquatic

with culms spongy, submerged or floating. Ligule
a fringed membrane. Primary branches of the
inflorescence unbranched, single or digitate; inter-
nodes and pedicels clavate and flat. Pedicellate
spikelets bisexual or staminate. Sessile spikelets
bisexual. Callus truncate. Lower glume of sessile
spikelet two-keeled, flattened on the back,
extended into a flattened tail-like awn; upper
glume thinner and boat-shaped. 2n ¼ 20.
One sp., V. cuspidata (Roxb.) Griff., tropical
Africa and India.
ZEA + TRIPSACUM
Plants monoecious. Staminate and pistillate
spikelets on the same inflorescence axis, with sta-
minate distal to pistillate, or staminate and pistil-
late spikelets in different inflorescences, the
staminate terminal on the plant. Pistillate spike-
lets unpaired, sessile only.
Zea and Tripsacum are sisters in all molecular
phylogenies (e.g., Mathews et al. 2002; Teerawata-
nanon et al. 2011) and are derived from the same
ancestral polyploidy event (Estep et al. 2014).
373. Tripsacum L. Fig. 65
Tripsacum L., Syst. Nat. (ed. 10): 1253, 1261, 1379 (1759).
Perennials, caespitose or rhizomatous, 0.5–5 m Fig. 65. Panicoideae-Andropogoneae-Tripsacinae. Trip-

tall. Leaves without auricles, the ligule a fringed sacum dactyloides. A Rhizome with shoot bases. B Leaves
membrane. Inflorescence branched or and inflorescences with staminate (a) and pistillate spike-
unbranched. Staminate spikelets with both lets (b). C Staminate spikelet. D Pistillate spikelet. (From
Barkworth et al. 2003, drawn by H. Paźdirková)
glumes on the abaxial side of the spikelet, over-
lapping. Pistillate spikelets embedded in the
internode, the node truncate, with a central peg.
Fruit encased in the hard glume plus internode. 374. Zea L.
2n ¼ 36, 72, 90, 108.
Zea L., Sp. Pl. 2: 971–972 (1753).
Sixteen spp., North, Central and South Euchlaena Schrad., Index Sem. (G€ ottingen) 1832: 3
America. (1832).
Annuals or perennials, caespitose or rhizoma- The question is unlikely to be resolved with only
tous, 1–5 m tall. Ligule membranous. Main culm herbarium studies – molecular and developmen-
and primary culm branches ending in a branched tal investigations are needed.
inflorescence of paired staminate spikelets. Pistil-
late inflorescences axillary on the culm branches,
375. Eremochloa B€
use
closely surrounded by spathes; sessile spikelets
distichous and alternate; glume and rachis enlarg- Eremochloa B€
use, Pl. Jungh. 357 (1854).
ing at maturity to enclose fruit in a rock-hard
shining fruit case. 2n ¼ 10, 20. Perennials, caespitose or rhizomatous. Ligule
Seven spp., Mexico and Central America. membranous, or a fringed membrane, or a fringe
Cultivated plants (maize, Z. mays ssp. mays) of hairs. Inflorescence axis flattened with clavate
retain the terminal staminate inflorescence of their internodes and overlapping spikelets. Pedicellate
wild relatives (tassel). Pistillate inflorescences ter- spikelets rudimentary, or reduced only to an awn,
minate short axillary branches on the main axis, or lacking, leaving a barren pedicel. Sessile spike-
and consist of paired pistillate spikelets in multiple lets with the lower glume 2-keeled, setose.
orthostichies, each spikelet with a single flower at 2n ¼ 18.
maturity. Glumes and lemmas are soft and flexible. Twelve spp., India, China, southeast Asia,
Maize is one of the most valuable crop plants in Australia.
the world. Teerawatananon et al. (2011) place Eremo-
chloa with Garnotia with strong support, but the
two genera have little in common morphologi-
XXII.3. SUBTRIBE ROTTBOELLIINAE J. Presl (1830) cally. Other phylogenies place Garnotia with
Arundinella (GPWG II 2012).
Inflorescences unbranched; spikelets generally
sunken in the rachis. Lemmas hyaline, unawned.
11 genera, 99 species. 376. Glyphochloa Clayton
Molecular data place Rottboelliinae in a clade
distinct from Tripsacinae (McKain and Kellogg, Glyphochloa Clayton, Kew Bull. 35: 814 (1981).
unpubl. data). While it is possible that the two are
sisters, evidence for this is weak, and thus lemma Annuals. Ligule membranous. Axillary branch
awns may have been lost either once or twice in complexes present. Pedicellate spikelet sterile, the
the evolution of Andropogoneae. A subset of spe- pedicel fused to the inflorescence axis. Glumes of
cies of Rottboelliinae have pedicels fused to the pedicellate spikelet muticous or awned. Sessile
internode and an overlapping subset have sculp- spikelet callus truncate, with a central peg. Lower
tured glumes. It is unclear whether either charac- glume of sessile spikelet muticous or awned, vari-
ter is synapomorphic here. Molecular studies ously decorated with projections, glabrous or pubes-
(GPWG II 2012; Teerawatananon et al. 2011; cent; upper glume of sessile spikelet unawned.
McKain and Kellogg unpubl. data) support a Nine spp., India.
clade containing at least Eremochloa, Hackelo- Clayton and Renvoize (1986) link Glypho-
chloa, Hemarthria, Heteropholis, Mnesithea, chloa to Heteropholis and Coelorachis clarkei. It
Ophiuros, Phacelurus, and Thaumastochloa, is placed here solely on the basis of morphology.
although no single study includes all taxa. Veld-
kamp et al. (1986) synonymized Coelorachis, 377. Hackelochloa Kuntze
Hackelochloa, Heteropholis, Ratzeburgia, and
Rottboellia with Mnesithea on the basis of con- Hackelochloa Kuntze, Revis. Gen. Pl. 2: 776 (1891).
tinuous variation in most taxonomic characters.
They also question the value of pedicel fusion in Annuals. Ligule a fringed membrane. Axillary
delimiting genera. Molecular data clearly support branch complexes present. Pedicellate spikelets
merger of Coelorachis and Rottboellia, which are bisexual, staminate, or sterile, the pedicel fused
synonymized here, but the monophyly of Mne- to the inflorescence axis, the glumes muticous.
sithea sensu Veldkamp et al. (1986) is less certain. Sessile spikelet callus transverse, with a central
peg. Lower glume of sessile spikelet nearly orbicu- maturity. Glumes of pedicellate spikelet muti-
lar, with a complex reticulate pattern of projec- cous. Sessile spikelet callus truncate, with a cen-
tions and pits. Embryo very large. 2n ¼ 14. tral peg. Lower glume of sessile spikelet broadly
Two spp., tropics. winged, constricted about the middle.
2n ¼ 18, 36.
One sp., M. myuros L., India.
378. Jardinea Steud.
This genus name has been applied to several
Jardinea Steud., Syn. Pl. Glumac. 1: 360 (1854). species now placed in Coelorachis, Rottboellia,
and Glyphochloa; see Clayton (1981) for discus-
Caespitose perennials. Ligule a fringed mem- sion. Manisuris myuros is placed in this subtribe
brane. Inflorescence with primary branches, on the basis of morphology alone.
these sometimes themselves branched. Pedicel-
late spikelets smaller than the sessile ones, 381. Mnesithea Kunth
reduced or vestigial, bisexual, staminate or ster-
ile. Sessile spikelet bisexual, with a pointed cal- Mnesithea Kunth, Révis. Gramin. 1: 153 (1829).
lus. Lower glumes dorsiventrally compressed, Ratzeburgia Kunth, Révis. Gramin. 2: 487 (1831).
sculptured, prickly, the apex acuminate or
extended into a tail-like awn. Upper glumes Caespitose perennials. Ligule membranous or a
laterally compressed. Lemma of sessile spikelet fringed membrane. Axillary branch complexes
keeled. Leaf epidermis with costal short-cells present. Inflorescence unbranched, the axis dis-
paired. articulating or not, internodes expanded at distal
Three spp., tropical Africa. end, spikelets appressed to or embedded in the
The glume compression is similar to that in inflorescence axis. Spikelets two or three per
Loxodera. No molecular data are available; the inflorescence node, two sessile and one pedicel-
genus is placed here based on morphology. late. Pedicellate spikelet reduced, sterile, with
glumes or vestigial. Sessile spikelet callus with a
central peg, fitting in to the cupulate apex of the
379. Loxodera Launert internode below. First glume of sessile spikelet
smooth or sculptured; second glume much less
Loxodera Launert, Bol. Soc. Brot. II, 37: 80 (1963).
Lepargochloa Launert, Bol. Soc. Brot. II, 37: 82 (1963). firm, embedded in the rachis. 2n ¼ 18.
Three spp., India and Southeast Asia.
Caespitose perennials. Ligule membranous. Inflo- The pedicel is fused to the rachis in M. laevis,
rescence unbranched, terminal, the internodes but is free in the other two species.
clavate or columnar with broad scarious rim
bearing long white hairs, the spikelets partially 382. Phacelurus Griseb.
embedded in the rachis. Disarticulation zone
oblique. Pedicellate spikelet staminate, sterile or Phacelurus Griseb., Spic. Fl. Rumel. 2: 423 (1844) [1846].
reduced, the glumes muticous or short-awned. Pseudovossia A. Camus, Bull. Mus. Hist. Nat. (Paris) 26:
Sessile spikelet with a pubescent callus, the 665 (1920).
lower glume strongly veined.
Five spp., tropical Africa. Plants perennial. Ligule membranous. Inflores-
cence unbranched or with digitate primary
branches, the internodes thickened. Pedicellate
380. Manisuris L. spikelet generally reduced, with an elongate callus
in some species. Sessile spikelet bisexual, with a
Manisuris L., Mant. Pl. 2: 164 (1767). blunt glabrous callus. 2n ¼ 20, 40.
Seven spp., Old World tropics.
Plants perennial. Ligule membranous. Pedicellate The single species of Pseudovossia is included
spikelet next to sessile spikelet of internode above here, following Clayton and Renvoize (1986).
by fusion of pedicel, the two falling together at
383. Rottboellia L. f. eropholis as sisters. The latter two are also

strongly supported as sisters by McKain and Kel-
Rottboellia L. f., Suppl. Pl. 13, 114 (1781) [1782], nom. logg (unpubl. data).
conserv.
Coelorachis Brongn., Voy. Monde 2: 64, f. 14 1829 [1831].
Thyrsia Stapf, Fl. Trop. Afr. 9: 48 (1917). 384. Hemarthria R. Br.
Annuals or perennials, caespitose or rhizomatous. Hemarthria R. Br., Prodr. 207 (1810).

Ligule membranous or a fringed membrane, or a
fringe of hairs. Axillary branch complexes present. Perennials (one species annual), caespitose or sto-
Inflorescence unbranched or sometimes with digi- loniferous. Ligule a fringe of hairs or a fringed
tate primary branches; internodes club-shaped or membrane. Axillary branch complexes present.
flattened, generally shorter than spikelets, with Inflorescence rachis internodes disarticulating
sunken spikelets, the peduncle widened at the transversely or obliquely. Pedicellate spikelets
apex. Spikelets in pairs or in groups of three with bisexual or reduced. Sessile spikelets bisexual,
two sessile and one pedicellate spikelet per node. glumes subulate tipped, not rugose, warty or pitted,
Pedicels free or fused wholly or partly to internode. lower glume hardened, upper glume membranous.
Pedicellate spikelet staminate or sterile. Sessile 2n ¼ 18, 20 (18 + 2B), 36, 54.
spikelet callus truncate, with or without a central Fourteen spp., Old World tropics, maybe
peg; lower glume thickened, often decorated, with Americas.
two winged keels, the apex generally emarginate or
obtuse. Lower flower of sessile spikelet staminate 385. Heteropholis C. E. Hubb. Fig. 66
or sterile. 2n ¼ 18, 20, 36, 40, 54.
Thirty-three spp., tropics. Heteropholis C. E. Hubb., Hooker’s Icon. Pl. 36: pl. 3548
Species of Rottboellia and Coelorachis are (1956).
clearly intermixed in molecular phylogenies
(McKain and Kellogg, unpubl. data) so the two Annuals or perennials, often decumbent. Ligule
are united here. The phylogeny suggests that membranous or a fringed membrane. Inflores-
fusion of the pedicel to the inflorescence inter- cence internodes clavate, the peduncle widened
node is homoplasious, supporting the suggestion at the apex. Pedicellate spikelets staminate or
of Veldkamp et al. (1986). The morphological reduced to glumes, the glumes leathery or indu-
distinction between Rottboellia and Mnesithea is rate. Sessile spikelet callus cuneate, truncate, gla-
weak, having to do with the predominant number brous or pubescent, with a central peg. Lower
of sessile spikelets at each inflorescence internode glume of sessile spikelet smooth or decorated
(two or three in Rottboellia, three in Mnesithea). with raised patterns or rugosities. Embryo large.
Thyrsia is recognized by Watson and Dallwitz Six spp., Old World.
(1992) and placed in Phacelurus by Clayton and
Renvoize (1986), but molecular data (McKain and 386. Ophiuros C. F. Gaertn.
Kellogg, unpubl. data) show that it is derived
within Rottboellia. Ophiuros C. F. Gaertn., Suppl. Carp. 1: 3, pl. 181, f. 3.
(1805).
OPHIUROS + THAUMASTOCHLOA + HEMARTHRIA
+ HETEROPHOLIS Annuals or perennials, caespitose, the stem base
Pedicellate spikelet generally absent or sometimes with corms. Ligule membranous. Axil-
reduced, the pedicel sometimes visible but always lary branch complexes present. Pedicellate spike-
fused to the inflorescence axis. Sessile spikelet let absent, the pedicel sometimes visible. Spikelet
sunken in internode, producing a rat-tail inflores- callus truncate, with a cylindrical peg. Lower
cence. glume obscurely pitted in many specimens.
These four genera form a strongly supported Lower flower of sessile spikelet staminate or
clade (GPWG II 2012), with Hemarthria and Het- sterile.
sometimes only one. Callus truncate, with or with-

out a central peg. Lower glume of sessile spikelet
sculptured. Lower flower of sessile spikelet sterile.
Eight spp., Australia and New Guinea.
XXII.4. SUBTRIBE ISCHAEMINAE J. Presl (1830)
Dimeriinae Hack. ex C.E. Hubb. (1934).
Lemma awned from a sinus, the awn generally

geniculate. Chromosome base number often
reduced from x ¼ 10 to x ¼ 9 or x ¼ 7.
Three genera, 173 spp.
The monophyly of Ischaeminae is supported
by molecular data (Estep et al. 2014; Teerawata-
nanon et al. 2011). The subtribe lacks obvious
morphological synapomorphies although the
lower glume often has a winged keel.
388. Andropterum Stapf
Andropterum Stapf, Fl. Trop. Afr. 9: 38 (1917).
Perennials. Ligule a fringe of hairs. Inflorescence

unbranched, the internodes and pedicels cuneate.
Pedicellate spikelets bisexual, the lower glume
with a broad asymmetrical wing. Sessile spikelet
squashed between the internode and pedicel.
Lower glume of sessile spikelet two-keeled with
strong central groove. Upper glume of sessile
Fig. 66. Panicoideae-Andropogoneae-Rottboelliinae. Het- spikelet laterally compressed.
eropholis sulcata. A Base of plant. B Leaf. C Inflorescences. One sp., A. stolzii (Pilg.) C. E. Hubb., tropical
D Portion of inflorescence. E–K Sessile spikelet. E Lower Africa.
glume. F Upper glume. G Upper lemma. H Its palea. I
Flower. J, K Caryopsis. L–O Pedicelled spikelet. L Lower
glume. M Upper glume. N Lemma. O Palea. (From Clayton DIMERIA + ISCHAEMUM
and Renvoize 1982, drawn by S. Ross-Craig) Leaf epidermis with intercostal cells with sev-
eral papillae per cell.
Four spp., tropical Africa to China and
Australia.
389. Dimeria R. Br.
387. Thaumastochloa C. E. Hubb. Dimeria R. Br., Prodr. 204 (1810).

Nanooravia Kiran Raj & Sivad., Nordic J. Bot. 31: 162
Thaumastochloa C. E. Hubb., Hooker’s Icon. Pl. 34: pl. (2013).
3313 (1936).
Caespitose annuals or perennials. Ligule membra-
Annuals. Ligule a fringed membrane. Axillary nous, or a fringed membrane. Inflorescence with
branch complexes present. Inflorescence becoming one or two branches, or up to 9 in a digitate
stiff and curved, breaking off in its entirety. Distal arrangement, the rachis not disarticulating. Spike-
end of peduncle widened. Spikelets 10 or fewer, lets solitary (not paired), on short pedicels, strongly
laterally compressed, the callus generally pubes-

cent. Glumes often winged on the keel. Lower
flower sterile. Upper flower with lemma awned;
palea present or absent. Stamens 2. 2n ¼ 14.
Fifty-nine spp., India, southeast Asia, Austra-
lia, Polynesia.
Molecular data show that Dimeria is derived
from within Ischaemum (Estep et al. 2014; Teer-
awatananon et al. 2011), apparently by suppres-
sion of the sessile spikelet. In D. woodrowii
Stapf the inflorescence is shed as a unit. The
newly described monospecific genus Nanooravia
is based on Dimeria santapaui, in which the
spikelets are not strongly compressed, and in
which the glumes bear auricles at the apex. Segre-
gation of the species in a new genus appears
premature, and it is retained here in Dimeria.
390. Ischaemum L. Fig. 67
Ischaemum L., Sp. Pl. 2: 1049 (1753).
Annuals or perennials, caespitose, rhizomatous

or stoloniferous. Leaves often cordate, with or
without a pseudopetiole, with or without auricles.
Ligule membranous, a fringed membrane or a
fringe of hairs. Inflorescence unbranched, or
with two branches, or with multiple digitate
branches, spikelets borne on one side of the
branch, the rachis and pedicel flattened and
together forming a U-shape, the distal end of the
Fig. 67. Panicoideae-Andropogoneae-Ischaeminae. Ischae-
internode often cupulate. Pedicellate spikelet mum afrum. A Habit. B Portion of inflorescence showing
bisexual, staminate or reduced. Sessile spikelet pedicelled spikelet. C Portion of inflorescence showing ses-
with the distal flower bisexual, the proximal sile spikelet. D Lower glume of sessile spikelet. (From Clay-
flower staminate. Lower glume dorsiventrally ton and Renvoize 1982, drawn by A. Davies)
compressed, often winged; upper glume laterally
compressed apparently because of compression
between rachis and pedicel, often awned. Lemmas charum group, often form a larger clade in molec-
membranous. 2n ¼ 18, 20, 40, 54, 56, 68. ular analyses (Kellogg 2012), although this may
Eighty-seven spp., tropics. be an artifact of taxon sampling. The subtribe is
retained here pending further data, and may need
to be extensively revised in the future.
XXII.5. SUBTRIBE SACCHARINAE Griseb. (1846)
391. Asthenochloa B€
use
Lemma awned from the apex or more often from
a sinus, the awn generally geniculate. Asthenochloa B€
use, Pl. Jungh.: 367 (1854).
Sixteen genera, 177 species.
This subtribe has no obvious morphological Decumbent annuals. Ligule membranous. Primary
synapomorphy. The genera formerly included in inflorescence branches themselves branched, each
or allied to Sorghum, together with the Sac- branch with a sessile spikelet and two pedicellate
ones, the latter represented only by pedicels. Callus Decumbent annuals. Ligule membranous or
of spikelet pubescent. Lower glume with a biden- a fringed membrane. Inflorescence branches
tate apex, the upper glume keeled. Stamens 2. branched. Callus of sessile spikelet pubescent.
One sp., A. tenera B€ use, Southeast Asia. Glumes awnless. 2n ¼ 20.
Two spp., Indo-Malesian.
Other than the glabrous lodicules this genus
392. Cleistachne Benth. appears similar to Sorghum.
Cleistachne Benth., Hooker’s Icon. Pl. 14: pl. 1379 (1882).
395. Sorghastrum Nash
Annuals, often with prop roots. Leaf blades pseu-
dopetiolate or not. Ligule membranous. Inflores- Sorghastrum Nash, Man. Fl. N. States: 71 (1901).
cence branches branched, spikelets unpaired.
Spikelet callus pubescent, obtuse. Glumes with Annuals or perennials, rhizomatous or caespi-
long hairs. Lodicules ciliate. 2n ¼ 36. tose. Leaves with or without auricles. Ligule
One sp., C. sorghoides Benth., tropical Africa, membranous, a fringed membrane or a fringe
India. of hairs. Inflorescence branches branched,
The pedicellate spikelet is generally inferred to spikelet-bearing only at their distal ends, other-
have been lost, resulting in unpaired spikelets. No wise slender. Sessile spikelet callus obtuse or
developmental studies have been done to confirm pungent, pubescent, the lower glume generally
this hypothesis. Liu et al. (2014) suggest that C. pubescent, the upper generally glabrous. Pedi-
sorghoides is of allopolyploid origin, with genomes celled spikelet usually reduced to a pedicel, but if
from a Sorghastrum-like ancestor and another present then resembling the sessile one.
genus more distantly related. Although they trans- 2n ¼ 20, 40, 60.
fer C. sorghoides to Sorghum, this placement is not Twenty-one spp., Africa and the Americas.
strongly supported. The species is retained here in
its own genus pending additional data. 396. Sorghum Moench
Sorghum Moench, Methodus: 207 (1794).

393. Leptatherum Nees Vacoparis Spangler, Austral. Syst. Bot. 16: 297 (2003).
Hemisorghum C. E. Hubb. ex Bor, Grass. Burma, Ceylon,
Leptatherum Nees, Proc. Linn. Soc. Lond. 1: 92 (1841). India & Pakistan 686 (1960).
Polliniopsis Hayata, Icon. Pl. Formosan. 7: 76, f. 45 (1918).
Mat-forming annuals, rooting from the nodes. Ligule membranous or a fringed membrane. Inflo-
Ligule membranous. Inflorescence slender, the rescence branches branched. Spikelets in pairs or
branches themselves branched, without hairs. in triads at the ends of branches, or ultimate
Rachis internodes longer than spikelets, slender, branches with 3 to 7 spikelet pairs. Pedicellate
flexuous. Glumes of sessile and pedicellate spike- spikelet staminate or sterile, often much smaller
lets indurate; lower glume of sessile spikelet than sessile one, the glumes often papery, with or
grooved. Callus of spikelet stubby, bearing a tuft without lemmas. Sessile spikelet ovate, with callus
of hairs; spikelets otherwise glabrous. Stamens 2. obtuse; glumes leathery, their apices obtuse, the
Three spp., Asia and Australia. lower with two keels; upper lemma awned or awn-
Chen et al. (2009) have clearly documented less, the apex obtuse. 2n ¼ 10, 20, 40.
the phylogenetic distinction between this genus Eight spp., Africa, India, southeast Asia,
and Microstegium (subtribe Andropogoninae), Northern Australia.
confirming the tentative hypothesis proposed by The molecular phylogenies of Dillon et al.
Mathews et al. (2002) and Spangler et al. (1999). (2004, 2007) place the two species of Vacoparis
(¼ Sorghum sects. Chaetosorghum and Heterosor-
394. Pseudosorghum A. Camus ghum) sister to the species of Sorghum sect. Sor-
ghum. See also discussion in Kellogg (2012). Data
Pseudosorghum A. Camus, Bull. Mus. Hist. Nat. (Paris) 26: from Liu et al. (2014) are also mostly consistent
662 (1920).
with this interpretation, although there is some This group may or may not form a clade
disagreement among gene trees. S. mekongense separate from the rest of the Saccharinae. Molec-
had been segregated as Hemisorghum, but molecular data remain insufficient and phylogenies are
ular data place it firmly in a clade with Sorghum unresolved (Hodkinson et al. 2002).
(McKain and Kellogg, unpubl. data).
399. Eriochrysis P. Beauv.
397. Sarga Ewart
Eriochrysis P. Beauv., Ess. Agrostogr. : 8. (1812); tax.:
Sarga Ewart, Proc. Roy. Soc. Victoria 23(2): 296–297 Filgueiras (1997).
(1911), rev.: Spangler (2003). Leptosaccharum (Hack.) A. Camus, Bull. Soc. Bot. France
70: 736 (1923).
Annuals or perennials. Nodes with a ring of stiff
hairs. Ligule a fringed membrane. Inflorescence Caespitose perennials. Ligule membranous, or a
branches branched or not. Pedicellate spikelet fringed membrane or a fringe of hairs. Inflores-
staminate, sterile, or reduced to a rudiment. Ses- cence covered with golden or reddish hairs, the
sile spikelet with the callus obtuse, acute or primary branches branched or unbranched. Fer-
sharply pointed, the lower glume apex often tile flower of the sessile spikelet lacking a palea.
thickened, keeled or winged. 2n ¼ 10, 20, 30, 40. Pedicellate spikelet pistillate. 2n ¼ 20.
Nine spp., Australia, southern Indonesia, Twelve spp., tropical America and Africa.
Mexico and Central America. E. filiformis (Hack.) Filgueiras was formerly
Species of Sarga are often placed in Sorghum. placed in its own genus, as Leptosaccharum fili-
However, Sarga appears to be phylogenetically formis (Hack.) Camus. Descriptions of the species
distinct from Sorghum, no more closely related in the literature are partially inaccurate; a cor-
to it than to other genera in Saccharinae. While rected description and illustration are provided
Liu et al. (2014) argue for retention of these spe- by Filgueiras (1997).
cies in Sorghum (as sect. Parasorghum), the
placement is not strongly supported. 400. Eulalia Kunth
Eulalia Kunth, Révis. Gramin. 1: 160 (1829).

398. Spathia Ewart Pseudopogonatherum A. Camus, Ann. Soc. Linn. Lyon II,
68: 204–205 (1921) [1922].
Spathia Ewart, Fl. N. Territory: 26 (1917).
Caespitose annuals. Ligule a fringed membrane. tous. Ligule membranous, or a fringed membrane
Inflorescence of digitate branches, enclosed by a or a fringe of hairs. Inflorescence covered with
spathe, the proximal spikelet pair on each branch silky hairs, unbranched or branched, if the latter
staminate. Sessile spikelet with a pubescent cal- then the branches digitate or on an elongate cen-
lus, the lower glume with a line of hairs, the upper tral axis. Spikelet with a blunt callus. Lower glume
glume pubescent, the flower bisexual with 3 with 2 keels, upper glume with 1 keel, the apex
stamens. Pedicellate spikelets staminate, the flow- muticous or awned. 2n ¼ 20, 40.
ers with 2 stamens. Thirty-seven spp., Old World tropics.
One sp., S. neurosa Ewart & Archer, Australia. Eulalia species vary with respect to many
characters used elsewhere to define genera, and
the genus is likely to be polyphyletic (Hodkinson
THE SACCHARUM GROUP et al. 2002).
Sessile and pedicellate spikelets similar, her-

maphrodite; callus of the spikelet with long 401. Imperata Cirillo
hairs, these generally longer than the glumes.
Imperata Cirillo, Pl. Rar. Neapol. 2: 26 (1792).
Glumes papery to membranous, not indurate.
Rhizomatous perennials. Leaves auriculate or

not, the blades pseudopetiolate or not. Ligule
membranous, or a fringed membrane, or a fringe
of hairs. Inflorescence branches branched, cov-
ered with long silvery hairs, “sessile” spikelets
short pedicellate or not. Top of pedicel a round
crater. Glumes with long silvery hairs. Proximal
flower of the spikelet reduced to a lemma, distal
flower with the lemma reduced, awnless, palea
present, lodicules absent, stamens 1 or 2.
2n ¼ 20, 40, 50, 60.
Thirteen spp., tropical and subtropical. I.
cylindrica (L.) Raeusch. is a particularly nasty
weed that is widespread in tropical areas.
402. Lasiorhachis (Hack.) Stapf
Lasiorhachis (Hack.) Stapf, Hooker’s Icon. Pl. 32: t. 3124

(1927).
Andropogon subg. Lasiorhachis Hack., Monogr. Phan. 6:
471 (1889).
Caespitose perennials. Primary panicle branches

themselves branched. Pedicellate spikelet her-
maphrodite, staminate, sterile, or absent. Sessile
spikelet with lower glume with two keels. Lemma
with a short bristle. Ovary with an apical linear,
hairy structure originating lower than the styles.
Three spp., Madagascar.
Clayton and Renvoize (1986) synonymize
Lasiorhachis with Saccharum, but there seems Fig. 68. Panicoideae-Andropogoneae-Saccharinae. Mis-
no really good reason to place it there versus canthus violaceus. A Leaf. B Inflorescence. C Portion of
with any other genus of the group. The variation inflorescence. D Spikelet. E Tip of upper lemma. (From
in sex expression of the pedicellate spikelet Clayton and Renvoize 1982, drawn by A. Davies)
encompasses that found throughout the tribe.
The species is distinguished from most other brane. Inflorescence with primary branches
species of Saccharum by its more highly branched borne along a central axis, not digitate, the
inflorescences, shorter hairs and lack of awns. branch axes disarticulating late in development
if at all, spikelets disarticulating below the glumes
leaving the pedicels and rachis intact. “Sessile”
403. Miscanthus Andersson Fig. 68 spikelet on a short pedicel. 2n ¼ 30, 35–43, 48,
57, 76, 95, 96, 114.
Miscanthus Andersson, Öfvers. F€orh. Kongl. Svenska About thirty spp., Africa to southeast Asia.
Vetensk.-Akad. 12: 165 (1855). Molecular data show clearly that Narenga,
Miscanthidium Stapf, Fl. Trop. Afr. 9: 89 (1917).
Sclerostachya (Andersson ex Hack.) A. Camus, Fl. Indo- Sclerostachya, and Miscanthidium are part of the
Chine 7: 243 (1922). same clade as Miscanthus s.s. and thus should be
Narenga Bor, Indian Forester 66: 267 (1940). placed into synonymy (Hodkinson et al. 2002).
However, at least five species of Miscanthus, includ-
Perennials, caespitose or rhizomatous, generally ing the widely cultivated species M. sinensis, form a
over 1 m tall. Leaf blades with or without a pseu- separate clade, which may or may not be related.
dopetiole. Ligule membranous or a fringed mem- Miscanthus x giganteus is cultivated as a biofuel
crop. Miscanthidium and Miscanthus are reported cules absent. Stamens 2. Intercostal epidermal
to hybridize with Saccharum (Watson and Dallwitz cells with papillae overarching the stomata.
1992 onward). The distinctive intercostal papillae Four genera, 31 spp.
also appear in Andropogoninae. Sun et al. (2010) Apocopis, Germainia and Pogonatherum are
have revised the Chinese species of Miscanthus and supported as a clade in the molecular phylogenies
provide a detailed discussion of variation in mor- of Teerawatananon et al. (2011) and Estep et al.
phological characters within the genus. (2014), with Germainia apparently derived from
within Apocopis. Lophopogon is placed here based
solely on morphology.
404. Saccharum L.
Saccharum L., Sp. Pl. 1: 54 (1753). 405. Pogonatherum P. Beauv. Fig. 69

Erianthus Michx., Fl. Bor.-Amer. 1: 54 (1803).
Pogonatherum P. Beauv., Ess. Agrostogr.: 56 (1812).
Large rhizomatous perennials, often several
meters tall. Leaves auriculate or not. Ligule mem- Highly branched perennials. Leaf blades with
branous, or a fringed membrane or a fringe of pseudopetioles. Ligule membranous or a fringed
hairs. Inflorescence with a thick strong central membrane. Inflorescence unbranched, subtended
axis, the branches themselves branched, with by a spathe-like leaf. Spikelets laterally com-
long hairs, disarticulating at the nodes. Lemmas pressed, tiny (<3 mm long). Sessile spikelet with
of some species lacking awns. 2n ¼ 40, 60, 68, a pubescent callus, the lower glume truncate or
76–78, 80, 90, 46–128, 110, 112, 116–117, 144. obtuse, awned or not, the upper glume awned.
Thirty-seven species. Pedicellate spikelet pistillate. 2n ¼ 20.
The generic limits of Saccharum, Erianthus, Three spp., tropical Asia.
and Miscanthus are not clear. Saccharum s.s. Stamen number is reduced to 1 in some
includes only six species, two of which (S. robus- plants.
tum and S. spontaneum) are wild and four of
which are cultivated and clonal (S. officinarum, S.
edule, S. barberi, and S. sinense); the latter two are 406. Apocopis Nees
of hybrid origin (D’Hont et al. 2002). Welker et al.
Apocopis Nees, Proc. Linn. Soc. Lond. 1: 93 (1841).
(2015) show that the type of Erianthus (E. gigan-
teus) falls in the Saccharum clade and thus that the
Annuals or perennials, caespitose. Ligule mem-
circumscription of Saccharum should be broad-
ened to include at least some species of Erianthus.
unbranched, or with unbranched branches, digi-
Hodkinson et al. (2002) present data showing that
tate. Pedicellate spikelet present and poorly
Saccharum s.s. is monophyletic, and that this clade
developed, or absent and represented only by
does not include either Narenga or Sclerostachya,
the pedicel; pedicel fused to the lower glume of
which instead fall into Miscanthus. Only two spe-
the sessile spikelet. Sessile spikelet with a pubes-
cies of Erianthus (E. rockii and E. contortus) were
cent callus; glumes with strong parallel veins,
sampled in that study, and these also fell in the
sometimes winged. 2n ¼ 20, 40.
Miscanthus clade. These two species should per-
Sixteen spp., tropical Asia.
haps be moved to Miscanthus, although more spe-
cies sampling is definitely needed. For a more
complete discussion, see Kellogg (2012). 407. Germainia Balansa & Poitr.
Germainia Balansa & Poitr., Bull. Soc. Hist. Nat. Toulouse

7: 344 (1873).
XXII.6. SUBTRIBE GERMAINIINAE Clayton
(1972) Annuals or perennials, caespitose or stolonifer-
ous. Ligule membranous or a fringed membrane.
Glumes with apices truncate. Upper lemma awned Inflorescence unbranched, or with unbranched
from a sinus, the awn generally geniculate. Lodi- branches, digitate. Sessile spikelet staminate or
408. Lophopogon Hack.
Lophopogon Hack., Die Nat. Pflanzenfam. II, 2: 26

(1887).
Annuals or perennials, caespitose. Ligule a

fringed membrane. Inflorescence with 2 branches
appressed to form an ovate head. Sessile and ped-
icellate spikelets similar, overlapping, the distal
ends of the internodes oblique. Proximal spikelet
pairs with both sessile and pedicellate spikelets
staminate. Glumes stiff, truncate, the apex with
three teeth; lower glume with distinct hair tufts,
upper glume awned.
Two spp., India.
XXII.7. SUBTRIBE ANDROPOGONINAE J. Presl

(1830)
Anthistirinae J. Presl (1830).
Upper lemma awned from the apex or from a

sinus, the awn generally geniculate.
Twenty-seven genera, 526 spp.
This subtribe has no obvious morphological
synapomorphy. Subtribes Andropogoninae and
Anthistirinae as delimited by Clayton and
Renvoize (1986) are not monophyletic. However,
several studies (e.g., Estep et al. 2014; Mathews
et al. 2002; Skendzic et al. 2007) have shown that
the two groups plus the Bothriochloa clade
together are monophyletic. The Andropogoninae
as delimited here correspond roughly to the “core
Fig. 69. Panicoideae-Andropogoneae-Germainiinae. Pogo- Andropogoneae” identified in early molecular
natherum paniceum. A Habit. B Spikelet pair. C Sessile phylogenetic studies (Kellogg 2000; Mathews
spikelet. D Lower glume. E Upper glume. F Upper lemma. et al. 2002; Spangler et al. 1999).
G Upper palea. H Stamen of upper flower. I Pistil. J Lower
lemma. K Lower palea. L Stamen of lower flower. M Pedi-
celled spikelet. (From Wu et al. 2007, with permission from
the Missouri Botanical Garden Press, St. Louis, and Science 409. Bhidea Stapf ex Bor
Press, Beijing)
Bhidea Stapf ex Bor, Kew Bull. 3: 445 (1948) [1949].
Annuals. Ligule membranous. Inflorescence

sterile. Pedicellate spikelet hermaphrodite, with unbranched or with 2 branches, the rachis flat-
or without a proximal sterile flower. tened. Callus pubescent, inserted in hollowed
Ten spp., Southeast Asia and Australia. internode tip. Pedicellate spikelet staminate, the
The bisexual pedicellate spikelet is unusual in lower glume winged, muticous or awned. Sessile
Andropogoneae, but also appears in Trachypogon. spikelet staminate in proximal spikelet pairs,
bisexual in distal ones, the lower glume 2-keeled Pedicellate spikelet hermaphrodite, callus sharp,
with large wings, the upper glume with three lobes, pubescent, obliquely attached, the lower glume
the central one extended into an awn. with two keels, the lemma pubescent. “Sessile”
Three spp., India. spikelet actually short pedicellate, staminate or
Clayton and Renvoize (1986) segregate this sterile, awnless. 2n ¼ 20, 40.
from Andropogon sect. Andropogon. No molecu- Four spp., Africa and tropical America.
lar data are available.
THE MONOCYMBIUM + ANADELPHIA + ELYMAN-
THE HOMOZEUGOS + AGENIUM + TRACHYPOGON DRA MORPHOGROUP
CLADE Upper glume with a long awn.
This group was found to be monophyletic by Molecular data place Elymandra in Andropo-
Kellogg and Watson (1993) and Guala (2000), but goninae (McKain and Kellogg, unpublished).
representatives have not been included in any
recent molecular phylogenetic study.
413. Anadelphia Hack.
410. Agenium Nees Anadelphia Hack., Bot. Jahrb. Syst. 6: 240 (1885).
Andropogon sect. Pobeguinea Stapf, J. Bot. (Morot) 19:
Agenium Nees, Nat. Syst. Bot.: 447 (1836). 100 (1905).
Monium Stapf, Fl. Trop. Afr. 9: 399 (1919).
Caespitose perennials. Leaves with blades disarti- Pobeguinea (Stapf) Jacq.-Fél., Rev. Int. Bot. Appl. Agric.
Trop. 30: 172 (1950).
culating, the ligule membranous or a fringed
membrane. Inflorescence unbranched, or with
Annuals or perennials. Axillary branch complexes
digitate unbranched branches. Pedicellate spike-
present. Inflorescence axes unbranched, with few
lets larger than the sessile ones, staminate, the
spikelets widely separated on the axis. Pedicellate
first glume pubescent, the lemma without an
spikelet the same size as the sessile one, much
awn. Pedicels deeply pigmented. Sessile spikelets
reduced, or absent. Sessile spikelet with a sharp
staminate in proximal spikelet pairs, bisexual in
callus; lower glume 2-pronged or truncate; awn of
distal pairs. Callus sharp, pubescent, obliquely
upper glume often nearly as long as the spikelet and
attached. Lower glume deeply grooved, the groove
spreading, upper lemma with two lobes. 2n ¼ 20.
translucent. Distal flower lacking a palea.
Fourteen spp., Africa.
Four spp., South America.
Pobeguinea and Monium are recognized as
distinct by Watson and Dallwitz (1992 onward)
411. Homozeugos Stapf but not by Clayton and Renvoize (1986). The ratio-
nale for the generic delimitation is provided by
Homozeugos Stapf, Hooker’s Icon. Pl. 31: t. 3033 (1915). Clayton (1966a). The invalid generic name Diecto-
mis Beauv. (1812) nom rejic. non Kunth (1815) is
Caespitose perennials. Leaves auriculate or not, based on D. fasciculata, a species now placed in
the ligule membranous or a fringed membrane. Anadelphia. Diectomis Kunth is based on D. fasti-
Inflorescence unbranched, or with unbranched giata and is placed near (or possibly in) Andropo-
branches, digitate. All spikelets bisexual. Callus gon. Thus the validly published Diectomis Kunth is
sharp, pubescent, obliquely attached. Glumes not a synonym of Anadelphia.
pubescent. Distal flower with or without a palea.
Six spp., Central Africa.
414. Elymandra Stapf
412. Trachypogon Nees Elymandra Stapf, Fl. Trop. Afr. 9: 407 (1919).
Pleiadelphia Stapf, Hooker’s Icon. Pl. 30: t. 3121 (1927).
Trachypogon Nees, Fl. Bras. Enum. Pl. 2(1): 341 (1829).
Annuals or perennials. Axillary branch com-
Caespitose perennials. Ligule membranous. Inflo- plexes present. Inflorescences unbranched or
rescence unbranched, or with unbranched with two branches with up to 18 spikelet pairs.
branches, digitate, the axes not disarticulating. Lower 1–10 sessile spikelets staminate, obliquely
attached to the axis. Lower glume of sessile spike- awned. Sessile spikelet callus pubescent, the
let pubescent to villous. Pedicellate spikelet with a glumes pubescent.
stipe-like callus. 2n ¼ 20. One sp., E. abyssinica (Hochst. ex A. Rich.)
Six spp., all African except one in Brazil. Andersson, eastern Africa and Viet Nam.
Clayton (1966b) synonymizes Pleiadelphia
with Elymandra, observing that the only difference
417. Hyperthelia Clayton
is the unbranched inflorescence of the former.
Hyperthelia Clayton, Kew Bull. 20: 438 (1967).
415. Monocymbium Stapf
Annuals or perennials, caespitose. Leaves auricu-
Monocymbium Stapf, Fl. Trop. Afr. 9: 386 (1919). late or not, with or without a pseudopetiole. Lon-
ger inflorescence branch bearing only three
Caespitose perennials. Ligule membranous or a spikelets near the tip, one sessile and hermaphro-
fringed membrane. Axillary branch complexes dite, and the other two pedicellate and staminate;
present. Inflorescence unbranched, wholly sur- shorter inflorescence branch bearing five spike-
rounded by a reddish-brown spathe. Pedicellate lets, the two proximal ones staminate, the distal
spikelet with glumes truncate, unawned. Sessile ones in a set of three as on the long branch.
spikelet with the callus pubescent, both flowers Long flattened, leaf-like appendage often present
lacking paleas. 2n ¼ 20. at the base of the branches. Lower glume of sessile
Three spp., tropical and southern Africa. spikelet strongly grooved, the upper glume
mucronate or awned. Both flowers of the sessile
THE EXOTHECA + HYPERTHELIA GROUP spikelet lacking a palea. 2n ¼ 20, 40.
Ligule membranous. Axillary branch com- Seven spp., African savannas, introduced to
plexes present. Inflorescence with two branches, tropical America.
one spikelet-bearing to near the base and relatively Remaining genera of Andropogoninae:
shorter, the longer one spikelet-bearing only in the Leaves with papillae present on the intercostal epi-
distal portion, barren in the proximal half. Callus dermal cells, each papilla consisting of one oblique
of hermaphrodite spikelet sharp-pointed. swelling per cell overarching the stomata.
Skendzic et al. (2007) place Hyperthelia in a The single oblique papilla is not common else-
clade with Hyparrhenia hirta and Andropogon where in the tribe, but does occur in Apocopis,
distachyos, suggesting that it might be merged Apluda, Loxodera, and Parahyparrhenia. The
with one of the latter two genera. However, the oblique papilla is apparently lacking in Andropogon
number of species sampled is not large enough to and Schizachyrium; Themeda has several papillae
provide confidence about its placement. Chloro- per cell.
plast DNA sequences also place Exotheca near
Hyparrhenia, but data are not available for
Hyperthelia (McKain and Kellogg, unpubl. data). 418. Cymbopogon Spreng.
416. Exotheca Andersson Cymbopogon Spreng., Pl. Min. Cogn. Pug. 2: 14 (1815).
Exotheca Andersson, Nova Acta Regiae Soc. Sci. Upsal. 2: Perennials, or rarely annual, caespitose. Leaves
253 (1856). aromatic, auriculate or not. Ligule membranous
or a fringed membrane. Axillary branch com-
Caespitose perennials. Leaves auriculate. Axillary plexes present. Inflorescence a pair of strongly
branch complexes present. The four proximal reflexed branches, one slightly longer than the
spikelets on each branch staminate and sessile or other, the lower one with a pair of staminate
nearly so, forming an involucre. Distal to the spikelets at its base. Pedicellate spikelet generally
“involucre” is a set of three spikelets, one her- staminate, rarely sterile. Sessile spikelet with a
maphrodite and sessile, the other two staminate pubescent callus, inserted in hollowed internode
and pedicellate. Pedicellate spikelet with glumes tip, staminate in the proximal spikelet pair (in
most species), bisexual in distal spikelet pairs. 421. Iseilema Andersson

Lower glume, flat or concave, 2-keeled, the keels
sometimes winged, both flowers lacking paleas. Iseilema Andersson, Nova Acta Regiae Soc. Sci. Upsal. 2:
2n ¼ 20, 22, 40, 60. 250 (1856).
Fifty-nine spp., Old World tropics and sub-
tropics. Annuals or perennials, caespitose. Axillary
The reflexed branches are diagnostic, but do branch complexes present. Ligule membranous,
not occur in ser. Proceri (Clayton and Renvoize or a fringed membrane, or a fringe of hairs.
1986) in which the branches are erect. The genus Leaves and spathes gland bearing, aromatic.
is strongly supported as monophyletic in molec- Inflorescence subtended by a boat-shaped spathe,
ular phylogenies, and is not obviously closely unbranched, with four staminate spikelets (inter-
related to any other genus (Estep et al. 2014). preted as two pairs) at the base, with 1 or 2 sessile
bisexual spikelets and a pair of pedicellate stami-
nate spikelets distally. Pedicels of proximal four
419. Eremopogon Stapf spikelets curved stiffly upward to form basket,
often with a tuft of hairs at the base of the basket.
Eremopogon Stapf, Fl. Trop. Afr. 9: 182 (1917). Pedicellate spikelets with glumes flat, strongly
veined. Sessile spikelets with distal flower lacking
Perennials, rarely annual. Axillary branch a palea. 2n ¼ 6, 8, 18, 28, 36.
complexes present. Inflorescences unbranched. Twenty-four spp., Indomalayan region, Aus-
Pedicellate spikelets staminate or sterile, awnless. tralia.
Sessile spikelets with a pubescent callus, the lower The odd inflorescence morphology of this
glume with a large circular pit in the center, both genus suggests a connection with Themeda.
flowers lacking paleas. 2n ¼ 40. Molecular data show that the two are closely
Four spp., Old World tropics. related (Skendzic et al. 2007), but Estep et al.
The large circular pit on the lower glume also (2014) suggest that they do not form a clade.
occurs in some species of Bothriochloa and
Dichanthium, and led Clayton and Renvoize
(1986) to include this genus in Dichanthium. 422. Pseudodichanthium Bor
However, molecular data place Eremopogon sep-
arate from either of these genera and instead Pseudodichanthium Bor, Indian Forester 66: 271 (1940).
sister to Iseilema (Estep et al. 2014). Eremopogon
lacks the hyaline groove in the pedicels that is Annuals. Ligule a fringed membrane. Inflores-
characteristic of Bothriochloa and Capillipedium. cence unbranched. Pedicellate spikelets stami-
nate or sterile, morphologically similar to the
sessile ones and larger, lower glume winged, the
420. Heteropogon Pers. pedicel cuneate, the distal end cupuliform. Proxi-
mal 2 to 3 sessile spikelets staminate and awnless,
Heteropogon Pers., Syn. Pl. 2: 533 (1807). distal ones bisexual, awned. Callus rounded.
Lower glume winged, the apex emarginate, both
Annuals or perennials, caespitose, monoecious or flowers lacking a palea.
andromonoecious. Ligule membranous or a One sp., P. serrafalcoides (Cooke & Stapf)
fringed membrane. Axillary branch complexes Bor, India.
present. Inflorescence unbranched. Pedicellate
spikelets staminate or sterile, with an elongated
callus that looks superficially like a pedicel, the 423. Themeda Forssk. Fig. 70
lower glume winged. Sessile spikelets terete, with
a long, sharp pubescent callus, staminate in the Themeda Forssk., Fl. Aegypt.-Arab. 178 (1775).
proximal portion of the inflorescence, pistillate or
sometimes hermaphrodite in the distal portion. Annuals or perennials, mostly caespitose. Axil-
2n ¼ 20, 22, 40, 44, 50, 60, 80. lary branch complexes present. Ligule membra-
Six spp., tropical. nous, or a fringed membrane or absent.
the three genera are interfertile at the tetraploid

level (Harlan and de Wet 1963). Molecular phy-
logenies show that Old World polyploid species
of Bothriochloa contain genomes donated by
Dichanthium and Capillipedium consistent
with the cytogenetic data (Estep et al. 2014). In
this one small group one sees many of the inflo-
rescence forms of the entire Andropogoneae,
supporting the idea that transitions among
them are easy.
424. Capillipedium Stapf
Capillipedium Stapf, Fl. Trop. Afr. 9: 169 (1917).
Annuals or perennials, caespitose. Leaves gener-

ally without auricles. Ligule membranous, a
fringed membrane, or a fringe of hairs. Inflores-
cences delicate and open, the slender branches
themselves branched. Inflorescence internodes
and pedicels of pedicellate spikelets with two
strong marginal veins and a hyaline groove,
often described as a “translucent line”, between.
Each branch with a long naked proximal portion
and fewer than eight pairs of spikelets. Pedicellate
spikelets staminate or sterile. Sessile spikelets
with a pubescent callus, both flowers generally
lacking paleas. 2n ¼ 20, 40, 60.
Fig. 70. Panicoideae-Andropogoneae-Andropogoninae.
Eighteen spp., Africa, Asia and Australia.
Themeda triandra. A Habit. B Inflorescence with
spatheole. C Spikelet triad, with one awned fertile spikelet
and 2 pedicelled sterile spikelets. D Two pairs of stami- 425. Bothriochloa Kuntze
nate spikelets. E Upper lemma and awn base. (From Wu
et al. 2007, with permission from the Missouri Botanical Bothriochloa Kuntze, Revis. Gen. Pl. 2: 762 (1891).
Garden Press, St. Louis, and Science Press, Beijing)
Perennials, caespitose or rhizomatous. Ligule
Inflorescence subtended by a spathe, unbranched, membranous or a fringed membrane. Inflores-
with four staminate spikelets (interpreted as two cence with digitate primary branches, or the
pairs) at the base, with 1 or 2 sessile bisexual branches borne on an elongated axis, each
spikelets and a pair of pedicellate staminate spi- branch with 8 or more pairs of spikelets. Inflo-
kelets distally. Callus pubescent, its attachment rescence internodes and pedicels of pedicellate
oblique. Sessile spikelet with glumes truncate, spikelets with two strong marginal veins and a
indurate; distal lemma apex mucronate in some hyaline groove, often described as a “translucent
species. 2n ¼ 20, 40, 60, 80. line”, between. Pedicellate spikelets staminate or
Twenty-nine spp., Old World tropics and sterile. Sessile spikelets with a pubescent callus,
subtropics. the lower glume often with a deep circular pit in
the center, with two keels. 2n ¼ 20, 30, 40, 50,
THE CAPILLIPEDIUM + BOTHRIOCHLOA + 60, 120.
DICHANTHIUM GROUP Thirty-seven spp., tropical.
While diploid members of Bothriochloa, The number of spikelets per inflorescence
Capillipedium and Dichanthium are intersterile, branch is not distinctive within the
Andropogoneae, but does distinguish Bothrio- 428. Parahyparrhenia A. Camus

chloa from Euclasta and Capillipedium.
Parahyparrhenia A. Camus, Bull. Mus. Natl. Hist. Nat. II,
22: 404 (1950).
426. Dichanthium Willemet
Annuals or perennials. Leaf blades narrow, fili-
Dichanthium Willemet, Ann. Bot. (Usteri) 18: 11 (1796). form, auriculate or not. Ligule membranous or a
fringe of hairs. Inflorescence unbranched or with
Annuals or perennials, caespitose. Leaves generally two branches, spikelet-bearing to near the base,
without auricles. Ligule membranous or a fringed or the proximal portion without spikelets, the
membrane. Inflorescence unbranched, or with dig- rachis flattened. Pedicellate spikelets staminate
itate branches. Proximal pair(s) of spikelets gener- or sterile, with glumes muticous or awned. Sessile
ally staminate. Sessile spikelet with the callus spikelet callus pubescent, acute to sharp-pointed,
obtuse, pubescent. Glumes ovate to elliptical, mul- obliquely attached to the rachis and extending
tiveined, overlapping; both flowers with paleas beyond it; the lower glume deeply grooved, the
minute or absent. 2n ¼ 20, 40, 50, 60. upper glume mucronate or awned.
Twenty-two spp., Old World tropics. Six spp., Africa, India, Thailand.
This genus is morphologically similar to
THE DIHETEROPOGON + PARAHYPARRHENIA + Diheteropogon.
P S E U D A N T H I S T I R I A GROUP
Flower of sessile spikelet single, subtended by
two hyaline bracts, interpreted as lemmas. 429. Pseudanthistiria (Hack.) Hook. f.
Diheteropogon is strongly supported as a
Pseudanthistiria (Hack.) Hook. f., Fl. Brit. India 7: 219
member of the large clade including Andropogon, (1897) [1896].
Schizachyrium, and Hyparrhenia (Estep et al.
2014), but no molecular data are available on Trailing annuals, rooting from the nodes. Leaf
Pseudanthistiria. The two lemmas presumably blades lanceolate. Ligule membranous or a
represent the lemmas of the proximal and distal fringed membrane. Axillary branch complexes
flowers, but could also be interpreted as the present. Inflorescence unbranched, with one
lemma and palea of the distal flower with the proximal spikelet pair and a distal triplet of one
proximal flower being entirely suppressed. sessile and two pedicellate spikelets. Pedicellate
Molecular data obtained as this book went to spikelets staminate, with muticous glumes. Ses-
press suggest that Parahyparrhenia may be mis- sile spikelet with the callus oblique, pubescent.
placed here and instead be sister to Eriochrysis 2n ¼ 20.
(McKain and Kellogg, unpublished). Four spp., India to Thailand.
427. Diheteropogon (Hack.) Stapf THE DIECTOMIS + HYPARRHENIA + ANDROPOGON

+ SCHIZACHYRIUM GROUP
Diheteropogon (Hack.) Stapf, Hooker’s Icon. Pl. 31: t. 3093 This clade has no obvious morphological syn-
(1922). apomorphy, but the latter three genera have been
included in several molecular phylogenies, and
Annuals or perennials, caespitose. Leaf blade consistently form a well-supported group along
broad or reduced to the midrib. Ligule membra- with Diheteropogon (Estep et al. 2014).
nous or a fringed membrane. Axillary branch
complexes present. Inflorescences with two
branches. Pedicellate spikelet staminate or sterile. 430. Diectomis Kunth
Sessile spikelet with the callus pubescent, acute to
Diectomis Kunth, Mém. Mus. Hist. Nat. 2: 69 (1815), nom.
sharp-pointed, inserted into a deep crater-like conserv., non Beauv. (1812).
apex of the internode below; lower glume
grooved. 2n ¼ 20, 40. Caespitose annuals. Ligule membranous. Axillary
Four spp., Africa. branch complexes present. Inflorescences
unbranched, the pedicels and rachis with long Remaining two genera:
hairs, with a translucent line down the center. Leaf epidermal papillae absent, an apparent loss.
Pedicellate spikelet much larger than the sessile
one; lower glume flat, papery, reddish, awned.
432. Andropogon L.
Sessile spikelet with a hairy callus, the lower
glume deeply grooved. Andropogon L., Sp. Pl. 2: 1045 (1753).
One sp., D. fastigiata (Sw.) P. Beauv., pan-
tropical. Perennials, a few species annual, caespitose or
Clayton and Renvoize (1986) and Clayton less often rhizomatous. Leaves auriculate or not.
et al. (2006 onward) include D. fastigiata in Ligule membranous, or a fringed membrane or a
Andropogon. Consistent with this hypothesis fringe of hairs. Axillary branch complexes pres-
Skendzic et al. (2007) place it in a clade with ent. Inflorescence unbranched or digitate. Pedi-
Schizachyrium semitectum and Andropogon ger- cellate spikelets staminate or sterile, or reduced
ardii, suggesting that it might be merged with only to the glumes, or lacking, leaving only the
one of the latter two genera. It is kept separate pedicel. Sessile spikelets with a pubescent callus
here because species sampling of molecular phy- inserted into the hollowed internode tip; lower
logenies is insufficient to place it reliably. Its glume muticous, with the margins raised or
unbranched inflorescence and pedicellate spike- keeled, hollowed or grooved between the keels;
lets with broad glumes are distinctive. upper glume mucronate or awned. 2n ¼ 20, 40,
60, 100, 120, 180.
431. Hyparrhenia Andersson ex E. Fourn. One hundred twenty-two spp., worldwide.
This genus may be paraphyletic (Skendzic
Hyparrhenia Andersson ex E. Fourn., Mexic. Pl. 2: 51, 67 et al. 2007) or monophyletic (Estep et al. 2014),
(1886). but will require much more detailed sampling to
Dybowskia Stapf, Fl. Trop. Afr. 9: 382 (1919). assess relationships. Clayton and Renvoize (1986)
divide the genus into 4 sections, Andropogon,
Annuals or perennials, caespitose. Leaves auricu- Leptopogon (ca. 55 species), Piestium, and Noto-
late or not. Ligule membranous or a fringed solen. Individual species and species complexes
membrane. Axillary branch complexes present. have been studied in detail (e.g., Hodge and Kel-
Inflorescence a pair of erect, spreading or logg 2014; Nagahama et al. 2014) but the genus as
deflexed branches, one slightly longer than the a whole would benefit from more comprehensive
other, the base of each inflorescence branch assessment of species limits.
often with one or two pairs of staminate spikelets.
Pedicellate spikelets staminate or sterile. Sessile
spikelets with a hairy callus, attached obliquely to 433. Schizachyrium Nees
the rachis; lower glume convex or flat. 2n ¼ 20,
30, 40, 44, 45, 60. Schizachyrium Nees, Fl. Bras. Enum. Pl.: 331 (1829).
Fifty-eight spp., Africa.
The two-branched inflorescences of this Perennials, a few species annual, caespitose or
genus are similar to those of Cymbopogon, but less often rhizomatous. Ligule membranous, or
the two genera can be distinguished by the con- a fringed membrane. Axillary branch complexes
nection of the sessile spikelet to the rachis. In present. Inflorescence unbranched, the inter-
Cymbopogon the spikelet is inserted into a deep nodes slender or clavate, with a fimbriate rim
crater-like node, whereas in Hyparrhenia the cal- distally. Pedicellate spikelets staminate or sterile,
lus of the spikelet is obliquely attached. Although or reduced only to the glumes, or lacking, leaving
they have similar inflorescences, Cymbopogon only the pedicel, the glumes muticous or awned.
and Hyparrhenia appear not to be particularly Sessile spikelets with a pubescent callus inserted
closely related. The single species of Dybowskia into the hollowed internode tip, the glumes muti-
is included within Hyparrhenia, following Clay- cous or awned; lower glume flat, convex or
ton (1969). grooved, both flowers lacking paleas. 2n ¼ 20,
30, 40, 50.
Sixty-four spp., tropics and temperate gins of C4 have occurred; the number could be
regions, worldwide. between 2 and 5 origins (GPWG II 2012).
The molecular phylogenetic study of
Skendzic et al. (2007) indicates that Schizachyr-
434. Apochloa Zuloaga & Morrone
ium is polyphyletic, but those authors included
only six of the sixty species. More recent data Apochloa Zuloaga & Morrone, Syst. Bot. 33: 288 (2008).
(Estep et al. 2014) indicate that it may be mono-
phyletic and sister to Andropogon. Tussock forming perennials. Leaves ending in a
sharp point, often without a clear sheath-blade
XXIII. TRIBE PASPALEAE J. Presl (1830) junction. Ligule a fringed membrane or lacking.
Inflorescence branches branched, spreading.
Trachideae Pilg. ex Potztal (1957). Upper lemma and palea shining, smooth. C3.
Arthropogoneae Pilg. ex Butzin (1972). 2n ¼ 20, 40.
Fifteen spp., Guyana Highlands and eastern
This tribe has no obvious morphological synapo- Brazil.
morphy. This genus was segregated from Panicum by
31 genera, 664 species. Sede et al. (2008). All included species were for-
Paspaleae as circumscribed here correspond to merly in Panicum sect. Lorea, which was found to
the group known as the x ¼ 10 Paniceae in several be polyphyletic.
molecular phylogenies (Aliscioni et al. 2003; Gius-
sani et al. 2001; Morrone et al. 2012; Vicentini et al.
2008). Paspaleae excluding Reynaudia are well 435. Arthropogon Nees
supported as being monophyletic (Morrone et al.
2012), and are sister to Andropogoneae (GPWG II Arthropogon Nees, Fl. Bras. Enum. Pl. 2(1): 319 (1829).
2012), a relationship supported by the synapomor- Achlaena Griseb., Cat. Pl. Cub.: 228 (1866).
Altoparadisium Filg., Davidse, Zuloaga & Morrone, Ann.
phy of the chromosome base number. However, no Missouri Bot. Gard. 88: 363 (2001).
obvious morphological character distinguishes
Paspaleae from Paniceae s.s. Caespitose perennials. Ligule membranous or a
Within Paspaleae are three well-supported fringed membrane or a fringe of hairs. Inflores-
clades, recognized here as subtribes following cence branches branched, spreading or upright.
Morrone et al. (2012). Most of the taxa occur in Spikelets laterally compressed. Callus with long
the New World, except for a few species of Hyme- hairs. Glumes awned, the awns about 10 times
nachne, and Baptorachis from Africa. Many spe- longer than the glume itself, the lower glume
cies in Paspaleae were formerly classified in a shorter than the spikelet or absent. Lower
broadly construed genus Panicum, but are clearly lemma longer than the upper one. Upper lemma
unrelated to Panicum s.s. and palea laterally compressed, the upper lemma
hyaline, not enclosing the apex of the palea. C4,
with a single bundle sheath. Mesophyll with or
XXIII.1. SUBTRIBE ARTHROPOGONINAE Butzin (1972) without distinctive isolated bundle sheath cells.
Six spp., Brazil and West Indies.
This subtribe has no obvious morphological Arthropogon s.s., Achlaena, and Altoparadi-
characters. sium form a strongly supported clade (Morrone
14 genera, 71 species. et al. 2012; GPWG II 2012); the long awns on the
This group was first identified as a clade by glumes are distinctive, so the three genera are
Giussani et al. (2001), who called it the “ambigu- united here. In addition, members of the clade
ous clade” because of its morphological heteroge- have a callus with long hairs and the upper
neity. The clade contains a mix of C3 and C4 taxa, lemma and palea laterally compressed. The for-
and the topology of the phylogeny makes it mer Arthropogon lanceolatus was segregated as
impossible to determine exactly how many ori- Canastra based on its phylogenetic position
(Giussani et al. 2001); Canastra and Arthropogon
are confirmed to be unrelated by GPWG II Stoloniferous perennials. Leaf blades broad, flat,
(2012). the bases cordate; ligules membranous. Primary
branches of inflorescence unbranched, with the
spikelets borne on one side, crowded. Glumes
436. Canastra Morrone, Zuloaga, Davidse & Filg. more or less equal. Lower lemma longer than
upper one. Upper flower obovoid, the rachilla
Canastra Morrone, Zuloaga, Davidse & Filg., Novon 11: internode below it elongated, smooth. Margins
429–436 (2001).
of upper lemma involute. C3.
One sp., P. gymnocarpon (Elliott) Nash,
Caespitose perennials. Ligule a fringed mem-
southeastern U.S.
brane. Leaf blades narrowed at the base, involute.
Inflorescence branches branched, dense. Callus
glabrous. Glumes awned, the awns about 10 440. Stephostachys Zuloaga & Morrone
times longer than the glume itself. Lower lemma
longer than the upper one. Upper lemma cartilag- Stephostachys Zuloaga & Morrone, Taxon 59: 1539 (2010).
inous, the apex not enclosing the apex of the Panicum sect. Megista Pilg., Notizbl. Bot. Gart. Berlin-
Dahlem 11: 243 (1931).
palea. C3. Mesophyll radiate, with fusoid cells.
Two spp., Brazil.
Rhizomatous perennials, 1–3 m tall. Ligule mem-
branous, laciniate. Base of leaf blades cordate to
rounded. Inflorescence branches unbranched,
437. Homolepis Chase
arranged in pseudoverticels of 10 to 60 branches
Homolepis Chase, Proc. Biol. Soc. Wash. 24: 146 (1911). each. Upper glume and lower lemma similar in
length. Lower flower staminate. Apices of upper
Annuals or perennials, caespitose. Ligule mem- lemma and palea with stomata and papillae. C3.
branous. Leaf blades generally cordate. Inflores- 2n ¼ 40.
cence branches branched. Glumes enclosing the One sp., S. mertensii (Roth) Zuloaga & Mor-
flowers. Upper lemma thin, not indurate, the rone, Central and South America.
margins lying flat on the palea. Hilum linear. C3. Additional description of this genus is
Midrib complex; mesophyll with fusoid cells. provided by Zuloaga et al. (2010).
2n ¼ 22, 24, 40.
Five spp., Mexico to Brazil. CYPHONANTHUS + ONCORACHIS
Inflorescence branches branched. C4, with a
single bundle sheath.
438. Oplismenopsis Parodi These two genera form a strongly supported
clade in the analysis of Morrone et al. (2012), and
Oplismenopsis Parodi, Notas Mus. La Plata, Bot. 2: could perhaps be merged into a single genus.
2 (1937).
441. Cyphonanthus Zuloaga & Morrone
Perennial with floating culms. Ligule a fringed
membrane. Inflorescence with the primary Cyphonanthus Zuloaga & Morrone, Taxon 56: 526 (2007).
branches unbranched. Spikelets with the lower
glume toward the rachis axis (adaxial). Lower Rambling caespitose perennials. Ligule a fringe of
glume with an awn; upper glume extended into hairs. Lower glume absent or minute, upper
an arista. Lower lemma with a short awn. Hilum glume gibbous, as long as the spikelet. Rachilla
linear. C3. not thickened. Upper lemma and palea open at
One sp., O. najada (Hack. & Arechav.) Parodi, the apex, pubescent at the apex and base, covered
Argentina. In water. with macrohairs and simple papillae. C4, with a
single bundle sheath.
439. Phanopyrum (Raf.) Nash One sp., C. discrepans (D€oll) Zuloaga &
Morrone, Central America and northern South
Phanopyrum (Raf.) Nash, Fl. S.E. U.S.:104, 1327 (1903). America to Brazil.
This species was segregated from Panicum on lower lemma with 5 to 9. Upper glume and lower
the basis of morphological, anatomical and lemma enclosing the upper lemma and palea.
molecular data. It corresponds to Panicum discre- Upper lemma and palea indurate, smooth,
pans and was placed in its own section (Panicum shiny, often with two prickle hairs at the apex.
sect. Discrepentia) by Zuloaga (1987), highlight- C4, with one bundle sheath, or two sheaths in C.
ing its differences from other species of Panicum. prionitis and C. petersonii. 2n ¼ 18, 36, 20, 40.
Seven spp., Mexico, Central America, South
America, Caribbean.
442. Oncorachis Morrone & Zuloaga
Coleataenia is supported as monophyletic
Oncorachis Morrone & Zuloaga, Taxon 58: 372 (2009). by molecular data only; no obvious morpho-
logical synapomorphies link the species,
Rhizomatous perennials. Leaf blades rigid. Ligule although the obliquely attached spikelets pro-
a fringe of hairs or lacking. Lower glume present, vide a distinctive, if subtle, character. This
1/6 to 2/3 the length of the spikelet, upper glume clade was first identified by Aliscioni et al.
not gibbous. Rachilla thickened above the glumes (2003), although reevaluation of the data has
and below the lower flower. Upper lemma and subsequently excluded Stephostachys from the
palea indurate, covered with macrohairs and sim- group. Information on nomenclature, distribu-
ple papillae. C4, with a single bundle sheath. tion and morphology of the species is pre-
Two spp., central and eastern Brazil. sented by Zuloaga et al. (2010). “Panicum”
This genus was erected to accommodate Strep- beyeri is not mentioned, however, and it is
tostachys ramosa Zuloaga & Soderstr. and S. unclear where it belongs. Various names are
macrantha (Trin.) Morrone & Zuloaga, which are connected to Panicum [unranked] Agrostoidea;
more closely related to Cyphonanthus than they are their history and, where necessary, typification
to other species of Streptostachys (Sede et al. 2009a). are presented by McNeill et al. (2010).
COLEATAENIA + TRISCENIA
444. Triscenia Griseb.
Triscenia is in an unresolved relationship to
Coleataenia in the analyses of Morrone et al. Triscenia Griseb., Pl. Wright. 2: 534 (1862).
(2012), and is strongly supported as being
embedded within Coleataenia in GPWG II Perennials. Leaf blades narrow, without clear
(2012). Because the two genera differ in photo- lamina and midrib. Ligule a fringed membrane.
synthetic pathway, their precise relationship Inflorescence branches unbranched. Glumes
affects inference about the evolutionary origins shorter than the spikelet. Upper lemma thin, not
of C4, and would repay further investigation. hardened, the margins flat on the palea. C3.
One sp., T. ovina Griseb., Cuba.
443. Coleataenia Griseb., Abh. K€onigl. Ges. Wiss.
ottingen 24: 308 (1979).
G€ KERATOCHLAENA + MESOSETUM + TATIANYX
Hilum linear. C4, NADP-ME.
Panicum [unranked] Agrostoidea Nash in Small, Fl. S. E. This clade is strongly supported in the molec-
U. S.: 85–86 (1903). ular analyses of Morrone et al. (2012).
Panicum [unranked] Tenera Hitchc. & Chase, Contr. U. S.
Natl. Herb. 15: 29, 99 (1910).
Panicum sect. Prionitia Zuloaga, Grass Syst. Evol.: 294 445. Keratochlaena Morrone
(1987).
Sorengia Zuloaga & Morrone, Taxon 59: 1541 (2010). Keratochlaena Morrone, Darwiniana 47: 231 (2009).
Sclerochlamys Morrone & Zuloaga, Taxon 58: 373 (2009),
Perennials, caespitose or rhizomatous. Ligule nom. illegit.
membranous, ciliate or not. Inflorescence
branches branched. Spikelets solitary, pedicellate, Caespitose perennials. Leaf blades rigid, pungent.
obliquely attached to the pedicel. Lower glume Inflorescence branches unbranched. Rachilla
with 1 to 3 prominent veins, upper glume and internodes thickened. Lower glume 1/3–2/3 the
length of the spikelet. Upper glume and lower

lemma coriaceous. Upper lemma and palea cov-
ered with compound papillae.
One sp., K. rigidifolia (Filg., Morrone &
Zuloaga) Morrone & Zuloaga, northern Brazil.
Sclerochlamys was erected to accommodate
Streptostachys rigidifolia Filg., Morrone &
Zuloaga, which is more closely related to Mesose-
tum than it is to other species of Streptostachys
(Sede et al. 2009a). The name proved to be a later
homonym for Sclerochlamys F. Muell., so was
replaced with the name Keratochlaena.
446. Mesosetum Steud.
Mesosetum Steud., Syn. Pl. Glumac. 1: 118 (1855)

[1854].

or stoloniferous. Ligule a fringe of hairs. Inflores-
cence unbranched. Pedicel apex oblique; spikelets
laterally compressed, with the lower glume
toward the rachis axis (adaxial); upper lemma
and palea smooth and gaping. 2n ¼ 16.
Twenty-seven spp., Central America and
northern South America.
Mesosetum is strongly supported as being sis-
ter to Tatianyx (GPWG II 2012); both genera are
C4 but reportedly differ in the number of bundle
sheaths (Watson and Dallwitz 1992 onward).
447. Tatianyx Zuloaga & Soderstr. Fig. 71 Fig. 71. Panicoideae-Paspaleae-Arthropogoninae. Tatia-
nyx arnacites. A Habit. B Pair of spikelets, showing
oblique attachment of pedicels. C Spikelet, upper glume
Tatianyx Zuloaga & Soderstr., Smithsonian Contr. Bot. side, showing oblique callus. D Lower palea, outer surface.
59: 56 (1985). E Lower lemma. F Upper glume. G Lower glume. H
Stamens. I Upper lemma. J Upper palea showing gynoe-
Caespitose perennials. Sheaths scarious, pubes- cium and lodicules. K Lodicules. L Upper anthecium,
cent. Ligule a fringe of hairs. Inflorescence lateral view. M Same, palea side. N Caryopsis, embryo
side. O Caryopsis, showing linear hilum. (Reprinted
branches branched, spreading. Spikelet attach- from Zuloaga and Soderstom 1985; courtesy of Smithso-
ment oblique to pedicel, the callus pubescent. nian Institution)
Glumes and lower lemma villous, enclosing the
upper lemma and palea. Upper lemma indurate.
One sp., T. arnacites (Trin.) Zuloaga & XXIII.2. SUBTRIBE OTACHYRIINAE Butzin
Soderstr., Brazil. (1970)
Tatianyx is strongly supported as being sister
to Mesosetum (GPWG II 2012), but Kerato- Inflorescence branches generally with spikelets
chlaena was not included in that analysis. The arranged along one side. Lower glume shorter
microhairs are described as unusual, but require than the upper one. Deletion of 9 bp in the chlo-
further examination with good quality material roplast gene ndhF.
(Watson and Dallwitz 1992 onward).
Four genera, thirty-eight species.

Otachyriinae are strongly supported as mono-
phyletic (Morrone et al. 2012), but the genera
included do not share any obvious morphological
synapomorphies; the shared characters appear
elsewhere in the Panicoideae. Otachyriinae
include all species formerly placed in “Panicum”
sect. Laxa, which is not monophyletic. The type
species of the former Laxa group is unrelated to
the others and is now Steinchisma laxa, here
placed in synonymy in Otachyrium. Other species
of the Laxa group have been transferred to Hyme-
nachne or placed in the new genus Rugoloa
(Acosta et al. 2014). Detailed descriptions of the
species in the former “Panicum” sect. Laxa,
including photos of leaf anatomy and epidermal
structure, are provided by Zuloaga et al. (1992).
448. Anthaenantia P. Beauv.
Anthaenantia P. Beauv., Ess. Agrostogr.: 48, 151, t. 10

(1812).
Leptocoryphium Nees, Fl. Bras. Enum. Pl. 2(1): 83 (1829).
Perennials, caespitose or rhizomatous. Ligule a

branched, spreading. Spikelets dorsiventrally to
laterally compressed, densely pubescent. Lower
glume lacking. Upper glume as long as the spike-
let. Upper glume and lower lemma with papillose-
based trichomes along the veins. Upper lemma
cartilaginous, not enclosing the palea, the apex
acute. C4, with one bundle sheath. 2n ¼ 40.
Five spp., southeastern U.S., Central America
and South America.
Anthaenantia is strongly supported as sister
to all other members of Otachyriinae (Acosta
et al. 2014; Morrone et al. 2012).
449. Hymenachne P. Beauv. Fig. 72
Hymenachne P. Beauv., Ess. Agrostogr.: 48 (1812).

Dallwatsonia B. K. Simon, Austrobaileya 3: 678 (1992).
Perennials. Culms filled with aerenchyma. Ligule

membranous. Inflorescence branches branched,
short and appressed to the main axis. Upper Fig. 72. Panicoideae-Paspaleae-Otachyrinae. Hymenachne
amplexicaulis. A Culms with rhizome and inflorescence. B
glume and lower lemma as long as the spikelet, Spikelet from the adaxial side (upper image) and from the
the latter acute, acuminate or with a short awn. abaxial side (lower image), showing small lower glume,
Upper lemma membranous, clasping the palea acute upper glume, and the attenuate lower lemma. (From
proximally but not at tip. C3. 2n ¼ 24. Barkworth et al. 2003, drawn by H. Paźdirková)
Twelve spp., wet sites in the tropics. ing, with the spikelets paired, borne on one side
Hymenachne falls into two clades, one Old and crowded, or on short higher-order branches.
World and one New World, and may not be Lower glume shorter than 1/2 the length of the
monophyletic (Acosta et al. 2014). spikelet. C3. Mesophyll of leaf often with fusoid
cells. 2n ¼ 20, 40.
Three spp., Central and South America,
450. Otachyrium Nees Caribbean.
Rugoloa corresponds to a set of three species
Otachyrium Nees, Fl. Bras. Enum. Pl. 2(1): 271–272 formerly placed in “Panicum” sect. Laxa, and was
(1829).
Steinchisma Raf., Bull. Bot., Geneva 1: 220 (1830). identified in a phylogenetic study by Acosta et al.
Plagiantha Renvoize, Kew Bull. 37: 323 (1982). (2014).
Cliffordiochloa B. K. Simon, Austrobaileya 3: 674 (1992).
Fasciculochloa B.K. Simon & C.M. Weiller, Austrobaileya
4: 374, f. 1–4 (1995).
XXIII.3. SUBTRIBE PASPALINAE Griseb. (1846)
Annuals or perennials, caespitose. Ligule mem-
Reimarochloinae Caro (1982).
branches branched or not. Spikelets often paired,
with both glumes shorter than spikelet and more Inflorescences unbranched, or with unbranched
or less equal in length. Lower flower staminate. branches, spikelets borne unilaterally on the
Lower palea enlarged at maturity, hardened, branches.
winged and enclosing upper flower. Lemmas Eleven genera, 554 spp.
entire, pointed, awnless, cartilaginous. C3 or C3/ Paspalinae is moderately well supported by
C4 intermediates. 2n ¼ 18, 20. molecular data, but has no obvious morphologi-
Sixteen spp., SE US to South America, Aus- cal synapomorphies (Morrone et al. 2012). The
tralia. majority of the species belong to the large genus
Otachyrium s.s. and Steinchisma form a clade Paspalum.
in molecular analyses and together are sister to
Plagiantha; all three share a distinctive hardened ACOSTIA + AXONOPUS
palea. Because of this morphological synapomor- Lower glume absent or minute. Upper glume
phy, the three are combined here, although not all as long as the spikelet.
combinations are yet available. In addition, Ota-
chyrium s.s. may be paraphyletic (Acosta et al. 452. Acostia Swallen
2014), also supporting combining the group. Sen-
dulsky and Soderstrom (1984) cite a single chro- Acostia Swallen, Bol. Soc. Argent. Bot. 12: 109 (1968).
mosome count for Otachyrium as n ¼ 9, but
molecular data (Morrone et al. 2012) place it in Perennials. Ligule a fringed membrane. Inflores-
the x ¼ 10 clade, strongly supported as sister to cence branches few, sparse, unbranched,
Steinchisma. Steinchisma is one of the few grass appressed to the main inflorescence axis, the spi-
genera that includes both C3 species and C3/C4 kelets borne on one side. Upper glume pubescent,
intermediates. extended to a slender acuminate tip. Lower lemma
similar to the upper glume but with a shorter tip,
the lower palea absent. Upper lemma smooth, the
451. Rugoloa Zuloaga margins inrolled on the palea. C4.
One sp., A. gracilis Swallen, Ecuador.
Rugoloa Zuloaga, Plant Syst. Evol. 300: 2164 (2014). Acostia is sister to Axonopus s.l., based on
morphological data (Morrone et al. 2012). The
Rhizomatous perennials. Ligule membranous. photosynthetic pathway has been verified by sta-
Leaves with blades rounded to cordate at the ble isotope analysis and anatomy (C. Osborne
base. Inflorescence branches branched, ascend- and O. Morrone, unpubl. data).
453. Axonopus P. Beauv.
Axonopus P. Beauv., Ess. Agrostogr.: 12 (1812); phylog.,

tax.: Lopez and Morrone (2012).
Centrochloa Swallen, J. Wash. Acad. Sci. 25: 192 (1935).
Baptorhachis Clayton & Renvoize, Kew Bull., Addit. Ser.
13: 298, 377 (1986).
Ophiochloa Filg., Davidse & Zuloaga, Novon 3: 360, f. 1–3
(1993).

or stoloniferous. Leaves auriculate or not, the
blade disarticulating or not, muticous or sharp-
pointed at the apex. Ligule membranous, or a
cence unbranched, or with two to many,
unbranched branches, subdigitate or borne on
an elongate axis. Spikelets borne in two rows on
one side of the rachis branch. Upper glume
awned or not. Lower lemma about as long as the
upper glume, adaxial; upper lemma with margins
inrolled on the palea. Stamens 2 or 3. C4, with one
bundle sheath. 2n ¼ 20, 40, 60, 80.
One hundred-four spp., tropics of the West-
ern Hemsiphere, one species in Africa.
Axonopus is a paraphyletic group from
which Ophiochloa and Centrochloa are derived
(Morrone et al. 2012). Baptorhachis is placed sis-
ter to Ophiochloa on the basis of morphological
data; in both genera the inflorescence axis is
broad and leaf like, often folding to enclose the
spikelets. Zanotti et al. (2010) have described
spikelet development in A. suffultus and find
that it differs from that in Paspalum.
454. Anthaenantiopsis Mez ex Pilg. Fig. 73

Fig. 73. Panicoideae-Paspaleae-Paspalinae. Anthaenan-
tiopsis rojasiana. A Plant. B Ligule region. C Spikelet,
Anthaenantiopsis Mez ex Pilg., Notizbl. Bot. Gart. Berlin- lateral view. D Upper glume. E Lower lemma. F Lower
Dahlem 11(104): 237–238 (1931). palea with lodicules. G Upper lemma and palea, lateral
view. H Caryopsis, hilar view. I Same, scutellar view.
Caespitose perennials with short rhizomes. Leaf (From Zuloaga et al. 2012, drawn by A. Bestard)
blades linear to filiform. Ligule membranous.
Inflorescence branches unbranched or branched, Four spp., South America.
narrow. Spikelets solitary or in twos or threes, Anthaenantiopsis is sister to Aakia and the
covered with long hairs. Lower glume minute. two sister to Osvaldoa in molecular analyses
Upper glume and lower lemma acuminate. Lower (Aliscioni et al. 2003; Lizarazu et al. 2014; Mor-
flower with a palea. Upper lemma with margins rone et al. 2012; Zuloaga et al. 2007) but has no
involute, apex acute. Upper lemma and palea with obvious morphological synapomorphy. Hopia is
prickle hairs and bicellular microhairs near the strongly supported as sister to the clade (GPWG
apex. C4, with a single bundle sheath. 2n ¼ 20, 40. II 2012).
455. Aakia J. R. Grande mary branches unbranched, spreading, unilat-

eral. Spikelets solitary or paired, laterally
Aakia J. R. Grande, Phytoneuron 22: 1 (2014). compressed. Lower glume with tubercle-based
hairs. Callus of upper flower with wing or scars
Caespitose perennials. Ligule membranous. on base of lemma. C3. 2n ¼ 60.
Leaf blades broadly lanceolate. Inflorescence Nine spp., Central and South America, Mada-
branches branched, spreading. Spikelets with gascar.
sparse hairs on the upper glume and lower Sede et al. (2009b) provide evidence that Echi-
lemma. Lower glume reduced, without veins. nolaena is polyphyletic. However, their phyloge-
Lower flower lacking a palea, sterile. Upper nies cannot rule out the possibility that the two
lemma and palea covered with simple papillae, clades of Echinolaena plus Ocellochloa plus “Pan-
with long macrohairs near the apex. C4, with a icum” venezuelae might form a monophyletic
single bundle sheath. group, which could then be recognized as an
One species, A. tuerckheimii (Hack.) J. R. expanded Echinolaena.
Grande, Mexico, Guatemala, Belize, and Nicaragua.
Aakia corresponds to the former “Panicum”
sect. Tuerckheimiana (Hitchc.) Zuloaga. Lizarazu 458. Gerritea Zuloaga, Morrone & T. J. Killeen
et al. (2014) provide morphological and molecu-
lar evidence that firmly place this speces in Pas- Gerritea Zuloaga, Morrone & Killeen, Novon 3: 213, f. 1–2
(1993).
paleae, unrelated to Panicum s.s.
Caespitose perennials. Leaf blades with a pseudo-
456. Osvaldoa J.R. Grande petiole, disarticulating from the sheaths. Ligule a
Osvaldoa J.R. Grande, Phytoneuron 22: 5 (2014). branched, the spikelets pedicellate. Glumes with
long papillose-based hairs, the lower glume only
Caespitose perennials, with short rhizomes. slightly shorter than the upper one. Palea of
Ligule membranous. Leaves with blades linear- upper flower gaping, not enclosed by the lemma
lanceolate, the base rounded. Inflorescence at the tip. C3, mesophyll with radiate chloren-
branches branched, upright. Spikelets in pairs chyma.
or solitary. Lower flower with a palea. Upper One sp., G. pseudopetiolata Zuloaga, Morrone
lemma and palea with papillae all over, the apex & T. J. Killeen, Bolivia.
with microhairs. C4, with a single bundle sheath.
2n ¼ 20.
One sp., O. valida (Mez) J.R. Grande, Brazil, 459. Hopia Zuloaga & Morrrone
Uruguay, Argentina. Hopia Zuloaga & Morrone, Taxon 56: 150–153 (2007).
Osvaldoa corresponds to the former “Pani-
cum” sect. Valida Zuloaga & Morrone. Lizarazu Perennials, with stolons and short rhizomes. Lig-
et al. (2014) provide morphological and molecu- ule membranous. Inflorescence branches un-
lar evidence that firmly place this speces in branched, erect, the spikelets borne on all sides.
Paspaleae, unrelated to Panicum s.s. The chromo- Lower glume shorter than the spikelet; upper
some base number of x ¼ 10 is also consistent glume and lower lemma approximately the same
with this placement. length. Lower flower with a palea. Upper palea
winged near the apex. Upper lemma and palea
457. Echinolaena Desv. with simple papillae at the apex and base, with
multicellular microhairs. C4, with a single bundle
Echinolaena Desv., J. Bot. Agric. 1: 75 (1813). sheath. 2n ¼ 20, 40.
One sp., H. obtusa (Kunth) Zuloaga & Mor-
Annuals or perennials, caespitose or rhizoma- rone, southwestern US and Mexico.
tous. Leaf blades cordate, with or without a pseu- This genus corresponds to the former “Pani-
dopetiole. Ligule a fringed membrane or a fringe cum” sect. Obtusum, with the single species “Pan-
of hairs. Inflorescence unbranched or with pri- icum” obtusum. Molecular phylogenies place it
sister to the Anthaenantiopsis clade (GPWG II Thrasya Kunth, Nov. Gen. Sp. (quarto ed.) 1: 120–121
2012; Morrone et al. 2012). (1815) [1816].
Reimarochloa Hitchc., Contr. U. S. Natl. Herb. 12: 198
(1909).
460. Ichnanthus P. Beauv. Spheneria Kuhlm., Comm. Lin. Telegr., Bot. 67(11): 57, t. 4
(1922).
Ichnanthus P. Beauv., Ess. Agrostogr.: 56 (1812). Thrasyopsis Parodi, Bol. Soc. Argent. Bot. 1: 293 (1946).
Annuals or perennials, caespitose or rhizoma- Annuals or perennials, rhizomatous, stolonif-

tous. Ligule membranous, or a fringed membrane erous or caespitose. Leaves auriculate or not,
or a fringe of hairs. Inflorescence branches the blades with or without a pseudopetiole.
branched or not. Spikelets laterally compressed. Ligule membranous, or a fringed membrane
Glume keeled, acute to acuminate. Callus of or a fringe of hairs. Inflorescence unbranched,
upper flower expanded into two wings or scars or of unbranched primary branches, these dig-
on base of lemma. 2n ¼ 18, 20, 40, 54. itate or spread on an elongated axis, the
Thirty-six spp., New World tropics, or pan- branch axis flattened and sometimes winged.
tropical. Spikelets single or in pairs, flattened on one
The disparate chromosome base numbers side and convex on the other, generally round
could indicate polyphyly. The genus has not to ovate in outline (lanceolate or top-shaped in
been sampled extensively in molecular phyloge- species segregated as Reimarochloa or Sphe-
nies. neria). Lower glume lacking or minute and
abaxial; upper glume and lower lemma gener-
ally the same length as the spikelet; upper
461. Ocellochloa Zuloaga & Morrone glume rarely lacking. Upper lemma and palea
generally leathery to indurate, harder than the
Ocellochloa Zuloaga & Morrone, Syst. Bot. 34: 688 (2009).
glume(s) and lower lemma. C4, with one bun-
dle sheath.
Stoloniferous perennials, rooting at the lower Three hundred seventy-two spp., tropical
nodes. Leaves with short pseudopetioles. Ligule regions throughout the world.
membranous or a fringed membrane. Inflores- Many species are serious weeds, but some are
cence branches unbranched, ascending; spikelets also good forage. Apomixis is common. The
paired. Upper glume and lower lemma consider- inclusion of several segregate genera is justified
ably longer than the upper flower and enclosing by molecular data. All the segregates are like
it, the rachilla internode below the upper flower Paspalum in having unbranched primary inflo-
elongate. Upper lemma and palea smooth and rescence branches and in lacking the lower
shiny. C3. 2n ¼ 20. glume; see also Denham and Zuloaga (2007).
Twelve spp., Mexico and Central America to
Argentina.
This genus corresponds to the former “Pani- 463. Renvoizea Zuloaga & Morrone
cum” sect. Stolonifera; the justification for the
segregation is provided by Sede et al. (2009b). Renvoizea Zuloaga & Morrone, Syst. Bot. 33: 294 (2008).
Ocellochloa is sister to one species of Echino-
laena; see that genus for discussion. Many species Perennials, caespitose and rhizomatous. Leaves
of Ocellochloa have deep craters on the lower with the sheaths fibrous, persistent, the blades
lemma but not the upper one. ending in a sharp point, without a clear sheath-
blade junction. Ligule a fringed membrane or
lacking. Inflorescence branches branched, dense
462. Paspalum L. and contracted. Upper lemma and palea shining,
smooth. C3.
Paspalum L., Syst. Nat. (ed. 10) 2: 846, 855, 1359 (1759). Ten spp., Brazil.
This genus was segregated from Panicum by purposes, there is no obvious morphological
Sede et al. (2008). All the included species were character that unites the members of the
formerly in “Panicum” sect. Lorea, which was group. Relationships within the group are not
found to be polyphyletic. Renvoizea is in an unre- fully resolved, although the major clades
solved position with respect to other Paspaleae described below are monophyletic. Chloroplast
(Morrone et al. 2012). data place Digitaria as sister to all other mem-
bers of this clade, whereas data from the nuclear
gene phytochrome B place Echinochloa in that
464. Streptostachys Desv.
position (Vicentini et al. 2008). It is not clear
Streptostachys Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: whether one set of data or the other better
190 (1810). reflects the organismal history, or whether a
complex history of polyploidy and/or gene
Caespitose perennials. Base of leaf blade cordate flow could account for the discrepancy. Photo-
or clasping the stem. Ligule a fringe of hairs. synthetic pathway is variable within this tribe
Primary inflorescence branches stiff, the spikelets and all C4 subtypes are found.
borne in pairs. Spikelet with thickened rachilla.
Lower glume 2/3–4/5 the length of the spikelet.
Upper lemma and palea covered with bicellular INCERTAE SEDIS
microhairs and papillae on all epidermal cells.
Hilum linear. C3. Mesophyll with radiate chloren- 465. Acritochaete Pilg.
chyma, the cells with invaginated cell walls, with
or without fusoid cells. Acritochaete Pilg., Bot. Jahrb. Syst. 32: 53 (1902).
Two spp., Trinidad to NE Paraguay.
Streptostachys was recently re-circumscribed Decumbent annual. Ligule membranous. Primary
by Sede et al. (2009a) to remove the species now branches of the inflorescence distantly spaced,
known as Keratochlaena rigidifolia. Streptosta- unbranched, bearing spikelets on only one side.
chys is sister to Axonopus s.l. (GPWG II 2012; Upper glume and lower lemma with awns more
Morrone et al. 2012). than 1 cm long; the awn of the lower lemma
coiled. Upper lemma papery, acuminate or with
XXIV. TRIBE PANICEAE R. Br. in Flinders a minute awn. C3.
(1814) One sp., A. volkensii Pilg., tropical Africa.
Molecular data place this species sister to
Boivinellinae (GPWG II 2012).
Cenchreae Rchb. (1828).
Spinificeae Dumort. (1829).
Digitarieae J .J. Schmitz & Regel (1841). 466. Dichanthelium (Hitchc. & Chase) Gould Fig. 74
Melinideae Hitchc. (1920).
Boivinelleae A. Camus (1925). Dichanthelium (Hitchc. & Chase) Gould, Brittonia 26: 59
Anthephoreae Pilg. ex Potztal (1957). (1974).
Cyphochlaeneae Bosser (1965). Panicum subg. Dichanthelium Hitchc. & Chase, Contr. U.
Neurachneae S.T. Blake (1972). S. Natl. Herb. 15: 20, 142 (1910).
Panicum sect. Clavelligerae Stapf, Fl. Trop. Afr. 9: 639, 641
Base chromosome number x ¼ 9. The tribe has no (1920).
other obvious morphological synapomorphy.
Seventy-two genera, 1254 species. Perennials. Basal leaves broad, cordate, generally
Paniceae sensu stricto are clearly monophy- forming a winter rosette. Ligule membranous or a
letic, based on many molecular and morpholog- fringed membrane. Inflorescence branches them-
ical sets of data (Aliscioni et al. 2003; Christin selves branched. Spikelets generally obovate.
et al. 2008; Giussani et al. 2001; Gómez-Martinez Glumes of very different sizes, often pubescent.
and Culham 2000; Morrone et al. 2012; Vicentini Margins of upper lemma tucked in on palea. C3.
et al. 2008), and share a base chromosome num- 2n ¼ 18, 36.
ber of x ¼ 9. Unfortunately for identification One hundred twenty spp., America, Africa.
cioni et al. 2003; Giussani et al. 2001; Morrone

et al. 2012; GPWG II 2012). The species currently
in “Panicum” sect. Clavelligerae have not been
transferred formally to Dichanthelium, but their
placement there is well supported.
467. Echinochloa P. Beauv.
Echinochloa P. Beauv., Ess. Agrostogr. 53, 161 (1812),

nom conserv.
Annuals or perennials. Ligule often absent. Spike-

lets paired or on short second-order branches,
hemispherical, flat on one side and rounded on
the other. Lower glume shorter than lower
lemma. Upper lemma with an incurved beak-
like apex. Palea tip reflexed. C4, with a single
bundle sheath. 2n ¼ 36, 54, 72.
Forty spp., world wide in tropical and warm
temperate regions.
Chloroplast data put Echinochloa in Biovinel-
linae (Giussani et al. 2001; Morrone et al. 2012),
but sequences of the nuclear gene phytochrome B
place it at the base of the x ¼ 9 Paniceae (Vice-
ntini et al. 2008), and the nuclear gene PEPC
places it sister to the Melinidinae (Christin et al.
2007). Because of these conflicting placements, I
leave Echinochloa incertae sedis. Many species of
Echinochloa are problematical weeds.
468. Holcolemma Stapf & C. E. Hubb.
Holcolemma Stapf & C. E. Hubb., Bull. Misc. Inform. Kew

1929: 244 (1929).
Annuals or perennials. Ligule a fringed membrane.

Inflorescence branches appressed; terminal spike-
let of each branch subtended by a bristle. Lower
lemma sulcate, saccate on either side of the groove.
Lower palea with winged keels, expanded and
Fig. 74. Panicoideae-Paniceae. Dichanthelium oligo- indurate at maturity, clasping the upper flower.
santhes subsp. scribnerianum. A Plant. B Spikelet. (From Four spp., East Africa, India, Sri Lanka, Aus-
Barkworth et al. 2003, drawn by H. Paźdirková) tralia.
Watson and Dallwitz (1992 onward) report
this genus as being C3, with radiate Isachne-type
Cleistogamous flowers are common particu- mesophyll. No molecular data are available for
larly in late season flowering in North American Holcolemma. Morrone et al. (2012) use morpho-
species. Monophyly of Dichanthelium has been logical data to place it in Cenchrinae, sister to the
confirmed by all molecular studies to date (Alis- Central American genus Ixophorus, with which it
shares a winged lower palea. If this placement is Glumes unequal, inflated at the base, the upper
accurate, then Holcolemma would be the only C3 glume with stiff tubercle-based hairs above the
group in that subtribe, or indeed in the larger clade middle. Lower lemma inflated, grooved and folded
that also includes Melinidinae and Panicinae. along the abaxial side. Upper flower on a short
stalk, the lemma and palea hardened, smooth.
One sp., T. annua S.W.L. Jacobs, Western
469. Homopholis C. E. Hubb.
Australia, near Kalumburu.
Homopholis C. E. Hubb., Bull. Misc. Inform. Kew 1934:
126 (1934). 472. Thyridachne C. E. Hubb.
Walwhalleya Wills & J. J. Bruhl, Austral. Syst. Bot. 19: 327
(2006).
Thyridachne C. E. Hubb., Kew Bull. 4: 363 (1949).
Perennials, caespitose, rhizomatous or stolonifer-

Annuals. Leaves ending in a sharp point. Ligule
ous. Leaves auriculate or not. Ligule membranous
membranous. Inflorescence branches branched,
or a fringed membrane. Inflorescence branches
erect. Spikelet sickle-shaped in outline; upper
branched, spreading. Spikelets pedicellate, the
glume gibbous. Lower lemma trilobed, channeled
glumes muticous to mucronate. Glumes longer
on the abaxial side. Upper lemma membranous to
than the flowers, with 7 veins, pubescent. Upper
cartilaginous. Lodicules absent. C3.
lemma and palea leathery, often finely muricate
One sp., T. tisserantii C. E. Hubb., Zaire and
all over; upper lemma with a small beak. C3.
Central African Republic.
Four spp., Australia.
No molecular data are available for this spe-
Walwhalleya was segregated from Homopho-
cies; a combined morphological and molecular
lis based on a cladistic analysis of morphological
analysis places it in various positions in Paniceae
characters. However, molecular phylogenies
s.s. (Morrone et al. 2012).
strongly support the sister relationship between
the two and there seems little reason to separate
them (GPWG II 2012; Morrone et al. 2012).
S A C C I O L E P I S , T R I C H A N T H E C I U M , AND THE
“P A N I C U M ” SECT. M O N T I C O L A E GROUP
470. Hylebates Chippin.
Sacciolepis, Trichanthecium, and the “Panicum”
Hylebates Chippin., J. S. African Bot. 11: 127 (1945). sect. Monticolae group form a well-supported
clade in molecular phylogenies (Aliscioni
Prostrate annuals. Base of leaf blades wrapping et al. 2003; GPWG II 2012; Morrone et al.
around the stem. Ligule a fringe of hairs. Inflores- 2012; Zuloaga et al. 2011). The latter groups
cence with branched branches. Spikelets minute, were formerly placed in Panicum, and are
<2 mm long. Glumes thin, membranous, the lower morphologically homogeneous. However,
one tiny and without veins. Lemmas and paleas Sacciolepis falls in the middle of the group and
thin and membranous, the lower lemma extended is morphologically quite distinct, so is kept
to a narrow awn-like apex, the upper one with flat separate here.
margins covering the palea. C3. 2n ¼ 18.
Two spp., eastern and southern tropical Africa.
The photosynthetic pathway of H. cordatus 473. “Panicum” sect. Monticolae + sect. Verru-
and H. chlorochloë has been confirmed by stable cosa + sect. Ovalifoliae
isotope ratio (C. Osborne, unpubl. data).
“Panicum” [group] Verrucosa Hitchc., North American
Flora 3(2): 200, 205. (1915).
“Panicum” sect. Monticolae Stapf, Fl. Trop. Afr. 9: 641,
471. Thedachloa S.W.L. Jacobs 649 (1920).
“Panicum” sect. Ovalifoliae Stapf, Fl. Trop. Afr. 9: 641,
Thedachloa S.W.L. Jacobs, Telopea 10: 635–637. 650 (1920).
Stoloniferous annuals. Ligule a fringe of hairs. Ligule membranous. Inflorescence branches

Inflorescence dense, ovoid, the branches branched. branched, spreading, diffuse, branches naked for
1/2 to 2/3 their length. Lower glume minute and diffuse, branches naked for 1/2 to 2/3 their
without veins, or as long as the spikelet, with length. Spikelets sometimes with an elongated
three veins. Upper lemma and palea covered internode between the lower and upper glumes.
with simple papillae and microhairs, sometimes Glumes more or less equal in length, the lower
transversely rugose. C3. one ½ the length of the spikelet to the same
Nine spp., New World. length. Upper lemma and palea covered with
This “group” represents the paraphyletic simple papillae and microhairs. C3. 2n ¼ 18, 36,
residue left when Trichanthecium and Sacciolepis 54.
are recognized (Zuloaga et al. 2011); relationships Thirty-eight spp., Africa, South America,
among most species are poorly supported. Central America, Caribbean.
In “Panicum” sect. Monticolae, including Trichanthecium is sister to Sacciolepis (Alis-
“P.” trichoides Sw., “P.” sellowii Nees and cioni et al. 2003; Zuloaga et al. 2011). The species
“P.” millegrana Poir., and the Verrucosa group, were formerly assigned to Panicum, but Tri-
including “P.” verrucosum Muhl. and chanthecium has a membranous ligule, gener-
“P.” brachyanthum Steud., the upper lemma ally globose spikelets, glumes and lemmas with
and palea are transversely rugose, a character fewer veins, microhairs on the upper lemma,
that occurs commonly in Paniceae. “Panicum” and the C3 photosynthetic pathway, all charac-
sect. Monticolae is not monophyletic, and may ters unlike Panicum s.s. T. schwackeanum is
include “P.” bartlettii Swallen, formerly placed reported to have 2n ¼ 20 chromosomes, a
in sect. Parvifolia. The position of “P.” tri- count that needs to be confirmed (Zuloaga
chanthum Nees (former sect. Parvifolia) is also et al. 2011).
uncertain.
474. Sacciolepis Nash

XXIV.1. SUBTRIBE ANTHEPHORINAE
Sacciolepis Nash, Man. Fl. N. States 89 (1901). Benth. (1881)
Digitariinae Butzin (1972).

Annuals or perennials. Ligule membranous,
with a ciliate margin or not. Inflorescence
open and spreading, or narrow and spike-like, Lower glume tiny or absent. Characteristic dele-
with short dense lateral branches. Spikelets tion of 18 bp from the chloroplast gene ndhF (Mor-
laterally compressed, but the upper lemma rone et al. 2012). C4 with a single bundle sheath.
and palea dorsiventrally compressed. Glumes Seven genera and 296 species.
pubescent, ribbed, the upper one gibbous. This tribe appears sister to all other members
Upper lemma indurate to cartilaginous. C3. of the x ¼ 9 clade in molecular phylogenies of
2n ¼ 16, 18, 36, 45. chloroplast genes (Giussani et al. 2001) but sister
Twenty-seven spp., pantropical but with most to the clade of Melinidinae, Cenchrinae, and Pani-
species in Africa. cinae in phylogenies constructed from phyb (Vice-
ntini et al. 2008), and within the latter clade in trees
constructed from PEPC (Christin et al. 2007). Mor-
475. Trichanthecium Zuloaga & Morrone rone et al. (2012) place the subtribe in an unre-
solved position in Paniceae. Although its position
Trichanthecium Zuloaga & Morrone, Syst. Bot. Monogr. is uncertain, the group is clearly monophyletic.
94: 13 (2011).
Panicum [group] Parvifolia Hitchc. & Chase, Contr. U.S.
Within this subtribe, Anthephora, Chaetopoa,
Natl. Herb. 17: 462, 506 (1915). Chlorocalymma, Tarigidia, and Trachys all have
Panicum sect. Verruculosa Stapf, Fl. Trop. Afr. 9: 639 bristles or bracts surrounding bisexual flowers,
(1920). similar to an involucre. Although Clayton and
Renvoize (1986) say that Anthephora “imitates
Annuals or perennials. Ligule membranous. Cenchrus”, molecular data clearly place it here,
Inflorescence branches branched, spreading, unrelated to the bristle clade.
476. Anthephora Schreb.
Anthephora Schreb., Beschr. Gr€as. 2: 105 (1810).
Annuals or perennials, rhizomatous or caespi-

tose. Ligule membranous. Inflorescence narrow,
cylindrical, spike-like. Primary branches modified
to form stiff clusters of spikelets and involucral
bracts. Bracts broad, ovate, subtending 3 to 11
spikelets. Lower glume absent, upper glume awl-
like. 2n ¼ 18, 36, 40.
Eleven spp., African Arabia, Americas.
477. Chaetopoa C. E. Hubb.
Chaetopoa C. E. Hubb., Hooker’s Icon. Pl. 37: t. 3646

(1967).
Annuals. Ligule membranous. Inflorescence

branches unbranched, short, with a group of ster-
ile pedicellate spikelets surrounding a sessile,
bisexual spikelet. Lower glume awl-like or absent,
the upper one short; upper lemma awned.
Two spp., Tanzania.
Molecular and morphological data indicate
that Chaetopoa is either derived from within
Anthephora or sister to Trachys (Morrone et al.
2012); neither position is well supported.
478. Chlorocalymma Clayton
Chlorocalymma Clayton, Kew Bull. 24: 461 (1970).
Annuals. Ligule membranous. Inflorescence with

2 or 3 primary branches, the axis of each broad Fig. 75. Panicoideae-Paniceae-Anthephorinae. Digitaria
and leaf-like, enclosing the spikelets. Second-order catamarcensis. A Plant. B Spikelet, seen from the side of
branches forming spiny involucres around spike- the lower glume. C Fertile lemma and palea, viewed from
lets. Lower glume tiny, upper glume short, lower the abaxial side. D Fertile anthecium, viewed from the
adaxial side, showing part of the palea and the lodicules
lemma winged. 2n ¼ 18. of the sterile flower. E Caryopsis, scutellar view. (From
One sp., C. cryptacanthum Clayton, Tanzania. Zuloaga et al. 2012, drawn by S. G. Gómez)
479. Digitaria Haller Fig. 75 unequal lengths. Lower glume tiny or absent.
Lemma margins flat and exposed on palea.
Digitaria Haller, Hist. Stirp. Helv. 2: 244 (1768). 2n ¼ 18, 30, 36, 45, 54, 60, 70, 72, 76, 108.
Two hundred seventy-seven spp., warm
Annuals or perennials, caespitose, rhizomatous or regions of the world.
stoloniferous. Ligule membranous or a fringed DNA sequence data suggest that Digitaria
membrane or a fringe of hairs. Inflorescence might be para- or polyphyletic with other Anthe-
branches unbranched, clustered together on the phorinae derived from within it (GPWG II 2012;
main axis. Spikelets in pairs, with pedicels of Morrone et al. 2012). However, species sampling
is insufficient to confirm this. Many Digitaria branches; branch axis disarticulating at the nodes.
species are weeds. Digitaria exilis (fonio) and D. Glumes shorter than spikelet; lower lemma broad.
iburua are grown for grain in parts of Africa. Two spp., Tanzania, India, Sri Lanka, Burma.
480. Megaloprotachne C. E. Hubb.

XXIV.2. SUBTRIBE BOIVINELLINAE Pilg. (1940)
Megaloprotachne C. E. Hubb., Bull. Misc. Inform. Kew
1929: 320–321 (1929). Microcalaminae Butzin (1970).
Annuals. Ligule a fringed membrane. Inflores- Plants sprawling or trailing. Leaves broad, lance-
cence with unbranched branches, with the spike- olate, with cordate or pseudopetiolate bases.
lets borne on one side. Spikelets in pairs. Lower Fourteen genera, 113 species.
glume obtuse, with hyaline tip, as long as the These genera form the “forest shade clade” of
spikelet. Lemma margins flat, overlapping the Aliscioni et al. (2003), Giussani et al. (2001), and
palea. Morrone et al. (2012). Most have paired spikelets.
One sp., M. albescens C. E. Hubb., Southern All except the polyploid members of Alloteropsis
Africa. are C3 and some are reported to have mesophyll
The only really distinctive character of with radiate chlorenchyma; in Watson and Dall-
Megaloprotachne appears to be the long lower witz (1992 onward) this is described as “Isachne-
glume. type” in some genera but not in all. Morrone et al.
(2012) include Echinochloa in this subtribe based
on sequences of a chloroplast gene (ndhF) and
481. Tarigidia Stent
morphology; however, it is unrelated to other
Tarigidia Stent, Bull. Misc. Inform. Kew 1932(3): 151
Boivinelleae based on nuclear gene sequences
(1932). (Christin et al. 2007; Vicentini et al. 2008) and is
therefore considered incertae sedis in Paniceae in
Annuals or perennials. Ligules membranous. this treatment.
Inflorescence of short, appressed or divergent
unbranched branches that are shed at maturity. 483. Acroceras Stapf
Primary branches winged. Spikelets nearly sessile.
Lower glume present, 1/4–2/3 the length of the Acroceras Stapf, Fl. Trop. Afr. 9: 621 (1920).
spikelet. Lemma margins lying flat on the palea. Commelinidium Stapf, Fl. Trop. Afr. 9: 627 (1920).
Two spp., one in southern Africa and one in
Puerto Rico. Annuals or perennials. Leaves with or without
The two species of Tarigidia are thought to be pseudopetioles. Ligule a fringed membrane.
independent intergeneric hybrids between Digi- Inflorescence of unbranched branches. Spikelets
taria and Anthephora (Loxton 1974; Vega et al. paired, both pedicellate, but often in long-and-
2010); morphological phylogenetic data are con- short combinations. Upper glume and both
sistent with this hypothesis, but molecular data lemmas with the apex crested, blunt, hard and
are not available. laterally compressed, crest of the upper lemma
greenish. 2n ¼ 36.
Twenty spp., Africa, Madagascar, India, SE
482. Trachys Pers. Asia.
Trachys Pers., Syn. Pl. 1: 85 (1805). 484. Alloteropsis J. Presl
Annuals or perennials. Ligule membranous or Alloteropsis J. Presl, Reliq. Haenk. 1(4–5): 343, pl. 47
a fringe of hairs. Inflorescence with primary (1830).
branches unbranched, the rachis broad and
winged, often enclosing the spikelets, which are Perennials. Ligule a fringe of hairs or a fringed
borne in clusters of 2 to 6 on the underside of the membrane. Inflorescence of unbranched
branches, the spikelets borne on one side. Spike-

lets clustered on the branches. Lower glume
much smaller than the upper one. Proximal
lemmas with or without awns, distal lemmas
awned. C3 and C4. 2n ¼ 18, 54.
Five spp., Old World tropics.
Alloteropsis is monophyletic, and the C3 path-
way in this group is derived (GPWG II 2012;
Ibrahim et al. 2009).
485. Amphicarpum Kunth Fig. 76
Amphicarpum Kunth, Révis. Gramin. 1: 28 (1829).
Annuals or perennials. Ligule a fringe of hairs.

Plants with cleistogamous spikelets borne
underground or at lower nodes of the culm. Inflo-
rescence branches branched, the visible above-
ground inflorescence sterile. Glumes one, if two
then the lower one much smaller than the upper.
Upper lemma coriaceous to indurate, acuminate,
with margins lying flat on the palea. 2n ¼ 18.
Two spp., southeastern US.
Amphicarpum is strongly supported as being
sister to Entolasia (GPWG II 2012), and differs
primarily because of geography (North America
versus African and Australian), the presence
of cleistogamous spikelets, and the number of
orders of branching in the inflorescence (Entola-
sia has unbranched branches). The two genera
could perhaps be combined.
486. Cyphochlaena Hack.
Cyphochlaena Hack., Oesterr. Bot. Z. 51: 465 (1901).
Annuals or perennials, trailing. Ligule a fringed

membrane. Inflorescence branches unbranched.
Spikelets paired; sessile one staminate or sterile;
pedicellate one pistillate or hermaphrodite. Pistil-
late spikelets laterally compressed. Lower glume
with an apical awn. Upper glume and lower lemma
coriaceous. Upper lemma translucent. Staminate
spikelet similar to pistillate or reduced. C3.
Two spp., Madagascar. Fig. 76. Panicoideae-Paniceae-Boivinellinae. Amphicar-
Cyphochlaena is sister to the clade of Poeci- pum amphicarpon. A Plant with aerial and subterranean
lostachys plus Oplismenus (GPWG II 2012). All spikelets. B Subterranean spikelet. C Subterranean flower.
D Aerial spikelets. E Aerial flower. (From Barkworth et al.
three share laterally compressed spikelets. 2003, drawn by A. Sudkamp)
487. Cyrtococcum Stapf eaten and dispersed by fruit-eating birds

(Davidse and Morton 1973).
Cyrtococcum Stapf, Fl. Trop. Afr. 9: 15 (1917).
Annuals or perennials. Ligule membranous or 490. Mayariochloa Salariato, Morrone & Zuloaga
a fringed membrane. Inflorescence branches
branched, spreading. Spikelets laterally com- Mayariochloa Salariato, Morrone & Zuloaga, Syst. Bot. 37:
110 (2012); phylog.: Salariato et al. (2012).
pressed and asymmetrically gibbous. Glumes
shorter than the flowers, obtuse. Upper lemma
Rhizomatous perennials. Ligule a fringe of hairs.
crested.
Leaves with the junction between sheath and
Fifteen spp., tropical areas, Old World.
blade indistinct; leaf blade apex obtuse. Inflores-
cence sparse, the branches branched but with few
488. Entolasia Stapf spikelets. Rachilla extension present. Upper
lemma crested and pilose at the tip. C3.
Entolasia Stapf, Fl. Trop. Afr. 9: 739 (1920). One sp., M. amphistemon (C. Wright) Salar-
iato, Morrone & Zuloaga, Cuba.
Annuals or perennials, rambling. Ligule a fringe The genus Mayariochloa was described to
of hairs. Inflorescences branches generally accommodate the species formerly known as
unbranched. Upper flower much shorter than Scutachne amphistemon (C. Wright) Hitchc. &
the spikelet. Upper lemma and palea with short Chase. The type of Scutachne, S. dura (Griseb.)
appressed trichomes, also sometimes with a tiny Hitchc. & Chase, is unrelated (Salariato et al.
crest reminiscent of Acroceras. 2n ¼ 18. 2012). Scutachne s.s. falls in Melinidiinae and
Six spp., Tropical Africa and Australia. is C4; here it is placed in synonymy under Uro-
Entolasia is strongly supported as being sister chloa.
to Amphicarpum (GPWG II 2012), and differs
primarily because of geography (African and
Australian versus North American) and the pres- 491. Microcalamus Franch.
ence of cleistogamous spikelets in Amphicarpum.
Microcalamus Franch., J. Bot. (Morot) 3: 282 (1889).
The two genera could perhaps be combined.
Perennials, decumbent. Leaves pseudopetiolate,
489. Lasiacis (Griseb.) Hitchc. the blade broadly ovate to elliptical. Ligule a fringed
membrane; contraligule present, a fringe of hairs.
Lasiacis (Griseb.) Hitchc., Contr. U.S. Natl. Herb. 15: 16 Inflorescence branches branched. Spikelets later-
(1910). ally compressed. Flowers with a pubescent callus,
the upper lemma crested at the tip.
Perennials (one species annual), the culms One sp., M. barbinodis Franch., Cameroon to
generally not self-supporting, woody. Leaf blades Gabon.
linear to ovate. Ligule membranous or a fringed
membrane. Inflorescence branches themselves
branched. Spikelets not compressed, obliquely 492. Oplismenus P. Beauv.
attached to pedicel. Lower glume saccate. Glumes
and lower lemma become black at maturity, and Oplismenus P. Beauv., Fl. Oware 2: 14 (1810), nom. con-
the epidermis filled with oil globules. Upper serv.
lemma and palea apex obtuse, with tiny hollows
near the apex bearing woolly pubescence. Meso- Annuals or perennials, decumbent, trailing. Ligule
phyll with adaxial palisade layer. 2n ¼ 18, 36. a fringed membrane. Inflorescence branches
Sixteen spp., tropics of the New World, unbranched. Spikelets laterally compressed,
Madagascar. paired, but sessile one often reduced. Lower
In L. grisebachii (Nash) Hitchc. and L. rusci- glume awned, upper glume awned or not, the awn
folia (Kunth) Hitchc. the rachilla is extended with a sticky secretion. Upper lemma smooth and
beyond the distalmost flower. The spikelets are shining, crested at the tip. 2n ¼ 18, 36, 54, 72, 90.
Eleven spp., tropical and subtropical regions 496. Pseudechinolaena Stapf

worldwide.
Pseudechinolaena Stapf, Fl. Trop. Afr. 9: 494 (1919).
493. Ottochloa Dandy Annuals. Leaf blades lanceolate, pseudopetiolate.

Ligule a fringed membrane. Inflorescence
Ottochloa Dandy, J. Bot. 69: 54 (1931). branches unbranched. Spikelets paired, laterally
compressed. Upper glume gibbous, with tubercle-
Slender decumbent perennials. Leaf blades based, hooked hairs developing after fertilization.
disarticulating from sheaths. Ligule membranous 2n ¼ 36.
or a fringed membrane. Inflorescence branches Six spp., one pantropical, the other five in
branched. Both glumes less than ½ the length of Madagascar.
the spikelet. Lower palea lacking. 2n ¼ 18.
Three spp., Old World tropics.
XXIV.3. SUBTRIBE NEURACHNINAE Clayton &
494. Parodiophyllochloa Zuloaga & Morrone Renvoize (1986)
Parodiophyllochloa Zuloaga & Morrone, Syst. Bot. 33: 69 Perennials. Ligule a fringe of hairs. Inflorescences
(2008). dense, capitate, unbranched, or the branches very
“Panicum” sect. Cordovensia Parodi, Physis (Buenos short. Spikelets with the lower flower reduced to a
Aires) 8: 68 (1925).
lemma.
Three genera and 19 species.
Perennials with short rhizomes. Ligules membra-
This group is almost entirely Australian,
nous. Inflorescence open, branched, with both
except for Cleistochloa, which also occurs in west-
chasmogamous and cleistogamous spikelets.
ern parts of the Pacific and New Guinea.
Lower glume 3/4 the length of the spikelet; lower
palea absent. Upper flower with simple papillae
over the lemma and palea; lemma apiculate. 497. Cleistochloa C. E. Hubb.
Six spp., Mexico to Argentina.
This genus was segregated from Panicum based Cleistochloa C. E. Hubb., Hooker’s Icon. Pl. 33: t. 3209
(1933).
on its morphology, leaf anatomy, and position in
Calyptochloa C. E. Hubb., Hooker’s Icon. Pl. 33: t. 3210
molecular phylogenies (Morrone et al. 2008). (1933).
Ancistrachne S. T. Blake, Pap. Dept. Biol. Univ. Queens-
land 1: 4 (1941).
495. Poecilostachys Hack. Dimorphochloa S. T. Blake, Pap. Dept. Biol. Univ. Queens-
land 1: 1 (1941).
Poecilostachys Hack., Sitzungsber. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Cl., Abt. 1 89: 131 (1884). Scrambling, caespitose or mat-forming, with wiry
culms. Leaf blades disarticulating. Ligule a fringe
Annuals or perennials. Leaf blades broad. Inflo- of hairs. Cleistogamous spikelets frequent in leaf
rescence branches unbranched. Spikelets paired, axils of some species. Lower glume lacking, or
laterally compressed. Lower glume awned or with minute, or less than ½ the length of the spikelet.
a short sharp point. Upper lemma and palea car- Upper glume and lower lemma pubescent, in
tilaginous, not indurate. Hilum long linear. C3. some species the hairs pustule-based and hooked.
Nineteen spp., Madagascar and Africa. Upper lemma awned or awnless. C3. Mesophyll
Poecilostachys is placed Incertae sedis by with radiate chlorenchyma.
Morrone et al. (2012) based on conflicting and Eight spp., Australia, Philippines, New Cale-
weakly supported positions in phylogenies. How- donia, New Guinea.
ever, it is strongly supported as sister to Oplisme- The genera synonymized here in Cleistochloa
nus by GPWG II (2012). form a clade in molecular phylogenies (GPWG II
2012; Morrone et al. 2012), have several morpho-

logical synapomorphies, and are generally similar
in habit. Morrone et al. (2012) included
Ancistrachne, Calyptochloa and Dimorphochloa
(¼ Cleistochloa rigida) in their analysis, while
(GPWG II 2012) included Ancistrachne Calypto-
chloa and Cleistochloa. Grassworld (http://grass-
world.myspecies.info/) maintains all four genera
as separate.
Remaining two genera:.
Spikelet with a pubescent callus. Glumes as long as
spikelet, hard, the upper hairy on margins; lemmas
less firm than the glumes.
Molecular phylogenies show that Thyridolepis
is sister to Neurachne (Christin et al. 2012; GPWG II
2012), and that the monospecific Paraneurachne is
derived within Neurachne (Christin et al. 2012).
Accordingly, the latter two genera are combined
here.
498. Neurachne R. Br.
Neurachne R. Br., Prodr. 196 (1810).

Paraneurachne S. T. Blake, Contr. Queensland Herb. 13:
20 (1972).
Caespitose or stoloniferous. Glumes muticous or

awned. Upper lemma acute or with a beaked tip.
C3, C4, or C3/C4 intermediate; generally with two
bundle sheaths. 2n ¼ 18, 36, 54.
Eight spp., Australia.
The two C4 species of Neurachne, N. munroi
(F. Muell.) F. Muell. and N. muelleri Hack., are
not sisters, nor is either sister to the C3-C4 Fig. 77. Panicoideae-Paniceae-Neurachninae. Thyridole-
N. minor intermediate, S. T. Blake (Christin pis mitchelliana. A Habit. B Plant with inflorescence. C
et al. 2012). Thus, the C4 pathway appears to Part of leaf flattened out, from the upper side showing
ligule, pseudopetiole and base of blade. D, E Spikelet from
have originated several times within this genus. lower part of inflorescence, front and back view. F, G
Spikelet from upper part of inflorescence, front view and
slightly oblique side view. H Lower glume from inside. I
499. Thyridolepis S. T. Blake Fig. 77 Upper glume from inside. J Flowers from the side, the
lower at right. K Lower flower from back. L Upper flower
Thyridolepis S. T. Blake, Contr. Queensland Herb. 13: 25 from back. M Upper flower from front. N Lodicules. O
(1972). Pistil and stamens. P Caryopsis, adaxial view, abaxial
view, and side view. (From Blake 1972)
Caespitose or decumbent. Glumes obtuse, the
lower one with a hyaline patch below a prominent CENCHRINAE + MELINIDINAE + PANICINAE CLADE
transverse ridge covered with stiff hairs. Upper These three subtribes form a strongly sup-
lemma with a beaked tip. C3. ported clade in GPWG II (2012), and were called
Three spp., Australia. the “C4, three subtypes clade” by Giussani et al.
(2001). They have no obvious morphological
character in common, but together they represent Beckeropsis Fig. & De Not, Agrostogr. Aegypt. 2: 49 (1853).
a single origin of the C4 pathway. Cenchrinae are Odontelytrum Hack., Oesterr. Bot. Z. 48: 86 (1898).
characterized by the NADP-ME subtype, with a
single bundle sheath around the vascular tissue. Annuals or perennials. Ligule generally a fringe of
Melinidinae and Panicinae both have a double hairs, occasionally membranous. Lateral (second-
bundle sheath, with the Melinidinae exhibiting ary) branches with an expanded base, deciduous
the PCK subtype and Panicinae the NAD-ME as a unit. Each branch terminating in a spikelet,
subtype. The clade appears to be sister to Homo- which is then subtended and more or less sur-
pholis in chloroplast phylogenies. rounded by bristles, the bristles corresponding to
higher-order branches. Bristles terete or flattened,
separate or more or less fused; forming a mem-
XXIV.4. SUBTRIBE CENCHRINAE Dumort. (1824)
branous cup in C. abyssinicus (Hack.) Morrone.
Setariinae Dumort. (1824).
2n ¼ 14, 18, 22, 34, 35, 36, 40, 44, 45, 52, 54, 68.
Pennisetinae Rchb. (1828). One hundred twenty-one spp., worldwide,
Snowdeniinae Butzin (1972). especially warm areas.
Pseudoraphidinae Keng & Keng f. (1990). “Pennisetum” and Cenchrus were formally
merged by Chemisquy et al. (2010), following
years of accumulating evidence that Cenchrus s.
Primary, secondary, and/or tertiary branches ter- s. is derived from within the former “Pennise-
minating in a sterile point or bristle. C4, with a tum”. Several species (e.g., C. ciliaris L.) have
single bundle sheath; NADP-ME subtype. always been difficult to place, and appear inter-
Twenty-three genera and 293 species. mediate between both “genera”. The species of
The bristles have been variously interpreted, Cenchrus + “Pennisetum” are strongly supported
but developmental and gene expression data as forming a clade with both morphological and
show that they are sterile branches (Doust molecular data; furthermore, allopolyploid taxa
and Kellogg 2002; Kellogg et al. 2004). The form from parents assigned to both genera sug-
bristle appears to have been lost in Alexfloydia, gesting gene flow via the polyploids. A few species
“Panicum” antidotale, Snowdenia, Stereochlaena, of Cenchrus s.l. have only one bristle per spikelet;
Whiteochloa, and Zuloagaea. although these were segregated as separate genera
The inflorescence branches are deciduous in (Odontelytrum, Beckeropsis), they are simply
the small genera Paratheria and Streptolophus, derived members of the clade. Cenchrus ameri-
making them similar to Cenchrus; it is possible canus (L.) Morrone (formerly Pennisetum glau-
that the three genera should be merged. cum L.) is cultivated as pearl millet, and C.
compressus (R. Br.) Morrone (formerly Pennise-
500. Alexfloydia B. K. Simon tum alopecuroides L.) is a common ornamental.
Many species of Cenchrus (e.g., C. ciliaris and C.
Alexfloydia B. K. Simon, Austrobaileya 3: 670 (1992). echinatus L.) are weedy. Aposporous apomixis is
common in the genus (Ozias-Akins et al. 2003),
Stoloniferous perennials, mat-forming. Ligule a but the number of origins is unclear (Akiyama
fringe of hairs. Inflorescence unbranched or et al. 2011).
branched, with 3 to 6 spikelets; the pedicels slen-
der. Spikelets laterally compressed. Upper lemma 502. Chamaeraphis R. Br.
cartilaginous, not indurate.
One sp., A. repens B. K. Simon, known from Chamaeraphis R. Br., Prodr. 193 (1810).
only a few localities in New South Wales, Austra-
lia. Listed as endangered. Caespitose perennials. Ligule membranous. Each
spikelet paired with a single bristle, the spikelet-
501. Cenchrus L. bristle pairs on alternate sides of the flattened
main inflorescence axis, deciduous. Lower
Cenchrus L., Sp. Pl. 2: 1049 (1753); phylog.: Akiyama et al. glume minute or absent; upper glume with 7
(2011); Chemisquy et al. (2010); Donadio et al. (2009). veins. Upper lemma smooth.
Pennisetum Rich., Syn. Pl. 1: 72 (1805).
One sp., C. hordeacea R. Br., Australia. flower at maturity. Upper lemma rugose, mucro-
This species is sister to Pseudoraphis in the nate. 2n ¼ 36, 54.
analysis of Morrone et al. (2012). One sp., I. unisetus (J. Presl) Schltdl., Mexico
and Central America.
Ixophorus is strongly supported as sister to
503. Dissochondrus (Hillebr.) Kuntze
Zuloagaea and Setariopsis (GPWG II 2012). The
Dissochondrus (Hillebr.) Kuntze, Revis. Gen. Pl. 2: 770 three genera have overlapping distributions in
(1891). Mexico and Central America. Ixophorus is an
allopolyploid, formed from the same allopoly-
Caespitose perennials. Leaves pseudopetiolate, ploid event that produced Z. bulbosa (Doust
the blades auriculate. Ligule membranous. Inflo- et al. 2007).
rescence narrow, the branches branched. Spike-
lets subtended by one or more bristles;
506. “Panicum” antidotale
disarticulation above the bristles and below the
glumes. Spikelets with both flowers fully devel- “Panicum” antidotale Retz., Observ. Bot. 4: 17 (1786).
oped and bisexual. Lemmas rugose. Leaf meso-
phyll with radiate chlorenchyma, and isolated Perennial with short woody rhizomes. Ligule a
bundle sheath cells similar to those in Arundi- fringe of hairs. Inflorescence branches branched,
nella. the higher-order branches contracted around the
One sp., D. biflorus Kuntze ex Hack., Hawaii. primary ones. Upper lemma and palea indurate,
shiny.
504. Hygrochloa Lazarides One sp., widespread in tropical areas of the
world.
Hygrochloa Lazarides, Brunonia 2: 86 (1979). Although this species looks like a conven-
tional Panicum, molecular data place it within
Monoecious annuals or perennials, growing in or Cenchrinae (Morrone et al. 2012).
near water. Leaf blades papillose above. Ligule a
fringe of hairs. Primary inflorescence branches
unbranched, ending in a point. Pistillate spikelets 507. Paratheria Griseb.
borne on lower branches, staminate spikelets
pedicellate on the main inflorescence axis. Paratheria Griseb., Cat. Pl. Cub. 236 (1866).
Upper lemma coriaceous, the margins lying flat
on the palea. Prostrate perennials, rooting at the nodes. Ligule
Two spp., Northern Australia. a fringed membrane or a fringe of hairs. Primary
branches of the inflorescence appressed to the
main axis, each with one spikelet and ending in
505. Ixophorus Schltdl. a sharp sterile tip. Inflorescence branches disarti-
culating as a unit, below the bristles. Cleistoga-
Ixophorus Schltdl., Linnaea 31: 420 (1861); devel., rev.: mous spikelets present at the base of the
Kellogg et al. (2004).
inflorescence. Glumes minute, without veins.
Lower and upper lemma acuminate to subulate,
Monoecious caespitose annuals or short-lived
the upper not indurate.
perennials. Leaves linear, conduplicate in bud,
Two spp., Madagascar, West Africa, West
the sheaths strongly flattened and keeled. Ligule
Indies and South America.
a fringed membrane. Inflorescence with the pri-
mary branches unbranched. Each spikelet accom-
panied by a bristle. Upper flower pistillate and 508. Paractaenum P. Beauv.
lower flower staminate. Lower paleas hyaline,
about as long as the upper glume, accrescent, Paractaenum P. Beauv., Ess. Agrostogr. 47 (1812).
forming a winged structure clasping the upper Plagiosetum Benth., Hooker’s Icon. Pl. 13: 33 (1877).
Small caespitose annuals. Ligule a fringed mem- 511. Setaria P. Beauv. Fig. 78
brane or a fringe of hairs. Primary inflorescence
branches deciduous, wedge-shaped in P. refrac- Setaria P. Beauv., Ess. Agrostogr. 51 (1812).
tum. Each spikelet subtended by one or more Paspalidium Stapf, Fl. Trop. Afr. 9: 582 (1920).
bristles, rachis ending in one bristle or a fan of
bristles. Lower lemma with more than nine veins. Annuals or perennials, caespitose, rhizomatous
Two spp., Australia. or stoloniferous. Leaf blades flat, folded, or pli-
Kellogg et al. (2009) suggest that this genus is cate, sagittate in a few species. Ligule a fringed
related to the Australian species of Setaria for- membrane or a fringe of hairs. Inflorescences lax,
merly classified in Paspalidium.
509. Pseudochaetochloa Hitchc.
Pseudochaetochloa Hitchc., J. Wash. Acad. Sci. 14: 492

(1924).
Caespitose perennials; dioecious. Ligule a fringe

of hairs. Primary inflorescence branches with 2 to
5 spikelets, with 7 to 12 bristles beneath each
spikelet, the bristles falling with the spikelets;
bristles with long hairs proximally. Glumes and
lower lemma membranous, the latter muticous.
Upper lemma and palea coriaceous.
One sp., P. australiensis Hitchc., Australia.
Pseudochaetochloa is similar to Pennisetum,
but has more elongate secondary branches and is
also dioecious. It falls in a clade with Spinifex and
Zygochloa, both of which are also dioecious
(Morrone et al. 2012).
510. Pseudoraphis Griff.
Pseudoraphis Griff., Not. Pl. Asiat. 3: 29 (1851).
Decumbent perennials. Ligule membranous. Pri-

mary inflorescence branches each with one or a
few spikelets and tipped with a bristle. Lower
glume minute, without veins; upper glume
mucronate to awned. Upper flower borne atop a
long rachilla internode. Lower lemma much
longer than the upper one, awned. 2n ¼ 16.
Ten spp., Asia and Australia.
Fig. 78. Panicoideae-Paniceae-Cenchrinae. Setaria lach-
Although one representative Pseudoraphis nea. A Plant. B Portion of an inflorescence branch. C
was included by Kellogg et al. (2009) in Cenchri- Spikelet, lateral view with base of bristle. D Upper
nae, its position remains unresolved. It is sup- lemma and palea, viewed from the adaxial side. E Upper
ported as sister to Chamaeraphis by Morrone lemma, viewed from the abaxial side. F Lower palea and
upper lemma and palea, viewed from the adaxial side. G
et al. (2012), but the position of the clade within Caryopsis, scutellar view. H Same, hilar view. (From
the subtribe is not clear. Zuloaga et al. 2012)
open and highly branched, or more or less con- unbranched, or with unbranched branches. Sterile
tracted and spiciform, or a set of unilateral branches (bristles) absent. Spikelets pedicellate.
racemes; bristles either associated with each Glumes minute. Lower lemma awned, as long as
mature spikelet or terminating the primary inflo- the upper lemma and palea. Upper lemma and
rescence branches or both. Disarticulation above palea membranous, not indurate, the lemma
the bristles and below the spikelets. Lower glume with margins flat and overlapping the palea.
generally shorter than the upper one. Upper Four spp., northeast tropical Africa.
lemma and palea crustaceous or coriaceous, gen- Although this genus lacks sterile branches in
erally ellipsoid, usually corrugate, granular or the inflorescence, sequences of the chloroplast
rugose, or rarely smooth. 2n ¼ 18, 36, 54, 63, 72. gene ndhF place it within Cenchrinae (Morrone
One hundred fifteen spp., world wide, in tem- et al. 2012).
perate and tropical areas.
Molecular phylogenies (Kellogg et al. 2009) and
514. Stenotaphrum Trin.
a recent monograph (Morrone et al. 2014) show that
Setaria and Paspalidium intergrade. Setaria is not Stenotaphrum Trin., Fund. Agrost. 175 (1820) [1822].
monophyletic even when Paspalidium is included.
However, to make it monophyletic would require Annuals or perennials, rhizomatous, stolonifer-
expanding it to include much of the bristle clade ous, or caespitose. Ligule a fringed membrane.
outside of Cenchrus. This is not warranted until a Inflorescence unbranched, or with very short pri-
better supported phylogeny is obtained. mary branches sunken into the broad flattened
inflorescence axis. Rachis of primary branches
512. Setariopsis Scribn. terminating in a point. Lower glume tiny, with
no veins. Upper lemma leathery but not indurate,
Setariopsis Scribn., Publ. Field Columb. Mus., Bot. Ser. yellow in fruit. 2n ¼ 18, 20, 36.
1(3): 288–289 (1896). Seven spp., tropical and subtropical.
Annuals or perennials. Ligule a fringed mem-

brane. Inflorescence branches unbranched. Each 515. Stereochlaena Hack.
spikelet subtended by a bristle; disarticulation
Stereochlaena Hack., Proc. Rhodesia Sci. Assoc. 7: 65
above the bristle and below the spikelet. Lower (1908).
glume small and ovate; upper glume saccate and
gibbous, becoming indurate. Lower lemma with Annuals or perennials. Ligule a fringed mem-
two abaxial pouches, becoming indurate, exceed- brane. Inflorescence branches unbranched, in
ing the upper flower. Upper lemma and palea pairs or digitate, the branches winged; bristles
indurate, apiculate. 2n ¼ 18. lacking. Spikelets paired, borne on one side of
Two spp., Mexico to Colombia. the branches. Lower glume tiny or absent;
Setariopsis is strongly supported as sister to upper glume minute to as long as the spikelet,
Ixophorus and Zuloagaea (GPWG II 2012). The awned or not. Lower lemma awned. Lodicules
three genera have overlapping distributions in absent.
Mexico and Central America. Four spp., Tanzania to southern Africa.
Stereochlaena is placed in the bristle clade on
the basis of morphological data alone (Morrone
513. Snowdenia C. E. Hubb.
et al. 2012).
Snowdenia C. E. Hubb., Bull. Misc. Inform. Kew 1929:
30–31 (1929). 516. Streptolophus Hughes
Stoloniferous or decumbent annuals or peren- Streptolophus Hughes, Bull. Misc. Inform. Kew 1923: 178
nials. Ligule membranous. Inflorescence (1923).
Annuals. Culm nodes hairy. Leaves pseudopetio- Six spp., Australia.

late, the blades cordate or sagittate. Ligule a
fringed membrane. Inflorescence with primary
519. Xerochloa R. Br.
branches short, themselves branched into cune-
ate higher-order branches. The branches divided Xerochloa R. Br., Prodr. 196 (1810).
into multiple branchlets with recurved tips that
form an involucre around the spikelets. Inflores- Annuals or perennials, erect or decumbent; culms
cence branches disarticulating as a unit, below herbaceous or woody. Ligule a fringed mem-
the bristles. Glumes small. brane. Inflorescence axes short and unbranched,
One sp., S. sagittifolius Hughes, Angola. each branch associated with a prophyll and end-
Streptolophus falls with Paratheria in the ana- ing in a point; inflorescences borne in a cluster
lyses of Morrone et al. (2012), although molecular and subtended by a spatheole. Lower glume
data are lacking for the former. In both genera the small; glumes and lower lemmas membranous.
branches disarticulate as a unit. The involucral Upper lemma acuminate.
branches around the spikelets make the plants Three spp., Australia.
superficially similar to Cenchrus.
520. Zuloagaea Bess

517. Uranthoecium Stapf
Zuloagaea Bess, Syst. Bot. 31: 666 (2006).
Uranthoecium Stapf, Hooker’s Icon. Pl. 31, t. 3073 (1916).
Caespitose perennials, with short tough rhi-
Caespitose annuals. Ligule a fringed membrane zomes, these often forming spherical corms. Lig-
or fringe of hairs. Primary inflorescence branches ule membranous. Inflorescence highly branched,
flattened, appressed to the main axis, short, end- spreading. Upper lemma and palea indurate, sca-
ing in a point, disarticulating between the spike- brous at the apex. 2n ¼ 18, 36, 54, 72.
lets. Glumes truncate, gibbous. Apex of lower One sp., Z. bulbosa (Kunth) Bess, Mexico,
lemma extended, long acuminate; apex of upper Central America, and northern South America.
lemma and palea prolonged, awl-shaped. Zuloagaea is strongly supported as sister to
One sp., U. truncatum (Maiden & Betche) Ixophorus and Setariopsis (GPWG II 2012). The
Stapf, Australia. three genera have overlapping distributions in Mex-
This species is closely related to Setaria reti- ico and Central America. Zuloagaea is one of only a
glumis (Domin) R. D. Webster (Kellogg et al. 2009) few genera of the bristle clade that lack bristles.
(= Paspalidium retiglume (Domin) Hughes).
Remaining two genera: Dioecious perennials
of dry sites, with dense capitate inflorescences. Both
518. Whiteochloa C. E. Hubb. form clumps connected by long rhizomes. Mem-
bers of the two are sisters in molecular phylogenies
Whiteochloa C. E. Hubb., Proc. Roy. Soc. Queensland 62: (Doust et al. 2007; Kellogg et al. 2009; Morrone
111 (1952).
et al. 2012), and the two genera could be merged.
Annuals or perennials, caespitose. Ligule a
fringed membrane. Inflorescence open, with lax, 521. Spinifex L.
branched branches, these ending in spikelets or
ending blindly in bristles. Spikelets laterally com- Spinifex L., Mant. Pl. 163, 300 (1771).
pressed or not compressed; rachilla internodes
prominent between the flowers. Lower glume Leaf blades hard, sometimes with a sharp point;
broad, ovate, clasping the spikelet; upper glume ligule a fringe of hairs. Inflorescences made up of
tuberculate-ciliate, with a hardened apex. Lower stiff branches all appearing to originate at a single
lemma sulcate. Upper flower on a short filiform point, each branch with its own spathe, the inflo-
internode. Upper lemma rugose, indurate, rescence disarticulating as a unit. Pistillate spike-
mucronate, yellow in fruit. lets solitary, each at the base of a long bare axis.
Staminate spikelets sessile on inflorescence axes. Lemmas awned or mucronate. Outer bundle
2n ¼ 18. sheath present (C. bromoides) or absent (C. festu-
Five spp., beaches of India, China, Australia. coides, C. cubanum). 2n ¼ 26.
Three spp., Mexico, West Indies, Brazil.
Chaetium bromoides (J. Presl) Benth. ex
522. Zygochloa S. T. Blake
Hemsl. falls within Urochloa in the phylogenetic
Zygochloa S. T. Blake, Pap. Dept. Biol. Univ. Queensland analyses of Salariato et al. (2010). Nonetheless,
1: 7 (1941). Chaetium is kept separate here provisionally
because C. bromoides is morphologically quite
Leaf blades flat, stiff, disarticulating from the different from other members of Urochloa
sheath. Ligule a fringe of hairs. Pistillate inflores- and because Salariato et al. were unable to include
cence branches each consisting of one spikelet the other two species of Chaetium. These may
plus three bracts, the branch ending in a sharp fall entirely outside the subtribe, based on
point; branches clustered into a globose head. their leaf anatomy. The type is C. festucoides
Upper lemma hardened. Staminate inflorescences Nees, which has not been studied phylogeneti-
similar to female, but branched. cally.
One sp., Z. paradoxa (R. Br.) S. T. Blake,
Australia. 524. Eccoptocarpha Launert
Eccoptocarpha Launert, Senckenberg. Biol. 46: 124 (1965).

XXIV.5. SUBTRIBE MELINIDINAE Pilg. (1940)
Annuals. Ligule a fringe of hairs. Inflorescence
Brachiariinae Butzin (1970). branches unbranched. Lower glume with glandu-
lar hairs, upper with stiff hairs. Upper glume and
Glumes and lemmas muticous to aristate. C4, with lower lemma with obvious net-like venation.
two bundle sheaths, PCK-subtype. Rachilla internode below upper flower slender,
Twelve genera and 176 species. elongate, sinuous. Upper lemma hairy at the tip,
Melinidinae are clearly monophyletic accord- hard.
ing to DNA sequences (Salariato et al. 2010), but One sp., E. obconiciventris Launert, Tanzania
the limits of the included genera are not clear. and Zambia.
This group is known in the literature as the PCK Molecular data are not available for Eccopto-
clade because it includes all Paniceae known to carpha, but it is placed sister to Moorochloa in the
exhibit the PCK subtype of C4 photosynthesis. analyses of Morrone et al. (2012).
However, not all species have been biochemically
typed; in addition, species with the other C4
subtypes appear to exhibit some PCK activity 525. Eriochloa Kunth
(Voznesenskaya et al. 2005) so it is not clear
Eriochloa Kunth, Nov. Gen. Sp. (quarto ed.) 1: 94–95, pl.
whether this is a good character. The bundle 30, 31 (1815).
sheath outline in most classical PCK species is
uneven, so may provide one anatomical synapo- Caespitose annuals. Culm nodes pubescent.
morphy. Ligule a fringe of hairs. Inflorescence branches
unbranched, pedicels and axis villous, the spike-
523. Chaetium Nees lets borne on one side. Spikelets with a globose
swelling (“bead”) at the base. Lower glume
Chaetium Nees, Fl. Bras. Enum. Pl. 2(1): 269–270 (1829). absent, upper glume and lower lemma villous,
muticous. Upper lemma rugose with a mucro or
Perennials. Ligule a fringed membrane or a fringe awn, margins tucked on to the palea. 2n ¼ 18.
of hairs. Inflorescence unbranched, or with One sp., E. distachya Kunth, South America.
unbranched branches. Spikelets with a pubescent Eriochloa is here restricted only to the type
callus. Glumes awned, the upper with 9–11 veins. species, E. distachya. Most species previously
classified within Eriochloa are part of Urochloa Two spp., seashores of the Old World tropics,
according to the extensive molecular phylogenetic Madagascar to Polynesia.
data of Salariato et al. (2010). Eriochloa s.s. appears
to be close to Moorochloa, Tricholaena and Meli-
529. Urochloa P. Beauv. Fig. 79
nis.
Urochloa P. Beauv., Ess. Agrostogr. 52, pl. 11, f. 1 (1812).
526. “Panicum” deustum Thunb. Brachiaria (Trin.) Griseb., Fl. Ross. 4: 469 (1853).
Scutachne Hitchc. & Chase, Proc. Biol. Soc. Wash. 24:
148–149, f. 13 (1911).
“Panicum” deustum Thunb., Prodr. Pl. Cap.: 19 (1794). Megathyrsus (Pilg.) B. K. Simon & S. W. L. Jacobs,
Austrobaileya 6: 572 (2003).
Caespitose perennials. Ligule a fringed mem-
brane. Inflorescence branches branched, often
with clavellate hairs. Glumes with apices acute.
Lemma and palea hardened.
One sp., Africa, Asia, South America.
“Panicum” deustum shares pubescent inflo-
rescence branches with Rupichloa; the two should
perhaps be combined in a single genus. They are
in an unresolved position in the phylogeny of
Morrone et al. (2012).
527. Rupichloa D. Salariato & Morrone
Rupichloa D. Salariato & Morrone, Taxon 58: 388–389

(2009).
Perennials, rhizomatous or caespitose. Ligule a

branched, with long pubescence. Lower glume
clasping the base of the spikelet, the upper
glume shorter than the upper lemma. Upper
lemma striate, verrucose, with a crested apex,
with flattened macrohairs. 2n ¼ 26, 28.
Two spp., Bahia and Minas Gerais, Brazil.
528. Thuarea Pers.
Thuarea Pers., Syn. Pl. 1: 110 (1805).
Mat-forming perennial. Ligule a fringe of hairs.

Inflorescence enclosed in the uppermost leaf,
unbranched with a flattened axis, with 1 or Fig. 79. Panicoideae-Paniceae-Melinidinae. Urochloa
paucispicata. A Plant. B Ligular region. C Portion of an
2 bisexual spikelets proximally, and staminate inflorescence branch. D Spikelet, viewed from the adaxial
spikelets distally. Staminate spikelets deciduous, side. E Upper lemma and palea, viewed from the adaxial
on bulbous pedicels. Bisexual spikelets persistent; side. F Apex of the lemma. G Upper lemma, viewed from
lower glume absent; upper lemma pubescent, the abaxial side, showing germination flap at base. H
Lodicules. I Perfect flower. J Gynoecium. K Ovary and
with margins lying flat on the palea. Staminate basal part of the styles. L Caryopsis, scutellar view. M
portion of the inflorescence folds onto the bisexual Caryopsis, hilar view. N Same, cross section. (From
portion at maturity and encloses the fruit. Zuloaga et al. 2012)
Annuals or perennials, caespitose. Leaf blade 531. Leucophrys Rendle

nodular in cross section. Ligule a fringed mem-
brane or a fringe of hairs. Inflorescence branches Leucophrys Rendle, Cat. Afr. Pl. 2: 193 (1899).
branched or unbranched, spikelets borne on one
side only, branch rachis often winged. Spikelets Caespitose perennials, the culms woody. Leaf
solitary or in pairs, in some cases with a globose blades disarticulating from the sheath. Ligule a
swelling at the base. Lower glume generally fringe of hairs. Inflorescence with unbranched
shorter than the spikelet, sometimes absent. branches, borne along a central axis. Glumes
Upper glume often pubescent, about as long as awnless, separated by an elongated rachilla
the spikelet. Upper lemma rugose or smooth, the internode, both as long as the spikelet. Upper
margins tucked on to the palea or lying flat on the glume and lower lemma with a transverse line of
palea, acute to mucronate, with or without dense hairs.
hairs at the apex. 2n ¼ 18, 36, 54, 72. One sp., L. mesocoma (Nees) Rendle, tropical
One hundred thirty-five spp., tropical and and southern Africa.
subtropical regions of the world. The type of Leucophrys, L. mesocoma, is
Most species of this genus were formerly placed in Brachiaria by Clayton et al. (2006
placed in Brachiaria, but were transferred into onward), but is shown to be distinct by Salariato
Urochloa by Webster (1987); this transfer has et al. (2010).
largely been supported by subsequent molecular
studies. Salariato et al. (2010, 2012) show con- 532. Melinis P. Beauv.
vincingly that Urochloa is paraphyletic and
includes (at least) Scutachne s.s. and Megathyrsus Melinis P. Beauv., Ess. Agrostogr.: 54 (1812).
as well as several species formerly classified in Rhynchelytrum Nees, Nat. Syst. Bot.: 378, 446 (1836).
Eriochloa. The latter genus is polyphyletic and Mildbraediochloa Butzin, Willdenowia 6: 288 (1971).
the type falls outside the Urochloa clade. While
it has been recognized traditionally by its promi- Annuals or perennials, culm nodes pubescent.
nent bead-like callus that character has clearly Ligule a fringe of hairs. Inflorescence much
arisen multiple times and is not synapomorphic. branched, with slender branches. Spikelets later-
In a survey of embryological characters of ally compressed. Lower glume tiny or absent;
grasses, Verboom et al. (1994) found that the upper glume awned or awnless, gibbous in some
outer integument in Urochloa (¼ “Panicum” ¼ species. Lemma of lower flower awned. Lemma of
Megathyrsus) maxima and U. oligotricha scarcely the upper flower much less firm than the glumes,
develops, forming a small rim around the base of awnless or awned from a sinus. 2n ¼ 36.
the ovule; this character may be more widespread Twenty-two spp., Africa, tropics and south.
and deserves further investigation.
530. Yvesia A. Camus 533. Moorochloa (Sm.) Veldkamp
Yvesia A. Camus, Bull. Soc. Bot. France 73: 687 (1927). Moorochloa (Sm.) Veldkamp, Reinwardtia 12: 138–139
(2004).
Tiny annuals, less than 25 cm tall. Ligule a fringe of

hairs. Inflorescence branches unbranched, capil-
branches unbranched, one-sided. Spikelets with
lary. Callus with long white hairs. Lower glume
the upper flower disarticulating, upper glume and
absent, upper glume similar to lower lemma, awn-
lower lemma villous. Callus inconspicuous.
less. Upper lemma mucronate.
Upper lemma and palea cartilaginous, smooth,
One sp., Y. madagascariensis A. Camus,
apex muticous. 2n ¼ 18, 36.
Madagascar.
Three spp., Africa and Asia, but now wide-
LEUCOPHRYS + MELINIS + MOOROCHLOA + TRICHOLAENA spread and weedy.
Disarticulation above the glumes. Upper Moorochloa is the name given for three
lemma and palea smooth and shiny. species formerly in Brachiaria. The type of
These four genera form a strongly supported Brachiaria is assigned to Urochloa, so a new
clade in Morrone et al. (2012). name was needed for these species.
534. Tricholaena Schrad.
Tricholaena Schrad., Mant. 2: 8, 163 (1824).

highly branched, the branches spreading. Spike-
lets laterally compressed. Lower glume apex trun-
cate. Upper glume 3- to 5-veined. Lemma of lower
flower without an awn. Lemma of upper flower
leathery to hard. 2n ¼ 36.
Four spp., Mediterranean region, Africa,
India.
XXIV.6. SUBTRIBE PANICINAE Fr. (1835)
Upper lemma and palea indurate. Upper palea,

and often the upper lemma, with papillae borne
on other papillae (compound papillae) near the
apex. C4, with two bundle sheaths; NAD-ME sub-
type.
Three genera, 111 species.
The photosynthetic subtype is probably syna-
pomorphic for Panicinae, although not all species
have been tested biochemically.
535. Panicum L. Fig. 80
Panicum L., Sp. Pl. 1: 55 (1753).

Yakirra Lazarides & R. D. Webster, Brunonia 7: 292
(1985).
Annuals or caespitose perennials. Ligule a fringed

membrane. Inflorescence branches branched or
unbranched. Spikelets solitary. Upper glume and
lower lemma with 7 to 13 veins. 2n ¼ 18, 36, 54,
72, 90.
About one hundred spp., pan-tropical.
Yakirra is placed in Panicum by molecular
data from GPWG II (2012) and Scataglini et al.
(2014) and is accordingly placed in synonymy
here. Its placement outside Panicum in the work
of Morrone et al. (2012) was based on an errone-
ous sequence. Panicum sect. Rudgeana, Arthra-
grostis and Yakirra all share an elongate rachilla
internode below the upper flower; the internode
bears flanges or auricles distally.
the lower glume. C Spikelet, view from the back of the

upper glume. D Upper lemma, viewed from the abaxial
Fig. 80. Panicoideae-Paniceae-Panicinae. Panicum ber- side. E Upper lemma and palea, viewed from the adaxial
gii. A Flowering stem. B Spikelet, view from the back of side. (From Zuloaga et al. 2012, drawn by J.E. Lacour)
536. Louisiella C.E. Hubb. & J. Léonard mineae and sources of potential new germplasm.
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IX. Subfamily Danthonioideae N.P. Barker & H.P. Linder (2001)
Perennials or more rarely annuals. Ligule a fringe lemma with three tufts of hairs on either side of the
of hairs. Inflorescence branches branched (rarely midrib; plants of Africa 538. Merxmuellera
2. Leaf blade disarticulating at the ligule, not leaving a flap
unbranched). Spikelets laterally compressed, with of blade tissue; lemma hairs evenly spread or in lines,
1 to many flowers. Lemma bilobed, awned from but not tufts; not African 3
the sinus, the base of the awn flat. Style branches 3 (2). Lemma hairs evenly spread, not in lines; plants
2, widely separated. Embryo sac with haustorial gynodioecious; New Guinea 546. Chimaerochloa
synergids. Epiblast absent, scutellar cleft present, 3. Lemma hairs in four vertical lines; plants bisexual; New
Zealand, Australia and Lord Howe Island
embryonic mesocotyl internode elongated, the 542. Chionochloa
embryonic leaf margins meeting. Stomata with 4 (1). Lemma lacking an awn, the apex shallowly triden-
dome-shaped or parallel-sided subsidiary cells. tate or entire 5
Photosynthetic pathway C3. 4. Lemma awned, the awn generally from the sinus
Seventeen genera, 281 spp. between two prominent lobes 8
5 (4). Lemma glabrous 549. Notochloe
This treatment follows the revised classifica- 5. Lemma pubescent 6
tion presented by Linder et al. (2010). 6 (5). Disarticulation of flowers from a horizontal break in
Although haustorial synergids are likely to be the middle of the rachilla internode, the dispersal unit
synapomorphic for this subfamily, a number of thus being the flower plus a short piece of the rachilla;
the genera remain to be investigated for this spikelets with no more than 4 flowers
550. Plinthanthesis
character (Verboom et al. 1994). Merxmuellera 6. Disarticulation at the base of the flower, the dispersal
rangei and M. papposa, formerly in Danthonioi- unit not including the rachilla internode below; spike-
deae, clearly belong in the Chloridoideae (Peter- lets often with more than 4 flowers 7
son et al. 2010) and have been transferred to the 7 (6). Glumes shorter to more often longer than the
genus Ellisochloa P. M. Peterson and N. P. Barker. spikelet; plants often stoloniferous; glumes often with
tuberculate hairs; inflorescence sometimes
The phylogeny of Linder et al. (2010) is consistent unbranched 554. Tribolium
with that of the GPWG II (2012), and more thor- 7. Glumes shorter than the spikelet; plants lacking stolons;
oughly sampled. The ligule is a fringe of hairs glumes without tuberculate hairs; inflorescence
except in Geochloa, where it is a fringed mem- branched 553. Schismus
brane; this may be synapomorphic for the genus. 8 (4). Lemma awn without a clear demarcation between
the basal column and the elongate apical portion;
lemma lobes minute or absent; plants over 1 m tall 9
8. Lemma awn clearly divided into a flattened column and
KEY TO THE GENERA OF DANTHONIOIDEAE a long slender apical portion; lemma lobes prominent;
plants often less than 1 m 10
1. Leaf blades disarticulating from the sheath 2 9 (8). Leaf blade with a prominent midrib; plants of South
1. Leaf blades persistent 4 America 547. Cortaderia
2(1). Leaf blade disarticulating 1–2 cm distal to the ligule, 9. Leaf blade with several wide prominent veins; plants of
leaving a small straight or recurved flap of blade tissue; New Zealand 545. Austroderia
348 IX. Subfamily Danthonioideae N.P. Barker & H.P. Linder (2001)
10 (8). Flowers 2 (rarely 1) per spikelet; rachilla extended morphies of the genus but is included within it
beyond the distal spikelet; lemma with setae in the on the basis of molecular data (Linder et al. 2010).
sinus between the lobes and the central awn 11
10. Flowers more than 2 per spikelet; rachilla not
extended; lemma with setae from the apices of the C A P E O C H L O A + G E O C H L O A CLADE
lobes, or lacking 12 Palea margins with tufts of long hairs.
11 (10). Spikelets less than 2.5 cm long 541. Pentameris
11. Spikelets more than 2.5 cm long
544. Pseudopentameris 539. Geochloa H. P. Linder & N. P. Barker
12 (10). Plants geophytic, with swollen rhizomes; basal
leaf sheaths wooly 539. Geochloa Geochloa H. P. Linder & N. P. Barker, Ann. Missouri Bot.
12. Plants usually not geophytic, swollen rhizomes lack- Gard. 97: 323 (2010).
ing; if present then basal leaf sheaths glabrous and
shining 13 Geophytic perennials with swollen rhizomes. Basal
13 (12). Pedicel of spikelet disarticulating slightly below
the base of the glumes, the disarticulation point
leaf sheaths wooly pubescent. Ligule a fringed mem-
marked by a tuft of hairs 543. Chaetobromus brane. Inflorescence compact. Spikelets with 2 to 7
13. Pedicel of spikelet either disarticulating at the base of flowers. Glumes as long as flowers or longer. Callus
the glumes or persistent and disarticulation above the of flower pubescent. Lemma with trichomes scat-
glumes; pedicel without a tuft of hairs 14 tered over the abaxial surface, lobes ending in setae;
14 (13). Leaves tough, with abundant sclerenchyma;
plants of Africa to western Asia 15
column of awn distinct, straight or twisted. 2n ¼ 48.
14. Leaves membranous; plants of North and South Amer- Three spp., southern Cape Province, South
ica, Europe, and the Pacific rim 16 Africa. Geochloa occurs in the fire-dominated
15 (14). Plants forming robust clumps, generally over 1 m fynbos, and flowers only in the years immediately
tall; if less, then geophytic with swollen rhizomes; following a fire (Linder and Ellis 1990a).
plants of the Cape Province of South Africa
540. Capeochloa
15. Plants wiry, generally much less than 90 cm tall, not 540. Capeochloa H. P. Linder & N. P. Barker
geophytic; plants of African mountains and western
Asia 551. Tenaxia Capeochloa H. P. Linder & N. P. Barker, Ann. Missouri
16 (14). Lemma hairs in two rows, generally in tufts; Bot. Gard. 97: 324 (2010).
hilum punctate; plants of Australia, New Guinea,
New Zealand, and South America
552. Rytidosperma Caespitose perennials. Basal leaf sheaths glabrous.
16. Lemma hairs not tufted; hilum elliptical to elongate; Inflorescence branches branched. Spikelets with
plants of North America, South America, Europe 2 to 4 flowers. Glumes as long as flowers or longer.
548. Danthonia Callus of flower pubescent. Lemma trichomes var-
iously distributed, lobes ending in setae or not;
column of awn distinct, twisted. Palea margins
GENERA OF DANTHONIOIDEAE with tufts of hairs. 2n ¼ 12, 36.
Three spp., southern Cape Province, South
538. Merxmuellera Conert Africa. Linder et al. (2010) suggest that C. arundina-
cea may be of hybrid origin, but include it in Capeo-
Merxmuellera Conert, Senck. Bot. 51: 129 (1970). chloa based on sequences of the nuclear ribosomal
RNA and its morphological similarity to C. cincta.
Caespitose perennials. Leaf blade disarticulating
1–2 cm distal to the ligule, leaving a small straight 541. Pentameris P. Beauv. Fig. 81
or recurved flap of blade tissue; leaf sheath
splitting. Inflorescence branches branched. Spi- Pentameris P. Beauv., Ess. Agrost. 92 (1812); tax.: Barker
kelets with >2 flowers. Glumes shorter than the (1993), Davidse (1988), Linder and Ellis (1990b).
Prionanthium Desv., Opusc. 64 (1831).
flowers. Callus of flower pubescent. Lemma with Pentaschistis (Nees) Spach., Hist. Nat. Veg. Phan. 13: 164
three tufts of trichomes on either side of the mid- (1846).
rib, lobes often ending in thin setae; column of Poagrostis Stapf., Fl. Cap. 7: 760 (1899).
awn indistinct.
Nine spp., mountainous parts of Africa and Annuals or perennials, caespitose, rhizomatous
Madagascar. M. stereophylla lacks the synapo- or stoloniferous. Multicellular glands often on
GENERA OF DANTHONIOIDEAE 349
to Pentameris s.s., sect. Dracomontanum, which

includes two species formerly assigned to “Pen-
taschistis”, and sect. Pentaschistis, which includes
species formerly assigned to “Pentaschistis” and
Prionanthium. The former genus Prionanthium is
distinctive in that it includes annual species with
multicellular glands on the leaves. However, it is
clearly derived from within Pentameris and is oth-
erwise morphologically similar.
542. Chionochloa Zotov
Chionochloa Zotov, New Zealand J. Bot. 1: 87 (1963); tax.:

Connor (1991).
Caespitose perennials, generally over 1 m tall.

Leaf blades with deep grooves between the
veins, disarticulating at or below the ligule. Inflo-
rescence branches branched. Spikelets with more
than 2 flowers. Glumes shorter or longer than the
flowers. Callus of flower pubescent. Lemma
pubescent with trichomes generally in four verti-
cal lines, the lobes acute, terminating in setae, the
awn base distinct or not. Palea margins with hair
tufts. Leaf epidermal microhairs at the bases of
grooves; silica cells round. 2n ¼ 42.
Twenty-five spp. New Zealand, SE Australia,
Lord Howe Island.
Fig. 81. Danthonioideae. Pentameris alticola. A Several
tillers, representing growth form of plant. B Junction of
leaf sheath and blade. C Detail of cushion-based hairs on CHAETOBROMUS + PSEUDOPENTAMERIS CLADE
blades. D Part of inflorescence showing nodes and spike- Callus of flowers elongate, almost twice as
lets. E Flowers. F Glumes. G Lemma. H Upper palea with long as the rachilla internode, acute, sharp-
rachilla extension. I Lodicule. J Pistil. (From Linder and
Ellis 1990b) pointed. Palea strongly sulcate so that the veins
nearly touch, the veins not reaching the apex.
leaves, inflorescences and glumes. Spikelets with 543. Chaetobromus Nees

2 flowers (rarely 1). Glumes extending beyond the
flowers. Callus of flowers pubescent. Lemma with Chaetobromus Nees in Lindley, Nat. Syst., ed. 2: 449
setae from the sinuses between the lateral lobes (1836).
and the central awn, awn with a flat twisted base.
Palea acute, the keels not reaching the apex. Peri- Perennials, caespitose or mat-forming, sometimes
carp sometimes free from the seed coat. 2n ¼ 14, rhizomatous. Inflorescence branches branched.
24, 26, 28, 40, 42, 52, 56, 91. Spikelets with a tuft of hairs at the point of dehis-
Eighty-four spp. Africa, Madagascar, intro- cence below the glumes, with more than 2 flowers,
duced to Australia. the most proximal flower smaller and less well
Linder et al. (2010) find that Pentameris s.s. developed than the others. Glumes longer than
and Prionanthium are both derived within a para- the flowers. Callus of flower pubescent. Lemma
phyletic “Pentaschistis”. Accordingly, they have pubescent, the lobes ending in setae, the central
combined the three genera into one, with the awn with a twisted base. 2n ¼ 12, 18, 36, 48, 52, 72.
name Pentameris taking priority. Three sections One sp., C. involucratus (Schrad.) Nees, SW
are recognized, sect. Pentameris, corresponding Africa.
544. Pseudopentameris Conert placed in Cortaderia were first recognized as a

separate clade by Barker et al. (2003).
Pseudopentameris Conert, Mitt. Bot. Staatss. M€
unch. 10:
303 (1971); tax.: Barker (1995).
546. Chimaerochloa H. P. Linder
Perennials, caespitose or rhizomatous. Inflores-
Chimaerochloa H. P. Linder, Ann. Missouri Bot. Gard. 97:
cence branches branched. Spikelets with 2 flowers, 346 (2010).
2.5–3.5 cm long. Glumes longer than the flowers.
Callus of flower pubescent. Lemma pubescent, Gynodioecious perennials, caespitose. Leaf blades
the lobes acute and generally shorter than the tough, fibrous, disarticulating from the sheath.
lemma itself; basal column of the awn clearly Inflorescence branches branched. Spikelets with
distinct, twisted. 2 to 7 flowers. Glumes shorter than the flowers.
Three spp., Cape region, South Africa. Callus of flower pubescent. Lemma pubescent,
the lobes acute, ending in setae, the awn with a
Remaining genera (Danthonia clade + Rytidos-
flat twisted base. 2n ¼ 72.
perma clade): Lodicules bearing microhairs.
One sp., C. archboldii H. P. Linder, New
D A N T H O N I A C L A D E (A U S T R O D E R I A + C O R T A D E R I A + Guinea.
DANTHONIA + NOTOCHLOE + PLINTHANTHESIS + This species is morphologically similar to
CHIMAEROCHLOA) Cortaderia, but chloroplast sequence data place
it in the Danthonia clade. It is possible that it is an
Linder et al. (2010) identify three possible
allopolyploid derivative of a Cortaderia species
synapomorphies for this clade: a gynodioecious
crossed with a Danthonia species (the latter as
breeding system, lemmas with five veins or fewer,
seed parent); data from a nuclear gene would be
and the lemma awn not differentiated into a flat-
required to test this hypothesis.
tened basal portion and a slender apical portion.
All three characters reverse within the clade, how-
ever, and are thus not useful for identification. 547. Cortaderia Stapf
545. Austroderia N. P. Barker & H. P. Linder Cortaderia Stapf, Gard. Chron. III, 22: 396 (1897).
Lamprothyrsus Pilger, Bot. Jahrb. 37, Beibl. 85: 58 (1906).
Austroderia N. P. Barker & H. P. Linder, Ann. Missouri
Bot. Gard. 97: 343 (2010). Caespitose perennials, gynoedioecious or pistil-
late only, generally several meters tall. Leaf blades
Gynodioecious perennials, caespitose or stolonif- with prominent midribs, leaf sheaths persistent
erous, generally several meters tall. Leaf blades but tending to fragment with age. Inflorescences
with several wide prominent veins; leaf sheaths plumose, the branches branched. Spikelets with
persistent, waxy, not fragmenting. Inflorescences more than 2 flowers. Glumes shorter or longer
plumose, the branches branched. Spikelets with than the flowers, with one vein. Callus of flower
more than 2 flowers. Glumes shorter or longer pubescent. Lemma pilose, the lobes absent or
than the flowers, with one vein. Callus of flower nearly so, although long setae sometimes present;
pubescent. Lemma pilose, the lobes absent or awn not clearly differentiated into a flattened
nearly so, although long setae sometimes present base and a slender apical portion. 2n ¼ 36, 72,
in place of the lobes; the base of the awn not 108, 136.
differentiated. 2n ¼ 90. Twenty spp., South America.
Five spp., New Zealand. Linder et al. (2010) describe the margins and
Austroderia corresponds to a set of species sometimes the entire distal part of the leaves as
formerly placed in Cortaderia. The two genera are “viciously scabrid”. C. pilosa may not be related
quite similar in terms of gross morphology, but to the other species of the genus, but is retained
differ in leaf anatomy as well as in geographic here on the basis of ambiguous molecular data as
distribution. The New Zealand species formerly described by Linder et al. (2010).
GENERA OF DANTHONIOIDEAE 351
548. Danthonia DC lobes, awn the same length as the lobes or longer.
Lodicules without microhairs.
Danthonia DC in Lam. & DC, Fl. Franç. ed. 3, 3: 32 (1805), Three spp., eastern Australia.
nom. cons.
Sieglingia Bernh., Syst. Verz. (Bernhardi): 20, 44 (1800).
R Y T I D O S P E R M A C L A D E (R Y T I D O S P E R M A + S C H I S -
MUS + TENAXIA + TRIBOLIUM)
Caespitose perennials, sometimes with cleistoga-
Lemma hairs often in scattered tufts. Lodi-
mous spikelets enclosed in the upper leaf sheaths.
cules with microhairs. Hilum punctate-ovate.
Leaf blades not disarticulating, membranous.
Inflorescence unbranched, or with branched
branches. Spikelets with 2 to 10 flowers. Callus of 551. Tenaxia N. P. Barker & H. P. Linder
flowers pubescent. Glumes shorter or longer than
the flowers. Lemma pubescent, at least on the mar- Tenaxia N. P. Barker & H. P. Linder, Ann. Missouri Bot.
Gard. 97: 350 (2010).
gins, with 7 to 9 veins, the lobes prominent, ending
in setae, the awn base flat, twisted, clearly distinct
from the apical portion. 2n ¼ 18, 24, 36, 48, 72, 98. Caespitose perennials with wiry stems. Leaves
Twenty-five spp. America and Europe. tough, sclerenchymatous, the sheaths persistent.
No morphological character can be used con- Inflorescence unbranched or with branched
sistently to recognize Danthonia; the four syna- branches. Spikelets with 2 to 7 flowers. Glumes
pomorphies indicated by Linder et al. (2010) as long as the flowers or longer. Callus of flower
(bisexual flowers, persistent leaf blades, 7 or pubescent. Lemma pubescent, with 7 to 9 veins,
more lemma veins, and orthophyllous leaves) the lobes with acute apices, terminating in setae or
are all homoplasious on the phylogeny. not, the awn with a twisted base. 2n ¼ 12, 36, 56.
Eight spp. Africa, India, Himalayas.
NOTOCHLOE + PLINTHANTHESIS CLADE Members of Tenaxia have been previously
Caespitose perennials, less than 1 m tall. Leaf assigned to either Merxmuellera or Danthonia.
sheaths persistent, shiny, white. Inflorescence Tenaxia has no obvious morphological synapo-
branches branched. Callus of flower pubescent, morphy, but forms a distinct and well-supported
short; rachilla with a horizontal disarticulation clade in molecular phylogenies. Phylogeography
zone surrounded by hairs. Leaf mesophyll with of T. disticha in South Africa has been investi-
empty spaces, formed “from large, empty cells gated by Linder et al. (2014).
that then disintegrate, leaving rather ragged cav-
ities” (Linder et al. 2010, p. 346). Fruit turbinate. SCHISMUS + TRIBOLIUM + RYTIDOSPERMA
Leaf blades membranous, not stiff and sceler-
enchymatous.
549. Notochloe Domin
552. Rytidosperma Steud.
Notochloe Domin, Feddes Repert. 10: 117 (1911).
Rytidosperma Steud., Syn. Pl. Glum. 1: 425 (1854).
Spikelets with 7 to 9 flowers. Glumes shorter than Monostachya Merr., Philipp. J. Sci. 5: 330 (1910).
the flowers. Lemma glabrous, with three small Notodanthonia Zotov, New Zealand J. Bot. 1: 104 (1963).
Erythranthera Zotov, New Zealand J. Bot. 1: 124 (1963).
apical teeth, awnless. Pyrrhanthera Zotov, New Zealand J. Bot. 1: 125 (1963).
One sp., N. microdon (Benth.) Domin, eastern Joycea H. P. Linder, Telopea 6: 611 (1996).
Australia. Austrodanthonia H. P. Linder, Telopea 7: 269 (1997).
550. Plinthanthesis Steud. Caespitose perennials. Inflorescence branches

branched or unbranched. Spikelets with 2 to 7
Plinthanthesis Steud., Syn. Pl. Glum. 1: 14 (1854). flowers, the callus wider than the internode that
bears it. Glumes shorter or more often longer
Inflorescences plumose. Spikelets with 2 to 4 than the flowers. Lemma with the central lobe
flowers. Glumes shorter or longer than the flow- ending in a twisted awn; lemma hairs rarely lack-
ers. Lemma pubescent below, with two short ing, generally in two transverse rows across the
abaxial side, these in one line of discrete tufts sections has a unique character. Plants in
about midway up the lemma and additional mar- Tribolium sect. Tribolium have hispid glumes,
ginal tufts somewhat lower. those in T. sect. Acutiflorae N. C. Visser &
Seventy-six spp., Australasia and South Spies ex H. P. Linder & Davidse bear stolons,
America. and those in T. sect. Uniolae N. C. Visser &
The circumscription of this genus was Spies ex H. P. Linder & Davidse have unbranched
addressed by Humphreys et al. (2010), who inflorescences with the spikelets all borne on one
show that it is monophyletic in the original side.
sense of Steudel (1853-1854).
553. Schismus P. Beauv. References
Schismus P. Beauv., Ess. Agrost.: 73 (1812). Barker, N.P. 1993. A biosystematic study of Pentameris
(Arundineae, Poaceae). Bothalia 23: 25-47.
Barker, N.P. 1995. A systematic study of the genus Pseu-
Annuals or perennials, caespitose, less than 35 cm dopentameris (Arundinoideae: Poaceae). Bothalia
tall. Inflorescence branches branched. Spikelets 25: 141-148.
with 3 to 7 flowers. Glumes shorter than the Barker, N.P., Linder, H.P., Morton, C.M., Lyle, M. 2003.
spikelet (rarely longer). Callus of flower pubes- The paraphyly of Cortaderia (Danthonioideae):
evidence from morphology and nuclear DNA
cent. Lemma with hairs scattered, or tufted, or in sequence data. Ann. Missouri Bot. Gard. 90: 1-24.
lines, with mucronate lobes, the central one gen- Connor, H.E. 1991. Chionochloa Zotov (Gramineae) in
erally shorter than the laterals. 2n ¼ 12, 24, 36, New Zealand. N. Z. J. Bot. 29: 219-282.
48, 72. Davidse, G. 1988. A revision of the genus Prionanthium
(Poaceae: Arundinae). Bothalia 18: 143-153.
Five spp., Africa. GPWG II 2012. Grass Phylogeny Working Group II. New
In Schismus schismoides (Stapf ex Conert) grass phylogeny resolves deep evolutionary relation-
Verboom and H. P. Linder, the central lobe ships and discovers C4 origins. New Phytol. 193:
forms a twisted awn and is thus longer than the 304-312.
Humphreys, A.M., Pirie, M.D., Linder, H.P. 2010. A plas-
laterals; this awn morphology is not shared with tid tree can bring order to the chaotic generic taxon-
other species in the genus. omy of Rytidosperma Steud. s.l. (Poaceae). Mol. Phyl.
Evol. 55: 911-928.
Linder, H.P., Ellis, R.P. 1990a. Vegetative morphology and
554. Tribolium Desv. interfire survival strategies in the Cape fynbos
grasses. Bothalia 20: 91-103.
Tribolium Desv., Opusc.: 64 (1831). Linder, H.P., Ellis, R.P. 1990b. A revision of Pentaschistis
Urochlaena Nees, Fl. Afr. Austr.: 437 (1841). (Arundineae: Poaceae). Contr. Bolus Herb. 12:
Karroochloa Conert & T€ urpe, Senck. Bot. 50: 290 (1969). 1-124.
Linder, H.P., Baeza, M., Barker, N.P., Galley, C., Hum-
phrey, A.M., Lloyd, K.M., Orlovich, D.A., Pirie, M.D.,
Annuals or perennials, caespitose and stolonifer- Simon, B.K., Walsh, N., Verboom, G.A. 2010. A
ous. Inflorescence unbranched, or with branched generic classification of the Danthonioideae. Ann.
Missouri Bot. Gard. 97: 306-364.
branches, generally narrow and spikelike or capi- Linder, H.P., Perl, F., Bouchenak-Khelladi, Y., Barker, N.
tate. Spikelets with 2 to 10 flowers. Glumes P. 2014. Phylogeographical pattern in the southern
shorter or longer than the flowers, glabrous or African grass Tenaxia disticha (Poaceae). Syst. Bot.
with hairs with tuberculate bases. Callus of flower 39: 428-440.
Peterson, P.M., Romaschenko, K., Johnson, G. 2010. A
pubescent. Lemma pubescent, the hairs either classification of the Chloridoideae (Poaceae) based
evenly spread or tufted in various patterns, the on multi-gene phylogenetic trees. Mol. Phyl. Evol. 55:
apex entire and acute or with three lobes, the 580-598.
lobes rounded, acute or acuminate, the central Steudel, E.G. 1853-1854. Synopsis Plantarum Gluma-
cearum. Stuttgart: J. B. Metzler.
one occasionally forming an awn. 2n ¼ 12, 24, 36. Verboom, G.A., Linder, H.P., Barker, N.P. 1994. Hausto-
Sixteen spp., southern Africa. rial synergids: An important character in the system-
The genus as a whole lacks a clear morpho- atics of danthonioid grasses (Arundinoideae:
logical synapomorphy, although each of the three Poaceae). Amer. J. Bot. 81: 1601-1610.
X. Subfamily Chloridoideae Kunth ex Beilschm. (1833)
Plants herbaceous. Annuals or perennials, caespi- that the presence of an epiblast might be a synap-
tose, rhizomatous or stoloniferous, erect or omorphy but noted that the character is highly
decumbent. Culms hollow or solid. Ligule mem- homoplasious in the subfamily.
branous, or a fringed membrane, or a fringe of Peterson et al. (2010a) have produced a com-
hairs. Leaf blades lacking pseudopetioles or auri- prehensive molecular phylogeny for the subfam-
cles. Inflorescences unbranched, or with ily, including 254 species in 99 genera. They
unbranched primary branches, or with branched delimit four tribes: Triraphideae, Eragrostideae,
primary branches. Spikelets laterally compressed, Zoysieae, and Cynodonteae, and I follow their
with 1 to many flowers, the proximal ones fertile, treatment here. More recently they suggest add-
the distal ones (if present) often reduced. Glumes ing Ellisochloa (formerly Merxmuellera rangei
2, disarticulation above the glumes. Lemmas and M. papposa) and the four species of Centro-
bilobed, awned from the sinus. Lodicules 2, podia as Centropodieae (Peterson et al. 2011).
fleshy. Caryopsis with the pericarp fused to the Tribes in 20th century classifications of the sub-
seed coat; hilum short, punctate; embryo with an family were distinguished by inflorescence char-
epiblast, with a scutellar cleft, with an elongated acters, with the primary branches of the
mesocotyl internode, with the embryonic leaf inflorescence themselves bearing branches in
margins not overlapping. Midrib anatomy sim- most Eragrostideae, and the primary branches
ple. Mesophyll radiate, with a double bundle unbranched in Cynodonteae. Phylogenetic data
sheath. All species C4 except for Merxmuellera show that the transition from one type of inflo-
rangei, the ancestral condition probably C3. Bicel- rescence to the other, which involves a change in
lular microhairs with the distal cell narrow and the numbers of orders of branches, is quite labile
elongate (“panicoid type”) to nearly spherical in evolutionary time. The most striking example
(“chloridoid type”). of this is in the Sporobolus clade, in which the
One hundred thirty-two genera, 1721 spp. former genus Spartina, with sturdy, unbranched
This subfamily has been shown consistently primary branches, is apparently derived from
to be monophyletic, with all species except Merx- within Sporobolus, with a highly branched and
muellera rangei sharing radiate chlorenchyma delicate inflorescence (Peterson et al. 2014b).
around the veins and C4 photosynthesis. Mem- Generic limits in Chloridoideae are often
bers of the subfamily have been biochemically problematical. The characters used to define
typed as having either the NAD-ME or PCK subgenera are mostly characters of the inflorescence
type of C4 photosynthesis (Brown 1977; Gutierrez and spikelet, but because these are evolutionarily
et al. 1974; Hattersley and Watson 1992). The labile they often do not indicate relationship.
presence of so-called chloridoid microhairs, in Phylogenetic studies such as those by Columbus
which the apical cell of the microhair is broad et al. (2007, 2010) and Peterson et al. (2010a,
and rounded, may also be synapomorphic but the 2010b, 2014a) indicate that several of the major
character is somewhat variable (Van den Borre genera are polyphyletic; these include “Chloris”
and Watson 1997). The GPWG (2001) suggested (with 63 species). Until these are dismembered,
354 X. Subfamily Chloridoideae Kunth ex Beilschm. (1833)
monophyletic groups (at any taxonomic level) 2 (1). Glumes lacking; lemma flabellate 623. Neostapfia
cannot be delimited by morphological data. Lep- 2. Glumes present; lemma oblong 3
3 (2). Lemma with five teeth, these often ending in awns
tochloa, known for some time to be polyphyletic, 624. Orcuttia
has recently been divided and its components 3. Lemma entire or erose, awns lacking 625. Tuctoria
reclassified (Peterson et al. 2012). Tridens has 4 (1). Spikelets embedded in the rachis 5
likewise recently been divided into Tridens and 4. Spikelets sessile or pedicellate, but not embedded in the
Tridentopsis (Peterson et al. 2014). Surprisingly, rachis 7
5 (4). Lower glume present, at least 1/3 the length of the
the large genus Eragrostis appears to be spikelet; upper glume attenuate, awned
monophyletic if several monotypic genera are 561. Lepturopetium
subsumed in it (Ingram and Doyle 2004). 5. The lower glume minute or absent; upper glume flat-
Muhlenbergia, Bouteloua, and Sporobolus have tened and extended into an attenuate awn-like
also been re-circumscribed and each is now appendage 6
6 (5). Glume with many veins; leaves lacking glands
monophyletic (Columbus 1999; Peterson et al. 662. Lepturus
2010b; Peterson et al. 2014b). 6. Glume with one to three veins; leaves with multicellular
glands abaxially 679. Oropetium
7 (4). Spikelets with awns on the glumes or lemmas or
TRIBES AND SUBTRIBES OF CHLORIDOIDEAE both 8
7. Spikelets lacking awns 27
XXV. Tribe Centropodieae 8 (7). Glumes awned or long acuminate 9
Core C4 Chloridoideae, incertae sedis 8. Glumes awnless 22
XXVI. Tribe Triraphideae 9 (8). Lemmas awnless 10
XXVII. Tribe Eragrostideae 9. Lemmas awned 12
Incertae sedis 10 (9). Spikelets falling with their pedicels; leaf blade apex
XXVII.1. Subtribe Cotteinae not muticous or pungent 557. Decaryella
XXVII.2. Subtribe Eragrostidinae 10. Spikelets falling entire but pedicels persistent; leaf
XXVII.3. Subtribe Uniolinae blade apex muticous to pungent 11
XXVIII. Tribe Zoysieae 11 (10). Lower glume absent 590. Zoysia
XXIX. Tribe Cynodonteae 11. Lower glume present 642. Perotis
Incertae sedis 12 (9). Awns twisted or flexuous 13
XXIX.1. Subtribe Boutelouinae 12. Awns straight 16
XXIX.2. Subtribe Orcuttiinae 13 (12). Inflorescence axis forming an arc; second glume
XXIX.3. Subtribe Gouiniinae with a divergent abaxial awn 630. Ctenium
XXIX.4. Subtribe Triodiinae 13. Inflorescence axis straight; second glume apex attenu-
XXIX.5. Subtribe Eleusininae ate or awned from the apex or from a sinus, the awn
XXIX.6. Subtribe Tripogoninae not divergent 14
XXIX.7. Subtribe Pappophorinae 14 (13). Leaf blade apex sharp-pointed 680. Tripogon
14. Leaf blade apex blunt 15
15 (14). Plants annual; ligule membranous, not fringed
KEY TO THE GENERA OF CHLORIDOIDEAE 559. Indopoa
15. Plants perennial; ligule a fringed membrane
647. Afrotrichloris
1. Inflorescence unbranched, with spikelets sessile or 16 (12). Ligule membranous 17
pedicellate, or if branched then dense and spike-like 16. Ligule a fringe of hairs or prominently fringed
Group I membrane 18
1. Inflorescence clearly branched, the primary branches 17 (16). Spikelets paired, pedicellate; inflorescence
branched, or unbranched and digitate or spread along dense and spike-like, the branches terminating in a
the main axis, not dense and spike-like 2 spikelet
2 (1). Primary branches of the inflorescence unbranched, 612. Muhlenbergia (former Lycurus)
digitate or spread along the main axis Group II 17. Spikelets solitary, sessile; inflorescence unbranched,
2. Primary branches of the inflorescence branched, at terminating in a barren tip 560. Kampochloa
least some spikelets borne on secondary or higher- 18 (16). Lemma broadest above the midpoint
order branches Group III 681. Neesiochloa
18. Lemma broadest at or about the midpoint 19
Group I – Inflorescences unbranched, with 19 (18). Lemma apex two lobed, awned from the sinus and
spikelets sessile or pedicellate; spike-like often from the lobes as well
panicles may also key here 608. Bouteloua (subg. Chondrosum)
19. Lemma apex entire 20
1. Ligules absent 2 20 (19). Glumes longer than the spikelet, borne on the
1. Ligules present 4 same side; spikelets paired 573. Tetrachaete
KEY TO THE GENERA OF CHLORIDOIDEAE 355
20. Glumes shorter than the spikelet, distichous; spikelets 37 (36). Inflorescence axis arcuate, ending in a barren tip
solitary 21 660. Harpochloa
21 (20). Annual; callus of spikelet sharp pointed; each 37. Inflorescence axis straight, ending in a spikelet 38
spikelet subtended by a hyaline bract 38 (37). Ligule membranous or a fringed membrane;
568. Viguierella inflorescence surrounded by a leaf sheath or spatheole
21. Perennial; callus of spikelet obtuse; spikelets without 650. Brachyachne
subtending bracts 583. Fingerhuthia 38. Ligule a fringe of hairs; inflorescence not surrounded
22 (8). Ligule membranous, not ciliolate; lemma with by a sheath 599. Lepturidium
three awns; flowers unisexual 608. Bouteloua 39 (27). Leaves stiff, divergent; plants perennial, strongly
22. Ligule a fringe of hairs or a fringed membrane; lemma rhizomatous and often also stoloniferous 40
with one awn; flowers bisexual 23 39. Leaves lax; plants annual or perennial, caespitose
23 (22). Plants stoloniferous, less than 15 cm tall 24 44
23. Plant caespitose, rhizomes short or absent, stolons 40 (39). Spikelets unisexual, plant dioecious 41
absent, or if present then plants more than 15 cm 40. Spikelets bisexual 42
25 41 (40). Female lemma tubular, fused to rachilla; glumes
24 (25). Inflorescence borne above the leaves; spikelets absent in female spikelet 611. Jouvea
turbinate; sterile flowers distal and proximal in the 41. Female lemma folded, not fused to rachilla; glumes
spikelet 602. Pommereulla present in female spikelet 609. Distichlis
24. Inflorescence surrounded by leaves, surpassed by the 42 (40). Inflorescence axis arcuate, ending in a barren tip
uppermost leaf blades; spikelets ovate to lanceolate; 660. Harpochloa
sterile flowers distal only 621. Munroa 42. Inflorescence axis straight, ending in a spikelet 43
25 (23). Spikelets three per node; plants of North and 43 (42). Lemmas with 5 or more veins 596. Aeluropus
Central America 610. Hilaria 43. Lemmas with 1–3 veins 588. Psilolemma
25. Spikelets one per node; plants of the Old World 26 44 (39). Leaves slender and thread-like; lemmas decidu-
26 (25). Lemma with three deep lobes 648. Astrebla ous, paleas and rachilla persistent
26. Lemma entire or with two shallow teeth 677. Eragrostiella
670. Tetrapogon 44. Leaves not thread-like; lemmas persistent, falling with
27 (7). Spikelets with one flower 28 the spikelet
27. Spikelets with more than one flower 39 579. Eragrostis (former Psammagrostis, Harpachne)
28 (27). Spikelets covered with stiff hooked prickles
615. Tragus
28. Spikelets variously smooth or pubescent; if bearing
prickles, then these not hooked 29 Group II – Inflorescence with two or more
29 (28). Glumes shorter than the lemmas 30 unbranched branches, either digitate or spread
29. Glumes as long as or longer than the lemmas, enclos-
ing the flower 32
along the main axis
30 (29). Plants annual; Australia 580. Heterachne 1. Primary branches digitate 2
30. Plants perennial; Africa 31 1. Primary branches spread along main inflorescence axis
31 (30). Leaves spine-tipped; glumes and lemmas gla- 26
brous; plants growing in sand 589. Urochondra 2 (1). Spikelets without awns 3
31. Leaves slender but not spine-tipped; glumes and 2. Spikelets with awns 10
lemmas with long hairs; plants growing in damp sites 3 (2). Leaf blades cylindrical in cross section, with sharp
584. Stiburus tips 669. Sclerodactylon
32 (29). Glumes hard, ending in a sharp point, wooly to 3. Leaf blades flattened, sometimes folded, their tips
scabrous 634. Dignathia sometimes acute but not sharp 4
32. Glumes membranaceous to coriaceous, not sharp- 4 (3). Culms with tuberous bases, the leaf sheaths densely
pointed, glabrous 33 wooly below; primary inflorescence branches disarti-
33 (32). Inflorescence with spikelets borne on all sides culating at maturity 564. Ochthochloa
34 4. Culm bases not tuberous, leaf sheaths pubescent or
33. Inflorescence with spikelets borne on one side only not, but not wooly; primary inflorescence branches
36 persistent 5
34 (33). Lower glume absent, the upper glume enclosing 5 (4). Leaf blades cordate at the base, with multicellular
the flower 590. Zoysia glands 673. Coelachyropsis
34. Both glumes present 35 5. Leaf blades narrow at the base, not cordate, lacking
35 (34). Plants perennial, rhizomatous or stoloniferous; multicellular glands 6
ligule a fringed membrane 641. Mosdenia 6 (5). Glumes longer than the flowers and enclosing them
35. Plants annual, caespitose; ligule membranous 7
642. Perotis 6. Glumes shorter than to as long as the flowers, not
36 (33). Spikelets dorsiventrally compressed enclosing them 8
664. Microchloa 7 (6). Spikelets dorsiventrally compressed, with one
36. Spikelets laterally compressed 37 flower; plants of Africa 632. Craspedorhachis
7. Spikelets laterally compressed, with 3 flowers (2 fertile); 25 (23). Spikelets dorsiventrally compressed
plants of Brazil 666. Rheochloa 658. “Enteropogon”
8 (6). Spikelets with one flower 653. Cynodon 25. Spikelets laterally compressed 668. Schoenefeldia
8. Spikelets with more than one flower 9 26 (1). Primary inflorescence branches stiff, divaricate,
9 (8). Hairs at apex of lemma slightly inflated at the tip ending in sharp point; plants forming spiny mounds,
657. Disakisperma shrubby 572. Cladoraphis
9. Hairs at apex of lemma not inflated 675. Eleusine 26. Primary inflorescence branches lax or not; if divari-
10 (2). Awns on lemmas only; glumes lacking awns 11 cate, then not ending a sharp point; plants not forming
10. Awns on glumes and lemmas 20 spiny mounds, not shrubby 27
11 (10). Lemmas of the distal flowers of the spikelet 27 (26). Spikelets without awns 28
broadly flaring, fan-shaped above 665. Oxychloris 27. Spikelets with awns 52
11. Lemmas of the distal flowers of the spikelet reduced, 28 (27). Lateral branches bearing no more than two spike-
not flaring or fan-shaped 12 lets; glumes longer than flowers, indurate 29
12 (11). Lemma divided about ¼ of its length into two 28. Lateral branches with four or more spikelets; glumes
large lobes, with prominent lateral veins extending shorter or longer than the flowers, coriaceous or mem-
into the lobes 663. Lintonia branous, not indurate 31
12. Lemma apex entire or bidentate, if the latter then the 29 (28). Inflorescence borne above the distalmost leaf, not
lobes less than ¼ the length of the lemma 13 surrounded by a sheath; axis of lateral branches flat-
13 (12). Lower glume tiny, less than 1/6 the length of the tened, broader distally than proximally
upper glume 667. Saugetia 635. Leptothrium
13. Lower glume well developed 14 29. Inflorescence surrounded by a leaf sheath, this often
14 (13). Glumes membranous to leathery, lemmas less inflated; axis of lateral branches terete 30
firm than the glumes, hyaline or thinly membranous 30 (29). Lower glume well developed; upper glume tuber-
639. Lophacme culate 566. Pseudozoysia
14. Glumes hyaline to membranous, less firm than the 30. Lower glume less than 0.2 the length of the upper
membranous, leathery or indurate lemmas 15 glume or absent; upper glume smooth or covered
15 (14). Rachilla extension naked 654. Daknopholis with hooked spines but not tuberculate
15. Rachilla extension bearing rudimentary flower(s) 16 615. Tragus
16 (15). Spikelet with 2 to 5 fertile flowers 17 31 (28). Spikelets partially embedded in the lateral
16. Spikelet with 1 fertile flower 18 branches, scarcely overlapping; the inflorescence dis-
17 (16). Lemma with hyaline wings 563. Neostapfiella articulating at the base and forming a tumbleweed
17. Lemma without wings 670. Tetrapogon dispersal unit
18 (16). Spikelet lanceolate, elliptical, or ovate 612. Muhlenbergia (former Schedonnardus)
658. “Enteropogon” 31. Spikelets not embedded in the lateral branches, often
18. Spikelet cuneate 19 overlapping at least half their length or more, often
19 (18). Spikelets dorsiventrally compressed densely packed; inflorescence axis not disarticulating
649. Austrochloris at the base 32
19. Spikelets laterally compressed 651. “Chloris” 32 (31). Glumes covered with stiff spines 615. Tragus
20 (10). Each inflorescence branch ending in a prominent 32. Glumes not covered with spines 33
sharp point; upper glume terminated by an awn bent 33 (32). First glume reduced to a short slender stub
at a sharp angle to the long axis of the glume 578. Catalepis
604. Dactyloctenium 33. First glume well developed 34
20. Racemes not ending in a sharp point; glume awn 34 (33). Spikelets disarticulating below the glumes and
straight or slightly curved, but not stiffly bent at a falling entire 35
sharp angle 21 34. Spikelets disarticulating above the glumes 38
21 (20). Upper glume with 5 to 7 veins 35 (34). Inflorescence branches generally shorter than the
561. Lepturopetium internodes 586. Tetrachne
21. Upper glume with 1 vein 22 35. Inflorescence branches generally longer than the
22 (21). Lemma with three awns from the tip internodes 36
671. Trichloris 36 (35). Glumes as long as or longer than the spikelet;
22. Lemma with a single awn 23 spikelet with one fertile flower
23 (22). Awn much shorter than the body of the lemma 593. Spartina (now part of Sporobolus s.l.)
24 36. Glumes shorter than the spikelet; spikelet with 2 or
23. Awn much longer than the body of the lemma 25 more fertile flowers 37
24 (23). Spikelets disarticulating between the glumes, the 37. Lemma cartilaginous or leathery; lower flowers of the
upper deciduous, the lower persistent; inflorescence spikelet sterile 587. Uniola
branches 2 to 6 652. Chrysochloa 37. Lemma membranous; lower flowers of the spikelet
24. Spikelets disarticulating below both glumes; inflores- fertile 676. Desmostachya
cence with as many as 40 branches 38 (34). Flowers unisexual 39
659. “Eustachys” 38. Flowers bisexual 41
KEY TO THE GENERA OF CHLORIDOIDEAE 357
39 (38). Spikelets with 1 fertile flower and 2 to 3 sterile 56 (55). Lemma awnless; awns on glumes only 57
ones 608. Bouteloua (former Pringleochloa) 56. Lemma with one or more awns 58
39. Spikelets with 4 or more fertile flowers 40 57 (56). Spikelets with 1 flower; glumes as long as or
40 (39). Palea as long as or longer than the lemma longer than the flower 593. Spartina (now part of
607. Allolepis Sporobolus s.l.)
40. Palea shorter than the lemma 619. Scleropogon 57. Spikelets with 4 or more flowers; glumes shorter than
41 (38). Callus of flower with dense straight hairs at least the flowers 674. Coelachyrum
¼ as long as the flower; leaf blades disarticulating from 58 (56). Lemma with multiple awns 59
the sheath 42 58. Lemma with one awn 64
41. Callus of flower glabrous, or pubescent but with hairs 59 (58). Awns much longer than spikelets, over 5 cm long
less than ¼ the length of the flower; leaf blades not 619. Scleropogon (pistillate inflorescences)
disarticulating 43 59. Awns about the same length as or shorter than spike-
42 (41). Spikelet less than 1 cm long 606. Odyssea lets, ca. 5 mm long or less 60
42. Spikelet 1–2 cm long 558. Halopyrum 60 (59). All flowers unisexual 608. Bouteloua
43 (41). Spikelets with one flower 44 60. At least some flowers bisexual 61
43. Spikelets with more than one flower 45 61 (60). Glumes awnless, although sometimes setaceous
44 (43). Ligule a fringed membrane; inflorescence con- 62
tracted, the branches upright 616. Polevansia 61. Glumes awned 63
44. Ligule membranous; inflorescence open, the branches 62 (61). Spikelets with 2 proximal sterile flowers
spreading 640. Trigonochloa 618. Blepharidachne
45 (43). Fruit flattened, concavo-convex, the surface 62. Spikelets with 2 proximal fertile flowers
rough 674. Coelachyrum 605. Neobouteloua
45. Fruit not flattened, the surface smooth 46 63 (61). Glumes glabrous, hyaline except for the midvein
46 (45). Upper glume and lemmas with 7 or 9 veins; palea and awn 612. Muhlenbergia
splitting at maturity 629. Vaseyochloa 63. Glumes with long hairs, coriaceous
46. Upper glume and lemmas with fewer than 7 veins; 678. Melanocenchris
palea not splitting 47 64 (58). Glumes longer than the spikelet, enclosing the
47 (46). Palea bowed outward away from flower; second flowers 65
glume with 5 veins 603. Brachychloa 64. One or both glumes shorter than the spikelet, not
47. Palea flat or curved inward toward flower; second enclosing the flowers 71
glume with 1 to 3 veins 48 65 (64). Plants less than 15 cm tall, growing in tufts
48 (47). Lemma covered with dense long hairs connected by stolons; inflorescence dense, capitate
600. Orinus 621. Munroa
48. Lemma with hairs only on the veins, or glabrous 49 65. Plants over 15 cm, not stoloniferous; inflorescence not
49 (48). Ligule apex acute to attenuate 656. Diplachne capitate 66
49. Ligule apex obtuse to truncate 50 66 (65). Spikelets falling entire along with their pedicels;
50 (49). Apex of lemma hairs slightly inflated at the tip glumes much more firm than the lemmas, covered
657. Disakisperma with long hairs or prickles 634. Dignathia
50. Apex of lemma hairs not inflated 51 66. Spikelets disarticulating above one or both glumes,
51 (50). Primary inflorescence branches two or more per pedicels and branches persistent; glumes less firm than
node 661. Leptochloa the lemmas, not pubescent or prickly 67
51. Primary inflorescence branches one per node 67 (66). Spikelets terete in cross section; leaf blades with
655. Dinebra multicellular glands 601. Pogoneura
52 (27). Primary inflorescence branches ending in a sterile 67. Spikelets laterally compressed; leaf blades lacking
point 53 glands 68
52. Primary inflorescence branches ending in a spikelet 68 (67). Spikelets not or scarcely overlapping on the
55 inflorescence branches, shorter than the internodes
53 (52). Leaves with abaxial glands; spikelet with 6 to 12 637. Gymnopogon
fertile flowers; Zambia 581. Richardsiella 68. Spikelets overlapping on the inflorescence branches,
53. Leaves without glands; spikelet with 1 fertile flower; longer than the internodes 69
New World 54 69 (68). Awn shorter than the lemma, generally less than
54 (53). Spikelets with a rachilla extension bearing 1 or 1 mm long 631. Trichoneura
more reduced flowers 608. Bouteloua 69. Awn the same length as or appreciably longer than the
54. Spikelets lacking a rachilla extension 593. Spartina lemma 70
(now part of Sporobolus s.l.) 70 (69). Spikelets disarticulating above the glumes; lemma
55 (52). Each primary inflorescence branch bearing one awn (3) 5–15 mm long 626. Gouinia
fertile spikelet plus two spikelets reduced to awns; 70. Spikelets disarticulating above the first glume and
lemma of fertile spikelet minute, pubescent below the second; lemma awn 15–25 mm long
633. Farrago 565. Pogonochloa
55. Primary inflorescence branches with various numbers 71 (64). Ligule a fringe of hairs or a fringed membrane
of spikelets, but not as in Farrago; lemma of fertile 72
spikelet well developed 56 71. Ligule membranous, not fringed 76
72 (71). Spikelets with two to many flowers, laterally 6 (5). Glumes shorter than the flowers 643. Triodia
compressed; leaf blade margins membranous 73 6. Glumes longer than the flowers 555. Ellisochloa
72. Spikelets with one flower, dorsiventrally compressed; 7 (5). Lemmas awned; plants of Australasia
leaf blade margins cartilaginous 75 645. Symplectrodia
73 (72). Plants aquatic, often floating; spikelets borne on 7. Lemmas mucronate; plants of California
both sides of primary inflorescence branches 620. Swallenia
562. Myriostachya 8 (2). Spikelets without awns 9
73. Plants terrestrial; spikelets borne on one side of pri- 8. Spikelets with awns 23
mary inflorescence branches 74 9 (8). Spikelets with one flower 10
74 (73). Inflorescence branches deciduous 9. Spikelets with more than one flower 13
579. Eragrostis (former Pogonarthria) 10 (9). Plants strongly stoloniferous, low, spreading; inflo-
74. Inflorescence branches persistent 646. Acrachne rescence dense, capitate or spike-like, nearly spherical,
75 (72). Lower glume ½ to ¾ the length of the upper surrounded by upper leaves
glume; upper glume apex obtuse to acute, not awned 592. Crypsis (now part of Sporobolus s.l.)
617. Willkommia 10. Plants rhizomatous or caespitose, erect; inflorescence
75. Lower glume about 1/10 the length of the upper glume; narrow or spreading, but not capitate, borne well
upper glume apex acuminate, awned above the upper leaves 11
614. Monelytrum 11 (10). Callus of flower glabrous 594. Sporobolus
76 (71). Clusters of fertile spikelets subtended by involu- 11. Flower with a bearded callus 12
cral bristles 612. Muhlenbergia (former Pereilema) 12 (11). Inflorescence narrow and dense like a bottle-
76. Bristles lacking 77 brush, the branches short and upright, closely pressed
77 (76). Plants prostrate, stoloniferous; inflorescences to the main axis 584. Stiburus
dense, capitate; glumes leaf-like, with 9–11 veins 12. Inflorescence broad to narrow, the branches long, not
655. Dinebra tightly appressed to the main axis 591. Calamovilfa
77. Plants upright; inflorescences not capitate; glumes (now part of Sporobolus s.l.)
similar in texture and morphology to lemmas, with 13 (9). Ligule membranous 14
one vein 78 13. Ligule a fringe of hairs or a membrane with long cilia
78 (77). Plants annual 79 16
78. Plants perennial 80 14 (13). Spikelets with one or two flowers, the lemmas
79 (78). Inflorescence branches not densely crowded; glabrous 612. Muhlenbergia
lemmas deciduous, palea persistent on the rachilla 14. Spikelets with more than two flowers; lemmas gener-
646. Acrachne ally with short hairs on keel and margins 15
79. Inflorescence dense, the primary branches crowded; 15 (14). Paleas widened or bowed below; caryopses not
lemmas persistent on the rachilla, falling with the rest deeply concave 683. Tridens
of the flower 636. Leptocarydion 15. Paleas not widened or bowed below; caryopses deeply
80 (78). Spikelets with fewer than 4 flowers; lemma awned concave to folded on the side of the hilum
from the abaxial side 638. Bewsia 627. Tridentopsis
80. Spikelets with 6 or more flowers; lemma awned from 16 (13). Glumes with 5 or more veins 17
the apex 567. Silentvalleya 16. Glumes with 1 or 3 veins 18
17 (16). Flowers bisexual; palea splitting at maturity
629. Vaseyochloa
Group III – Inflorescence branches themselves 17. Flowers unisexual, the plants dioecious; palea not
branched, at least some spikelets borne on splitting 607. Allolepis
secondary or higher-order branches, the 18 (16). Lemmas papery, laterally flattened, broadly
spikelets mostly pedicellate winged on the keel 580. Heterachne
18. Lemmas dorsiventrally or laterally compressed but
1. Ligule absent, the leaf and sheath intergrading not flat, papery or winged 19
623. Neostapfia 19 (18). Lemmas glabrous or evenly pubescent all over,
1. Ligule present 2 apex acute 20
2 (1). Leaves stiff, sharp-tipped, often strongly divaricate 19. Lemmas pubescent on the veins only, the apex emar-
3 ginate to bilobed, sometimes also mucronate 21
2. Leaves flexible, not sharp-tipped 8 20 (19). Primary inflorescence branches generally longer
3 (2). Spikelets with exactly one flower; reduced flowers than the internodes separating them, generally spread-
lacking 4 ing; lemma with 3 veins 579. Eragrostis
3. Spikelets with more than one flower, although some- 20. Primary inflorescence branches about the same length
times only one fertile 5 as the internodes separating them, upright and
4 (3). Spikelets 2–3 mm long; lemma apex obtuse, appressed to the main inflorescence axis; lemma with
mucronate 589. Urochondra 1 vein 595. Thellungia (now part of Sporobolus s.l.)
4. Spikelets 16–24 mm long; lemma apex two-lobed, 21 (19). Palea keels glabrous or ciliolate; cleistogamous
awned from the sinus 644. Monodia spikelets lacking 22
5 (3). Lemma divided into deep broad lobes 6 21. Palea keels long ciliate; lower leaf sheaths containing
5. Lemma apex acute to acuminate, not lobed 7 cleistogamous spikelets 628. Triplasis
TRIBES AND GENERA OF CHLORIDOIDEAE 359
22 (21). Paleas widened or bowed below; caryopses not 39. Disarticulation above the glumes 40
deeply concave 683. Tridens 40 (39). Callus of flower elongate, sharp
22. Paleas not widened or bowed below; caryopses deeply 556. Centropodia
concave to folded on the side of the hilum 40. Callus of flower short, blunt 41
627. Tridentopsis 41 (40). Lemma lobes about ½ the length of the lemma
23 (8). Awns two or more per lemma 24 42
23. Awns one per lemma, or lemmas unawned and awns 41. Lemma apex entire or with shallow lobes, less than ½
on glumes only 30 the length of the lemma 43
24 (23). Lemmas with two or three, rarely four, awns 42 (41). Awn pubescent 628. Triplasis
25 42 Awn glabrous 622. Erioneuron
24. Lemma with 5 awns or more 27 43 (41). Cleistogamous spikelets forming in the leaf
25 (24). Lemma awns two, one on each marginal lobe, sheaths 44
occasionally with 2 more subsidiary awns, the lemma 43. All spikelets chasmogamous 45
apex rounded between the awns 576. Kaokochloa 44 (43). Lemma with 5 veins; palea lacking wings
25. Lemma awns three, one on each marginal lobe and one 597. Cleistogenes
from the lemma apex 26 44. Lemma with 3 veins; palea with wings
26 (25). Plants with bisexual flowers; central lemma awn 582. Steirachne
recurved 571. Triraphis 45 (43). Lemma glabrous; upper lemmas of each spikelet
26. Plants with unisexual flowers, dioecious; central reduced to a cluster of awns
lemma awn straight 613. Sohnsia 579. Eragrostis (former Ectrosia)
27 (24). Glumes with one vein; lemma with 12 awns or 45. Lemma pubescent on the nerves or all over the proxi-
more 682. Pappophorum mal abaxial side; upper lemmas of each spikelet not
27. Glumes with seven veins or more; lemma with 5–11 reduced to awns 46
awns 28 46 (45). Awn about the same length as the lemma, or up to
28 (27). Awns 5, alternating with hyaline lobes 4 times longer 626. Gouinia
577. Schmidtia 46. Awns generally much shorter than the lemma
28. Awns 7 to 11, not alternating with hyaline lobes 29 683. Tridens
29 (28). Awns with antrorse prickles but not with long
hairs; rachilla disarticulating between the flowers
574. Cottea
29. Awns plumose; rachilla not disarticulating TRIBES AND GENERA OF CHLORIDOIDEAE
575. Enneapogon
30 (23). Spikelets with exactly one flower 31
30. Spikelets with two or more flowers 33 XXV. TRIBE CENTROPODIEAE P.M. Peterson, N.P.
31 (30). Ligule membranous; lemma with 3 veins Barker & H.P. Linder (2011)
612. Muhlenbergia
31. Ligule a line of hairs; lemma with 1 vein 32 Perennials, caespitose or rhizomatous. Inflores-
32 (31). Lemma with a long awn 598. Hubbardochloa cence branches branched. Callus of flower pubes-
32. Lemma awnless; awn only on the glume cent. Lemma with 9 veins, with hairs tufted, in a
594. Sporobolus transverse row, the apex with two deep lobes, awned
33 (30). Glumes awned 34 from the sinus, the awn straight or geniculate.
33. Glumes unawned 38
34 (33). Ligule membranous 35 Two genera, five species.
34. Ligule a line of hairs 36
35 (34). Glumes with divided awns; spikelets in pairs
612. Muhlenbergia (former Lycurus)
555. Ellisochloa P. M. Peterson & N. P. Barker
35. Glumes long acuminate; spikelets solitary
672. Apochiton Ellisochloa P. M. Peterson & N. P. Barker, Taxon 60: 1118
36 (34). Plants annual; awns on lemmas much longer than (2011).
the lemma itself 569. Habrochloa
36. Plants perennial; awn on lemma shorter than the Plants less than 50 cm tall. Ligule a fringe of hairs.
lemma itself 37 Leaf blades stiff, terete, the adaxial surface repre-
37 (36). Panicle lax and open, the branches generally
not spikelet-bearing to the base, most spikelets
sented only by a shallow groove, the apex some-
pedicellate times sharp-pointed. Inflorescence branches
612. Muhlenbergia (former Redfieldia) upright. Spikelets with 2 or 3 flowers. Glumes
37. Panicle dense, spike-like, the branches generally longer than the flowers, their apices acute to acu-
spikelet-bearing to the base, most spikelets sessile or minate. Lemma with 4 to 6 hair tufts. C3.
nearly so 583. Fingerhuthia
38 (33). Plants tall, reed-like 570. Neyraudia
Two spp., southern Africa.
38. Plants shorter, not reed-like 39 The two species of Ellisochloa, E. rangei (Pilg.)
39 (38). Disarticulation below the glumes, spikelets falling P. M. Peterson & N. P. Barker and E. papposa
entire 585. Entoplocamia (Nees) P. M. Peterson & N. P. Barker, were formerly
included in Merxmuellera, but multiple studies

have shown that they were misplaced in that
genus; they clearly belong here in the clade sister
to the remainder of the Chloridoideae. Merxmuel-
lera s.s. is a member of subfamily Danthonioideae.
556. Centropodia Rchb. Fig. 82
Centropodia Rchb., Consp. Regn. Veg. 212a (1828).
Plants caespitose or rhizomatous. Ligule a fringed

membrane or a fringe of hairs. Leaf blades disarti-
culating at the ligule. Inflorescence exserted or
surrounded by the subtending sheath. Spikelets
with 2 to 6 flowers plus additional reduced flowers
distally. Callus of flower sharp. Glumes shorter or
longer than the flowers, their apices acute to acu-
minate. Lemma with lines of hairs between the
veins, hairs tufted above, the awn straight or
twisted. Leaf epidermal papillae absent. Microhairs
panicoid or chloridoid. C4, anatomically similar to
NAD-ME subtype. 2n ¼ 24, 48.
Four spp., Africa, Middle East, northern India.
Remaining genera:. Photosynthetic pathway C4.
CORE C4 CHLORIDOIDEAE INCERTAE SEDIS
557. Decaryella A. Camus
Decaryella A. Camus, Bull. Soc. Bot. France 78: 177 (1931).
Decumbent annuals, less than 30 cm tall. Leaf blades

narrow, stiff. Ligule a fringed membrane. Inflores-
cence unbranched. Spikelets on long pedicels, falling
with the pedicel at maturity, laterally compressed,
with 1 or 2 fertile flowers, rachilla extension present
or absent. Glumes longer than the flower, leathery,
hairy, cuspidate-acuminate. Lemma obtuse or acute.
Leaf epidermal intercostal papillae absent. Fig. 82. Chloridoideae-Centropodieae. Centropodia
glauca. A Habit. B Ligule. C Spikelet. (From M€
uller 2007)
One sp., D. madagascariensis A. Camus,
Madagascar.
branches unbranched, appressed to the main
558. Halopyrum Stapf axis. Spikelets with 8 to 25 flowers. Callus of
flower with straight hairs extending almost half
Halopyrum Stapf, Hooker’s Icon. Pl. 25: t 2448 (1896). the length of the spikelet. Glumes shorter than the
spikelet. Lemma mucronate. Palea covered with
Perennials with robust stolons. Ligule a fringe of long hairs. Leaf epidermal papillae one per cell.
hairs. Leaf blades with filiform tips, disarticulat- One sp., H. mucronatum (L.) Stapf, coastal
ing from sheath. Inflorescence with primary areas of Africa, India, Sri Lanka.
The inflorescence and the single epidermal unbranched branches; branch axes hollowed, the
papilla per cell suggest a possible placement in spikelets embedded in the rachis. Spikelets laterally
Eleusininae. compressed, with 1 fertile flower and 1 sterile
lemma; rachilla hairy. Upper glume longer than the
flower. Fertile lemma hairy, two-lobed, awned from
559. Indopoa Bor
sinus. Leaf epidermal papillae one to several per cell.
Indopoa Bor, Kew Bull. 13: 225 (1958). Two spp., New Caledonia, Marshall Islands,
Cocos Islands.
Tiny annuals, less than 15 cm tall. Ligule membra-
nous. Inflorescence unbranched, with fewer than 10 562. Myriostachya (Benth.) Hook. f.
spikelets. Spikelets with 4 to 6 fertile flowers plus
additional distal sterile ones. Callus of flower pubes- Myriostachya (Benth.) Hook. f., Fl. Brit. India 7: 327 (1897).
cent. Glumes shorter than the spikelet. Lemma apex
with two lobes, awned from each lobe and from the Perennial with culms 1.5 to 3 m long, often floating.
sinus, the median awn very broad, twisted and curv- Ligule a fringe of hairs. Inflorescence branches
ing around the fruit. Fruit slender, needle-like, with unbranched, disarticulating from the elongate cen-
an apical appendage. Endosperm unusually hard. tral axis. Spikelets with 2 to 12 flowers plus addi-
Leaf epidermal papillae absent. tional distal sterile ones. Glumes shorter than the
One sp., I. paupercula (Stapf) Bor, India. spikelet, awned. Lemma glabrous, mucronate. Leaf
Indopoa was segregated from Tripogon and epidermal papillae absent.
its placement is uncertain. The awn morphology One sp., M. wightiana (Nees ex Steud.) Hook.
is reminiscent of Danthonioideae or early-diverg- f., southern India and Sri Lanka to southeast Asia.
ing Chloridoideae; the species has not been
included in a molecular phylogeny.
563. Neostapfiella A. Camus
560. Kampochloa Clayton Neostapfiella A. Camus, Bull. Soc. Linn. Lyon 5: 4 (1926).
Kampochloa Clayton, Kew Bull. 21: 103 (1967). Annuals, less than 30 cm tall, with or without
stolons. Ligule a fringed membrane. Inflores-
Perennials, less than 30 cm tall. Ligule membra- cence unbranched, or with 2 to 4 unbranched
nous. Inflorescence unbranched, ending in a bare digitate branches. Spikelets wedge-shaped, with
tip, the axis curved. Spikelet with one fertile flower, 2 fertile flowers, or the upper flower male or
one staminate or sterile one distally, and a group sterile. Callus of flower elongate, pubescent,
of lemmas with long awns terminating the rachilla. sharp pointed. Glumes shorter than the spikelet,
Glumes shorter than the spikelet, the lower glume thinner than the lemma. Lemma two-toothed,
terminating in a short awn from a sinus, the upper with a subapical awn. Leaf epidermal papillae
with a longer abaxial awn. Lemma with long hairs present or absent; when present, several per cell.
along the veins, the apex with two teeth, mucro- Three spp., Madagascar.
nate. Leaf epidermal papillae absent.
One sp., K. brachyphylla Clayton, Angola and
564. Ochthochloa Edgew.
Zambia.
Peterson et al. (2014a) speculate that Kampo- Ochthochloa Edgew., J. Asiat. Soc. Bengal 11: 26–27 (1842).
chloa could be derived from Ctenium, based on
morphological similarity. Stoloniferous perennials; culms with tuberous
bases. Leaf sheaths wooly near the base. Ligule a
561. Lepturopetium Morat fringed membrane. Primary branches of inflores-
cence themselves unbranched, digitate, disarticu-
Lepturopetium Morat, Adansonia II, 20: 377 (1981). lating at maturity. Spikelets with 4 to 8 fertile
flowers plus additional distal sterile ones. Glumes
Stoloniferous perennials. Ligule a fringed mem- shorter than the spikelet, unawned. Lemma with
brane. Inflorescence unbranched, or with 2 or 3 hairs on veins and keel, the apex muticous.
Pericarp free from the seed coat. Leaf epidermal Callus of flowers pubescent. Glumes shorter than
intercostal papillae present, one or more per cell. the spikelet, the apices acute to acuminate.
One sp., O. compressa (Forssk.) Hilu, NE Lemma glabrous, with an apical awn. Leaf epider-
Africa, Iran, India. mal papillae absent.
One sp., S. nairii Nair, Sreek., Vaj. & Bargh.,
southern India.
565. Pogonochloa C. E. Hubb. Clayton and Renvoize (1986) say that this is
“like Gouinia [in Cynodonteae], but upper glume
Pogonochloa C. E. Hubb., Hooker’s Icon. Pl. 35: t. 3421
(1940). as long as lemma”. Also the lemma is hairy in
Gouinia.
Perennials. Ligule membranous. Inflorescence of
numerous unbranched branches, these appressed 568. Viguierella A. Camus
to the central axis. Spikelets with two or three
flowers, the lowest bisexual, the next staminate or Viguierella A. Camus, Bull. Bi-Mens. Soc. Linn. 5: 11 (1926).
sterile, the most distal only an awn. Callus of flower
pubescent. Glumes longer than the spikelet, with Annuals. Ligule a fringe of hairs. Inflorescence
short awns. Lemma pubescent, awned from the narrow, bottlebrush like, generally unbranched.
apex, the awn flexuous, many times longer than Spikelet subtended by a small bract, with 1 flower
the flower. Leaf epidermal papillae absent. plus additional distal sterile ones, the entire spike-
One sp., P. greenwayi C. E. Hubb., Zambia let plus the pedicel disarticulating as a unit.
and Zimbabwe. Flower callus pubescent, sharp pointed. Glumes
pilose, awned, shorter than the spikelet. Lemma
awned. Leaf epidermal papillae absent.
566. Pseudozoysia Chiov. One sp., V. madagascariensis A. Camus,
Madagascar.
Pseudozoysia Chiov., Pl. Nov. Minus Not. Aethiopia 20
(1928). Clayton and Renvoize (1986) connect this
species with Scleropogon, in Cynodonteae, on
Tiny perennials, less than 15 cm tall. Ligule a fringe the basis of the subtending bract.
of hairs. Inflorescence surrounded by the subtend-
ing leaf sheath, the branches unbranched, each XXVI. TRIBE TRIRAPHIDEAE P.M. Petersen
branch with only 2 spikelets, deciduous from the (2010)
main axis. Spikelets with 1 flower, without rachilla Annuals or perennials. Ligule a fringe of hairs.
extension. Glumes longer than the flower, hard, Inflorescence branches branched. Glumes shorter
tuberculate, acute, the lower ovate, the upper sub- than the spikelet. Lemmas keeled, pubescent on
globose and saccate. Lemma awnless, hairless. the lateral veins. Leaf epidermal papillae absent.
One sp., P. sessilis Chiov., Somalia. Microhairs slender, with an elongate apical cell
Little information is available on this species. (“panicoid type”).
Anatomical and molecular data would be partic- Three genera, 13 species.
ularly useful.
569. Habrochloa C. E. Hubbard
567. Silentvalleya V.J. Nair, Sreek., Vajr. & Bhar-
gavan Habrochloa C. E. Hubbard, Hooker’s Icon. Pl. 37: t. 3645
(1967).
Silentvalleya V.J. Nair, Sreek., Vajr. & Bhargavan, J. Bom-
bay Nat. Hist. Soc 79: 654 (1982). Slender annuals. Spikelets with 3 to 5 flowers,
the flowers with a pubescent callus. Glumes as
Caespitose perennials. Ligule membranous. Inflo- long as the spikelet, with acuminate apices.
rescence branches unbranched, the branches Callus of flower reported to be pubescent.
arranged along a central axis. Spikelets with 6 or Lemma bilobed, the apex truncate, with a long
more flowers plus additional distal sterile ones. awn from the sinus.
One sp., H. bullockii C. E. Hubb., Central Africa. apices acute to acuminate. Lemma with two
Habrochloa is placed here following Peterson lobes, awned from the sinus. 2n ¼ 40.
(pers. comm.). Five spp., Asia, Africa.
570. Neyraudia Hook. f. Fig. 83 571. Triraphis R. Br.
Neyraudia Hook. f., Fl. Brit. India 7: 305 (1897). Triraphis R. Br., Prodr.: 185 (1810).
Caespitose or short-rhizomatous, reed-like per- Annuals or perennials, caespitose or rhizoma-

ennials. Leaf blades disarticulating or not. Inflo- tous. Spikelets with 4 to 24 flowers, with addi-
rescence large, plumose. Spikelets with several tional reduced flowers distally. Callus of flower
flowers, the proximal one sterile, the next 3 to pubescent. Glume apices entire or bifid, mucro-
8 fertile, and the distalmost reduced. Glume nate or awned. Lemmas with three lobes, each
ending in an awn; central lobe with two teeth.
Fruit linear, trigonous. 2n ¼ 20.
Eight spp., Africa, Australia, South America.
XXVII. TRIBE ERAGROSTIDEAE Stapf (1898)

Annuals or perennials, caespitose, rhizomatous, or
stoloniferous. Leaves with or without glands. Ligule
a fringe of hairs, less often membranous or a fringed
membrane. Inflorescence branched or unbranched.
Spikelets laterally compressed, with 1 to many flow-
ers, the distal flowers often reduced. Callus of flow-
ers generally pubescent. Glumes shorter than the
spikelet in most species, rarely the same length,
generally lacking awns. Flowers generally bisexual,
rarely unisexual and the plants then dioecious.
Lemma glabrous in most species, with three (to 13)
veins, the apex generally entire, rarely mucronate or
awned. Pericarp free from the seed coat or not. Leaf
epidermal papillae absent in most species (except E.
obtusiflora). Microhairs slender, with an elongate
apical cell (“panicoid type”), or with a broad apical
cell (“chloridoid type”), or with a broad apical cell
with internal membranes (“Enneapogon type”).
INCERTAE SEDIS
572. Cladoraphis Franch.
Cladoraphis Franch., Bull. Mens. Soc. Linn. Paris 1: 673

(1887).
Shrubby perennials. Leaves with blades mostly

Fig. 83. Chloridoideae-Triraphideae. Neyraudia mon- absent or strongly reduced. Ligule a fringe of hairs.
tana. A Leaves and inflorescence. B Spikelet with flowers Inflorescence axis and primary branches woody, stiff,
disarticulated above two glumes. C Flower, left abaxial ending in a sharp point, the primary branches
view, and right adaxial view. D Palea, left abaxial view, distributed along an elongate inflorescence axis. Sec-
right adaxially. (From Wu et al. 2007, with permission
from the Missouri Botanical Garden Press, St. Louis, and ondary branches short, not woody, but spine-tipped
Science Press, Beijing) like the primary ones. Spikelets with 3 to 12 flowers
plus additional reduced ones distally. Glumes shorter flowers, with additional reduced flowers distally.
than the spikelet, their apices acute. Lemma apex Glumes shorter than the spikelet, pubescent, the
acute. Pericarp free from seed coat. 2n ¼ 20, 40. apices acute. Rachilla disarticulating between the
Two spp., South Africa, Namibia. flowers. Lemmas with 7 to 11 lobes ending in awns,
plus two lateral lobes without awns, the awns with
antrorse prickles. 2n ¼ 20.
573. Tetrachaete Chiov.
One sp., C. pappophoroides Kunth, Southern
Tetrachaete Chiov., Annuario Reale Ist. Bot. Roma 8: 28 USA to Argentina.
(1903).
575. Enneapogon Desv. ex P. Beauv. Fig. 84
Tiny annuals, less than 25 cm tall. Ligule a fringed
membrane. Inflorescence surrounded by the sub- Enneapogon Desv. ex P. Beauv., Ess. Agrostogr. 81, 161
tending leaf sheath, the branches unbranched, (1812).
each branch with only 2 spikelets, deciduous
from the main axis. Spikelets with 1 flower, with- Annuals or perennials, caespitose or short rhizo-
out rachilla extension. Glumes much longer than matous. Ligule membranous, or a fringed mem-
the flower, awn-like, bending outward, with long brane, or a fringe of hairs. Inflorescence
feathery hairs. Lemma ovate, gibbous, leathery, unbranched, or with unbranched primary
with a long awn. Leaf epidermal papillae absent. branches, these digitate or arranged along a cen-
One sp., T. elionuroides Chiov., Ethiopia to tral axis. Spikelets with one fertile flower, with
Tanzania and Arabia. additional reduced flowers distally; upper flowers
Tetrachaete is placed in Eragrostideae follow- reduced and internodes contracted so the assem-
ing Soreng et al. (2012 and onward); the data bled lemmas look like a brush. Glumes shorter or
supporting the placement are unpublished. longer than the spikelet, the lower one with a
sharp tip, the upper one mucronate or awned.
Lemmas with 9 veins, each ending in an awn;
awns much longer than the lemma and plumose
XXVII.1. SUBTRIBE COTTEINAE Reeder (1965)
above. Leaf blade microhairs with an inflated
terminal cell, with internal membranes. 2n ¼ 18,
Lemmas pubescent, lobed, with five or more awns.
20, 36, 40, 60.
Spikelets with several flowers, incomplete flowers
Twenty-four spp., tropical and subtropical
distal. Glumes with multiple veins. Basal cell of
regions worldwide.
bicellular microhairs elongate, 60–180 mm.
Four genera, 28 spp.
Members of this subtribe have been placed in 576. Kaokochloa de Winter
the tribe Pappophoreae in the past, based on the
multiple awns of the lemma. Molecular data, Kaokochloa de Winter, Bothalia 7: 479 (1961).
however, are unequivocal and consistent in
showing this character as being convergent and Annuals. Leaf blades viscid. Inflorescence
the traditional Pappophoreae as being polyphy- branches branched. Spikelets with 3 to 6 flowers,
letic (Columbus et al. 2007; GPWG II 2012; Peter- with additional reduced flowers distally; rachilla
son et al. 2010a; Reutemann et al. 2011). disarticulating between the glumes, the lower
glume persisting on the plant and the upper fall-
ing with the flowers. Glumes as long as the spike-
574. Cottea Kunth let, their apices acute. Callus of flower pubescent.
Lemmas with 2 lobes ending in awns; apex
Cottea Kunth, Révis. Gramin. 1: 84 (1829).
rounded between the awns, curved inward.
One sp., K. nigrirostris De Winter, southwest-
Perennials. Culms swollen below, where associated
ern Africa.
with cleistogamous spikelets. Ligule a fringe of hairs.
Inflorescence branches branched, borne upright
and close to the axis. Spikelets with 4 to 8 fertile 577. Schmidtia Steud. ex J.A. Schmidt
Schmidtia Steud. ex J.A. Schmidt, Beitr. Fl. Cap. Verd. Ins.

144 (1852), nom. conserv.
Annuals or perennials, caespitose or short

rhizomatous. Ligule a fringe of hairs. Inflorescence
branches branched. Spikelets with 3 to 7 flowers,
with additional reduced flowers distally. Glumes
shorter than the spikelet. Lemmas with 6 lobes
alternating with 5 awns, the lateral awns only
slightly shorter than the central one. 2n ¼ 36.
Two spp., Africa and Pakistan.
The two species of Schmidtia are paraphyletic
in the analyses of Reutemann et al. (2011), with
Enneapogon derived from within them; the two
genera should perhaps be combined.
XXVII.2. SUBTRIBE ERAGROSTIDINAE J. Presl

(1830)
Lemma with (1) 3 (5) veins, the apex entire, awn-

less to rarely mucronate or short-awned. Bicellular
microhairs with short basal cells, 15–75 mm.
Five genera, 459 spp.
This subtribe is dominated by the large and
polymorphic genus Eragrostis. There is no obvi-
ous morphological synapomorphy for the clade,
although many species have branched inflores-
cences bearing unawned, laterally compressed
spikelets that have many flowers, and glumes
shorter than the adjacent lemmas.
578. Catalepis Stapf & Stent
Catalepis Stapf & Stent, Bull. Misc. Inform. Kew 1929: 11

(1929).

branches unbranched, these short and closely
spaced on the inflorescence axis, each bearing
no more than 9 spikelets. Spikelets with one
flower, rachilla extension present. Lower glume
tiny and subulate, upper glume longer than the
flower, the apex acuminate. Lemma with tufts of
hairs on the margins, awnless. Leaf epidermal
papillae absent.
One sp., C. gracilis Stapf & Stent, South
Africa.
Fig. 84. Chloridoideae-Eragrostideae-Cotteinae. Ennea- Grassbase (Clayton et al. 2006 onward) indi-
pogon desvauxii. A Plant. B Spikelet. C Flower. D Cleis- cates that this species lacks a rachilla extension,
togamous spikelet. (From Barkworth et al. 2003, drawn by but the original description says that it is present.
K. Klitz) This genus is placed here based on the
Acamptoclados Nash, Fl. S.E. U.S. 139, 1327 (1903).

Psammagrostis C.A. Gardner & C.E. Hubb., Hooker’s
Icon. Pl. 34: t 3361 (1938).
Ectrosiopsis (Ohwi) Ohwi ex Jansen, Acta Bot. Neerl. 1:
474 (1952).
Diandrochloa de Winter, Bothalia 7: 387 (1960).
Annuals or perennials, caespitose, rhizomatous, or

stoloniferous. Leaves with or without glands. Lig-
ule a fringe of hairs, less often membranous or a
fringed membrane. Inflorescence generally
exserted, but sometimes surrounded by a subtend-
ing leaf, the branches themselves branched in most
species, sometimes unbranched. Spikelets with
2 to many flowers, the distal flowers often reduced.
Glumes shorter than the spikelet in most species,
rarely the same length or longer, generally with
one vein, generally lacking awns. Flowers generally
bisexual, but in two species unisexual and the
plants then dioecious. Lemma glabrous in most
species, with three veins, the apex generally entire,
rarely mucronate or awned. Pericarp free from
seed coat or not. Leaf epidermal papillae absent
in most species (except E. obtusiflora). Microhairs
slender, with an elongate apical cell (“panicoid
type”), or with a broad apical cell (“chloridoid
type”), or with a broad apical cell with internal
membranes (“Enneapogon type”). 2n ¼ 20, 40, 50,
60, 80, 100, 108.
Four hundred thirty-seven spp., warm areas
worldwide.
The broad circumscription of Eragrostis sug-
gested here follows recommendations of Colum-
bus et al. (2007), Ingram and Doyle (2004), and
Peterson et al. (2010a) whose molecular phyloge-
netic studies identify a strongly supported clade
Fig. 85. Chloridoideae- Eragrostideae-Eragrostidinae. including Eragrostis and its segregates. Subclades
Eragrostis frankii. A Flowering plant. B Spikelets. (From
Barkworth et al. 2003, drawn by C. Roché) within this group do not correspond to previ-
ously recognized genera, and Eragrostis in its
traditional sense is paraphyletic. Ingram and
classification presented by Peterson et al. (2010a),
Doyle (2004) found that Acamptoclados, Dia-
although no molecular data are available. The
ndrochloa, Neeragrostis, and Pogonarthria should
species is superficially similar to Polevansia, in
be included within Eragrostis. Former Acampto-
Cynodonteae.
clados (¼ E. sessilispica) is an Eragrostis in which
the inflorescence disarticulates at its base and is
579. Eragrostis Wolf Fig. 85 dispersed whole. The seven species formerly in
Diandrochloa have a membranous ligule, but
Eragrostis Wolf, Gen. Pl. 23 (1776). other than that are like Eragrostis, and were
Ectrosia R. Br., Prodr. 185 (1810). placed in that genus by Koch (1978). Former
Harpachne A. Rich., Tent. Fl. Abyss. 2: 431 (1851).
Pogonarthria Stapf, Fl. Cap. 7: 316 (1898). Neeragrostis (¼ E. reptans) is dioecious, a char-
Neeragrostis Bush, Trans. Acad. Sci. St. Louis 13: 178 acter that appears elsewhere in Eragrostis. Former
(1903). Pogonarthria has inflorescences with the primary
branches unbranched; the shift from branched to persistent, their apices obtuse or acute. Lemma
unbranched inflorescences has occurred many with a winged keel and obtuse apex. Palea gib-
times in grass evolution so is likely not a good bous, with winged keels.
character on which to base a genus. Pogonarthria Three spp., Northern Australia.
also has lemmas that are acuminate or awned, a
character that it shares with former Psammagros-
tis, Ectrosiopsis, Harpachne, and Ectrosia. The 581. Richardsiella Elffers & Kenn.-O’Byrne
latter genus was placed in Eragrostis by Colum-
Richardsiella Elffers & Kenn.-O’Byrne, Kew Bull. 11: 455
bus et al. (2007) and Peterson et al. (2010a), and (1957).
Peterson et al. (2010a) confirmed the placement
of Psammagrostis and Harpachne. Species with Tiny annuals, less than 18 cm tall. Leaves with
deciduous glumes were segregated as Ectrosiop- abaxial glands. Ligule a fringe of hairs. Inflores-
sis, and those with an unbranched inflorescence cence axis and primary branches ending in a slen-
as Psammagrostis and Harpachne. der bristle, the primary branches curved when dry.
Although sampling within Eragrostis in molec- Spikelet with 6 to 12 flowers, with additional
ular phylogenies is reasonably good, the number reduced flowers distally. Glumes the same length
of species included remains a fraction of the total. as the spikelet, deciduous, with tubercle-based
There remains the possibility that some species hairs; apices of glumes acuminate and mucronate.
currently placed in Eragrostis should in fact be Lemma apex obtuse, mucronate. Pericarp free
placed elsewhere. For example, the species from seed coat. Leaf epidermal papillae absent.
described as E. walteri Pilg. has been widely cited One sp., R. eruciformis Ellfers & O’Byrne,
as being the only C3 species of Eragrostis, but Zambia.
recent molecular phylogenies (Ingram et al. 2011) This species is similar to Dinebra somalensis,
show that it is in fact a member of subfamily which also has leaf glands. The bristle-tipped
Arundinoideae and thus was misplaced here. Thel- branches are similar to those of Cladoraphis. No
lungia is excluded from Eragrostis by molecular molecular data are available for this species and it
data (GPWG II 2012; Ingram and Doyle 2004) and is placed in this subtribe following the classifica-
is placed in Zoysieae by Peterson et al. (2010a). tion of Peterson et al. (2010a).
The subgeneric classification of Eragrostis is
still under debate. Major characters proposed for
subdivision include the pattern of spikelet disar- 582. Steirachne Ekman
ticulation (Van den Borre and Watson 1994) or
the C4 photosynthetic subtype, as indicated by Steirachne Ekman, Ark. Bot. 10: 35 (1911).
leaf anatomy. A study of the latter character sug-
gests that PCK-type anatomy may be synapo- Perennials, with cleistogamous spikelets enclosed in
morphic for a clade corresponding to subgenus the sheaths. Ligule a fringed membrane or a fringe
Caesiae (Ingram 2010). However, allopolyploidy of hairs. Inflorescence branches unbranched. Spi-
is common in the genus and is likely to make any kelets with 6 to 10 flowers, with additional reduced
attempt at subgeneric classification difficult flowers distally. Rachilla internodes long, pubescent
(Ingram 2010). at apex. Glumes shorter than the spikelet, with a
single vein, their apices acuminate. Lemma awned.
Palea winged. Stamens 2.
580. Heterachne Benth. Two spp., Brazil, Guyana, and Venezuela.
Heterachne Benth., Hooker’s Icon. Pl. 13: t. 1250 (1877).
Annuals. Leaf blades stiff. Ligule a fringe of hairs. XXVII.3. SUBTRIBE UNIOLINAE Clayton (1982)
Inflorescence branches themselves branched,
with spikelets borne in clusters. Spikelets with 1 Ligule a fringe of hairs. Spikelets disarticulating
or 2 flowers, with additional reduced flowers dis- below the glumes. Lemma with 3 to 10 veins, the
tally, forming a fan-like cluster; rachilla zigzag. apex entire, awnless to mucronate. Basal cell of
Glumes shorter than the spikelet, deciduous or bicellular microhairs short, 15–35 mm. Leaf blade
with abaxial sclerenchyma of vascular bundles 585. Entoplocamia Stapf Fig. 86

continuous with columns of colorless cells.
Six genera, 15 spp. Entoplocamia Stapf, Fl. Cap. 7: 318 (1898).
This subtribe has no obvious morphological
synapomorphy. Annuals. Inflorescence with short unbranched
branches or with sessile spikelets. Proximal ster-
ile flowers 2, fertile flowers 4 to 20. Glumes
583. Fingerhuthia Nees ex Lehm. shorter than the spikelet, the lower one with an
acute apex, the upper acuminate and mucronate.
Fingerhuthia Nees ex Lehm., Sem. Hort. Bot. Hamburg.
1834: 7 (1834).
Perennials, caespitose or rhizomatous. Inflores-

cence branches branched. Spikelets with one fer-
tile flower, plus 1 to 3 sterile or staminate flowers
distally. Callus blunt. Glumes shorter than the
spikelet, apices acuminate, awned. Lemma
awned. 2n ¼ 20, 40.
Two spp., Afghanistan, Arabia, southern Africa.
This genus was placed in a clade with Uniola
and Tetrachne by Ingram and Doyle (2004).
584. Stiburus Stapf
Stiburus Stapf, Fl. Cap. 7: 696–697 (1900).
Caespitose perennials. Leaf blades narrow and

upright. Ligule a fringe of hairs. Inflorescence
dense, like a bottlebrush, the branches branched,
borne upright and closely appressed to the axis.
Spikelets with 1 to 3 flowers plus additional
reduced flowers distally. Glumes shorter than
the flowers; glumes and lemmas covered with
long hairs. Stamens 2. 2n ¼ 20.
Ingram and Doyle (2004) present molecular
evidence hinting at a relationship between Sti-
burus and Uniola, although there is little morpho-
logical resemblance. Their data contradict that of
Clayton et al. (2006 onward) and Clayton and
Renvoize (1986) who consider the two species of
Stiburus as part of Eragrostis.
ENTOPLOCAMIA + TETRACHNE + UNIOLA

Palea keels winged. Spikelets with multiple
flowers, the proximal ones sterile.
These three genera form a well-supported
clade in the study of Peterson et al. (2010a), but
Fingerhuthia is sister to Tetrachne in GPWG II
(2012) and Ingram and Doyle (2004). If the latter Fig. 86. Chloridoideae-Eragrostideae-Uniolinae. Ento-
topology is correct, then winged palea keels may plocamia aristulata. A Habit. B Ligule. C Inflorescence.
be lost in Fingerhuthia. (From M€uller 2007)
Lemma awned, the awn sharp. Lodicules absent. Ingram and Doyle (2004) identified a clade
Pericarp free from the seed coat. made up of Spartina, Calamovilfa, Sporobolus and
One sp., E. aristulata (Hack. & Rendel) Stapf, Thellungia. This was supported by Columbus et al.
southwestern Africa. (2007) who added Crypsis and placed Zoysia sister
to the rest of the group. Peterson et al. (2010a)
586. Tetrachne Nees placed Urochondra sister to all other species. Psilo-
lemma was included in this clade by Peterson et al.
Tetrachne Nees, Fl. Afr. Austral. Ill. 1: 375 (1841). (2014b). In their analyses, which are far more com-
prehensive than anything published previously,
Perennials. Inflorescence with unbranched Psilolemma, Urochondra, Zoysia, and Sporobolus
branches, these distantly spaced and not overlap- somalensis are variously placed outside a large
ping on the central axis. Proximal sterile flowers clade made up of the remaining species of Sporobo-
2, fertile flowers 4 to 5. Glumes shorter than the lus, plus Spartina and several smaller genera.
spikelet, their apices acute. Lemma unawned. The oldest name of the combined genus would
Pericarp free from the seed coat. 2n ¼ 20. be Spartina, which was published in April 1789 and
One sp., T. dregei Nees, South Africa and thus has priority over Crypsis, published in October
Pakistan. of the same year. However, Peterson et al. (2014c)
Tetrachne is sister to Entoplocamia in the have proposed conservation of the name Sporobo-
phylogeny of Peterson et al. (2010a). Watson lus and have made the necessary combinations. The
and Dallwitz (1992 onward) suggest that this spe- recommendations of Peterson et al. (2014b, c) are
cies hybridizes with Fingerhuthia, which is the adopted here but genera could not be renumbered
sister taxon in GPWG II (2012) and Ingram and and descriptions altered in time for publication.
Doyle (2004). Although Peterson et al. (2007) and Soreng
et al. (2012 and onward) divide Zoysieae into the
587. Uniola L. monotypic subtribe Zoysiinae Benth. and place
Uniola L., Sp. Pl. 1: 71 (1753).
all other genera in Sporobolinae Benth., this
opsis Yates, Southwest Naturalist 11: 382 (1966).
Leptochlo€ seems like an unnecessary subdivision. Accord-
ingly, the subtribal rank is dispensed with here.
Perennials, caespitose, rhizomatous, or stolonif-
erous. Ligule sometimes lacking. Inflorescence 588. Psilolemma S. M. Phillips
branches unbranched, overlapping each other
on the central axis. Proximal sterile flowers 2 to Psilolemma S. M. Phillips, Kew Bull. 29: 267 (1974).
8. Glumes shorter than the spikelet, their apices
acute to acuminate, mucronate. 2n ¼ 40. Stoloniferous perennials. Ligule a line of hairs. Leaf
Eight spp., southern U.S. to Ecuador. blades stiff, sharp-pointed. Inflorescence branches
U. condensata, one of two Uniola species widely spaced on central axis, with 2 or 3 spikelets
transferred to Leptochlo€opsis (Yates 1966), is sis- each (up to 5). Spikelets with 4 to 14 flowers, sub-
ter to U. paniculata in the study of Peterson et al. terete to laterally compressed. Glumes shorter than
(2010a); the two genera are combined here. the spikelet. Lemma glabrous, with 1 to 3 veins, the
apex obtuse. Pericarp easily separated from the
XXVIII. TRIBE ZOYSIEAE Benth. (1881) seed coat. Leaf epidermal papillae absent.
One sp., P. jaegeri (Pilger) S. M. Phillips, East
Ligule generally a fringe of hairs. Spikelets later-
Africa.
ally compressed, with one flower. Lower glume
Phillips (1974) notes similarity in habit
absent, or with no veins, or with only one vein.
between this species and Odyssea.
Lemma with a single vein, awnless. Pericarp free
from seed coat.
Nine genera, 235 species. 589. Urochondra C. E. Hubb.
Many members of this tribe are found in
sandy or saline, often moist sites (Peterson et al. Urochondra C. E. Hubb., Hooker’s Icon. Pl. 35: t 3457 (1947).
2007). The presence of one flower per spikelet may
be synapomorphic here, although the character is Perennials. Leaf blades stiff, sharp-pointed. Inflo-
homoplasious in the Chloridoideae as a whole. rescence dense, cylindrical, the branches
branched, borne upright and closely appressed to

the axis. Glumes shorter than the flower, their
apices acute to acuminate, awnless. Lemma
keeled, the apex obtuse, mucronate. Lodicules
absent. Fruit with a beak formed from fused
style branches. Pericarp swelling when wet. Leaf
epidermal papillae present, one per cell.
One sp., U. setulosa (Trin.) C. E. Hubb.,
Sudan, Somalia and Sindh (Pakistan).
This species is sister to all other members of
Zoysieae in the phylogeny of Peterson et al. (2010a).
Remaining genera:
Epidermal papillae lacking on the abaxial leaf
blade; sometimes present adaxially. Leaf blades
disarticulating from the sheaths in many but not
all genera.
590. Zoysia Willd.
Zoysia Willd., Ges. Naturf. Freunde Berlin Neue Schriften

3: 440–441 (1801), nom. conserv.
Perennials, caespitose or rhizomatous, if the lat-

ter then forming dense turf. Leaf blades with a
sharp pointed apex. Ligule a fringed membrane
or a fringe of hairs. Inflorescence unbranched,
in one species consisting of a single spikelet.
Fig. 87. Chloridoideae-Zoysieae. Sporobolus cryptandrus.
Spikelets short pedicellate; rachilla extension A Plant with inflorescence. B Spikelet, the glumes sepa-
absent. Lower glume absent; upper glume as rated from the floret. (From Gould and Box 1965)
long as the flower or longer and enclosing it, the
apex muticous to awned. Lemma hyaline, acute out rachilla extension. Flowers with a bearded
or with a tiny apical notch, with a small mucro. callus. Glumes shorter or longer than the flower,
Leaf epidermal papillae present adaxially. Micro- awnless. Lemma awnless, hairy or not. 2n ¼ 40.
hairs with partitioning membranes. 2n ¼ 40. Five spp., North America.
Eleven spp., warm regions of Asia and Aus- Molecular phylogenetic data (Columbus et al.
tralasia. 2007; GPWG II 2012; Peterson et al. 2010a) place
Species of Zoysia are widely cultivated for this sister to Spartina, or to a subset of Spartina
lawns. species (Fortune et al. 2006). This genus is now
Remaining genera: placed as a section and subsection of Sporobolus
Pericarp free from the seed coat. s.l., and the necessary nomenclatural combina-
tions have been made (Peterson et al. 2014b).
591. Calamovilfa (A. Gray) Hack. ex Scribn. &
Southw. 592. Crypsis Aiton
Calamovilfa (A. Gray) Hack. ex Scribn. & Southw., True Crypsis Aiton, Hort. Kew. 1: 48 (1789), nom. conserv.
Grasses: 113 (1890).
Stoloniferous annuals; culms prostrate or ascend-
Perennials, up to 2 m tall, caespitose or rhizoma- ing. Inflorescence branches branched, the inflo-
tous. Ligule sometimes a fringed membrane. rescence contracted and spike-like or capitate,
Inflorescence branches branched. Spikelets with- often surrounded by the subtending leaf.
Spikelets without a rachilla extension. Glumes Grain spherical to ellipsoid; pericarp swelling
shorter than or as long as the spikelets, muticous when wet and ejecting the seed. 2n ¼ 18, 24, 36,
or awned. Lemma muticous or awned. Lodicules 38, 54, 72, 80, 88, 90, 108, 126.
absent. Pericarp swelling when wet. 2n ¼ 16, 18, One hundred ninety-eight spp., warm regions
32, 36, 54. worldwide.
Ten spp., Mediterranean and Middle East, to Sporobolus in its traditional sense is paraphy-
China and central Africa. This genus is now letic, with Crypsis, Spartina, and Calamovilfa
placed as a section and subsection of Sporobolus derived from within it (Peterson et al. 2010a).
s.l., and the necessary nomenclatural combina- A recent comprehensive molecular phylogenetic
tions have been made (Peterson et al. 2014b). study has circumscribed the genus in a broad
sense and has formally transferred species from
the segregate genera to Sporobolus (Peterson et al.
593. Spartina Schreb.
2014b). Because Sporobolus is not the oldest name
Spartina Schreb., Gen. Pl. ed. 8: 43 (1789). in the clade, the transfers rest on a proposal to
conserve the name Sporobolus (Peterson et al.
Perennials, caespitose or rhizomatous. Inflores- 2014c). A well-supported subgeneric classifica-
cence branches unbranched, the central axis tion is also presented based on the phylogeny.
elongate or contracted. Branches ending in a
naked tip or in a spikelet. Spikelets tightly packed 595. Thellungia Stapf
in two rows along each branch, without a rachilla
extension. Glumes as long as the flowers or lon- Thellungia Stapf, Bull. Misc. Inform. Kew 1920: 97 (1920).
ger, the apex muticous to awned, the upper glume
much longer than the lower. Lemma keeled, the Perennials, ca. 1 to 1.5 m tall. Inflorescence with
apex entire or bidentate, awnless. Lodicules primary branches branched, about the same
absent. 2n ¼ 28, 40, 42, 60, 62, 84, 120, 122, 124. length as the internodes separating them, upright
Seventeen spp., coastal areas of the New and appressed to the main inflorescence axis. Spi-
World, Atlantic coast of Europe and Africa, kelets with 2 to 4 flowers. Glumes translucent,
inland wet areas and prairie in the U.S. shorter than the spikelet, awnless. Lemmas trans-
Diploids are unknown in Spartina, raising the lucent, awnless, glabrous, with one vein.
possibility that the genus is of allopolyploid ori- One sp., T. advena Stapf, Australia.
gin. Fortune et al. (2006) provide some evidence Although Clayton et al. (2006 onward) and
for this possibility in their study of the single- Clayton and Renvoize (1986) include this genus in
copy nuclear gene waxy. This genus is now placed Eragrostis, molecular data clearly place it in Zoy-
as a section of Sporobolus s.l., and the necessary sieae (Ingram and Doyle 2004; Peterson et al.
nomenclatural combinations have been made 2014b). The inflorescence structure and the one-
(Peterson et al. 2014b). veined lemmas are distinct from those in Eragrostis.
The species has been formally transferred to Spor-
594. Sporobolus R. Br. Fig. 87 obolus by Peterson et al. (2014b).
Sporobolus R. Br., Prodr. 169 (1810).

XXIX. TRIBE CYNODONTEAE Dumort. (1824)
Annuals or perennials, caespitose, rhizomatous, Members of this tribe have no obvious morpho-
or stoloniferous. Leaf blade apex with a short logical character in common, and the range of
mucro or sharp point. Ligule membranous, or a variation is almost as great as that of the entire
fringed membrane, or a fringe of hairs. Inflores- subfamily. As noted by Peterson et al. (2007), “[t]
cence branches branched, the proximal ones here are no definitive non-molecular characters
sometimes sterile. Spikelets sometimes more or that differentiate Cynodonteae from Eragrosti-
less terete, sometimes with 2 flowers. Glumes deae and/or Zoysieae.” Nonetheless, Cynodon-
shorter or longer than the spikelet, the lower teae are strongly supported as monophyletic by
glume awnless, the upper glume muticous to molecular data (Columbus et al. 2007; GPWG II
awned. Callus of flower glabrous. Lemma thin, 2012; Peterson et al. 2010a, 2012, 2014a).
shiny, with one or three veins, without an awn. Although there is some phylogenetic structure
among the early-diverging lineages, this is gener- because the word “cleistogene” is a technical mor-
ally poorly supported and as a result several phological term to describe spikelets enclosed in
genera are placed incertae sedis. the leaf sheaths, the generic name was in violation
Eighty-two genera, 900 species. of International Code of Botanical Nomenclature,
Article 68. However, Article 20.2 of the most recent
INCERTAE SEDIS code (McNeill et al. 2007) states only that “the
name of a genus may not coincide with a Latin
596. Aeluropus Trin. technical term.” Because the term cleistogene is
not Latin and hence not “a Latin technical term”,
Aeluropus Trin., Fund. Agrost. 143 (1820). the name Cleistogenes stands; in example 5, the
ICBN explicitly states that Kengia is illegitimate.
or stoloniferous. Ligule a fringed membrane or a 598. Hubbardochloa Auquier
fringe of hairs. Leaf blades stiff, with a sharp apex. Hubbardochloa Auquier, Bull. Jard. Bot. Belg. 50: 241 (1980).
Inflorescence unbranched, or with unbranched
branches appressed to the central axis. Spikelets Tiny annual, less than 15 cm tall. Leaf blades pseu-
laterally compressed, with 2 to 30 flowers, with dopetiolate. Ligule a line of hairs. Inflorescence
additional reduced flowers distally. Glumes branches delicate, branched. Spikelets with one
shorter than the spikelet, unawned. Lemma with flower. Callus with minute hairs. Lemma with a
5 or more veins, the apex mucronate. Leaf epider- long terminal awn. Palea absent. Lodicules absent.
mal papillae one per cell. Leaf epidermal papillae abundant, several per cell.
Eight spp., Mediterranean, Asia, Northern One sp., H. gracilis Auquier, Rwanda, Bur-
Africa. undi, Zambia.
Although this species has been put in a
monotypic subtribe by Peterson et al. (2010a), it 599. Lepturidium Hitchc. & Ekman
is in an unresolved position in molecular phylo-
genies and there seems little reason to create a Lepturidium Hitchc. & Ekman, Man. Grasses W. Ind.: 111
higher-level taxon to include it. Peterson et al. (1936).
(2012, 2014a) provide weak support for placing
Aeluropus sister to Triodiinae, whereas the Perennials, less than 30 cm tall. Ligule a fringe of
GPWG II (2012) places it sister to Eleusininae. hairs. Leaf blades stiff and leathery. Inflorescence
unbranched, flattened, surrounded by the sub-
tending leaf sheath, the spikelets borne on one
597. Cleistogenes Keng side. Spikelets laterally compressed, with 1 or
2 flowers, the proximal one fertile, the distal one
Cleistogenes Keng, Sinensia 5: 147 (1934).
Kengia Packer, Bot. Not. 113: 291 (1960), nom. superfl. sterile or absent. Glumes enclosing the flowers,
their apices acute. Callus of the flower pubescent.
Caespitose perennials. Upper leaf sheaths enclos- Lemma with two teeth and a short mucro. Leaf
ing cleistogamous flowers. Leaf blades stiff. Ligule epidermal papillae absent.
a fringe of hairs or a fringed membrane. Inflor- One sp., L. insulare Hitchc. & Ekman, Cuba.
escences unbranched or branched, the branches Clayton and Renvoize (1986) comment that
branched or not. Spikelets laterally compressed, Lepturidium is “barely distinct from Brachyachne
with 2 to 8 fertile flowers plus additional reduced [Eleusininae]”, but Peterson et al. (2010a) show
flowers distally. Callus of flower pubescent. that the two are unrelated.
Glumes shorter than the spikelet, their apices
muticous. Lemma with 5 veins, the apex entire 600. Orinus Hitchc.
or with two teeth, muticous or awned. Leaf epi-
dermal papillae one per cell. Orinus Hitchc., J. Wash. Acad. Sci. 23: 136 (1933).
Fourteen spp., Europe and Asia.
The names Cleistogenes and Kengia refer to Rhizomatous perennials, dune grasses. Leaf blades
the same genus. Packer (1960) argued that, with sharp tip. Ligule membranous. Inflorescence
branches unbranched, borne along a central axis. teae following Soreng et al. (2012 and onward); the
Spikelets laterally compressed, with 1 to 8 flowers data supporting this placement are unpublished.
plus additional reduced flowers distally. Rachilla
internode elongated between flowers. Glumes BRACHYCHLOA + DACTYLOCTENIUM + NEOBOU-
shorter than the spikelet, their apices acute to TELOUA + ODYSSEA
acuminate. Lemma apex muticous or mucronate. Spikelets laterally compressed.
Leaf epidermal papillae one per cell. These four genera form a well-supported
Five spp., Western Himalayas. clade in GPWG II (2012).
Peterson et al. (2012) provide weak support
for placement of Orinus sister to Triodia. 603. Brachychloa S. M. Phillips
Brachychloa S. M. Phillips, Kew Bull. 37: 145, 158 (1982).

601. Pogoneura Napper
Pogoneura Napper, Kirkia 3: 112 (1963). Annuals or perennials, caespitose or stolonifer-

ous. Ligule a fringed membrane. Inflorescence
Annuals. Leaf blades with multicellular glands branches unbranched, spread along the central
adaxially and on blade margins. Inflorescence axis, disarticulating or not. Spikelets with 4 to 7
branches unbranched, spread along a central axis. flowers plus additional reduced flowers distally.
Spikelets more or less terete, with 2 flowers, rachilla Glumes shorter than the flowers, their apices
extension lacking, or reduced flowers present dis- acute. Lemma glabrous, the apex with a minute
tally. Glumes longer than flowers and enclosing mucro from an equally minute sinus. Palea
them. Callus of flowers pubescent. Lemma with entire, gibbous. Pericarp free from seed coat.
hairs on lateral veins and keel, bidentate, awned Leaf epidermal papillae one per cell.
from sinus. Lodicules minute or absent. Two spp., Mozambique and Natal, South
One sp., P. biflora Napper, East Africa. Africa.
Peterson et al. (2014b) place Pogoneura here
in Cynodonteae, rather than its former placement 604. Dactyloctenium Willd.
in Zoysieae (Peterson et al. 2010a). The presence
of two flowers per spikelet is unusual in Zoysieae Dactyloctenium Willd., Enum. Pl. 2: 1029–1030 (1809).
and provides morphological evidence that the
genus belongs here. Annuals or perennials, caespitose or stolonifer-
ous. Leaf blades with a sharp pointed or muticous
602. Pommereulla L. f. apex. Ligule membranous, fringed or not. Inflor-
escences unbranched, or with unbranched pri-
Pommereulla L. f., Nov. Gram. Gen. 31 (1780). mary branches, digitate, ending in a naked
point. Spikelets with 1 to 9 flowers, with addi-
Tiny stoloniferous perennials, less than 15 cm tional reduced flowers distally. Glumes shorter
tall. Ligule a fringe of hairs. Inflorescence or longer than the flowers, the apex of the lower
unbranched. Spikelets turbinate, with 2 sterile one muticous or mucronate, the upper glume
proximal flowers, 2 or 3 fertile flowers, and 1 or with a terminal or subapical, sharply bent awn.
more distal rudimentary flowers. Lemmas fan- Callus of flower glabrous. Lemma acute, awnless,
shaped, 3-lobed, hairy below, with awn-tips from mucronate or awned, the awn recurved. Fruit
lobes and an awn from the middle of the back. rugose, pericarp free from seed coat. Leaf epider-
Leaf epidermal papillae present, one or two mal papillae one per cell. 2n ¼ 20, 36, 40, 48.
per cell. Thirteen spp., Natal (South Africa) to north
One sp., P. cornucopiae L. f., Southern India, India, Australia, one species a cosmopolitan weed.
Sri Lanka.
Watson and Dallwitz (1992 onward) indicate 605. Neobouteloua Gould
that the lemmas are spirally arranged in the spike-
let; if correct this phenotype is unique within the Neobouteloua Gould, Bol. Soc. Argent. Bot. 12: 106–108
Poaceae. Pommereulla is placed here in Cynodon- (1968).
Perennials, less than 30 cm tall. Ligule a fringed et al. (2012 and onward) restrict Boutelouinae to
membrane or a fringe of hairs. Inflorescence of the genus Bouteloua itself, and split the clade into
unbranched branches, on an elongated axis. Spi- several other subtribes, several of which are mono-
kelets with one fertile flower and a cluster of 2 to 4 generic. Their classification results in morphologi-
rudimentary ones. Glumes shorter than the flow- cally diagnosable units but does not then reflect
ers, the apex of the lower one muticous or mucro- the relationship among them.
nate, the upper one acuminate or setaceous.
Callus of flower pubescent. Lemma (both fertile
and sterile) pubescent with 3 awns, the median 607. Allolepis Soderst. & H. F. Decker
one recurved. Pericarp fused to seed coat. Leaf Allolepis Soderst. & H. F. Decker, Madroño 18: 34 (1965).
epidermal papillae one per cell.
Two spp., Argentina and Chile. Dioecious perennials, rhizomatous and stolonif-
erous. Ligule a fringed membrane. Inflorescence
606. Odyssea Stapf branches unbranched, borne along a central axis
and appressed to it. Spikelets laterally com-
Odyssea Stapf, Hooker’s Icon. Pl. 31: t. 3100 (1922). pressed, with 8 to 9 flowers and additional
reduced flowers distally. Glumes shorter than
Rhizomatous perennials, sometimes suffrutes- the spikelet, their apices acute. Lemma of female
cent. Ligule a fringe of hairs. Leaf blades hard, spikelet with wide hyaline margins. Palea convo-
stiff, sharp-pointed, disarticulating from the lute around gynoecium; keels winged. Leaf epider-
sheaths. Inflorescence branches unbranched. Spi- mal papillae absent. 2n ¼ 40.
kelets with 4 to 10 fertile flowers plus additional One sp., A. texana Soderst. & H. F. Decker,
distal sterile ones. Glumes shorter than the flow- southern USA and Mexico.
ers, thinner than the lemmas; proximal glume
apex acute, distal glume apex obtuse. Callus of
flower pubescent with straight hairs ca. ¼ the 608. Bouteloua Lag. Fig. 88
length of the flower. Lemma apex with two
Bouteloua Lag., Varied. Ci. 2, 4: 134 (1805), nom. conserv.
teeth, mucronate. Pericarp free from seed coat. Chondrosum Desv., Nouv. Bull. Sci. Soc. Philom. 2: 188
Leaf epidermal papillae absent. 2n ¼ 36. (1810).
Two spp., Africa. Pentarrhaphis Kunth, Nov. Gen. Sp. (quarto ed.) 1:
177–178, t. 60 (1815).
XXIX.1. SUBTRIBE BOUTELOUINAE Stapf (1917) Opizia J. Presl, Reliq. Haenk. 1(4–5): 293, pl. 41, f. 1–11
(1830).
Cathestecum J. Presl, ibid.: 294, t. 42 (1830).
Scleropogoninae Pilg., Die Nat. Pflanzenfam., ed. 2, 14d: Buchloe Engelm., Trans. Acad. Sci. St. Louis 1: 432 (1859),
167, 49 (1956). nom. conserv.
Muhlenbergiinae Pilg. ibid.: 168, 67 (1956). Pringleochloa Scribn., Bot. Gaz. 21: 137 (1896).
Monanthochloinae Pilg. ex Potztal, Willdenowia 5: 472 Cyclostachya Reeder & C. Reeder, Bull Torrey Bot. Club
(1969). 90: 195 (1963).
Hilariinae P. M. Peterson & Columbus, Aliso 23: 592 Buchlomimus J. & C. Reeder & Rzedowski, Brittonia 17: 29
(2007). (1965).
Traginae P. M. Peterson & Columbus, Aliso 23: 592 (2007). Soderstromia C.V. Morton, Leafl. W. Bot. 10: 327 (1966).
Annuals or perennials. Ligule generally a fringed Griffithsochloa G.J. Pierce, Bull. Torrey Bot. Club 105: 134
membrane. Inflorescences generally unbranched (1978).
or with unbranched branches.
Sixteen genera, 287 spp. Annuals or perennials, caespitose, rhizomatous
This clade appears with moderate support in or stoloniferous; plants hermaphrodite, monoe-
the analyses of GPWG II (2012) and is strongly cious or dioecious. Ligule membranous, or a
supported by Peterson et al. (2014a). Although fringed membrane or a fringe of hairs. Inflores-
there is no morphological synapomorphy, many cence unbranched or with unbranched branches,
members have inflorescences unbranched, or with on an elongate or contracted axis, the branches
unbranched branches; many members are also deciduous or persistent, with a naked tip, the
dioecious. As recognized here, Boutelouinae are latter straight or forked. Each branch with 1 to
much broader than in other classifications. Soreng 10 spikelets. Spikelets laterally or dorsiventrally
awns. Flowers bisexual or unisexual. Leaf epider-

mal papillae absent. Microhairs with an elongated
apical cell (“panicoid type”), rarely with a short
broad apical cell (“chloridoid type”). 2n ¼ 20, 40,
41, 42, 56, 60, 70, 98.
Fifty-eight spp., Canada to Argentina.
Molecular phylogenetic data (Columbus et al.
1998) show that Bouteloua in its traditional sense
is paraphyletic, with multiple monotypic genera
segregated from within it. Accordingly, Colum-
bus (1999) expanded the circumscription of Bou-
teloua to include the entire clade. Subsequent
molecular studies have reinforced the validity of
this conclusion. Bouteloua could be placed in its
own subtribe, Boutelouinae Stapf, but this then
forces recognition of several other monogeneric
subtribes; here I suggest a more broadly circum-
scribed Boutelouinae.
609. Distichlis Raf.
Distichlis Raf., J. Phys. Chim. Hist. Nat. Arts 89: 104

(1819).
Monanthochloë Engelm., Trans. Acad. Sci. St. Louis 1: 436
(1859).
Reederochloa Soderstr. & H.F. Decker, Brittonia 16: 334
(1964).
Rhizomatous or stoloniferous perennials; dioe-

cious. Leaves rigid and distichous. Ligule mem-
branous, with or without a fringe of hairs.
Inflorescence branched or not, or reduced to a
single spikelet. Spikelets laterally compressed,
with 2 to 15 flowers plus additional reduced flow-
ers distally. Glumes present or absent, shorter
than the spikelet. Lemma apex entire, awnless.
Microhairs with basal cell sunken in epidermis.
Subsidiary cells of stomata on the lemma each
bearing a single papilla. Vascular bundles of the
Fig. 88. Chloridoideae-Cynodonteae-Boutelouiinae. Boute- leaf with an even outline, suggesting C4, NAD-ME
loua spp. A–D B. repens. A Flowering culms. B Spikelet.
C Lemma. D Palea. E, F B. radicosa. E Whole plant. photosynthesis. 2n ¼ 38, 40, 42, 72.
F Lemma. (From Barkworth et al. 2003, drawn by C. Roché) Ten spp., nine amphitropical in the New
World, one in Australia.
compressed, subtended by bristles or not, with 1 The inclusion of Reederochloa and Monantho-
fertile flower plus 1 to 3 sterile flowers distally, chloë in Distichlis is supported by multiple char-
these composed only of lemmas with long awns. acteristics of morphology and anatomy, as well as
Lower glume absent to narrow and awl-like to molecular data (Bell and Columbus 2008). Clayton
lanceolate or ovate. Upper glume narrow, acute and Renvoize (1986) suggest that the presence of
to acuminate. Lemma glabrous or sparsely pubes- microhairs with the basal cell sunken in the epi-
cent, the apex acute or 3-lobed, the lobes gener- dermis may be synapomorphic for this genus. Bell
ally ending in awns, often with lobes between the and Columbus (2008) show clearly that this sort of
microhair also occurs in Eragrostis obtusiflora, the rachilla extending beyond the distalmost
and hint that the latter species may be sister to flower as an elongate thorn; glumes absent.
Distichlis rather than part of Eragrostis. Distichlis Lemma margins of pistillate flower fused nearly
could be placed in its own subtribe, Monantho- to the apex. Lodicules absent. Male inflorescence
chloinae Pilg. ex Potztal, but there seems little unbranched; lower glumes reduced or absent.
reason to do so. Leaf epidermal papillae absent.
Two spp., California to Central America.
610. Hilaria Kunth
612. Muhlenbergia Schreb.
Hilaria Kunth, Nov. Gen. Sp. (quarto ed.) 1: 116–117, pl.
37 (1815). Muhlenbergia Schreb., Gen. Pl. 1: 44 (1789).
Pleuraphis Torr., Ann. Lyceum Nat. Hist. New York 1: 148 Aegopogon Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 899
(1824). (1805).
Lycurus Kunth, Nov. Gen. Sp. (quarto ed.) 1: 141 (1815).
Annuals or perennials, caespitose, rhizomatous Pereilema J. Presl, Reliq. Haenk. 1: 233 (1830).
Schedonnardus Steud., Syn. Pl. Glumac. 1: 146 (1854).
or stoloniferous. Leaf blades with a sharp-pointed Chaboissaea E. Fourn., Mexic. Pl. 2: 112. (1886).
apex. Ligule membranous or a fringed mem- Redfieldia Vasey, Bull. Torrey Bot. Club 14: 133 (1887).
brane. Inflorescence branches contracted, Bealia Scribn., True Grasses 104 (1890).
unbranched, spread along a central axis. Spikelets Blepharoneuron Nash, Bull. Torrey Bot. Club 25: 88
3 per branch, crowded together, falling as a unit. (1898).
Schaffnerella Nash, N. Amer. Fl. 17: 141 (1912).
Central spikelet pedicellate, laterally compressed,
with one pistillate or bisexual flower, occasionally
Annuals or perennials. Ligule membranous, cili-
a second staminate or sterile one distally; glumes
ate or not. Inflorescence branched or
shorter than the spikelet, narrow, with 2 or more
unbranched, the branches erect or spreading.
lobes, awned; lemma apex entire to bifid, muti-
Spikelets generally solitary, but borne in 2 s or
cous to awned. Lateral spikelets sessile, dorsiven-
3 s in a few species. Spikelets with one flower,
trally compressed, with 1 to 5 flowers, staminate
sometimes two- or three-flowered, laterally com-
or sterile; glumes flabellate, asymmetric, with 1 or
pressed but not strongly so. Rachilla ending in a
more awns; lemma bilobed, awnless, mucronate
fertile flower or not. Callus glabrous or with short
or awned, the awn from the back. Leaf epidermal
hairs. Glumes awned or not, the lower one occa-
papillae one per cell. 2n ¼ 18, 36, 54, 72.
sionally absent. Lemma with 3 veins, entire or
Ten spp., southern U.S. to Guatemala.
with two teeth, generally with an awn, the latter
In subg. Hilaria the glumes are coriaceous
abaxial, apical or from a sinus; lemmas margins
and fused below, whereas in subg. Pleuraphis
pubescent or not. Leaf epidermal cells with inter-
they are papery and free; the two are strongly
costal papillae present, one per cell (several per
supported as sisters, supporting their combina-
cell in a few species). 2n ¼ 14, 16, 18, 20, 28, 30,
tion into a single genus. Although Hilaria has
32, 40, 42, 60.
been put in a monotypic subtribe by Peterson
One hundred seventy-six spp., most in the
et al. (2010a), it is part of the larger clade here
New World, but a few in Asia.
called Boutelouinae, based on the phylogeny (but
Muhlenbergia (as subtribe Muhlenbergiinae)
not the classification) of Peterson et al. (2012).
has been the subject of several recent phyloge-
netic studies (Columbus et al. 2010; Peterson et al.
611. Jouvea E. Fourn. 2010a, 2010b), which demonstrate that it is clearly
monophyletic. Peterson et al. (2010b) provide
Jouvea E. Fourn.Bull. Soc. Roy. Bot. Belgique 15: 475 (1876). phylogenies that sample about 70 % of the species
and provide strong evidence that Muhlenbergia
Dioecious perennials, stoloniferous. Ligule a as traditionally defined is paraphyletic, so a broad
fringed membrane or a fringe of hairs. Inflores- circumscription is adopted here. Placing the
cences in axillary clusters; pistillate inflorescence genus in its own subtribe requires recognizing
with a spathe, with one spikelet. Spikelet terete, several other monogeneric subtribes within the
with 2 to 5 flowers, these embedded in the rachilla, clade here recognized as Boutelouinae s.l. Clayton
and Renvoize (1986) suggest Muhlenbergia is with three awns, the central one straight. Leaf
allied to Sporobolus, but molecular data do not epidermal papillae absent.
support this placement. Some species can be con- One sp., S. filifolia (E. Fourn.) Airy Shaw,
fused with members of the Pooideae. Mexico.
In their comprehensive phylogenetic study of
Muhlenbergia, Peterson et al. (2010b) describe MONELYTRUM + TRAGUS + POLEVANSIA + WILL-
KOMMIA
five well-supported clades.
Inflorescence branches unbranched,
1. M. subg. Muhlenbergia. This clade includes rhizoma- appressed to an elongated axis. Spikelets dorsiven-
tous plants with leaf blades that are broad and flat,
and inflorescence branches that are branched and trally compressed, with one flower. Rachilla not
upright. All members of the clade share characteristic extended beyond flower. Lower glume obtuse,
C4 PCK leaf anatomy and all species of Muhlenbergia with no obvious veins, the apex unawned.
that have been biochemically typed as C4 PCK fall here. These genera clearly form a clade, based on
Although most members of this clade have conven- molecular (Peterson et al. 2010a, 2012) and mor-
tional spikelet arrangement, several species have
reduced staminate or sterile spikelets associated with phological characters and have been called sub-
each fertile spikelet; these were formerly placed in tribe Traginae. However, relationships among
Aegopogon and Pereilema, but clearly are derived them are poorly supported in molecular phyloge-
from within Muhlenbergia. nies; it is possible that the entire group should be
2. M. subg. Trichochloa Trin. This clade is strongly sup- included in a single genus.
ported but has few if any unique morphological charac-
ters. The plants are stout and caespitose, and the glumes
have one or no veins. Peterson and Herrera Arrieta 614. Monelytrum Hack.
(2001) describe distinctive leaf anatomy with inflated
cells adaxial to the primary vascular bundles. Monelytrum Hack., Verh. Bot. Vereins Prov. Brandenburg
3. M. subg. Clomena (P. Beauv.) Hack. This clade includes 30: 140 (1888).
plants that are caespitose, with leaves having lower
sheaths that are flat and papery at maturity. The
upper glumes have three veins and often three teeth. Stoloniferous perennials. Leaf blades with thick-
4. M. subg. Pseudosporobolus (Parodi) P. M. Peterson. Spe- ened margins and tubercle-based hairs; cordate.
cies of this clade have no evident morphological syna- Ligule a fringed membrane. Inflorescence
pomorphies and some have been segregated as separate branches with long hairs; branches each bearing
genera in the past. Whereas the primary inflorescence 2 to 4 fertile spikelets, and 1 to 3 awn-like struc-
branches are branched in some taxa, in M. paniculata
(formerly Schedonnardus) the primary branches are tures (rudimentary spikelets). Lower glume tiny,
unbranched; the central axis elongates and coils at matu- about 1/10 the length of the upper one. Upper
rity and the inflorescence is dispersed as a unit. Other glume hairy, similar to the lemma in length, long
taxa have spikelets with more than one flower. awned. Lemma with a short apical awn. Leaf epi-
5. M. subg. Bealia (Scribn.) P. M. Peterson. Most mem- dermal intercostal papillae one per cell.
bers of this clade are annuals. All have lemmas with
hairy margins and midveins. One sp., M. leuderitzianum Hack., southwest-
ern Africa.
Muhlenbergia ramulosa (Kunth) Swallen
remains incertae sedis as it is not clearly a mem-
615. Tragus Haller
ber of any of the five groups.
Tragus Haller, Hist. Stirp. Helv. 2: 203 (1768), nom. conserv.
613. Sohnsia Airy Shaw
Annuals or perennials. Ligule a fringed mem-
Sohnsia Airy Shaw, Kew Bull. 18: 272 (1965). brane or a fringe of hairs. Inflorescence branches
disarticulating at maturity, each bearing 1 to 5
Dioecious perennials. Ligule a fringed membrane. spikelets. Upper glume hardened, with prominent
Inflorescence branches branched. Spikelets with veins and rows of hooked prickles. Lemma awn-
2 or 3 flowers plus additional reduced ones dis- less. Leaf epidermal intercostal papillae present
tally. Glumes shorter than the spikelet, with one or absent, one per cell. 2n ¼ 20, 40.
vein, the apex acute. Callus of flower pubescent. Eight spp., Africa, South America, and
Lemmas with lines of hairs on veins and keel; Australia.
POLEVANSIA + WILLKOMMIA the lobes awned at the apex. Lodicules absent.

Ligule a fringed membrane. Inflorescence Anthers 1, 2, or 3. 2n ¼ 14.
branches flat. Lemma awnless or with a short Four spp., U.S.A. and Argentina.
awn from the apex. This genus is strongly supported as mono-
phyletic by molecular data (Peterson et al. 2010a),
616. Polevansia de Winter and has obvious synapomorphies. See also Hun-
ziker and Anton (1979).
Polevansia de Winter, Bothalia 9: 130 (1966).
619. Scleropogon Phil.
Perennials. Spikelets with a short blunt hairy cal-
Scleropogon Phil., Anales Univ. Chile 36: 205 (1870).
lus. Leaf epidermal intercostal papillae absent.
One sp., P. rigida de Winter, Lesotho and
South Africa. Dioecious or monoecious perennials, stolonifer-
ous. Ligule a fringe of hairs. Pistillate inflores-
cence unbranched, dense, head-like; each
617. Willkommia Hack. spikelet subtended by a bract; spikelets cylindri-
cal, the lemmas overlapping. Glumes shorter than
Willkommia Hack., Verh. Bot. Vereins Prov. Branden- the spikelet, their apices acuminate. Lemma mar-
burg 30: 145 (1888).
gins convolute, with three veins, each ending in
an awn more than 5 cm long. Palea with two
Annuals or perennials, stoloniferous or not. Spi-
awns. Lodicules absent. Staminate inflorescence
kelets with a sharp callus. Upper glume longer
with branched branches. Glumes muticous.
than the flowers. Leaf epidermal intercostal papil-
Lemma apex muticous, without awns. 2n ¼ 40.
lae one per cell.
One sp., S. brevifolius Phil., southern USA
Four spp., southern Africa, Texas, introduced
and Mexico, Argentina and Chile.
in Argentina.
BLEPHARIDACHNE + SCLEROPOGON + SWALLENIA 620. Swallenia Soderstr. & H.F. Decker
+ MUNROA + ERIONEURON
Inflorescence narrow, unbranched or with Swallenia Soderstr. & H.F. Decker, Madroño 17: 88 (1963).
branched branches. Plants bisexual or gynomo-
noecious. This clade corresponds to the former Rhizomatous perennials, less than 40 cm tall. Ligule
subtribe Scleropogoninae, which is not recog- a fringe of hairs. Leaf blades stiff, their apices sharp-
nized here because to do so would force recogni- pointed. Inflorescence branches branched. Spikelets
tion of several monogeneric subtribes. laterally compressed, with 3 to 7 flowers plus addi-
tional reduced flowers distally. Glumes shorter than
618. Blepharidachne Hack. the spikelet, their apices acuminate. Lemma pubes-
cent on the veins, the apex mucronate. 2n ¼ 20.
Blepharidachne Hack., Die Nat. Pflanzenfam. II, 2: 126 One sp., S. alexandrae (Swallen) Soderstr. &
(1887). H. F. Decker, California.
Caespitose perennials or annuals, decumbent and MUNROA + ERIONEURON

mat-forming, less than 20 cm tall. Leaf blades Ligule a fringe of hairs. Spikelets laterally
sharp-pointed, the upper ones generally extend- compressed. Callus of flower pubescent. Caryop-
ing beyond the inflorescence. Ligule a fringe of sis with embryo projecting below the base.
hairs or absent. Inflorescence branches branched, These two genera form a clade in molecular
with few spikelets, the peduncles short, sur- phylogenies (e.g., GPWG II 2012).
rounded by subtending leaf sheaths. Spikelets
laterally compressed, with 4 flowers, the proximal
621. Munroa Torr.
two staminate or sterile, the third pistillate or
bisexual, the fourth a rudiment with three awns. Munroa Torr., Pacif. Railr. Rep. 4(5): 158 (1857).
Glumes shorter than the spikelet, mucronate or Dasyochloa Willd. ex Rydb., Agric. Exp. Sta. Agric. Coll.
not. Lemmas hairy on margins, with three lobes, Colorado Bull. 100: 18, 37 (1906).
Stoloniferous annuals or perennials, decumbent 623. Neostapfia Burtt Davy

and less than 15 cm tall. Inflorescence
unbranched or with unbranched branches spread Neostapfia Burtt Davy, Erythea 7: 43 (1899).
along a central axis, with 1 to 3 spikelets, these
nearly sessile; peduncle short, so that the inflo- Inflorescence branches each bearing one spikelet
rescence is enclosed by the leaf sheath. Spikelets so that the whole appears unbranched, with
with 2 to 10 flowers, the terminal spikelet with aborted spikelets at the tip. Spikelets dorsiventrally
2 or 3 flowers, lower spikelets with more. Glumes compressed, with 5 flowers. Glumes absent. Lemma
present or absent, shorter than the spikelet or as flabellate, membranous, truncate. Lodicules min-
long as the flowers, the upper glume mucronate. ute. Leaf epidermal papillae absent. 2n ¼ 40.
Lemma with a tuft of hairs midway up margin, One sp., N. colusana (Burtt Davy) Burtt Davy,
apex entire or with 2 or 4 lobes, awned from sinus California.
(es). Caryopsis translucent. 2n ¼ 14, 16.
Five spp., temperate regions of North and 624. Orcuttia Vasey Fig. 89
South America.
The combination of Munroa and Dasyochloa Orcuttia Vasey, Bull. Torrey Bot. Club 13: 219 (1886).
is based on their morphological similarities; in
addition molecular data place the two as sisters Inflorescence unbranched. Spikelets with 4 to 40
(Columbus et al. 2007; Peterson et al. 2010a). See flowers. Glumes shorter than the spikelet. Lemma
also Valdés-Reyna and Hatch (1997).
622. Erioneuron Nash
Erioneuron Nash, Fl. S.E. U. S.: 143 (1903).
Caespitose or stoloniferous perennials. Leaf

blades with thickened margins. Inflorescence
densely capitate, with branched branches. Spike-
lets laterally compressed. Glumes shorter than or
about as long as the spikelet, their apices acumi-
nate and muticous. Lemma densely pubescent,
entire or bilobed, awned from a deep sinus.
2n ¼ 16, 32.
Three spp., southern U.S.A., Argentina.
XXIX.2. SUBTRIBE ORCUTTIINAE P.M. Petersen and

Columbus (2007); tax.: Reeder (1982)
Annuals. Leaves without ligules, the sheath and

blade intergrading. Spikelets with multiple flow-
ers, the lower ones bisexual, the upper undevel-
oped. Glumes present or absent; lodicules reduced
or absent. Microhairs with rounded, mushroom-
shaped apical cells.
Three genera, 9 spp. Plants of vernal pools in
California.
In addition to the synapomorphies listed
above, Orcuttiinae are strongly supported as
monophyletic by molecular phylogenetic ana- Fig. 89. Chloridoideae-Cynodonteae-Orcuttiinae. Orcut-
tia inaequalis. A Habit. B Flower showing five-lobed
lyses (Columbus et al. 2010; Peterson et al. lemma and palea. C Flowers and glumes. (From Bark-
2010a, 2012). worth et al. 2003, drawn by K. Klitz)
with five lobes or teeth, each ending in a mucro or These genera form a moderately well-sup-
awn. Lodicules absent. Leaf epidermal papillae ported clade in Peterson et al. (2012, 2014a), sister
one per cell. 2n ¼ 24–32. to the rest of the subtribe; they correspond to
Five spp., California and Mexico. Gouiniinae s.s. They do not share any obvious
Some members of this genus are reported to morphological characters.
lack an epiblast on the embryo (Reeder 1982).
626. Gouinia E. Fourn. ex Benth. & Hook. f.
625. Tuctoria Reeder
Gouinia E. Fourn. ex Benth. & Hook. f., Gen. Pl. 3: 1178
Tuctoria Reeder, Amer. J. Bot. 69: 1090 (1982). (1883).
Inflorescence unbranched, with spikelets arranged Caespitose perennials, sometimes several meters
in a dense spiral. Spikelets with 5 to 40 flowers. tall. Ligule membranous or a fringed membrane
Glumes shorter than the spikelet. Lemmas entire or a fringe of hairs. Inflorescence branches
or with tiny teeth; central mucro present or absent. unbranched, borne along a central axis. Spike-
Lodicules minute, fused to palea. Leaf epidermal lets with 2 to 6 flowers plus additional reduced
papillae absent. 2n ¼ 24, 40. flowers distally, or with a rachilla extension.
Three spp., California and Mexico. Glumes shorter than the flowers, the lower one
with a muticous or mucronate apex, the upper
one with a muticous, mucronate or awned apex.
XXIX.3. SUBTRIBE GOUINIINAE P.M. Peterson & Callus of flower pubescent. Lemma with a long
Columbus (2007) awn, the awn apical or from a sinus. Leaf epi-
dermal papillae generally absent. Microhairs
Cteniinae P. M. Peterson, Romaschenko & Herrera with a long narrow apical cell. 2n ¼ 40, 64, 76,
Arrieta, Taxon (2014).
80.
Farragoinae P. M. Peterson, Romaschenko & Herrera
Arrieta, Taxon (2014). Fourteen spp., Central and South America.
Gymnopogoninae P. M. Peterson, Romaschenko & Her-
rera Arrieta, Taxon (2014). 627. Tridentopsis P. M. Peterson
Perotidinae P. M. Peterson, Romaschenko & Herrera
Trichoneurinae P. M. Peterson, Romaschenko & Herrera Tridentopsis P. M. Peterson, Taxon 63: 284 (2014).
Annuals or perennials. Spikelets laterally com- Caespitose perennials. Culm nodes often with
pressed. long hairs. Ligule membranous or a fringed
Seventeen genera, 195 spp. membrane or a fringe of hairs. Inflorescence
This subtribe has no obvious morphological branches branched or not. Spikelets with 5 to
synapomorphies. As circumscribed here, it 15 flowers plus additional reduced flowers dis-
includes the Tridentinae recognized by Peterson tally. Glumes shorter than or almost as long as
et al. (2010a, 2012), and is substantially more the flowers, their apices muticous or mucronate.
inclusive than Gouiniinae of Peterson et al. Callus of flower glabrous or pubescent. Lemma
(2014a). As elsewhere in this subfamily, the deci- with midvein sometimes excurrent; veins and
sion of which clades to recognize at the subtribal keel with dense hairs, forming lines on the
level represents a trade-off between naming back of the lemma. Caryopsis dorsiventrally flat-
inclusive groups that indicate broad relationships tened and concave to folded, with a reticulate
versus naming much smaller groups that have surface. 2n ¼ 40.
morphological synapomorphies. The subtribe as Two spp., southwestern U.S., northern Mex-
recognized here is supported as monophyletic ico, Caribbean.
(Peterson et al. 2014a). The genus Tridentopsis was erected to accom-
modate the former Tridens muticus, which is
GOUINIA + SCHENCKOCHLOA + TRIDENTOPSIS + firmly placed in Gouiniinae by molecular data
TRIPLASIS + VASEYOCHLOA (Columbus et al. 2010; Peterson et al. 2010a,
Palea keels winged or expanded near the base. 2014a). Tridens flavus, a synonym of the type
Rachilla extension present. species of Tridens, is consistently placed with
Pappophorum [Pappophorinae] (Columbus et al. tally. Glumes longer than the flowers, the lower
2010; GPWG II 2012; Peterson et al. 2010a). glume sharply pointed, upper obliquely awned
from the back. Fertile lemma entire or with two
teeth, the awn terminal or subapical. Leaf epider-
628. Triplasis P. Beauv.
mal papillae absent. 2n ¼ 18, 36, 54, 160.
Triplasis P. Beauv., Ess. Agrostogr. 81 (1812). Twenty spp., warm regions of America,
Africa, and Madagascar.
Annuals or perennials. Lower leaf sheaths inflated, Ctenium is strongly supported as monophy-
containing cleistogamous spikelets. Ligule a fringed letic and is placed in a monogeneric subtribe by
membrane or a fringe of hairs. Inflorescence Peterson et al. (2014a). It is sister to Trichoneura.
branches branched. Spikelets with 2 to 5 flowers, Peterson et al. (2014a) speculate that Kampochloa
the upper ones reduced, the rachilla extended (Chloridoideae, incertae sedis) may be derived
beyond the distalmost flower. Glumes shorter from Ctenium, based on morphological similarity.
than the flowers, their apices acute. Callus of flower
pubescent. Lemma with lines of hairs along the 631. Trichoneura Andersson
margins and keel; with two lobes; awned from the
sinus, the awn pubescent. Palea with hairs on keels Trichoneura Andersson., Kongl. Svenska Vetenskapsa-
and at tip. Embryo projecting below base of fruit. kad. Handl., n.s. 1853: 148 (1855).
Leaf epidermal papillae absent. 2n ¼ 40.
Two spp., eastern U.S., Central America. Annuals or perennials. Ligule membranous.
Inflorescence branches unbranched, borne along
a central axis, the spikelets on one side of the
629. Vaseyochloa Hitchc. branch. Spikelets laterally compressed, with 4 to
9 flowers, wedge-shaped. Glumes longer than the
Vaseyochloa Hitchc., J. Wash. Acad. Sci. 23: 452 (1933).
adjacent lemmas, in some species longer than the
spikelet, mucronate or awned. Lemma margins
Caespitose perennials. Ligule a fringed membrane.
with long stiff cilia, the apex mucronate or with
Inflorescence branches branched. Spikelets with 5
a short awn. Fruit flattened dorsiventrally. Micro-
to 10 flowers. Glumes shorter than the flowers.
hairs more or less spherical. 2n ¼ 20.
Callus of flower pubescent. Lemma awnless, with
Eight spp., Africa and Arabia; Texas, Galapa-
7 to 9 veins, the proximal portion pubescent. Palea
gos.
splitting at maturity. Fruit rotund, with style
Trichoneura is strongly supported as mono-
branches persisting as two horns. Leaf epidermal
phyletic and is placed in a monogeneric subtribe
papillae one per cell. 2n ¼ 56, 60, 68.
by Peterson et al. (2014a). It is sister to Ctenium.
One sp., V. multinervosa (Vasey) Hitchc.,
Texas, USA.
CRASPEDORHACHIS + FARRAGO
Spikelets dorsiventrally compressed, with one
Remainder of Gouiniinae s.l. flower. Glumes coriaceous longer than the flower.
Lemma hyaline.
630. Ctenium Panz.
632. Craspedorhachis Benth.
Ctenium Panz., Ideen Revis. Gr€as.: 38, 61 (1813), nom.
conserv. Craspedorhachis Benth., Hooker’s Icon. Pl. 14: t. 1377
(1882).
Annuals or perennials, generally caespitose, some
species aromatic. Ligule membranous. Inflores- Caespitose perennials. Ligule a fringed mem-
cence unbranched, or with unbranched primary brane. Inflorescence branches unbranched, digi-
branches; axes curved, crescentic in section, end- tate. Spikelets enclosed in hollows in the branch
ing in a sterile point. Spikelets laterally com- axis, without a rachilla extension. Lower glume
pressed, with two awned sterile flowers below the asymmetrical about the keel, winged, persistent,
fertile one, plus additional reduced flowers dis- the upper glume deciduous. Lemma with long
hairs on veins, the apex obtuse. Leaf epidermal 635. Leptothrium Kunth
papillae absent. 2n ¼ 27.
Three spp., southern Africa. Leptothrium Kunth, Révis. Gramin. 1: 156 (1829).
Craspedorhachis is monophyletic and strongly Latipes Kunth, Révis. Gramin. 1: 53 (1829).
supported as sister to Farrago by Peterson et al.
(2014a). The chromosome number of 2n ¼ 27 was Perennials with wiry culms. Leaves with or with-
reported for C. rhodesiana Rendle by Moffett and out auricles. Ligule a fringe of hairs. Inflorescence
Hurcombe (1949), who noted that it was triploid. unbranched, or with unbranched branches.
The species may thus be a sterile hybrid. Branches flattened, wedge shaped, bearing 1 or
2 spikelets, deciduous. Spikelets with 1 flower.
Glumes longer than flower, hard, the lower flat,
633. Farrago Clayton recurved, acuminate, the upper enclosing the
flower. Lemma glabrous, awnless. Pericarp not
Farrago Clayton, Kew Bull. 21: 125 (1967). tightly fused to seed coat. Leaf epidermal papillae
one per cell, or none. 2n ¼ 20.
Annuals. Ligule a fringe of hairs. Inflorescence Two spp., Senegal to Pakistan; Caribbean,
branches unbranched, each branch with 1 fertile South America.
spikelet, plus two spikelets reduced to awns.
Glumes awned. Lemma a tiny scale, hairy. Leaf
epidermal papillae absent. 636. Leptocarydion Hochst. ex Stapf
One sp., F. racemosa Clayton, Tanzania.
Farrago is strongly supported as sister to Leptocarydion Hochst. ex Stapf, Fl. Cap. 7: 316 (1898).
Craspedorachis by Peterson et al. (2014a) and
the two genera placed in their own subtribe. Annuals. Ligule a fringed membrane. Inflores-
cence dense, the primary branches crowded on
DIGNATHIA + LEPTOTHRIUM + LEPTOCARYDION + the rachis. Spikelets with 6 to 14 flowers plus
GYMNOPOGON + BEWSIA + LOPHACME additional distal sterile ones. Callus of the flower
This clade is strongly supported in the ana- hairy. Glumes shorter than the spikelet. Lemma
lyses of Peterson et al. (2014a), and could be pubescent, the apex entire or bidentate, awned
recognized at the subtribal level as Gymnopogo- from the apex or from the sinus; stamens 2 or 3.
ninae P.M. Peterson, Romasch. & Y. Herrera. Leaf epidermal papillae several per cell.
Dignathia and Leptothrium are in a very weakly One sp., L. vulpiastrum (De Not.) Stapf, east-
supported clade sister to the other genera (Peter- ern and southern Africa.
son et al. 2014a); their sister relationship should Leptocarydion is strongly supported as sister
be considered provisional. to the clade consisting of Lophacme, Bewsia, and
Gymnopogon (Peterson et al. 2014a).
634. Dignathia Stapf
637. Gymnopogon P. Beauv.
Dignathia Stapf, Hooker’s Icon. Pl. 30: t. 2950 (1911).
Gymnopogon P. Beauv., Ess. Agrostogr. 41, 164 (1812);
Annuals or perennials. Ligule a fringed mem- tax.: Cialdella and Zuloaga (2011).
brane. Inflorescence branches unbranched,
borne along a central axis, deciduous, curved, Perennials (annuals), caespitose or rhizomatous.
bearing 1 or 2 spikelets and a terminal one con- Leaf blades stiffly distichous, apiculate at the
sisting only of glumes. Spikelets laterally com- apex. Ligule a fringed membrane. Inflorescence
pressed, with 1 or 3 flowers. Glumes longer than branches unbranched, digitate or borne along a
the flowers, hard, gibbous, covered with prickles central axis. Spikelets contiguous or distant along
or long hairs, their apices acuminate, muticous or the inflorescence branches, tightly appressed to
awned. Lemma apex with a short awn or mucro. the branches, generally with 1 flower, the proxi-
Leaf epidermal papillae one per cell. mal ones bisexual, the distal ones reduced;
Five spp., East Africa to NW India. rachilla extended, ending in an awn. Glumes
longer than the flowers, acuminate. Lemma PEROTIS + MOSDENIA + TRIGONOCHLOA

minutely two-lobed, muticous or awned from Leaf blades lanceolate to lance-ovate, clasping
sinus. Anthers 2 or 3. Leaf epidermal papillae the culm. Spikelets with 1 flower, rachilla exten-
several per cell. 2n ¼ 40. sion absent. Glumes longer than the flower. Cary-
Fourteen spp., southern USA and South opsis longitudinally grooved.
America; one species in India and SE Asia. These genera fall in a strongly supported clade
Gymnopogon is strongly supported as sister in the analyses of Peterson et al. (2012, 2014a), and
to the Lophacme-Bewsia clade (Peterson et al could be recognized at the subtribal level as Per-
2014a). otidinae P.M. Peterson, Romasch., & Y. Herrera.
The longitudinally grooved caryopsis may be
synapomorphic (Snow and Peterson 2012b).
638. Bewsia Goossens
Bewsia Goossens, S. African J. Sci. 37: 183 (1941). 640. Trigonochloa P. M. Peterson & N. Snow
Caespitose perennials. Ligule membranous. Inflo- Trigonochloa P. M. Peterson & N. Snow, Ann. Bot. 109:
rescence with unbranched branches, these 1327 (2012); tax.: Snow and Peterson (2012b).
distributed along and appressed to the main
axis. Spikelets with 2 to 4 flowers, with additional Annuals or perennials, rhizomatous or stolonifer-
reduced flowers distally. Glumes shorter than the ous, culms sometimes decumbent. Ligule mem-
flowers, the apices acuminate. Callus of flower branous. Inflorescence branches unbranched,
pubescent. Lemma pubescent on keel and mar- spread along the central axis. Spikelets laterally
gins, with an abaxial awn. Leaf epidermal papillae compressed, disarticulating above the glumes.
absent. 2n ¼ 30, 40. Lemma apex entire. Caryopsis trigonous, with a
One sp., B. biflora (Hack.) Goossens, central deep groove on the hilar side. Leaf epidermal
and southern Africa. cells with a single central papilla. 2n ¼ 36.
GPWG II (2012) and Peterson et al. (2012) Two spp., tropical Africa and Asia.
place this sister to Gymnopogon with strong sup- The leaves of Trigonochloa are thinly mem-
port, although the two genera do not have obvi- branous. The primary vascular bundles of T. uni-
ous shared morphological characters. flora have unusually large bundle sheath cells on
either side, a character first observed by Metcalfe
(1960); this character may be a synapomorphy for
639. Lophacme Stapf
the genus (Peterson et al. 2012), but data are
Lophacme Stapf, Fl. Cap. 7: 316 (1898). lacking for T. ruprestis. Micromorphology of the
lemma is distinctive and supports exclusion of
Annuals or perennials. Ligule a line of hairs, or Trigonochloa from Leptochloa; silica cells and
membranous, or lacking. Leaf blades stiff. Inflo- epidermal papillae are lacking in Trigonchloa.
rescence branches unbranched, digitate. Spikelets
MOSDENIA + PEROTIS
wedge-shaped, with 1 or 2 fertile flowers and distal
Inflorescence unbranched. Disarticulation
sterile flowers reduced to awns, these dispersed
below the glumes.
with the upper flower. Callus of flower pubescent.
Mosdenia and Perotis are sisters in the ana-
Glumes longer or shorter than the spikelet. Lemma
lyses of Peterson et al. (2014a).
hyaline or thinly membranous, the apex with two
teeth, with a long awn from the sinus. Pericarp
easily separated from seed coat. Leaf epidermal 641. Mosdenia Stent
papillae absent. Microhairs with an apical cell
that is long and narrow, or about as broad as long. Mosdenia Stent, Bothalia 1: 170 (1923).
Peterson et al. (2012, 2014a) place Lophacme Perennials, rhizomatous or stoloniferous, the sto-
in a well-supported clade with Gymnopogon and lons with overlapping scale leaves. Ligule a fringed
Bewsia. membrane. Spikelets densely crowded along the
rachis, spreading, dorsiventrally compressed. Plectrachne Henrard, Vierteljahrsschr. Naturf. Ges.

Glumes with apices acuminate, awnless. Lemma Z€
urich. 74: 132 (1929).
delicate, the apex obtuse, awnless, without hairs.
Leaf epidermal papillae absent. 2n ¼ 40. Spikelets with 4 to 15 flowers. Glumes shorter than
One sp., M. leptostachys (Ficalho & Hiern) flowers. Lemmas becoming hard, with three teeth
Clayton, South Africa. or lobes. Leaf epidermal papillae absent abaxially.
Sixty-six spp., Australia.
642. Perotis Aiton Fig. 90
644. Monodia S. W. L. Jacobs
Perotis Aiton, Hort. Kew. 1: 85 (1789).
Lopholepis Decne., Arch. Mus. Hist. Nat. 1: 147 (1839), Monodia S. W. L. Jacobs, Kew Bull. 40: 659 (1985).
nom. nov. pro seq. Holboellia Hook.
Toliara Judz., Adansonia III, 31: 274 (2009). Spikelets with one flower; rachilla extended beyond
the flower. Lemma convolute, with two lobes,
Annuals or perennials, most caespitose. Leaf awned from the sinus, with a line of hairs on each
blades stiff, with a sharp apex. Ligule membra- margin. Leaf epidermal papillae one per cell.
nous or a fringe of hairs. Inflorescence sometimes One sp., M. stipoides S. W. L. Jacobs, Western
partially enclosed by the subtending leaf. Spike- Australia.
lets more or less terete. Glumes longer than the
flowers, with long awns. Lemma apex entire, awn-
less. 2n ¼ 36, 40. 645. Symplectrodia Lazarides
Fifteen spp., Old World tropics.
Symplectrodia Lazarides, Nuytsia 5: 273 (1985).
Both Lopholepis and Toliara are derived from
within Perotis (Peterson et al. 2014a). The place-
ment of Toliara was predicted by Judziewicz Spikelets with only one flower fertile, the others
(2009) on the basis of morphology; the two sterile and clumped; rachilla internodes unusu-
genera are quite similar, but P. (¼ T.) arenacea ally long. Lemma entire, with an apical awn; palea
lacks the distinctive awned glumes of Perotis. The fused to rachilla below. Leaf epidermal papillae
spikelet of P. (¼ L.) ornithocephala is described one to several per cell.
as “one of the oddest shaped spikelets in the Two spp., Australia.
Gramineae” (Clayton and Renvoize 1986); the
upper glume is constricted so as to resemble a
cartoon of a bird’s head. XXIX.5. SUBTRIBE ELEUSININAE Dumort. (1829)
Annuals or perennials. Inflorescence

XXIX.4. SUBTRIBE TRIODIINAE Benth. (1881) unbranched, or with primary branches only, or
with branched branches. Spikelets laterally or
Perennials with hard, needle-like, sharp-pointed dorsiventrally compressed or nearly terete.
leaves. Ligule a fringe of hairs. Inflorescence Lemma with 3 veins. Pericarp fused to or free
branches branched. Spikelets laterally com- from seed coat. Leaf epidermal papillae generally
pressed. Mesophyll “Triodia-type” with PCR tis- present, generally one per cell.
sue only on sides of major bundles and draping Thirty genera, 229 species.
over minor bundles. This subtribe has no obvious morphological
Three genera, seventy species. synapomorphy, but is well supported by molecular
This subtribe is sufficiently cohesive that all data (GPWG II 2012; Peterson et al. 2010a, 2012).
genera could easily be placed in a single genus.
646. Acrachne Wight & Arn. ex Chiov.
643. Triodia R. Br. Fig. 91 Acrachne Wight & Arn. ex Chiov., Annuario Reale Ist. Bot.
Roma 8: 361 (1908).
Triodia R. Br., Prodr.: 182 (1810).
Annuals. Ligule membranous or a fringed mem-

brane. Inflorescence branches unbranched, with
overlapping spikelets, digitate or spread along a
central axis. Spikelets laterally compressed, with
2 to 25 flowers, with additional reduced flowers
distally. Glumes shorter than the spikelet, the
lower one mucronate, the upper one awned.
Lemma awned from the apex, deciduous; paleas
persistent; rachilla tough. Pericarp free from the
seed coat. Leaf epidermal papillae absent.
Three spp., Old World tropics.
647. Afrotrichloris Chiov.
Afrotrichloris Chiov., Ann. Bot. (Rome) 13: 371 (1915).
Perennials. Ligule a fringed membrane. Inflores-

cence unbranched. Spikelets with one fertile
flower and a cluster of sterile awned lemmas.
Glumes divergent, acuminate. Lemmas hairy,
deeply divided into two lobes, awned from
sinus. Pericarp free from seed coat. Leaf epider-
mal papillae one per cell.
Two spp., Somalia.
648. Astrebla F. Muell.
Astrebla F. Muell., Fl. Austral. 7: 602 (1878).
Caespitose perennials. Ligule a fringed membrane

or a fringe of hairs. Inflorescence unbranched or
with two branches. Spikelets laterally compressed
or nearly terete, with 1 to 6 flowers plus several
reduced ones distally. Glumes shorter than or
about the same length as the spikelet, their apices
muticous or mucronate. Callus of flowers pubes-
cent. Lemmas with silky hairs, three-lobed, the
central lobe or all three lobes awned. Pericarp
free from seed coat. 2n ¼ 40.
Four spp., Australia.
649. Austrochloris Lazarides
Austrochloris Lazarides, Austral. J. Bot., Suppl. Ser. 5: 33

(1972).
Caryopsis. (From Wu et al. 2007, with permission from

Fig. 90. Chloridoideae-Cynodonteae-Gouiniinae. Perotis the Missouri Botanical Garden Press, St. Louis, and Sci-
rara. A Habit. B Spikelet showing long awns on glumes. C ence Press, Beijing; drawn by Shi Weiqing)
ile lemma; rachilla hairy. Glumes much longer

than flower, divergent, their apices acute to acu-
minate. Callus of flower pubescent. Fertile lemma
with an obtuse apex, with subapical awn. Pericarp
free from seed coat.
One sp., A. dichanthoides (Everist) Lazarides,
Australia.
650. Brachyachne (Benth. & Hook. f.) Stapf
Brachyachne (Benth. & Hook. f.) Stapf, Hooker’s Icon. Pl.

31: t. 3099 (1922).
Annuals or perennials, stoloniferous or not, often

decumbent or prostrate. Ligule membranous, or a
cence unbranched, or of digitate unbranched
branches, these flattened and hollowed, sur-
rounded by a leaf sheath or spatheole. Spikelets
laterally compressed, with one flower and a
rachilla extension. Glumes longer than the flower
and enclosing it, broad, with obtuse apices. Callus
of flower pubescent or glabrous. Lemma apex
acute to bidentate, muticous or mucronate. Peri-
carp fused to seed coat.
Ten spp., wet areas of tropical Africa, south-
east Asia, Australia.
651. “Chloris” Sw.
“Chloris” Sw., Prod. 1, 25 (1788).

Pterochloris (A. Camus) A. Camus, Bull. Mus. Hist. Nat.
(Paris) 29: 349 (1957).
Annuals or perennials, caespitose or stolonifer-

ous. Ligule membranous, or a fringed mem-
brane, or a fringe of hairs. Inflorescence
unbranched, or more often with digitate pri-
mary branches. Spikelets laterally compressed,
with 1 or 2 flowers plus additional reduced
flowers distally, or with a naked rachilla exten-
Fig. 91. Chloridoideae-Cynodonteae-Triodiinae. Triodia sion. Callus of flower glabrous or more often
scariosa. A Habit. B Inflorescence. C–G All adaxial view. C pubescent. Glumes shorter or longer than the
Apex of lower glume. D Lower glume. E Upper glume. F spikelet, their apices muticous or mucronate.
Lemma. G Palea. H Ligule. (From Jessop et al. 2006) Lemma with a prominent keel, muticous or
awned from tip or abaxial side or a sinus. Leaf
Perennials. Ligule a fringe of hairs. Inflorescence epidermal papillae several per cell. Pericarp free
with 2 or 3 unbranched branches, the branch axes from or fused to seed coat. 2n ¼ 14, 20, 26, 30,
hollowed and flattened. Spikelets dorsiventrally 36, 40, 72, 80.
compressed, with one fertile flower plus one ster- Sixty-three spp., warm regions of the world.
Peterson et al. (2010a) show that four species Peterson et al. (2010a) show an unresolved
of Chloris form a clade, whereas Columbus et al. relationship among species of Cynodon and those
(2010) suggested polyphyly for “Chloris”, based of Brachyachne. Species limits and crossability of
on a sample of three species, all different from the species were investigated by Harlan et al. (1969).
ones that Peterson et al. (2010a) investigated.
Both studies show that Trichloris should be
654. Daknopholis Clayton
excluded from “Chloris”. Although Daknopholis
is placed Incertae sedis in this treatment, it may Daknopholis Clayton, Kew Bull. 21: 102 (1967).
perhaps be included in “Chloris”, with which it
shares many morphological characters. Ptero- Stoloniferous annuals. Ligule a fringe of hairs.
chloris is included in “Chloris” here, following Inflorescence branches unbranched, flattened,
Clayton and Renvoize (1986). borne on a contracted axis. Spikelets with one
flower, rachilla extension short. Glumes shorter
652. Chrysochloa Swallen than lemmas, unawned, papery, their apices
obtuse to truncate. Lemma two-lobed, awned or
Chrysochloa Swallen, Proc. Biol. Soc. Wash. 54: 44 (1941). sometimes awnless. Pericarp free from seed coat.
Leaf epidermal papillae several per cell.
Annuals or perennials, caespitose or stolonifer- One sp., D. boivinii (A. Camus) Clayton,
ous. Leaves with blunt apices. Ligule a fringed Indian Ocean islands, Madagascar, and eastern
membrane. Inflorescence unbranched, or of coast of Africa.
flattened unbranched branches. Spikelets later- Daknopholis is sometimes placed in synon-
ally compressed, with 2 flowers, the lower ymy under “Chloris”.
bisexual, the upper staminate or sterile. Glumes
about the same length as lemmas; lower glume 655. Dinebra Jacq.
persistent, the apex acute; upper glume decidu-
ous, the apex emarginate or acute, muticous to Dinebra Jacq., Fragm. Bot. 77, pl. 121, f. 1 (1809); phylog.,
awned. Lemma midvein and margins pubes- tax.: Peterson et al. (2012); Snow and Peterson (2012a).
cent, with a short point or awn from near the Drake-brockmania Stapf, Bull. Misc. Inform. Kew 1912:
top of the abaxial side. Pericarp fused to seed 197 (1912).
Heterocarpha Stapf & C. E. Hubb., Bull. Misc. Inform. Kew
coat. 2n ¼ 14. 1929: 263 (1929).
Four spp., tropical Africa.
Annuals or perennials, caespitose, stoloniferous
653. Cynodon Rich. or not. Leaf blades sometimes with multicellular
glands abaxially. Ligule membranous or a fringed
Cynodon Rich., Syn. Pl. 1: 85 (1805) nom. conserv.; phy- membrane. Inflorescence branches unbranched,
log.: Jewell et al. (2012). sub-digitate or borne along a central axis, often
deciduous. Spikelets overlapping on the
Annuals or perennials, caespitose, rhizomatous or branches, laterally compressed, with 1 to 18 flow-
stoloniferous. Ligule membranous or a fringed ers, the distal ones reduced. Glumes shorter or
membrane. Inflorescence branches unbranched, longer than flowers, the apex acute, acuminate or
flattened, borne on a short central axis. Spikelets extended into a long tail. Lemma pubescent
laterally compressed, with 1 flower; rachilla exten- on veins, veins sometimes 5 or 7, the apex mucro-
sion present or absent; distal incomplete flowers nate or not. Caryopsis trigonous. Pericarp free
reduced or absent. Glumes shorter than or about from or fused to seed coat. Leaf epidermal papil-
the same length as the spikelet, their apices acute lae present, one per cell. Glandular microhairs
or acuminate, muticous or mucronate. Lemma present. 2n ¼ 20.
apex obtuse or acute, awnless. Leaf epidermal Twenty-seven spp., temperate to tropical
papillae one to several per cell. Pericarp fused to areas nearly worldwide.
seed coat. 2n ¼ 16, 18, 27, 36, 40, 54. Molecular data place Dinebra in a clade with
Twelve spp., Old world tropics, one species 24 species formerly placed in Leptochloa (Colum-
pantropical. bus et al. 2007; Peterson et al. 2010a, 2012), plus
Fig. 92. Chloridoideae-Cynodonteae-Eleusininae. Diplachne fusca. A Habit. B Ligule. C Spikelet. D Lower glume. E
Upper glume. F Lemma. G Palea. H Flower with separate lodicules. I Caryopsis. (From Cope 1982)
the monospecific Drake-brockmania and Hetero- long as the flowers, their apices acute. Lemma
carpha. Phillips (1974) includes Heterocarpha often with a dark spot near the base, the apex
haareri in Drake-brockmania on the basis of “an acute to truncate or bifid, mucronate to awned.
inflorescence of short, broad, deciduous spikes.” 2n ¼ 20.
The number of veins in the upper glume varies in Two spp., temperate to tropical areas
Dinebra from 1 in species formerly classified in throughout the world.
Leptochloa, to as many as 17 in the former Drake-
Brockmania somalensis.
657. Disakisperma Steud.
656. Diplachne P. Beauv. Fig. 92 Disakisperma Steud., Syn., Pl. Glumac. 1: 287 (1854);
phylog., tax.: Peterson et al. (2012), Snow and Peterson
Diplachne P. Beauv., Agrostogr. 80, pl. 16, f. 9 (1812); (2012a), Snow et al. (2013).
phylog., tax.: Peterson et al. (2012), Snow and Peterson
(2012a). Perennials. Ligule membranous, truncate, erose
or ciliate. Inflorescence with unbranched
Annuals or perennials, caespitose or rhizoma- branches, these subdigitate or spread along a
tous. Ligule membranous, relatively long and central axis. Spikelets with 4 to 13 flowers.
acute. Inflorescence with unbranched branches, Glumes shorter than the flowers, their apices
these spread along a central axis. Spikelets with 6 acute to obtuse. Lemma entire or bifid, unawned
to 20 flowers. Glumes shorter than or about as or mucronate. Caryopsis flattened dorsiventrally,
weakly concave; pericarp weakly fused to the seed the rachilla terminating in a club-shaped lemma.
coat. 2n ¼ 40, 60, 80. Glumes shorter or longer than the spikelets, their
Four spp., North and South America, Africa. apices obtuse to acute, the upper one often with a
Disakisperma species have lemmas with subapical awn. Callus of flower glabrous to
macrohairs that are somewhat inflated at the tip pubescent. Lemma mucronate or awned, dark
(“clavicorniculate”) (Snow 1996, and pers. brown at maturity. Pericarp fused to seed coat.
comm.). Based on this character and molecular Leaf epidermal papillae one or several per cell.
data, Coelachyrum yemenicum (Schweinf.) S.M. 2n ¼ 40.
Phillips has recently been transferred to Disakis- Sixteen spp., tropics, mostly New World.
perma, as D. yemenica (Chiov.) P. M. Peterson & E. petraea (the type species) and E. paspa-
N. Snow. loides are sisters in the study of Peterson et al.
(2010a), indicating that some component of this
genus is monophyletic. However, E. disticho-
658. “Enteropogon” Nees
phylla is unrelated.
“Enteropogon” Nees, Intr. Nat. Syst. Bot.: 448 (1836).
660. Harpochloa Knuth
Annuals or perennials. Ligule membranous, or a
fringed membrane, or a fringe of hairs. Inflores- Harpochloa Knuth, Révis. Gramin. 1: 92 (1829).
cence unbranched, or of unbranched branches,
these digitate or spread along a central axis. Spi- Perennials. Leaves stiff. Ligule a fringed mem-
kelets laterally or dorsiventrally compressed, with brane. Inflorescence unbranched, ending in a
one fertile flower plus additional reduced flowers bare tip, the axis strongly curved. Spikelet later-
distally, the latter forming a clump (rudiment). ally compressed, with one fertile flower plus an
Glumes shorter or longer than the spikelet, their additional reduced flower distally. Glumes as long
apices muticous or mucronate, the upper one as the spikelet, with acute apices, the lower one
sometimes awned. Lemma indurate, the veins persistent, the upper deciduous. Lemma pubes-
forming prominent ridges, the apex entire or cent, awnless, the apex notched. Pericarp fused to
bifid, awned from the apex or from a sinus. seed coat. Adaxial mesophyll cells colorless and
Fruit concavo-convex in section, with a longitudi- interrupting the outer bundle sheath. 2n ¼ 40.
nal groove. Pericarp free from seed coat. 2n ¼ 20. Two spp., Central and southern Africa.
Seventeen spp., tropics. Roodt and Spies (2003) also report 2n ¼ 36
“Enteropogon” is either para- or polyphyletic, and 54 for Harpochloa.
but taxon sampling in existing molecular trees is
insufficient to draw any conclusions (Columbus
et al. 2007; Peterson et al. 2010a); the type species 661. Leptochloa P. Beauv.
(E. melicoides) has not been sampled. Peterson
Leptochloa P. Beauv., Ess. Agrostogr.: 71 , pl. 15, f. 1
et al. (2010a) indicate that Saugetia is unrelated to (1812); phylog.: Peterson et al. (2012), Snow and Peterson
the sampled species of “Enteropogon”. (2012a).
659. “Eustachys” Desv. Perennials, caespitose, rhizomatous or stolonifer-

ous. Ligule short, membranous. Inflorescence
“Eustachys” Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: branches unbranched, sub-digitate or spread
188 (1810). along a central axis. Spikelets laterally com-
pressed, with 1 to 7 flowers plus additional
Perennials, caespitose, short-rhizomatous, or sto- reduced flowers distally. Glumes shorter or lon-
loniferous. Ligule a fringed membrane or a fringe ger than the flowers, generally muticous. Callus of
of hairs. Inflorescence unbranched, or with flower pubescent or glabrous. Lemma apex awn-
unbranched branches, the branch axes flattened less, or with 1 to 3 awns. Fruit smooth, pericarp
or hollowed. Spikelets laterally compressed, with free from, adherent to or fused to seed coat.
1 flower plus additional reduced flowers distally, 2n ¼ 20, 40, 60.
Five spp., southern U.S. to South America, nia sister to (Peterson et al. 2010a) or within
Australia. (GPWG II 2012) “Chloris”.
Molecular data show that Leptochloa as tradi-
tionally circumscribed is polyphyletic (Columbus
664. Microchloa R. Br.
et al. 2007; Peterson et al. 2010a, 2012), consistent
with previous data from morphology and anat- Microchloa R. Br., Prodr. 208 (1810).
omy (Snow 1997). Monophyly is restored once Rendlia Chiov., Ann. Bot. (Rome) 13: 53 (1914).
Dinebra, Diplachne and Disakisperma are re-
defined, and Trigonochloa is recognized. Annuals or perennials. Ligule a fringed mem-
brane. Inflorescence unbranched, curved, cres-
662. Lepturus R. Br. centic in outline, hollowed, the spikelets borne
on one side. Spikelets dorsiventrally compressed,
Lepturus R. Br., Prodr.: 207 (1810). with one fertile flower, in some species with addi-
tional reduced flowers distally. Callus of flowers
Annuals or perennials, caespitose or stolon- pubescent. Glumes extending beyond the flowers,
iferous. Leaf blades with or without a pseudo- their apices obtuse or acute. Lemma acute to
petiole, with sharp-pointed apices. Ligule bilobed, sometimes with a short mucro. Pericarp
membranous or a fringed membrane or absent. fused to seed coat. Leaf epidermal papillae absent.
Inflorescence unbranched, disarticulating at 2n ¼ 40.
nodes; spikelets embedded in the rachis. Spikelets Six spp., Africa, 1 pantropical.
dorsiventrally compressed, with one or two flow- Clayton and Renvoize (1986) place Rendlia
ers, with additional reduced flowers distally or altera (Rendle) Chiov. in synonymy with Micro-
with a sterile rachilla extension. Lower glume chloa, whereas Watson and Dallwitz (1992
minute or absent; upper glume covering the onward) exclude it.
spikelet, muticous or awned. Pericarp free from
seed coat. Leaf epidermal papillae several per cell. 665. Oxychloris Lazarides
Microhairs short and stout, spherical. 2n ¼ 14,
18, 26, 36, 42, 52, 54. Oxychloris Lazarides, Nuytsia 5: 283 (1985).
Sixteen spp., beaches of Indian and Pacific
oceans. Annuals or perennials. Ligule a fringed mem-
brane. Inflorescence with 3 to 6 unbranched
663. Lintonia Stapf branches, these digitate, flattened and hollowed.
Spikelets laterally compressed, with one flower
Lintonia Stapf, Hooker’s Icon. Pl. 30: t. 2944 (1911). plus additional reduced flowers distally, the callus
long, sharp, and pubescent. Glumes shorter than
Stoloniferous perennials. Ligule membranous. the spikelet, the lower one with an obtuse apex,
Inflorescence with several unbranched branches, the upper with an emarginate apex. Lemma of
digitate or along a short central axis. Primary fertile flower with tufts of hairs, winged, with a
branches slender, sinuous. Spikelets laterally subapical awn; lemmas of sterile flowers broadly
compressed, with 4 to 10 flowers plus several expanded, flabellate. Lodicules absent. Pericarp
reduced ones distally. Callus below proximal fused to seed coat.
flower pubescent. Glumes shorter than the spike- One sp., O. scariosa (F. Muell.) Lazarides,
let. Lemmas with clavate hairs on sides, bilobed Australia.
with prominent hardened veins extending into
the lobes, awned from back, branching. Lodicules 666. Rheochloa Filg., P. M. Peterson & Y. Herrera
cylindrical. Androecium and gynoecium on a
short stipe. Pericarp free from seed coat. 2n ¼ 30. Rheochloa Filg., P.M. Peterson & Y. Herrera, Syst. Bot. 24:
Two spp., Eastern Africa. 123 (1999).
Clayton and Renvoize (1986) note similarity
between the lemma venation in this genus and Decumbent perennials. Ligule a fringed mem-
that in Astrebla, but molecular data place Linto- brane. Inflorescence branches unbranched,
digitate. Spikelets laterally compressed, with 669. Sclerodactylon Stapf

three flowers, the proximal two fertile and the
distal one reduced. Glumes longer than the flow- Sclerodactylon Stapf, Bull. Misc. Inform. Kew 1911: 318
ers, coriaceous, the apices acute. Lemma hyaline, (1911).
with an obtuse ciliate apex. Pericarp fused to seed
coat. Leaf epidermal papillae several per cell. Stoloniferous perennials. Ligule a line of hairs. Leaf
One sp., R. scabriflora Filg., P.M. Peterson & blades woody, elliptical or circular in cross section,
Y. Herrera, Brazil. with sharp tips. Inflorescence branches two or
more, unbranched. Spikelets laterally compressed,
with 5 to 40 flowers plus additional reduced flowers
667. Saugetia Hitchc. & Chase distally. Glumes shorter than the spikelet, their
apices acute, the upper glume mucronate. Palea
Saugetia Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 378 winged. Pericarp free from seed coat.
(1917).
One sp., S. macrostachyum (Benth.) A.
Camus, Madagascar, islands of the Indian
Perennials. Leaf blades dominated by the midrib,
Ocean, east African coast.
the lateral flattened portion narrow. Ligule a
fringed membrane. Inflorescence unbranched,
or of digitate unbranched branches. Spikelets 670. Tetrapogon Desf.
partially embedded in the inflorescence axis, dor-
siventrally compressed, with one flower plus an Tetrapogon Desf., Fl. Atlant. 2: 388 (1799).
additional reduced flower distally, with elongated
rachilla internodes above glumes and between Annuals or perennials, caespitose or short rhizo-
flowers. Callus of spikelet long, pubescent. matous. Ligule a fringed membrane. Inflorescence
Glumes shorter than the spikelet, the proximal unbranched, or with 2 or 3 unbranched branches.
one much smaller than the distal one, their apices Spikelets laterally compressed, wedge-shaped,
acuminate. Lemma two toothed, awned from the with 1 to 7 flowers plus additional reduced flowers
sinus. Pericarp free from seed coat. Leaf epider- distally. Callus of flower pubescent. Glumes
mal papillae absent. shorter than to longer than the spikelet, muticous
Two spp., West Indies. or mucronate. Lemma entire or two-toothed, with
Peterson et al. (2010a) place S. fasciculata long hairs on veins and keel, with a subapical awn.
sister to Tetrapogon, contradicting the decision Pericarp free from seed coat or not. 2n ¼ 20.
of Clayton et al. (2006 onward) and Clayton and Five spp., Africa, Middle East and India.
Renvoize (1986) to sink Saugetia into “Enteropo-
gon”.
671. Trichloris E. Fourn. ex Benth.
668. Schoenefeldia Kunth Trichloris E. Fourn. ex Benth., J. Linn. Soc. Bot. 19: 102
(1881).
Schoenefeldia Kunth, Révis. Gramin. 1: 86 (1829).
Perennials, stoloniferous or not. Ligule a fringe of
Annuals or perennials. Ligule a fringed mem- hairs. Inflorescence of unbranched branches, dig-
brane. Inflorescence of unbranched branches, itate. Spikelets dorsiventrally compressed, with 1
digitate. Spikelets laterally compressed, with one or 2 flowers plus an additional reduced flower
flower, or with a rachilla extension ending in an distally. Callus of flower pubescent. Glumes
awn. Callus of flower pubescent. Glumes narrow, shorter than the spikelet, their apices acuminate,
setaceous, longer than flower, acuminate or the upper one awned. Lemma with two teeth,
awned. Lemma with two teeth, awned from awned from sinus and both teeth. Palea keels
sinus, the awns from adjacent lemmas entangled. prolonged. Fruit concavo-convex in section. Peri-
Pericarp free from seed coat. carp free from seed coat.
Two spp., Africa, Madagascar and India. Two spp., South America.
The fruit and pericarp characters suggest that reduced flowers distally. Glumes shorter than the
this genus might belong in the following clade. spikelet, their apices obtuse to acute, muticous or
mucronate, or the upper awned. Lemma glabrous
APOCHITON + COELACHYROPSIS + COELACHYRUM or with hairs or prickles on margins and keel, the
+ ELEUSINE apex muticous to mucronate. Leaf epidermal
Fruit trigonous or concavo-convex, rugose or papillae absent. Microhair apical cell short and
sculpted in all but A. burtii, the pericarp free from broad. Macrohairs crispate.
the seed coat. Four spp., Africa, Arabia, Pakistan.
Molecular data are not available for the type
672. Apochiton C. E. Hubb. species, C. brevifolium Hochst. & Nees; however,
it has crispate macrohairs, which it shares with C.
Apochiton C. E. Hubb., Hooker’s Icon. Pl. 34: t. 3319 poiflorum and C. stoloniferum (Snow 1996). This
(1936). macrohair type may thus be synapomorphic for
Coelachyrum. Coelachyrum and Coelachyropsis
Annuals. Ligule membranous. Inflorescence share rugose fruits, with the pericarp free from
branches branched, the spikelets on filiform ped- the seed coat, and are combined by Clayton et al.
icels. Spikelets laterally compressed, with 3 to 5 (2006 onward) and Clayton and Renvoize (1986).
flowers plus additional reduced flowers distally. However, Petersen et al. (2010a) indicate that
Glumes shorter than the spikelet, their apices Coelachyrum, represented by C. poiflorum, is sis-
mucronate or awned. Callus of flower pubescent. ter to Eleusine and should not include Coelachyr-
Lemma apex acute, entire, awned. Palea with two opsis, represented by C. lagopoides.
awns. Microhairs with a broad apical cell.
One sp., A. burtii C. E. Hubb., Tanzania.
675. Eleusine Gaertn.
673. Coelachyropsis Bor Eleusine Gaertn., Fruct. Sem. Pl. 1: 7 (1788); phylog.: Liu
et al. (2011); Neves et al. (2005).
Coelachyropsis Bor, Ann. Naturhist. Mus. Wien 75: 23
(1971) [1972]. Annuals or perennials, caespitose or with short
rhizomes. Leaf sheaths keeled. Ligule membra-
Annuals, less than 20 cm tall. Leaf blades cordate at nous, or a fringed membrane. Inflorecence
the base, with multicellular glands. Ligule a fringed unbranched, or with the primary branches
membrane. Inflorescence branches unbranched, unbranched and digitate. Spikelets laterally com-
digitate. Spikelets laterally compressed, with 4 to pressed, with 3 to 15 flowers plus additional
10 flowers plus additional reduced flowers distally. reduced flowers distally. Glumes shorter than
Glumes shorter than the spikelet, their apices acu- the spikelet, keeled, their apices acute to obtuse,
minate, mucronate. Lemma pubescent on midvein awnless. Lemma muticous or mucronate. Leaf
and margins, the apex mucronate. epidermal papillae absent. 2n ¼ 18, 20, 36, 40.
One sp., C. lagopoides Bor, southern India Ten spp., tropical Africa, South America, and
and Sri Lanka. one widespread weed.
Eleusine coracana (L.) Gaertn. (finger millet)
674. Coelachyrum Hochst. & Nees is widely cultivated in Africa, where it provides
particularly drought-resistant grain.
Coelachyrum Hochst. & Nees, Linnaea 16: 221 (1842).
Cypholepis Chiov., Annuario Reale Ist. Bot. Roma 8: 357
(1908).
XXIX.6. SUBTRIBE TRIPOGONINAE Stapf (1917)
Annuals or perennials, caespitose or stoloniferous.
Ligule membranous. Inflorescence branches This subtribe is monophyletic in the phylogenies
branched or not, if unbranched then digitate or presented by Peterson et al. (2010a) and GPWG II
borne along a central axis. Spikelets laterally com- (2012) but has no obvious morphological synap-
pressed, with 5 to 16 flowers plus additional omorphy.
Five genera, 57 spp. 679. Oropetium Trin. Fig. 93
Oropetium Trin., Fund. Agrost.: 98 (1822); tax.: Phillips

676. Desmostachya (Stapf) Stapf (1975).
Desmostachya (Stapf) Stapf, Fl. Cap. 7: 316 (1898). Perennial or annual cushion plants. Ligule mem-
branous, or a fringed membrane or a fringe of
Rhizomatous perennials. Ligule a fringed mem- hairs. Leaves with multicellular glands abaxially.
brane. Leaf blades coriaceous. Inflorescence Inflorescence unbranched, straight, sinuous or
branches unbranched, spread along a central coiled, the spikelets sunken in the rachis. Rachis
axis. Spikelets laterally compressed, overlapping, tough or disarticulating. Spikelets dorsiventrally
with as many as 18 flowers. Glumes ovate, their
apices acute. Lemma and palea similar in length,
their apices entire. Leaf epidermal papillae
absent. 2n ¼ 20.
One sp., D. bipinnata (L.) Stapf, North Africa,
Middle East, India, Indo-China.
677. Eragrostiella Bor
Eragrostiella Bor, Indian Forester 66: 269–270 (1940).
Perennials. Leaf blades filiform. Ligule a fringed

membrane or a fringe of hairs. Inflorescence
unbranched. Spikelets laterally compressed,
with 7 to 50 flowers, the distal flowers reduced.
Glumes shorter than the spikelet, the proximal
one persistent and the distal one deciduous.
Lemma disarticulating, leaving behind the per-
sistent palea.
Six spp., Eastern Africa, India, Sri Lanka,
Burma, Australia.
Eragrostiella is derived from within a para-
phyletic Tripogon (Peterson et al. 2010a).
678. Melanocenchris Nees
Melanocenchris Nees, Proc. Linn. Soc. Lond. 1: 94 (1841).
Annuals or perennials, less than 30 cm tall. Ligule

a fringe of hairs. Inflorescence a densely capitate
cluster of unbranched branches, these ending in a
forked bristle. Spikelets wedge-shaped, dorsiven-
trally compressed, with 1 flower plus several
reduced flowers distally. Glumes next to each
other, shorter than or reaching the apex of the Fig. 93. Chloridoideae-Cynodonteae-Tripogoninae. Oro-
flowers, coriaceous, with long hairs, awned; upper petium minimum. A Habit. B Ligule. C Part of rachis of
glume with broad wings basally. Lemma with inflorescence. D Upper part of inflorescence. E Lower
glume. F Upper glume. G Flower. H Lemma. I Palea. J
clavate hairs, with 1 or 3 awns. Pistil, stamens and lodicules. K Caryopsis. (From Clayton
Three spp., tropical Africa, India, Sri Lanka. et al. 1974, drawn by S. Ross-Craig)
compressed, with one flower. Callus of flower Annuals, less than 30 cm tall. Ligule a fringed
pubescent. Lower glume absent, upper glume membrane. Spikelets spherical, with 7 to 10 flow-
covering the flower. Lemma awned. Pericarp sep- ers plus additional reduced flowers distally. Cal-
arable from seed coat. Leaf epidermal papillae lus of flower pubescent. Glumes shorter than the
absent. 2n ¼ 18, 20, 36. spikelet, their apices awned. Lemma fan-shaped,
Six spp., Africa and India. pubescent, the apex bifid with a short awn from
GPWG II (2012) place Oropetium sister to the sinus. Palea bent inward over the pistil, with
Tripogon and the pair sister to Melanocenchris, tufts of hair at the bend. Pericarp free.
with strong support. One sp., N. barbata (Nees) Pilg., NE Brazil.
680. Tripogon Roem. & Schult. 682. Pappophorum Schreb. Fig. 94
Tripogon Roem. & Schult., Syst. Veg. 2: 34 (1817). Pappophorum Schreb., Gen. Pl. 2: 787 (1791).
Annuals or perennials, caespitose or rhizoma- Perennials, generally 0.5–2 m tall. Ligule a fringe
tous. Ligule membranous, or a fringed mem- of hairs. Spikelets laterally compressed, with 1 to
brane or a fringe of hairs. Leaf blade apex 4 flowers plus additional reduced flowers distally,
sharp-pointed. Inflorescence unbranched. Spi- the distal rachilla internodes contracted so the
kelets laterally compressed, with 2 to 50 flowers assembled lemmas look like a brush. Callus of
plus additional reduced flowers distally, in a flower pubescent. Glumes as long as or extending
few species the most proximal flower sterile. beyond the flowers, with 1 vein, their apices muti-
Glumes shorter than the spikelet, their apices cous to awned. Lemma with 11–26 awns. Micro-
mucronate or awned. Lemma apex incised, with hairs salt-secreting. 2n ¼ 40, 60, 100.
a mucro or 1 or 3 awns. Anthers 1, 2, or 3. Nine spp., North and South America.
2n ¼ 20.
Forty-four spp., tropics, most in Old World.
683. Tridens Roem. & Schult.
Tridens Roem. & Schult., Syst. Veg. 2: 34, 599 (1817).

XXIX.7. SUBTRIBE PAPPOPHORINAE Dumort. (1829)
Perennials, caespitose or rhizomatous. Ligule mem-
Inflorescence branches branched. Callus of flower branous or a fringed membrane or a fringe of hairs.
pubescent. Inflorescence branches branched or not. Spikelets
Three genera, ten spp. with 3 to 15 flowers plus additional reduced flowers
This subtribe has no obvious morphological distally. Glumes shorter than or almost as long as
synapomorphy. Although Neesiochloa and Pap- the flowers, their apices muticous or mucronate.
pophorum have long been placed with Enneapo- Callus of flower glabrous or pubescent. Lemma
gon, Cottea and Schmidtia in the tribe emarginate or with two lobes, mucronate or awned
Pappophoreae based on the laciniate and multi- between the lobes; veins and keel with dense hairs,
awned lemmas, the two groups are not closely forming lines on the back of the lemma. Leaf epi-
related, and the traditional Pappophoreae are dermal papillae absent. 2n ¼ 16, 32, 40, 72.
polyphyletic (Ingram and Doyle 2004; Peterson Fifteen spp., New World, Angola.
et al. 2010a; Reutemann et al. 2011). The former Tridens muticus is firmly placed
in the Tridentinae clade by both Columbus et al.
(2010) and Peterson et al. (2010a, 2014a) and is
681. Neesiochloa Pilg. now recognized as Tridentopsis. Tridens flavus is
consistently placed with Pappophorum (Colum-
Neesiochloa Pilg., Repert. Spec. Nov. Regni Veg. 48: 119 bus et al. 2010; GPWG II 2012; Peterson et al.
(1940).
2010a, 2014a).
References 395
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Ruggell. Giussani, L.M. 2011. Phylogenetic relationships
Metcalfe, C.R. 1960. Anatomy of the monocotyledons. I. within Pappophoreae s.l. (Poaceae: Chloridoideae):
Gramineae. Oxford: Clarendon Press. additional evidence based on ITS and trnL-F
Moffett, A.A., Hurcombe, R. 1949. Chromosome numbers sequence data. South Afr. J. Bot. 77: 693–702.
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M€uller, M.A.N. 2007. Grasses of Namibia. Windhoek, grass subfamily Chloridoideae. I. Basic chromosome
Namibia: Ministry of Agriculture, Water and For- numbers. Taxon 52: 557–566.
estry. Snow, N. 1996. The phylogenetic utility of lemmatal
Neves, S.S., Swire-Clark, G., Hilu, K.W., Baird, W.V. 2005. micromorphology in Leptochloa s.l. and related
Phylogeny of Eleusine (Poaceae: Chloridoideae) genera in subtribe Eleusininae (Poaceae, Chloridoi-
based on nuclear ITS and plastid trnT-trnF deae, Eragrostideae). Ann. Missouri Bot. Gard. 83:
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Sida 17: 645–666. Nat. 11: 372–394.
XI. Subfamily Micrairoideae Pilg. (1956)
Annuals or perennials. Culms solid or hollow. KEY TO THE GENERA OF MICRAIROIDEAE

Ligule a fringe of hairs. Spikelets generally with
2 flowers; rachilla extension absent. Disarticula- 1. Plants floating; inflorescence unbranched
tion above the glumes. Ovary glabrous. Styles 2, 691. Limnopoa
1. Plants terrestrial, if in water then not floating; inflores-
not fused. Hilum short or long linear. Embryo cence branched 2
without an epiblast, with a scutellar cleft, with 2 (1). Primary branches of the inflorescence unbranched,
an elongate mesocotyl internode, the embryonic ending in a sharp point, the spikelets borne on one
leaf margins meeting. side 688. Heteranthoecia
Nine genera, 188 species. 2. Primary branches of the inflorescence branched, end-
ing in a spikelet, not one-sided 3
Micrairoideae are strongly supported as 3 (2). Plants mat-forming, moss-like, with leaves densely
monophyletic in recent molecular analyses clustered and apparently spirally arranged
(GPWG II 2012; Sánchez-Ken and Clark 2010; 684. Micraira
Sánchez-Ken et al. 2007). The subfamily was initi- 3. Plants variously caespitose or trailing, but not mat-
ally recognized to accommodate the peculiar forming, the leaves not densely clustered or spirally
arranged 4
Australian genus Micraira, but has since been 4 (3). Lemma membranous or chartaceous or hyaline 5
expanded to include Isachneae and Eriachneae, 4. Lemma coriaceous or indurate 7
two tribes formerly placed in Panicoideae on the 5 (4). Ligule absent 689. Hubbardia
basis of spikelet morphology (Clayton and 5. Ligule present 6
Renvoize 1986). The origin of C4 photosynthesis 6 (5). Spikelets with 1 flower; disarticulating below the
glumes 692. Sphaerocaryum
in Eriachneae is independent of other origins in 6. Spikelets with 2 flowers, rarely 1; disarticulating above
the PACMAD clade (GPWG II 2012), and aspects the glumes 687. Coelachne
of leaf anatomy and gene expression are quite 7 (4). Upper glume twice as long as the lower one
distinct from other C4 taxa (Prendergast et al. 686. Pheidochloa
1987; Sinha and Kellogg 1996). 7. Glumes nearly equal in length 8
8 (7). Spikelets dorsiventrally compressed or not com-
Micraira and Eriachne are successive sisters pressed; lemma glabrous or puberulent, never awned
to the remainder of the subfamily (GPWG II 690. Isachne
2012). Although the subfamily is often divided 8. Spikelets laterally compressed; lemma often pubescent
into tribes, this seems unnecessary. If they are with long hairs, often awned 685. Eriachne
to be recognized, then Micraireae include only
Micraira, Eriachneae include Eriachne and Phei-
dochloa, and Isachneae include the rest.
400 XI. Subfamily Micrairoideae Pilg. (1956)
GENERA OF MICRAIROIDEAE with 2 flowers. Glumes longer than the flowers, sepa-
rated by a long rachilla internode, the second glume
684. Micraira F. Muell. about twice the length of the first, their apices acute.
Callus of flower elongate, sharp-pointed, pubescent.
Micraira F. Muell., Fragm. 5: 208 (1867). Lemma margins involute, the apex acute, awned, the
awn flexuous. Stamens 2. Hilum long linear. Leaf
Tiny moss-like perennials, in tufts connected by vasculature with a double bundle sheath, the outer
stolons. Ligule membranous, or a fringed mem- one parenchymatous and carbon reducing, densely
brane, or a fringe of hairs, or absent. Leaves packed with chloroplasts, the inner one made up of
arranged in a spiral. Inflorescence branches
thick-walled cells. Plants C4 NADP-ME.
branched. Spikelets laterally compressed, with
Two spp., Asia, Australasia.
2 flowers, with a rachilla extension. Glumes shorter
or longer than the flowers, their apices emarginate Remaining genera:
to acuminate, muticous to awned. Lemma mem- Mesophyll radiate around the vascular bundles,
branous or hyaline, the apex emarginate to obtuse, similar to Kranz anatomy but the vein spacing dis-
muticous or mucronate. Palea divided into two. tant, characteristic of C3 plants.
Lodicules absent. Stamens 2. 2n ¼ 20.
Fifteen spp., Australia. 687. Coelachne R. Br.
The spiral vegetative phyllotaxis in Micraira
was documented by Philipson (1935). The paleas Coelachne R. Br., Prodr. Fl. Nov. Holl.: 187 (1810).
of some species have many veins.
Annuals or perennials. Ligule a fringe of hairs or
685. Eriachne R. Br.
absent. Inflorescence branches branched. Spike-
Eriachne R. Br., Prodr.: 183 (1810).
lets dorsiventrally compressed, with two dissimi-
lar flowers separated by a long rachilla internode;
Annuals or perennials, caespitose or rhizomatous. rachilla extension lacking. Glumes shorter than
Ligule a fringe of hairs. Inflorescence branches the spikelet, their apices generally obtuse. Lemma
branched. Spikelets laterally compressed, with slightly inflated, lacking veins, the apex obtuse or
acute, unawned. Stamens 2 or 3. Hilum short.
2 flowers. Glumes shorter or longer than the flow-
Leaf epidermal papillae absent, or consisting of
ers, their apices obtuse to attenuate, muticous or one symmetrical swelling per cell. 2n ¼ 40.
rarely awned. Callus of flower short, blunt, pubes- Twelve spp., Old World tropics.
cent. Lemma margins involute, the apex obtuse to
acuminate, mucronate or awned, the awn straight 688. Heteranthoecia Stapf
or curved. Stamens 2 or 3. Hilum long linear. Leaf
Heteranthoecia Stapf, Hooker’s Icon. Pl. 30: t. 2937
vasculature with a double bundle sheath, the outer (1911).
one parenchymatous and carbon reducing, densely
packed with chloroplasts, the inner one made up of Annuals. Ligule a fringe of hairs. Inflorescence
thick-walled cells. Plants C4 NADP-ME. branches unbranched, spread along a central axis,
Forty-eight spp., Asia, Australia, Pacific. the branches flattened, ending in a barren tip, spi-
Spikelets in some specimens of Eriachne have kelets borne in two rows along abaxial side. Spike-
a noticeable internode between the glumes; rarely lets dorsiventrally compressed, with 2 flowers.
a rachilla extension bearing a rudimentary flower Glumes shorter than the spikelet, the lower decidu-
may be present. Paleas may be two-pronged or ous, the upper persistent, their apices acute. Lemma
awned at the apex. papery, with club shaped-hairs, the apex obtuse.
686. Pheidochloa S. T. Blake Leaf epidermal papillae present. Hilum short.
One sp., H. guineensis (Franch.) Robyns,
Pheidochloa S. T. Blake, Proc. Roy. Soc. Queensland 56: 20 tropical Africa. Plants of wet places.
(1944). 689. Hubbardia Bor
branches branched. Spikelets laterally compressed, Hubbardia Bor, Kew Bull. 5: 385 (1951).
GENERA OF MICRAIROIDEAE 401
Annuals. Leaves flimsy. Ligule absent. Inflores- One sp., H. heptaneuron Bor, western India
cence branches branched. Spikelets dorsiventrally (Karnataka and Maharashtra).
compressed, with 2 flowers, the lower sterile, the Spikelet morphology has led previous work-
upper bisexual. Glumes longer than the flowers, ers to place Hubbardia in Panicoideae, but mor-
pubescent with tubercle-based hairs, their apices phological (Zuloaga et al. 2011) and molecular
obtuse. Lower sterile flower reduced to a lemma (Sánchez-Ken, unpublished) data place it here.
with an obtuse apex. Lemma of upper flower hya- Although Hubbardia was once thought to be
line above, the apex obtuse; palea absent. Leaf extinct, it has been rediscovered; it remains rare
epidermal papillae absent. Hilum short. and endangered (Yadav et al. 2009, 2010).
690. Isachne R. Br. Fig. 95
Isachne R. Br., Prodr.: 196 (1810).
Annuals or perennials, caespitose, rhizomatous or

stoloniferous. Leaf blades disarticulating at the
ligule. Ligule a fringed membrane, or a fringe of
hairs, or absent. Inflorescence branches branched.
Spikelets dorsiventrally compressed, with 1 or
2 flowers, the proximal one staminate or bisexual,
the distal one bisexual. Glumes shorter or longer
than the spikelet, their apices generally obtuse to
acuminate. Lemma apex obtuse to acute, awnless.
Hilum long-linear. Leaf epidermal papillae one
per cell, or absent. 2n ¼ 20, 50, and 60.
One hundred-three spp., Africa, Asia and
Australasia. Species of Isachne are often aquatic.
691. Limnopoa C. E. Hubb.
Limnopoa C. E. Hubb., Hooker’s Icon. Pl. 35: t. 3432

(1943).
Floating annuals, less than 25 cm tall. Ligule a

fringe of hairs. Inflorescence unbranched, the
spikelets in pairs, one sessile, one pedicellate,
borne on one side of the inflorescence axis. Spi-
kelets dorsiventrally compressed, with 2 flowers,
the proximal one staminate, the distal one bisex-
ual. Glumes shorter than the flowers, the lower
one with an acute apex, the upper one obovate
and with an obtuse apex. Lemma pubescent, the
apex obtuse. Hilum long-linear. Leaf epidermal
papillae one per cell.
One sp., L. meeboldii (C.E.C. Fisch.) C.E.
Hubb., tropical Asia.
692. Sphaerocaryum Nees ex Hook. f.
Fig. 95 Micrairoideae. Isachne pulchella. A Habit. B Sphaerocaryum Nees ex Hook. f., Fl. Brit. India 7: 246
Spikelet. C Abaxial and adaxial views of lower lemma (1897).
and palea. D Abaxial and adaxial views of upper lemma
and palea. E Glumes. (From Wu et al. 2007, with permis-
sion from the Missouri Botanical Garden Press, St. Louis, Annuals, less than 15 cm tall. Ligule a fringe of
and Science Press, Beijing; drawn by Si Weiqing) hairs. Leaf blades ovate. Inflorescence branches
402 XI. Subfamily Micrairoideae Pilg. (1956)
branched. Spikelets not compressed, with 1 flower; Prendergast, H.D.V., Hattersley, P.W., Stone, N.E. 1987.
disarticulating below the glumes. Glumes as long New structural/biochemical associations in leaf
blades of C4 grasses (Poaceae). Aust. J. Plant Phys.
as the flower, translucent, their apices obtuse; 14: 403–420.
lower glume lacking veins, upper glume with 1 Sánchez-Ken, J.G., Clark, L.G. 2010. Phylogeny and a new
vein. Lemma translucent, pubescent, the apex tribal classification of the Panicoideae s.l. (Poaceae)
obtuse. Hilum short. Leaf epidermal papillae one based on plastid and nuclear sequence data and
structural data. Amer. J. Bot. 97: 1732–1748.
per cell. 2n ¼ 20. Sánchez-Ken, J.G., Clark, L.G., Kellogg, E.A., Kay, E.E.
One sp., S. pulchellum (Roth.) Merr., India 2007. Reinstatement and emendation of subfamily
and southeast Asia. Micrairoideae (Poaceae). Syst. Bot. 32: 71–80.
Sinha, N.R., Kellogg, E.A. 1996. Parallelism and diversity
in multiple origins of C4 photosynthesis in the grass
family. Amer. J. Bot. 83: 1458–1570.
References Wu, Z., Raven, P.H., Hong, D. (eds.) 2007. Flora of China
Illustrations. Beijing: Science Press.
Yadav, S.R., Chandore, A.N., Nimbalkar, M.S., Gurav, R.
Clayton, W.D., Renvoize, S.A. 1986. Genera graminum: V. 2009. Reintroduction of Hubbardia heptaneuron
grasses of the world. London: Her Majesty’s Statio- Bor, a critically endangered endemic grass in West-
nery Office. ern Ghats. Curr. Sci. 96: 880.
GPWG II 2012. Grass Phylogeny Working Group II. New Yadav, S.R., Chandore, A.N., Gund, S.M., Nandikar, M.,
grass phylogeny resolves deep evolutionary relation- Lekhak, M. 2010. Relocation of Hubbardia hepta-
ships and discovers C4 origins. New Phytol. 193: neuron Bor, from its type locality. Curr. Sci. 98: 884.
304–312. Zuloaga, F.O., Morrone, O., Scataglini, M.A. 2011. Mono-
Philipson, W.R. 1935. A grass with spiral phyllotaxis: graph of Trichanthecium (Poaceae, Paniceae). Syst.
Micraira subulifolia. Bull. Misc. Inf. (Kew) 1935: Bot. Monographs 94: 1–101.
324–326.
XII. Subfamily Arundinoideae Burmeist. (1837)
Perennials; caespitose, rhizomatous, or stolonif- each with only one or two species, still lack
erous, tall and reed-like to minute and moss-like. molecular data, so there remains a possibility
Ligule membranous or a fringe of hairs. Inflores- that several are still misplaced here.
cence branches branched. Spikelets laterally com- A few species currently placed elsewhere may
pressed. Callus of flower pubescent. Style also belong in Arundinoideae. The recent discov-
branches 2, generally free. Hilum short or long- ery that “Eragrostis” walteri is not an Eragrostis at
linear. Epiblast absent, scutellar cleft present, all (Ingram et al. 2011) (see below) is an example.
mesocotyl internode elongated, embryonic leaf
margins meeting or overlapping. Leaf epidermal
papillae absent.
Nineteen genera, 46 spp. KEY TO THE GENERA OF ARUNDINOIDAE
The circumscription of this subfamily has
1. Perennial reeds, often 2 m or more tall; plants strongly
varied enormously over the years. Clayton and rhizomatous 2
Renvoize (1986) included in it genera now 1. Plants rarely over 1 m tall; generally caespitose 3
assigned to Stipeae (Pooideae), Danthonioideae, 2 (1). Ligule membranous, with short cilia; lemma villous;
Micrairoideae, Panicoideae and Aristidoideae, proximal flowers of the spikelet bisexual
while Watson and Dallwitz (1992 onward) used 694. Arundo
2. Ligule with a minute membrane and long cilia; lemma
an equally broad interpretation, including Steyer- glabrous (but the callus with long hairs); proximal
markochloeae, Cyperochloa, and Spartochloa (all flowers of the spikelet staminate or sterile
now Panicoideae s.l.), as well as taxa now placed 703. Phragmites
in Micrairoideae and Aristidoideae. Kellogg and 3 (1). Spikelets arranged in several dense globular clusters
Campbell (1987) showed that the subfamily as spread along the inflorescence axis; plants annual
697. Elytrophorus
defined at the time was a polyphyletic assemblage 3. Spikelets not clustered; plants annual or perennial 4
whose members were variously sister to other 4 (3). Spikelets with one flower 5
well-defined subfamilies. Polyphyly of the tradi- 4. Spikelets with more than one flower 8
tional Arundinoideae was confirmed by molecu- 5 (4). Inflorescence unbranched; plants annual; lemma
lar work of Barker et al. (1995) and Clark et al. with a transverse line of hairs 708. Danthonidium
5. Inflorescence branches branched; plants perennial;
(1995). In subsequent years, the subfamily has lemma glabrous or pubescent but the hairs not in a
been relieved of most of its disparate elements transverse line 6
and is reduced to the 19 genera described here 6 (5). Lemma with 3 awns; plants of Australia
(GPWG 2001; GPWG II 2012). However, the 707. Amphipogon
group remains morphologically heterogeneous, 6. Lemma with one awn; plants of Africa or India 7
7 (6). Palea unawned; plants of Ethiopia
as evidenced by the large number of genera for 700. Leptagrostis
a small number of species, and there is no obvi- 7. Palea with two apical setae; plants of India and Sri
ous morphological synapomorphy. Nine genera, Lanka 696. Dichaetaria
404 XII. Subfamily Arundinoideae Burmeist. (1837)
8 (4). Lemma with 9 veins, hairs in lines or tufts, bilobed, ending in awns, principal awn from the sinus,
with a central geniculate awn from the sinus 9 flattened and geniculate.
8. Lemma not as above 11
9 (8). Lemma broad and deeply cup-shaped; plants of
Six spp., Angola, Tanzania, Malawi, Mozam-
Australia 710. Monachather bique.
9. Lemma longer than broad, not cup-shaped; plants of
Africa 10
694. Arundo L.
10 (9). Culms slender, decumbent; leaf blades membra-
nous; plants of Yemen and Ethiopia Arundo L., Sp. Pl. 1: 81 (1753); tax.: Hardion et al. (2012).
711. Phaenanthoecium
10. Culms upright, the plants tufted; leaf blades succulent; Reeds over 2 m tall, rhizomatous. Leaves cauline,
plants of southern Africa 709. Dregeochloa
11 (8). Spikelets lacking awns 12
the blades disarticulating from the sheaths. Ligule
11. Spikelets with awns 14 membranous, with short cilia. Inflorescence
12 (11). Spikelets with 7 to 15 flowers; plants of southern branches branched. Spikelets with 2 to 5 flowers,
Africa 698. “Eragrostis” walteri with a rachilla extension. Glumes as long as the
12. Spikelets with 5 or fewer flowers; plants not African flowers, their apices acute to acuminate. Lemma
13
13 (12). Leaf blades not pseudopetiolate; plants of Europe,
villous proximally, the apex entire or bidentate,
Turkey, China, Japan 701. Molinia mucronate or awned from the sinus and from the
13. Leaf blades pseudopetiolate; plants of India and Sri lobes. Hilum short. 2n ¼ 24, 100, 110.
Lanka 706. Zenkeria Three spp., southern Europe and northern
14 (11). Lemma with hairs in tufts 693. Alloeochaete Africa to China, introduced worldwide.
14. Lemma glabrous, if pubescent then hairs not tufted
15
15 (14). Awn shorter than the lemma 16 695. Crinipes Hochst.
15. Awn longer than the lemma 17
16 (15). Leaves slender, stiff, the basal sheaths hard and Crinipes Hochst., Flora 38: 279 (1855).
fibrous; ligule a fringe of hairs; plants of Africa
705. Styppeiochloa Caespitose. Leaves stiff, the blades disarticulating
16. Leaves broad, flat, flexuous, the basal sheaths mem- from the sheaths. Ligule a fringe of hairs. Inflo-
branous; ligule membranous; plants of Japan but rescence branches branched. Spikelets with 2 or 3
cultivated elsewhere 699. Hakonechloa
17 (15). Spikelets with 4 to 7 flowers 702. Nematopoa
flowers, sometimes with reduced flowers distally
17. Spikelets with 2 or 3 flowers 18 or with a rachilla extension. Callus of flower long
18 (17). Plants <40 cm tall; spikelets <3 mm long and narrow. Glumes shorter than flowers, with 1
704. Piptophyllum vein, their apices acuminate, mucronate or short
18. Plants 60–150 cm tall; spikelets >4 mm long awned. Lemma with hairy margins, the apex
695. Crinipes
entire or bidentate, with an awn from the apex
or the sinus. Hilum short.
One sp., Sudan, Ethiopia, Uganda.
GENERA OF ARUNDINOIDEAE
696. Dichaetaria Nees ex Steud.
693. Alloeochaete C. E. Hubb.
Dichaetaria Nees ex Steud., Syn. Pl. Glumac. 1: 145 (1855).
Alloeochaete C. E. Hubb., Hooker’s Icon. Pl. 35: t. 3418
(1940). Caespitose. Ligule a fringe of hairs; contraligule
present. Inflorescence branches branched. Spike-
Caespitose. Leaf blades disarticulating from the lets with 1 flower, plus reduced flowers distally.
sheath, the sheath wooly-tomentose. Ligule a Glumes shorter than the spikelet, their apices
fringe of hairs. Inflorescence branches branched. acute, the lower persistent, the upper deciduous.
Spikelets with 4 to 10 flowers, the proximal flower Callus of flower elongate. Lemma leathery, awl-
staminate or sterile. Glumes shorter than the shaped, glabrous, with two teeth, awned from the
flowers, their apices obtuse to acuminate, mucro- sinus. Palea with 2 setae. Fruit beaked. Hilum
nate to awned. Callus of flower blunt or elongate. long-linear.
Lemma with 5 veins, with a tuft of hairs on One sp., D. wightii Nees ex Steud., India and
each side, the apex with two teeth or lobes, these Sri Lanka.
GENERA OF ARUNDINOIDEAE 405
697. Elytrophorus P. Beauv.

Elytrophorus P. Beauv., Ess. Agrostogr.: 67, pl. 14, f. 2 (1812).
Caespitose annuals. Ligule membranous or a

fringed membrane. Inflorescence with the primary
branches forming dense globular clusters of spike-
lets; outer spikelet of a cluster with the lower
glume enlarged and most or all flowers reduced
to sterile lemmas. Spikelets with 3 to 7 flowers.
Glumes as long as the flowers, their apices acu-
minate, awn-tipped. Callus of flower glabrous.
Lemma keeled, the apex attenuate, entire, with a
short awn. Palea keels winged. Lodicules 1 or 2.
Stamens 1 to 3. Hilum short. 2n ¼ 24.
Two spp., tropical Africa, Australia, India, China.
698. “Eragrostis” walteri Pilg.
“Eragrostis” walteri Pilg.
Caespitose. Ligule a fringe of hairs. Inflorescence

branches branched. Spikelets with 7 to 15 flowers.
attenuate. Lemma apex mucronate.
One sp., southern Africa.
For many years this species was thought to be
the only C3 member of Eragrostis. However, it
clearly belongs in Arundinoideae, probably sister
to Elytrophorus (Ingram et al. 2011).
699. Hakonechloa Makino ex Honda Fig. 96

Hakonechloa Makino ex Honda, J. Fac. Sci. Univ. Tokyo,
Sect. 3, Bot. 3: 113 (1930).
Short rhizomatous. Ligule membranous, with a

ciliate margin. Abaxial ligule present, a line of
hairs. Leaf blades resupinate. Inflorescence
branches branched. Spikelets with 5 to 10 flowers,
the rachilla internodes pubescent. Glumes shorter
than the flowers, their apices acuminate. Callus of
flower elongate. Lemma with 3 veins, the margins
ciliate, the apex with two teeth, with a short awn
from the sinus. 2n ¼ 50.
One sp., H. macra (Munro) Honda, Japan,
cultivated elsewhere.
700. Leptagrostis C. E. Hubb. Fig. 96. Arundinoideae. Hakonechloa macra. A Plant

with rhizome and inflorescence. B Spikelet with glumes
and flowers. (From Barkworth et al. 2003, drawn by C.
Leptagrostis C. E. Hubb., Bull. Misc. Inform. Kew 1937: 63
Roché)
(1937).
Caespitose, less than 25 cm tall. Ligule a fringe of ultimate ones bearing long hairs. Spikelets with
hairs. Inflorescence branches branched. Spikelets 3 to 11 flowers, the proximal one staminate or
with 1 flower, with a rachilla extension. Glumes sterile; rachilla extension present. Glumes shorter
with acuminate apices, the lower one shorter than than the flowers, their apices acute. Callus of
the flower, the upper one longer. Callus of flower flower elongate, with long hairs. Lemma narrow,
blunt. Lemma with two lobes, awned from the the apex long-attenuate, awnless but sometimes
sinus, the awn about 1 mm long. mucronate. Hilum short. 2n ¼ 36, 44, 46, 48, 50,
One sp., L. schimperiana (Hochst.) C. E. 52, 54, 84, 96, 120.
Hubb., Ethiopia. Four spp., in marshes and wet sites worldwide.
No information is available on the fruit or leaf In addition to the numbers cited above, aneu-
anatomy of this species. ploid chromosome counts are reported for Phrag-
mites australis (e.g., 2n ¼ 53, Connor et al. 1998;
701. Molinia Schrank 2n ¼ 42–59, Gervais et al. 1993), suggesting meiotic
irregularity. Interspecific hybridization has been
Molinia Schrank, Baier. Fl. 1: 100, 334 (1789). documented within the genus (Chu et al. 2011).
Caespitose, with only the uppermost culm inter-

704. Piptophyllum C. E. Hubb.
nodes elongated. Ligule a fringe of hairs. Leaf
blades disarticulating from the sheath. Inflores- Piptophyllum C. E. Hubb., Kew Bull. 12: 53 (1957).
cence branches branched. Spikelets with 2 to 5
flowers. Glumes shorter than the flowers, their Caespitose. Ligule a fringe of hairs. Leaf blades
apices acute. Callus of flower truncate, sometimes disarticulating from the sheaths. Inflorescence
glabrous. Lemma papery, the apex acute, awnless. branches branched. Spikelets with 2 flowers,
Fruit with pericarp easily separated from the seed with a rachilla extension. Glumes shorter than
coat. Hilum long-linear. 2n ¼ 18, 36, 50, 54, 72, 90. the flowers, hyaline, with 1 vein, their apices
Three spp., Europe, Turkey, China, Japan, acute, the upper glume mucronate. Lemma with
cultivated elsewhere. 5 to 9 veins, the apex with two lobes and three
awns, the principal awn from the sinus, flexuous,
702. Nematopoa C. E. Hubb. the base flat and twisted. Hilum short.
One sp., P. welwitschii (Rendle) C.E. Hubb.,
Nematopoa C. E. Hubb., Kew Bull. 12: 51 (1957). Angola.
Caespitose. Ligule a fringe of hairs. Leaf blades

705. Styppeiochloa de Winter
thread-like. Inflorescence branches branched.
Spikelets with 4 to 7 flowers. Callus of flower Styppeiochloa de Winter, Bothalia 9: 134 (1966).
short. Glumes shorter than the flowers, hyaline,
without veins, their apices acute. Lemma pubes- Caespitose, the basal sheaths hard and fibrous.
cent, with 1 vein, with 2 lobes, awned from the Ligule a fringe of hairs. Leaf blades sharp-pointed.
sinus, the awn flexuous. Hilum short. Inflorescence branches branched. Spikelets with
One sp., N. longipes (Stapf & C.E. Hubb.) C.E. 2 to 6 flowers. Glumes shorter than the flowers,
Hubb., south tropical Africa. their apices acuminate, mucronate or awned. Cal-
lus of flower short, blunt. Lemma apex with
703. Phragmites Adans. 2 lobes, with short awns from the lobes and the
sinus. Hilum long-linear. 2n ¼ 24, 48.
Phragmites Adans., Fam. Pl. 2: 34, 559 (1763); phylog.: Three spp., Africa and Madagascar.
Chu et al. (2011); Lambertini et al. (2006, 2012).
Reeds, generally over 2 m tall, rhizomatous. Leaf 706. Zenkeria Trin.

blades disarticulating from the sheath. Ligule Zenkeria Trin., Linnaea 11: 150 (1837).
membranous, or a fringed membrane, or a fringe Caespitose. Ligule a fringe of hairs. Leaf blades
of hairs. Inflorescence branches branched, the pseudopetiolate, with leathery margins.
GENERA OF ARUNDINOIDEAE 407
Inflorescence branches branched. Spikelets with One sp., D. gammiei (Bhide) C. E. Hubb.,
2 flowers, with or without a rachilla extension. western India.
Glumes shorter than the flowers, with 1 vein, their
apices acute to acuminate. Callus of flower with 709. Dregeochloa Conert
thick hairs. Lemma with 5 to 11 veins, the apex
entire, obtuse to acuminate. Stamens sometimes Dregeochloa Conert, Senckenberg. Biol. 47: 335 (1966).
2.
Five spp., India and Sri Lanka. Perennials, short rhizomatous, less than 25 cm
tall. Ligule a fringe of hairs. Leaves with blades
somewhat succulent, with a sharp pointed apex.
707. Amphipogon R. Br.
Inflorescence branched or unbranched. Spikelets
Amphipogon R. Br., Prodr.: 175 (1810). with 4 to 10 flowers. Glumes shorter or longer
Diplopogon R. Br., Prod. 176 (1810). than the flowers, their apices acute. Callus of
flower elongate. Lemma leathery, with a trans-
Rhizomatous or caespitose. Leaves convolute, verse row of hairs, the hairs tufted, the apex
often disarticulating from the sheaths, the apices with the lobes ending in awns or not. Ovary
sharp pointed. Ligule a fringe of hairs. Inflores- pubescent. Pericarp free from the seed coat, brit-
cence unbranched or capitate, sometimes sur- tle. Hilum short. 2n ¼ 42.
rounded by an involucre of narrow bracts. Two spp., southern Africa.
Spikelets with 1 flower, without a rachilla exten-
sion. Glumes shorter or longer than the lemma,
710. Monachather Steud.
with or without awns. Callus of flower short or
long. Lemma with three veins, each ending in an Monachather Steud., Syn. Pl. Glumac. 1: 247 (1854).
awn or tooth. Palea with two awns. Hilum short.
Fruit with pericarp free from the seed coat. Leaf Caespitose perennials; basal internodes enlarged
epidermal cells with evenly spaced papillae. Micro- and covered with wooly hairs. Ligule a fringed
hairs with apical cell nearly as wide as long membrane. Inflorescence branches branched.
(“chloridoid”). Spikelets with 5 or 6 flowers, with a rachilla
Nine spp., Australia. extension. Callus of flower short. Glumes laterally
The narrow involucrate bracts subtending the compressed, longer than the flowers, their apices
inflorescence in A. setaceus (R. Br.) T. D. Macfarl. acute. Lemma broad, cup-shaped, leathery, with a
are presumed to be the glumes of sterile spikelets. transverse line of hairs, the apex deeply divided
into lobes. Palea appressed against the inner
DANTHONIDIUM + D R E G E O C H L O A + M O N A C H A T H E R + curve of the lemma, the keels winged. Hilum
PHAENANTHOECIUM short. 2n ¼ 72.
Lemma with 9 veins, hairs in lines or tufts, One sp., M. paradoxa Steud., Australia.
bilobed, with a central geniculate awn from the
sinus. 711. Phaenanthoecium C. E. Hubb.
708. Danthonidium C. E. Hubb. Phaenanthoecium C. E. Hubb., Bull. Misc. Inform. Kew

1936: 329 (1936).
Danthonidium C. E. Hubb., Hooker’s Icon. Pl. 34: t. 3331
(1937). Decumbent, caespitose perennials. Ligule a fringe
of hairs. Inflorescence unbranched. Spikelets with
Annuals. Ligule a fringe of hairs. Inflorescence 5 to 9 flowers. Callus of flower blunt. Glumes
unbranched. Spikelets with 1 flower, with a shorter than the flowers, their apices obtuse or
rachilla extension. Glumes longer than the flower, notched. Lemma with the margins with tufts of
their apices acuminate, awn-tipped. Callus of hair, the apex with the lobes ending in awns.
flower blunt. Lemma leathery, with a transverse Hilum long-linear.
row of hairs, covering the palea, the apex with One sp., P. koestlinii (Hochst. ex A. Rich.)
lobes ending in awns. C. E. Hubb., northern Africa.

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Index
References to main entries of accepted names in bold-faced print, to synonyms in upright print, to illustrations in italics.
A A. scabrivalvis, 230 Aristidoideae, 267

Aakia, 321 Amphicarpum, 329 Arrhenatherum, 231
Acamptoclados, 366 A. amphicarpon, 329 Arthragrostis, 342
Achlaena, 314 Amphipogon, 407 Arthraxon, 291
Achnatherum s.s., 217 Anadelphia, 308 Arthropogon, 314
“Achnatherum” stillmanii, 216 Anatherostipa, 215 Arthropogoneae, 314
Aciachne, 214 Ancistrachne, 331 Arthropogoninae, 314
“Acidosasa”, 162 Ancistragrostis, 237 Arthrostylidiinae, 170
Acostia, 319 Andropogon, 313 “Arthrostylidium”, 171
Acrachne, 384 A. sect. Pobeguinea, 308 Arundinaria, 166
Acritochaete, 323 A. subg. Lasiorhachis, 305 A. amabilis, 152
Acroceras, 328 Andropogoneae, 289 Arundinaria clade, 166
Actinocladum, 171 Andropogoneae s.s., 291 Arundinarieae, 159
Aegilops, 224 Andropogoninae, 307 Arundinella, 290
Aegopogon, 376 Andropterum, 301 A. nepalensis, 291
Aeluropus, 372 Anemanthele, 217 A. parviflora, 291
Afrotrichloris, 385 Aniselytron, 254 Arundinelleae, 289
Agenium, 308 Anisopogon, 210 Arundinellinae, 290
Agnesia, 191 Anomochloa, 131 Arundinoideae, 403
Agropyron, 225 A. marantoidea, 132 Arundo, 404
Agropyropsis, 248 Anomochlooideae, 131 Arundoclaytonia, 284
Agrostidinae, 234 Anthaenantia, 318 Asthenochloa, 302
Agrostis, 236 Anthaenantiopsis, 320 Astrebla, 385
Agrostopoa, 254 A. rojasiana, 320 Athroostachys, 172
Aira, 241 Anthephora, 327 Atractantha, 172
Airinae, 240 Anthephoreae, 323 “Aulonemia”, 172
Airopsis, 235 Anthephorinae, 326 Australopyrum, 225
Alexfloydia, 333 Anthistirinae, 307 Austrochloris, 385
Alloeochaete, 404 Anthochloa, 257 Austrodanthonia, 351
Allolepis, 374 Anthosachne, 225 Austroderia, 350
Alloteropsis, 328 Anthoxanthinae, 233 Austrofestuca, 257
Alopecurinae, 254 Anthoxanthum, 234 Austrostipa, 217
Alopecurus, 254 A. odoratum, 235 Avellinia, 233
Altoparadisium, 314 Antinoria, 240 Avena, 231
Alvimia, 171 Apera, 254 Avenella, 241
Ambylopyrum, 224 Aphanelytrum, 257 Aveninae, 231
Amelichloa, 218 Apluda, 291 Avenula, 241
Ammochloa, 243 Apochiton, 392 Axonopus, 320
A. involucrata, 243 Apochloa, 314
A. palaestina, 243 Apoclada, 174 B
Ammochloinae, 243 Apocopis, 306 Bambusa, 182
Ammophila, 236 Arberella, 191 B. beechyana, 152
A. arenaria, 4 Arctagrostis, 259 B. chungii, 182
“Ampelocalamus”, 163 Arctophila, 255 B. multiplex, 48
“Ampelocalamus” calcareus, 168 Arctopoa, 255 Bambuseae, 169
Ampelodesmos, 213 Aristella, 217 Bambusinae, 176
Amphibromus, 230 Aristida, 268 Bambusoideae, 151
DOI 10.1007/978-3-319-15332-2, # Springer International Publishing Switzerland 2015
410 Index
Baptorhachis, 320 Capeochloa + Geochloa clade, 348 Cleistachne, 303

“Bashania”, 163 Capillipedium, 311 Cleistochloa, 331
BDG complex, 182 The Capillipedium + Bothriochloa + Cleistogenes, 372
Bealia, 376 Dichanthium Group, 311 Cliffordiochloa, 319
Beckeropsis, 333 Castellia, 245 Coelachne, 400
Beckmannia, 255 Catabrosa, 251 Coelachyropsis, 392
Bellardiochloa, 255 Catabrosella, 251 Coelachyrum, 392
BEP clade, 139 Catalepis, 365 Coelorachis, 300
Bergbambos, 160 Catapodium, 249 C. aurita, 27
Bewsia, 383 Cathariostachys, 186 Coix, 293
Bhidea, 307 Cathestecum, 374 Colanthelia, 172
Bistigmatic clade, 136 Celtica, 217 Coleanthinae, 251
Blepharidachne, 378 Cenchreae, 323 Coleanthus, 251
Blepharoneuron, 376 Cenchrinae, 333 C. subtilis, 252
Boissiera, 223 Cenchrinae + Melinidinae + Panicinae Coleataenia, 316
Boivinelleae, 323 clade, 332 Colpodium, 253
Boivinellinae, 328 Cenchrus, 333 Commelinidium, 328
Bonia, 176 Centotheca, 287 Connorochloa, 225
Borinda, 164 C. lappacea, 288 Cornucopiae, 255
Bothriochloa, 311 Centotheceae, 286 Cortaderia, 350
Bouteloua, 374 Centrochloa, 320 Corynephorus, 241
B. sp., 14 Centropodia, 360 Cottea, 364
Boutelouinae, 374 C. glauca, 360 Cotteinae, 364
Brachiaria, 339 Centropodieae, 359 Craspedorhachis, 381
Brachiariinae, 338 Cephalostachyum, 184 Criciuma, 174
Brachyachne, 386 Chaboissaea, 376 Crinipes, 404
Brachychloa, 373 Chaetium, 338 Critesion, 227
Brachyelytreae, 207 Chaetobromus, 349 Crypsis, 370
Brachyelytrum, 208 Chaetobromus + Pseudopentameris Cryptochloa, 192
B. erectum, 208 clade, 349 Cteniinae, 380
Brachypodieae, 222 Chaetopoa, 327 Ctenium, 381
Brachypodium, 222 Chaetopogon, 236 Ctenopsis, 245
B. distachyon, 222 Chamaeraphis, 333 Cutandia, 248
Briza, 235 Chandrasekharania, 282 Cyathopus, 237
Brizinae, 234 Chascolytrum, 235 Cyathorhachis, 292
Bromeae, 223 Chasmanthieae, 284 Cyclostachya, 374
Bromidium, 237 Chasmanthium, 284 Cymbopogon, 309
Bromuniola, 284 Chasmopodium, 291 Cynodon, 387
Bromus, 223 Chevalierella, 285 Cynodonteae, 371
B. berteroanus, 223 Chikusichloa, 148 Cynosurinae, 243
Brylkinia, 219 Chimaerochloa, 350 Cynosurus, 244
Brylkinieae, 219 Chimonobambusa, 163 C. echinatus, 244
Buchloe, 374 Chimonocalamus, 161 Cyperochloa, 287
Buchlomimus, 374 C. delicatus, 161 Cyperochloeae, 286
Buergersiochloa, 190 C. pallens, 161 Cyphochlaena, 329
B. bambusoides, 190 Chionachne, 292 Cyphochlaeneae, 323
Buergersiochloeae, 189 Chionochloa, 349 Cypholepis, 392
Buergersiochloinae, 190 Chloridoideae, 353 Cyphonanthus, 315
Burmabambus, 166 “Chloris”, 386 Cyrtochloa, 176
Butania, 166 Chlorocalymma, 327 Cyrtococcum, 330
Chondrosum, 374
C Chrysochloa, 387 D
“Calamagrostis”, 237 Chrysopogon, 292 Dactylidinae, 247
Calamovilfa, 370 Chusquea, 169 Dactylis, 247
Calderonella, 286 C. latifolia, 169 D. glomerata, 247
Calotheca, 235 Chusqueinae, 169 Dactyloctenium, 373
Calyptochloa, 331 Cinna, 255 Daknopholis, 387
Cambajuva, 171 Cinninae, 254 Dallwatsonia, 318
Campeiostachys, 225 Cladoraphis, 363 Danthonia, 351
Canastra, 315 Clausospicula, 292 Danthonia clade, 350
Capeochloa, 348 Clavinodum, 165 Danthoniastrum, 210
Index 411
Danthonidium, 407 Dregeochloa, 407 Euclasta, 293

Danthonioideae, 347 “Drepanostachyum”, 163 Eulalia, 304
Danthoniopsis, 282 Drymochloa, 245 Eulaliopsis, 293
Dasyochloa, 378 Dryopoa, 245 “Eustachys”, 389
Dasypyrum, 225 Dupontia, 255 Euthryptochloa, 209
Davidsea, 184 Duthiea, 210 Exotheca, 309
Decaryella, 360 D. oligostachya, 211 The Exotheca + Hyperthelia
Decaryochloa, 186 Duthieeae, 209 Group, 309
Dendrocalaminae, 176 Dybowskia, 313
Dendrocalamopsis, 182 F
Dendrocalamus, 182 E “Fargesia”, 164
Dendrochloa, 185 Eccoilopus, 295 Farrago, 382
Deschampsia, 241 Eccoptocarpha, 338 Farragoinae, 380
D. antarctica, 242 Echinaria, 250 Fasciculochloa, 319
D. caespitosa, 4 Echinochloa, 324 Ferrocalamus, 161
Desmazeria, 249 Echinolaena, 321 Festuca, 245
Desmostachya, 393 Echinopogon, 238 F. ovina, 246
Deyeuxia, 238 Ectrosia, 366 F. sect. Amphigenes, 245
D. zangxiensi, 238 Ectrosiopsis, 366 F. sect. Montanae, 245
Diandrochloa, 366 Ehrharta, 145 F. sect. Pseudoscariosa, 245
Diandrolyra, 195 E. erecta, 146 F. sect. Scariosae, 245
Diarrhena, 221 Ehrharteae, 145 F. sect. Subbulbosae, 246
D. obovata, 221 Ehrhartoideae, 143 F. sect. Subulatae, 244
Diarrheneae, 221 Ekmanochloa, 192 F. sect. Subuliflorae, 245
Dichaetaria, 404 Eleusine, 392 F. subg. Drymanthele, 245
Dichanthelium, 323 Eleusininae, 384 F. subg. Obtusae, 245
D. oligosanthes subsp. Elionurus, 293 F. subg. Schedonorus, 246
scribnerianum, 324 Ellisochloa, 359 Festucella, 259
Dichanthium, 312 Elymandra, 308 Festucoideae, 199
Dichelachne, 238 Elymus, 225 Festucopsis, 226
Didymogonyx, 172 Elytrigia, 225 Filgueirasia, 173
Diectomis, 312 Elytrophorus, 405 Fimbribambusa, 177
The Diectomis + Hyparrhenia + Elytrostachys, 173 Fingerhuthia, 368
Andropogon + Schizachyrium Enneapogon, 364 Froesiochloa, 192
Group, 312 E. desvauxii, 365
Digitaria, 327 “Enteropogon”, 389 G
D. catamarcensis, 327 Entolasia, 330 Gaoligongshania, 168
Digitarieae, 323 Entoplocamia, 368 Garnotia, 290
Digitariinae, 326 E. aristulata, 368 Garnotieae, 289
Dignathia, 382 Eragrostideae, 363 Gastridium, 238
Diheteropogon, 312 Eragrostidinae, 365 Gaudinia, 233
The Diheteropogon + Parahyparrhenia Eragrostiella, 393 Gaudiniopsis, 258
+ Pseudanthistiria Eragrostis, 366 “Gelidocalamus”, 162
Group, 312 E. frankii, 366 Geochloa, 348
Dilophotriche, 283 “Eragrostis” walteri, 405 Germainia, 306
Dimeria, 301 Eremitis, 190 Germainiinae, 306
Dimeriinae, 301 Eremium, 227 Gerritea, 321
Dimorphochloa, 331 Eremocaulon, 174 Gigantochloa, 183
Dinebra, 387 Eremochloa, 298 Gilgiochloa, 283
Dinochloa, 181 Eremopoa, 257 Glaziophyton, 170
D. macclellandii, 14 Eremopogon, 310 Glyceria, 219
Diplachne, 388 Eremopyrum, 226 G. multiflora, 220
D. fusca, 388 Eriachne, 400 Glyphochloa, 298
Diplopogon, 407 Erianthecium, 235 Gouinia, 380
Disakisperma, 388 Erianthus, 306 Gouiniinae, 380
Dissanthelium, 257 Eriochloa, 338 Gouldochloa, 284
Dissochondrus, 334 Eriochrysis, 304 Graphephorum, 231
Distichlis, 375 “Eriocoma group”, 218 Greslania, 177
DMNS clade, 181 Erioneuron, 379 Griffithsochloa, 374
Douglasdeweya, 225 Erythranthera, 351 Guadua, 175
Drake-brockmania, 387 Euchlaena, 297 G. ciliata, 175
412 Index
Guaduella, 139 Hubbardia, 400 Lamarckia, 248

G. oblonga, 140 Hubbardochloa, 372 Lamprothyrsus, 350
Guaduinae, 174 Humbertochloa, 144 Lasiacis, 330
Gymnachne, 236 Hyalopoa, 251 Lasiorhachis, 305
Gymnopogon, 382 Hydrochloa, 149 Lasiurus, 294
Gymnopogoninae, 380 Hydrothauma, 288 Latipes, 382
Gynerieae, 286 Hygrochloa, 334 Lecomtella, 288
Gynerium, 286 Hygroryza, 148 Leersia, 147
G. sagittatum, 286 H. aristata, 148 L. denudata, 147
Hylebates, 325 L. drepanothrix, 147
H Hymenachne, 318 L. friesii, 147
Habrochloa, 362 H. amplexicaulis, 318 L. hexandra, 147
Hackelochloa, 298 Hyparrhenia, 313 L. tisserantii, 147
Hainardia, 249 Hyperthelia, 309 Leleba, 182
H. cylindrica, 249 Hypogynium, 294 Lepargochloa, 299
Hakonechloa, 405 Hypseochloa, 238 Leptagrostis, 405
H. macra, 405 Hystrix, 225 Leptaspis, 135
Halopyrum, 360 Leptatherum, 303
Harpachne, 366 I Leptocanna, 185
Harpochloa, 389 Ichnanthus, 322 Leptocarydion, 382
Haynaldia, 225 Imperata, 304 Leptochloa, 389
Helicotrichon Indocalamus, 168 Leptochlo€ opsis, 369
H. subg. Pratavenastrum, 241 Not “Indocalamus”, 164 Leptocoryphium, 318
Helictotrichon, 232 “Indocalamus” wilsonii, 168 Leptophyllochloa, 233
Hellerochloa, 245 Indopoa, 361 Leptosaccharum, 304
Hemarthria, 300 “Indosasa”, 167 Leptothrium, 382
Hemisorghum, 303 Isachne, 401 Lepturidium, 372
Henrardia, 226 I. pulchella, 401 Lepturopetium, 361
Hesperostipa, 215 Ischaeminae, 301 Lepturus, 390
Heterachne, 367 Ischaemum, 302 Leucophrys, 340
Heteranthelium, 226 I. afrum, 302 Leucopoa, 245
Heteranthoecia, 400 Ischnochloa, 294 “Leucopoa grade”, 245
Heterocarpha, 387 Ischurochloa, 182 Leymus, 227
Heteropholis, 300 Iseilema, 310 Libyella, 257
H. sulcata, 301 Ixophorus, 334 Limnas, 256
Heteropogon, 310 Limnodea, 256
Hickelia, 186 J Limnopoa, 401
H. africana, 187 Jansenella, 283 Lindbergella, 257
Hickeliinae, 186 “Jarava”, 218 Lingnania, 182
Hierochloe, 234 Jardinea, 299 Lintonia, 390
Hilaria, 376 Jouvea, 376 Lithachne, 192
Hilariinae, 374 Joycea, 351 Littledalea, 223
“Himalayacalamus”, 164 L. tibetica, 224
Hitchcockella, 187 K Loliinae, 244
HK clade, 180 Kampochloa, 361 “intermediate grade,” 244
Holcolemma, 324 Kaokochloa, 364 Loliolum, 247
Holcus, 242 Karroochloa, 352 Lolium, 246
Holttumochloa, 180 Kengia, 372 Lophacme, 383
Homolepis, 315 Kengyilia, 225 Lophatherum, 285
Homopholis, 325 Keratochlaena, 316 Lopholepis, 384
Homozeugos, 308 Kerriochloa, 294 Lophopogon, 307
The Homozeugos + Agenium + Kinabaluchloa, 180 Lophopyrum, 229
Trachypogon clade, 308 Klemachloa, 182 Lorenzochloa, 215
Hookerochloa, 259 Koeleria, 232 Loudetia, 283
Hopia, 321 Kuruna, 168 Loudetiopsis, 283
Hordelymus, 227 Louisiella, 342
Hordeum, 227 L Loxodera, 299
H. brevisubulatum subsp. “Lachnagrostis”, 239 Luziola, 149
turkestanicum, 227 Lagurus, 232 Luziola + Zizaniopsis clade, 148
HSAQN clade, 259 Lakshmia, 294 Lycochloa, 220
Index 413
Lycurus, 376 Monocladus, 176 O

Lygeeae, 208 Monocymbium, 309 Ocellochloa, 322
Lygeum, 209 M. cerisiiforme, 11 Ochlandra, 185
The Monocymbium + Anadelphia + Ochthochloa, 361
M Elymandra Group, 308 Odontelytrum, 333
Maclurochloa, 183 Monodia, 384 Odyssea, 374
Maclurolyra, 195 Monospatha, 166 Oldeania, 160
Macrochloa, 212 Monostachya, 351 “Oligostachyum”, 165
Maltebrunia, 147 Moorochloa, 340 Olmeca, 175
Manisuris, 299 Mosdenia, 383 Oloptum, 217
Matudacalamus, 172 Muhlenbergia, 376 Olyra, 192
Mayariochloa, 330 M. subg. Bealia, 377 O. micrantha, 193
Megalachne, 230 M. subg. Clomena, 377 Olyreae, 189
Megaloprotachne, 328 M. subg. Muhlenbergia, 377 Olyrinae, 191
Megastachya, 287 M. subg. Pseudosporobolus, 377 Oncorachis, 316
Megathyrsus, 339 M. subg. Trichochloa, 377 Ophiochloa, 320
Melanocenchris, 393 Muhlenbergiinae, 374 Ophiuros, 300
Melica, 220 Mullerochloa, 181 Opizia, 374
Meliceae, 219 Munroa, 378 Oplismenopsis, 315
Melinideae, 323 Myriocladus, 174 Oplismenus, 330
Melinidinae, 338 Myriostachya, 361 Orcuttia, 379
Melinis, 340 O. inaequalis, 379
Melocalamus, 177 N Orcuttiinae, 379
Melocanna, 184 Nanooravia, 301 Oreobambos, 178
Melocanninae, 184 Nardeae, 208 O. buchwaldii, 179
Menstruocalamus, 163 Narduroides, 245 Oreocalamus, 163
Merostachys, 173 Nardus, 208 Oreochloa, 250
M. sp., 173 N. stricta, 209 Oreopoa, 257
M. speciosa, 173 Narenga, 305 Orinus, 372
Merxmuellera, 348 Nassella, 219 Oropetium, 393
Mesosetum, 317 Nastinae, 186 O. minimum, 393
Metasasa, 162 Nastus s.s., 187 Ortachne, 215
Metcalfia, 210 Not “Nastus”, 177 Orthoclada, 285
Mibora, 250 Neesiochloa, 394 O. africana, 285
Miborinae, 250 Nematopoa, 406 Orthoraphium, 213
Micraira, 400 Neobouteloua, 373 Oryza, 147
Micrairoideae, 399 Neohouzeaua, 184 Oryzeae, 146
Microbriza, 236 Neololeba, 181 Oryzidium, 288
Microcalaminae, 328 Neomicrocalamus, 177 Oryzinae, 146
Microcalamus, 330 Neomolinia, 221 Oryzopsis, 213
Microchloa, 390 Neosinocalamus, 182 Osvaldoa, 321
Microlaena, 145 Neostapfia, 379 Otachyriinae, 317
Micropyropsis, 246 Neostapfiella, 361 Otachyrium, 319
Micropyrum, 245 “Neotrinia”, 212 Otatea, 175
Microstegium, 294 “Neotrinia group” of “Achnatherum”, Ottochloa, 331
Mildbraediochloa, 340 212 Oxychloris, 390
Miliinae, 254 Nephelochloa, 256 O. scariosa, 11
Milium, 256 Neurachne, 332 Oxyrhachis, 296
Miscanthidium, 305 Neurachneae, 323 Oxytenanthera, 178
Miscanthus, 305 Neurachninae, 331
M. violaceus, 305 Neurolepidinae, 169 P
Mnesithea, 299 Neurolepis, 170 PACMAD clade, 267
Mniochloa, 194 Neuropoa, 257 Paniceae, 323
Molineriella, 242 Neyraudia, 363 Panicinae, 341
Molinia, 406 N. montana, 363 Panicoideae, 271
Monachather, 407 Nicoraepoa, 259 Panicoideae s.s. clade, 287
Monanthochloë, 375 Nipponocalamus, 165 Panicum, 341
Monanthochloinae, 374 Notochloe, 351 P. bergii, 341
Monelytrum, 377 Notochloe + Plinthanthesis clade, 351 “P.” deustum, 339
Monium, 308 Notodanthonia, 351 P. [group] Parvifolia, 326
414 Index
Panicum, 341 (cont.) Phaenosperma, 209 Pommereulla, 373

“P.” [group] Verrucosa, 325 Phaenospermateae, 209 Pooideae, 199
P. sect. Clavelligerae, 323 Phalaridinae, 233 Porteresia, 147
“P.” sect. Cordovensia, 331 Phalaris, 233 Potamophila, 148
P. sect. Megista, 315 P. canariensis, 234 Pringleochloa, 374
“P.” sect. Monticolae, 325 Phanopyrum, 315 Prionanthium, 348
“P.” sect. Monticolae + sect. Pharoideae, 135 Prosphytochloa, 147
Verrucosa + sect. Ovalifoliae, Pharus, 135 Psammagrostis, 366
325 P. glaber, 136 Psammochloa, 213
“P.” sect. Ovalifoliae, 325 Pheidochloa, 400 Psathyrostachys, 228
P. sect. Prionitia, 316 Phippsia, 252 Pseudanthistiria, 312
P. sect. Verruculosa, 326 Phleum, 256 Pseudarrhenatherum, 231
P. subg. Dichanthelium, 323 Pholiurus, 257 Pseudechinolaena, 331
P. [unranked] Agrostoidea, 316 Phragmites, 406 Pseudobambusa, 178
P. [unranked] Tenera, 316 Phuphanochloa, 182 Pseudobromus, 247
“Panicum” antidotale, 334 Phyllorachideae, 144 Pseudochaetochloa, 335
Pappophorinae, 394 Phyllorachis, 144 Pseudocoix, 186
Pappophorum, 394 P. sagittata, 145 Pseudodanthonia, 211
P. bicolor, 395 “Phyllostachys”, 165 Pseudodichanthium, 310
P. vaginatum, 395 Phyllostachys clade, 162 Pseudopentameris, 350
Pappostipa, 215 Pinga, 178 Pseudophleum, 256
Parabambusa, 178 Piptatheropsis, 215 Pseudopogonatherum, 304
Paracolpodium, 252 Piptatherum s.s., 216 Pseudoraphidinae, 333
Paractaenum, 334 P. sect. Virescentia, 217 Pseudoraphis, 335
Parafestuca, 233 Piptochaetium, 216 Pseudoroegneria, 228
Parahyparrhenia, 312 Piptophyllum, 406 “Pseudosasa”, 162
Paraneurachne, 332 Piresia, 193 Pseudosclerochloa, 252
Parapholiinae, 248 Piresiella, 194 Pseudosorghum, 303
Parapholis, 249 Plagiantha, 319 Pseudostachyum, 185
Paratheria, 334 Plagiosetum, 334 Pseudovossia, 299
Pariana, 191 Plectrachne, 384 Pseudoxytenanthera, 178
P. radiciflora, 191 Pleiadelphia, 308 Pseudozoysia, 362
Parianeae, 189 “Pleioblastus”, 165 Psilolemma, 369
Parianinae, 190 Pleuraphis, 376 Pterochloris, 386
Parodiochloa, 257 Pleuropogon, 220 Ptilagrostis s.s., 214
Parodiolyra, 193 Plinthanthesis, 351 P. pelliotii, 214
Parodiophyllochloa, 331 Poa, 257 “Ptilagrostis” kingii, 216
Pascopyrum, 225 P. compressa, 4 Puccinellia, 253
Paspaleae, 314 P. lilloi, 257 Puccinelliinae, 251
Paspalidium, 335 P. sp., 14 Puelia, 139
Paspalinae, 319 Poagrostis, 348 Puelioideae, 139
Paspalum, 322 Pobeguinea, 308 Pyrrhanthera, 351
Patis, 216 Podagrostis, 239
Pennisetinae, 333 Podophorus, 230 Q
Pennisetum, 333 Poeae s.l., 229 Qiongzhuea, 163
Pentameris, 348 Poecilostachys, 331
P. alticola, 349 Pogonachne, 295 R
Pentapogon, 239 Pogonarthria, 366 Racemobambos, 189
Pentarrhaphis, 374 Pogonatherum, 306 R. hepburnii, 189
Pentaschistis, 348 P. paniceum, 307 Racemobambosinae, 188
Pereilema, 376 Pogoneura, 373 Raddia, 194
Periballia, 242 Pogonochloa, 362 Raddiella, 193
Peridictyon, 228 Pohlidium, 286 Ratzeburgia, 299
Perotidinae, 380 Poidium, 235 Redfieldia, 376
Perotis, 384 Poinae, 254 Reederochloa, 375
P. rara, 385 Polevansia, 378 Rehia, 194
Perrierbambus, 188 Polliniopsis, 303 Reimarochloa, 322
Petriella, 145 Polyanthus, 165 Reimarochloinae, 319
Peyritschia, 233 Polypogon, 239 Reitzia, 194
Phacelurus, 299 Polytoca, 292 Relchela, 236
Phaenanthoecium, 407 Polytrias, 295 Rendlia, 390
Index 415
Renvoizea, 322 Scutachne, 339 S. hirtigluma subsp. patula, 268

Rettbergia, 170 Secale, 228 Stipeae, 211
Reynaudia, 289 Sehima, 295 Stipella, 218
Rheochloa, 390 Sesleria, 250 Stipellula, 218
Rhipidocladum, 174 S. autumnalis, 251 Streblochaete, 221
R. sect. Didymogonyx, 173 Sesleriinae, 250 Streptochaeta, 132
Rhizocephalus, 258 Setaria, 335 S. spicata subsp. spicata, 133
Rhombolytrum, 235 S. lachnea, 335 Streptogyna, 140
Rhynchelytrum, 340 Setariinae, 333 S. crinita, 141
Rhynchoryza, 148 Setariopsis, 336 Streptolophus, 336
Rhytachne, 296 Shibataea, 162 Streptostachys, 323
Richardsiella, 367 Shibataea clade, 161 Styppeiochloa, 406
Robynsiochloa, 291 Sieglingia, 351 Sucrea, 194
Roegneria, 225 Silentvalleya, 362 Suddia, 144
Rostraria, 233 Simplicia, 258 Swallenia, 378
Rottboellia, 300 Sinarundinaria, 164 Swallenochloa, 170
Rottboelliinae, 298 “Sinobambusa”, 166 Sylvipoa, 259
Rugoloa, 319 Sinocalamus s.s., 182 Symplectrodia, 384
Rupichloa, 339 Sinochasea, 211
Rytidosperma, 351 Sirochloa, 188 T
Rytidosperma clade, 351 Sitanion, 225 Taeniatherum, 228
Snowdenia, 336 Tarigidia, 328
S Snowdeniinae, 333 Tatianyx, 317
Saccharinae, 302 Soderstromia, 374 T. arnacites, 317
Saccharum, 306 Soejatmia, 183 Teinostachyum, 186
The Saccharum group, 304 Sohnsia, 377 Temburongia, 180
Sacciolepis, 326 Sorengia, 316 Temochloa, 180
Sacciolepis, Trichanthecium, and the Sorghastrum, 303 Tenaxia, 351
“Panicum” sect. Monticolae Sorghum, 303 Tetrachaete, 364
group, 325 Spartina, 371 Tetrachne, 369
Sarga, 304 Spartochloa, 287 Tetragonocalamus, 182
Sarocalamus, 165 Spathia, 304 Tetrapogon, 391
Sartidia, 268 Sphaerobambos, 181 Tetrarrhena, 145
Sasa, 167 Sphaerocaryum, 401 Thamnocalamus, 167
Sasamorpha, 167 Spheneria, 322 Thaumastochloa, 301
Saugetia, 391 Sphenopholis, 232 Thedachloa, 325
Saxipoa, 259 Sphenopus, 248 Thelepogon, 296
Schaffnerella, 376 Spinifex, 337 Thellungia, 371
Schedonnardus, 376 Spinificeae, 323 Themeda, 310
Schedonorus, 246 Spodiopogon, 295 T. triandra, 311
Schismus, 352 Sporobolus, 371 Thinopyrum, 229
Schizachne, 220 S. cryptandrus, 370 Thrasya, 322
Schizachyrium, 313 Stapletonia, 186 Thrasyopsis, 322
Schizostachyidinae, 184 Steinchisma, 319 Thuarea, 339
Schizostachyum, 185 Steirachne, 367 Thyridachne, 325
S. dumetorum, 185 Stenostachys, 228 Thyridolepis, 332
Schmidtia, 364 Stenotaphrum, 336 T. mitchelliana, 332
Schoenefeldia, 391 Stephanachne, 211 Thyrsia, 300
Sclerachne, 292 Stephostachys, 315 Thyrsostachys, 183
Sclerochlamys, 316 Stereochlaena, 336 Thysanolaena, 287
Sclerochloa, 253 Steyermarkochloa, 281 Thysanolaeneae, 286
Sclerodactylon, 391 S. angustifolia, 282 Timouria, 218
Scleropoa, 249 Steyermarkochloeae, 281 Toliara, 384
Scleropogon, 378 Stiburus, 368 Torreyochloa, 231
Scleropogoninae, 374 “Stillmania”, 216 Torreyochloinae, 230
Sclerostachya, 305 Stipa, s.s., 213 Tovarochloa, 257
Scolochloa, 240 S. sect. Anatherostipa, 215 Trachideae, 314
S. festucacea, 240 S. sect. Obtusae, 215 Trachypogon, 308
Scolochloinae, 240 S. sect. Stipella, 218 Trachys, 328
Scribneria, 242 S. subgen. Macrochloa, 212 Traginae, 374
Scrotochloa, 136 Stipagrostis, 268 Tragus, 377
416 Index
Tribolium, 352 Tristachya, 283 W

Trichanthecium, 326 T. decora, 284 Walwhalleya, 325
Trichloris, 391 Tristachyideae, 282 Wangenheimia, 245
Tricholaena, 341 Triticeae, 223 Whiteochloa, 337
Trichoneura, 381 Triticum, 229 Willkommia, 378
Trichoneurinae, 380 Tuctoria, 380
Trichopteryx, 283 Tzvelevia, 257 X
Tridens, 394 Xerochloa, 337
Tridentopsis, 380 U
Trigonochloa, 383 Uniola, 369 Y
Trikeraia, 214 Uniolinae, 367 Yakirra, 341
Trilobachne, 292 Uranthoecium, 337 Yushania, 40, 166
Triniochloa, 221 Urelytrum, 296 Yvesia, 340
Triodia, 384 Urochlaena, 352
T. scariosa, 386 Urochloa, 339 Z
Triodiinae, 384 U. paucispicata, 339 Zea, 297
Triplachne, 239 Urochondra, 369 Zenkeria, 406
Triplasis, 381 Zeugiteae, 284
Triplopogon, 296 V Zeugites, 286
Tripogon, 394 Vacoparis, 303 Zingeria, 253
Tripogoninae, 392 Vahlodea, 242 Zizania, 148
Tripsacinae, 296 Valiha, 188 Zizaniinae, 147
Tripsacum, 297 Vaseyochloa, 381 Zizaniopsis, 149
T. dactyloides, 297 Ventenata, 258 Zonotriche, 283
Triraphideae, 362 Vetiveria, 292 Zotovia, 145
Triraphis, 363 Vietnamosasa, 180 Zoysia, 370
Triscenia, 316 Viguierella, 362 Zoysieae, 369
Trisetaria, 232 Vossia, 297 Zuloagaea, 337
Trisetokoeleria, 233 Vulpia, 245 Zygochloa, 338
Trisetum, 232 Vulpiella, 250
T. barbinode, 232

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Edited by K.

Prof. Dr. Klaus Kubitzki

ISBN 978-3-319-15331-5 ISBN 978-3-319-15332-2 (eBook)

Library of Congress Control Number: 2015935331

# Springer International Publishing Switzerland 2015

Printed on acid-free paper

Springer is part of Springer ScienceþBusiness Media (www.springer.com)

problems can be resolved by current data. Problems outstanding are discussed

St. Louis, MO Elizabeth A. Kellogg

Poaceae – General Information

Fruit and Seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Bistigmatic clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

PACMAD Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

XII. Subfamily Arundinoideae Burmeist. (1837) . . . . . . . . . . . . . . . . . . . . . . . . 403

Rhizomatous perennials, bisexual or monoe- VEGETATIVE MORPHOLOGY AND ANATOMY

known patterns of development; for example, the TRANSITION TO FLOWERING

Inflorescence maturation, maturation of each (2009), the architecture of the inflorescence is

factor (AGAMOUS) that specifies both carpel

OVULE et al. (1994). The distinction between hemiana-

MEGAGAMETOPHYTE although only 16 are shown in the figure. Prolifera-

embryo both root and shoot apical meristems

Fig. 6. Bambusa multiplex. Seedling in two stages of

CHROMOSOME NUMBER possible, it will only be provided by genomic data

MORPHOLOGY AND ANTHER STRUCTURE ever, simultaneous microsporogenesis, with cyto-

In addition, successful pollination has been tion.) Phase V, gametophytic interactions,

tetrahydro-b-carboline-3-carboxylic acid, an sequestered in the vacuole in epidermal cells of

Urochloa (¼ Panicum maximum = Megathyrsus Apomixis is strongly associated with poly-

Rhizomatous perennials, herbaceous. Ligule relationship in phylogenetic trees is an artifact.

This is one of the most curious of the grasses,

2. Streptochaeta Nees Figs. 11a, 11b

Leaves with auricles; ligule absent. Inflorescence

All remaining subfamilies have flowers arranged

teoles, the bract between bracteoles and axis removed. D

with a tiny beak; covered with hooked hairs. Lodi-

Scrotochloa Judziewicz, Phytologia 56: 299–304 (1984).

Ligule membranous, pubescent. Inflorescence

Stigmas two. Orders of stigmatic branching two.

Fig. 12. Pharoideae. Pharus glaber. A Flowering plant. B

Judziewicz, E.J., Clark, L.G. 2007. Classification and bio-

Rhizomatous perennials, herbaceous, the culms 7. Puelia Franch.

Early phylogenetic analyses provided little sup-

8. Streptogyna P. Beauv. Fig. 14

Streptogyna P. Beauv., Ess. Agrost. 80 (1812).

Rhizomatous or caespitose perennials. Leaf blades

GPWG II 2012. Grass Phylogeny Working Group II. New

EHRHARTOIDEAE INCERTAE SEDIS 11. Phyllorachis Trimen. Fig. 15

spikelets. Lower lemma of pistillate spikelet

II. TRIBE EHRHARTEAE Nevski (1937)

Annuals or perennials, rhizomatous, stoloniferous,

12. Ehrharta Thunb. Fig. 16

(Verboom et al. 2003). The early-diverging mem-

III. TRIBE ORYZEAE Dumort. (1824)

Annuals or perennials. Leaf blades with or without

III.1. SUBTRIBE ORYZINAE Griseb. (1853)

Fig. 16. Ehrhartoideae-Ehrharteae. Ehrharta erecta.

arsenic from the soil, making it potentially useful

14. Maltebrunia Kunth

18. Hygroryza Nees Fig. 18

19. Potamophila R. Br.

20. Rhynchoryza Baillon